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SEA OTTER STUDIES - 1970
by Karl Schneider March 1971
This report is a compilation of rough reports from the marine mammals staff files It summarizes most of the work accomplished on sea otters during 1970 The analysis of much of the data is incomplete and many of the conclusions are tentative This report is intended to present the data collected in a useful form and to indicate the progress and stage of thinking at the time it was prepared It is intended for staff use only
The contents of the report are as follmrs
Surveys
Alaska Peninsula - south side Alaska Peninsula - north side Afognak - Barren Islands Kenai Peninsula Prince Willia Sound Sitka - Cape Spencer
ants
Sexual
Identification of Hale Areas by Pup Counts Kanaga Island Tanaga Island
Identification of Male Areas from Ha1vests Delarof Isla1ds Adak Island Amchitka Island
Composition of Female Areas Segregation Hithin Female Areas Composition of Hale Areas Population Sex Ratio
Summary of Annual Cycle Magnitude of Breeding and Pupping Peaks Gestation Period Fetal Groth Rate Age of Sexual Maturity Frequency of 07ulation Maximum Breeding Ovulation Significance of Delayed Implantation Fetal Orientation
Sea Otter Studies - 2 - ]1archt 1971
Location of Pregnancy Birth Weight Sex Ratio at Birth Sex Ratio of the Population Litter Size In Utero Mortality Mortality at or near Parturition Recovery of the Uterus after Parturition Frequency of Breeding Conclusions Literature Cited
AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OFTHE ALASKA PENINSULA
March 1970
Between March 16 and March 27 1970 an aerial count of sea otters was rnade along the south side of the Alaska Peninsula from Shaw Island to Cape Lazaref on Unlmak Island Sanak Island the Sandman Reefs the Pavlof Islands northern Shumagin Islands and Sutwick Island were included A BLM Aero Commander N6392U was flown by Cal Ward at an altitude averaging 150 feet and an airspeed bullgtf 100 mphmiddot Observers were Karl Schneider next to the pilot and Jim Faro behind the pilot This is the same technique aircraft pilot and observers as were used in the 1969 Aleutian count The count on March 27 was made in a Gruman Goose 1r1ith Robert Wolfe substituting for Faro Pups with their mothers were not counted
Because of conditions of visibility and distribution of animals all of the counts made are considered to be low
In general the variability in factors influencing this type of count is so great that the results are not rei iable for population estimates or for determining short-term fluctuations in dense populations However aerial counts should be useful to determine general distribution and abundance and to follow large changes in population size Such changes are occurring where relatively dense populations are expanding into habitat that is either unshyoccupied or sparsely occupied Aerial counts are also the quickest way to familiarize new personnel with large areas of habitat
The following is a 1ist of definitions of terms used to describe counting conditions Application of these terms is completely subjective and based on the writers experience
Excellent- Surface of water relatively calm Usually overcast with 1ittle surface glare Single animals easy to spot at a distance
Very good - Surface may be slightly choppy but not enough confusing reflection to prevent spotting of animals off shore
Good - Off shore animals may be difficult to spot but single animals near shore and in kelp beds and small pods everywhere are readily spotted
Fair -Usually choppy or surface glare Single animals ~in kelp beds or in the lee of rocks or shore and most groups of animals relatively easy to see Other single animals and some pods in deep bays in the shadow of cliffs or off shore may be missed
Poor -Single animals difficult to spot and many pods may be missed but conditions still good enough to get a rough idea of the distribution of animals
The conditions encountered for each area are listed below
Description of Conditions During March 1970 Aerial Count of Sea Otters
March 20 Aniakchak Bay to Chignik Lagoon (1022 AM- 1240 PM) Conditions were generally fair with a heavy chop The south side of Sutwick Island and the area between the island and mainland were poor with heavy surf Visibility was very good inside Kujulik Bay however many animals were scattered throughout the area On two passes ~cross the bay 12 and 27 were seen therefore many were probably missed Similar scattering occured in Chignik Bay The overall count for the area is probably low
Pavlof Bay to Cold Bay (130PM-200PM) Conditions poor with heavy chop and squalls The count was superficial however a second superfi cia 1 count of this a rea was made on March 21 and only one otter was seen in Cold Bay
Cold Bay to Caee Lazaref (310PM -420PM) Conditions very poor fair in a few spots Heavy chop dense kelp
March 21 Northern Shumagin Islands (943AM- 1137 AM) Poo~ with chop on windward sides Fai_r on leeward sides (westerly wind) however air turbulence forced us away from high cliffs Heavy surf created very poor conditions on the south side of Unga
Pavlof Islands (1145 AM- 125ff PM) Conditions similar to those in the Shumagins
Chignik Lagoon to Pavlof Bay (305 PM- 535 PM) Conditions varying from fair to poor with chop and surface glare Count concentrated on points and better looking areas Most deep bays not completely surveyed
March 22 Sanak Islands (849AM- 1020 AM) Conditions fair to poor with chop on south side and surface glare on north side Animals scattered and very difficult to see Many otters on rocks Count is probably very low
Sandman Reefs ( 1025 AM~ 1115 AM) Conditions similar to Sanak With the exception of one pod of 450+ the animals were widely scattered and very difficult to pick out Only those very close to the aircraft could be seen Bad squalls moved in and the wind increased preventing a complete count of Deer Island and the eastern half of the reefs
March 23 Wide Bay to Aniakchak Bay (1123 AM- 140PM) Conditions generally good at first becoming fair after Cape Providence with a light chop and some surface glare Otters in Amber and Aniakchak Bays were widely scattered and many may have been missed
Port Heiden to Naknek_ Superficial counts were made from Ugashik Bay to Naknek on March 19 and from Port Heiden to Egegik on March 23 These were made under poor conditions by flying outside the surf line The water was choppy and full of silt No sea otters were seen
11arch 27 Shaw Island to Puale Bay (1030 AM - 1205 PM) Conditions very good to excellent High overcast and very little wind down to Katmai Bay After this the conditions deteriorated rapidly to fair and the count had to be stopped because of a combination of bad weather and lack of fuel As shown on the charts the count was concentrated on the points and islands however the good visibility allowed us to see well into the bays We feel certain no concentrations were missed
The numbers of sea otters counted are presented in the Table Specific locations and numbers are shown on the charts Where coverage of an area was not complete the approximate path of the aircraft is shown
Discussion of Results
Surveys along the Alaska Peninsula have been fragmentary and most of those made were done under less than ideal conditionsmiddot There are a number of reasons for this Weather tends to be poor along the south shore where squall5 form along the mountains Areas where an aircraft can refuel are north of the mountains and there are relatively few passes therefore range of the aircraft is a problem Also there is much shallow water off shore in the area from the Shumagin Islands to Sanak Island Coverage of this area is difficult and many animals are undoubtably missed
The present survey covered most of the area however conditions were such that most of the counts are probably quite low Despite these problems and a lack of continuity we have a fairly good picture of the recovery of sea otter populations in this area
Reports from various individuals are available from the 1930s and 1940s 11ajor surveys by the Fish and Wildlife Service in 1951 (Jones) 1957 (Lensink) 1962 and 1965 (Kenyon and Spencer) covered at least portions of the area In 1969 the southern Shumagin Islands were counted by the Alaska Department of Fish and Game and the present survey covered most of the rest of the area
Basically there are four distinct population nuclei in the area These centered around the Sanak Island-Sandman Reef area the Shumagin Island area the Sutwick Island area and the Augustine Island-Cape Douglas area The following discussion is based on information from reports by Lensink and Kenyon and from Alaska Department of Fish and Game surveys middot
~anak Island-Sandman Reefs
Small numbers of sea otters have been reported at Sanak Island since 1922 No reports came from the Sandman Reefs until 1942 In 1948 27 were sighted at
----Cherni Island The 1951 aerial survey showed 65 around Sanak and 97 in the Sandman Reefs In 1957 251 were seen around Sanak and 508 around the Sandman Reefs In addition two were seen in the Pavlof Islands however few were on the mainland In 1962 548 were counted in the Sanak area and 638 in the Sandman Reefs None were reported from the mainland or Pavlof lsland~however
much of the increase was in the northern area around Deer Island The 1962 survey was probably the best being made under excel lent conditions
AERIAL COUNT OF SEA OTTERS MARCH 1970
Partial or Location Count Date Camp 1ete Count Visibility
Cape Lazaref - Cold lsecty_
Cape Lazaref 3 320 Complete Poor
lkatan Peninsula 71 320 Complete Poor
False Pass - Amagat I 66 320 Camp 1ete Poor
Cold Bay 321 Partial Poor
TOTAL 141
Sanak Islands 239 322 Camp 1ete Fair to Poor
Sandman Reefs
Clubbing Rocks 12 322 Complete Fair to Poor
Cherni I amp Rocks 495+ 322 Complete fair to Poor
Goose I 7 322 Complete Fair to Poor
Hay I 18 322 Camp lete Fair to Poor
Hunt I 2 322 Complete Fair to Poor
Deer I 322 Partial Fair to Poor~
TOTAL 568+
Pavlof Islands
Inner 11iasik 2 321 Camp 1ete Fair to Poor
Outer lliasik 16 321 Camp 1ete Fair to Poor
Goloi 2 321 Complete Fair to Poor
Dolgoi 67 321 Complete Fair to Poor
Poperechnoi 29 321 Complete Fair to Poor
Ukolnoi 2 321 Complete Fair to Poor
yenWosnesenski 4 321 Complete Fair to Poor
TOTAL 122
Partial or Location Count Date Comp 1ete Count vis ib j 1i ty
Northern Shumagin Islands
Unga 184 321 Complete Fair to Poor
Popof 52 321 Complete Fair to Poor
Korovin 46 321 Complete Fair to Poor
Karpa __2t 321 Complete Fair to Poor
TOTAL 286
Cold ~ to Beaver ~ 0 320-21 Partial Poor
Beaver Bay to Kupreanof Pen
Beaver Bay 2 321 Partial Fair to Poor
Cape A 1 iaks in 4 321 Partial Fair to Poor
Guillemot I 16 321 Partial Fair to Poor
__1Kupreanof Peninsula 321 Partial Fair to Poor
TOTAL 23
Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor
AntJrrCastle Cape to-1-Ji e ~
Castle Cape-Castle Bay 16 321 Partial Fair to Poor
Chignik Bay 118 320 Complete Fair
Nakchamik I 5 320 Complete Fair
Hook Bay 13 320 Complete Fair
Cape Kum 1i un 88 320 Complete Fair
Kujul ik Bay 1199 320 Complete Very Good
Univikshak I 62 320 Complete Fair
Cape Kuml ik 54 320 Complete Fair to Poor
Sutwick I 14 320 Complete Fair to Poor
Cape Agutka 323 Complete Fair~
TOTAL 1766
Partial or Location Count Date Comp 1ete Count vis i b i 1i ty
Cape Kunmik 13 323 Complete Fair
Naka 1i kok Bay amp offshore islands 16 323 Complete Fair
Chiginagak Bay 5 323 Complete Fair
Agripina amp lmuya Bay 105 323 Complete Good
Wide Bay to Puale Bay N 0 T S U R V E Y E D
Puale Bay to c Kubugakl i __]_ 327 Partial Fair
TOTAL 146
Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent
Cape Chiniak to Shaw _I
Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent
Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent
TOTAL 71
In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring
A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak
With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time
As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population
Shumagin Is 1 ands
This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population
There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion
In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good
There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied
There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas
Sutwi ck Area
Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather
In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved
~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather
The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys
There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years
to come
Augustine Island-Cape Douqlas
For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro
On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group
It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there
The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table
SUM
MAR
Y OF
SE
A OT
TER
SURV
EYS
(Com
pile
d by
p
op
ula
tio
n f
rom
L
ensi
nk
(196
2 T
hes
is
K
enyo
n 0
96
2 amp
196
5 R
epor
ts
amp M
anu
scri
pt)
an
d AD
FampG
Dat
a)
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
AU
GU
STIN
E -
C
DOUG
LAS
Aug
usti
ne
Isla
nd
A
ppro
xim
atel
y 50
(1
948)
Shaw
1
amp
Dou
glas
Are
a S
igh
tin
gs
from
Sh
aw
Is
to
Tux
edn
i B
ay
TOTA
L
40
___
_
41
52
-shy 52
18
_lQ
J
119
130+
-shy
130+
J
71
ALAS
KA
PEN
INSU
LA
c
Kul
iak
-
c
lgva
k
c
lgva
k -
c
Kun
mik
Ani
akch
ak
Bay
amp
Am
ber
Bay
Sutw
i ck
Are
a M
isce
llan
eous
R
epor
ts
Kuj
ul i
k B
ay
amp c
K
uml
ik
Sin
ce
1936
C
Kum
1 i u
n
Uni
viks
hak
Isla
nd
N
akch
amik
Is
lan
d
8
355 12
13
0 19 6
581
103
180
1
22
47
109
684 86
Not
S
urve
yed
7(+
)
139
197 14
1 2
53
101 62
5
I
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
ALAS
KA
PEN
INSU
LA
(Co
nt
)
Chi
gnik
B
ay
Cas
tle
Bay
amp
Cap
e
TOTA
L
SHUM
AGIN
Mai
nlan
d S
ho
re
Kup
rean
of
Pen
to
P
avlo
f B
ay
Ung
a Is
lan
d
Popo
f Is
lan
d
Kor
ovin
Is
lan
d
Kar
pa
Isla
nd
And
roni
ca
amp t
he
Hay
stac
ks
Nag
ai
Sp
ecta
cle
Ben
del
Tu
rner
Tw
ins
Few
R
epor
ts
Bef
ore
19
40
0
-shy
-shy
388
889 1 2 2 1
149
26
8
~-
7
- 949 4
338 I105
-~-middot
118
_lsect
1 9
12
23
184 52
46 4
75
232 8 27 6
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
SHUM
AGIN
(C
on
t)
Nea
r
Pen
insu
la
Big
Kon
i u
j i
Lit
tle K
oniu
j i
amp A
tkin
s
Sim
eono
f
Bir
d
Che
rnab
ura
TOTA
L
500
Est
imat
e (1
947)
l63
3 (
195 3
)
3 0
220
430
455
160
___L
ll
183
0
14 3
222
255
294 38
_J
l
1 35
2
150 15
296
290
329 76
_6
15
1 0
-shy 309
SANA
K -
SAND
MAN
Mai
nlan
d S
ho
re
Pav
lof
Bay
to
Un
imak
B
ay
Pav
lof
Isla
nd
s
Wos
nese
nski
Is
lan
d
Uko
l noi
Is
lan
d
Pop
erec
hnoi
Is
lan
d
Dol
goi
Isla
nd
2
141 4 2 29
67
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
SAN
AK
-SA
NDM
AN
Pav
lof
Isla
nd
s
Gol
oi
Isla
nd
Out
er
llia
sik
Inne
r ll
iasik
Sand
man
R
eefs
Che
rni
r-s I
and
(Co
nt
) (C
on
t)
Isla
nd
Isla
nd
C1u
bb i
ng
Roc
ks
Goo
se
Isla
nd
Dee
r Is
lan
d
amp O
ther
R
eefs
San
ak
Isla
nd
TOTA
L
2 16 2
27
(194
8)
271
259
495+
No
sig
hti
ng
s b
efo
re
1942
~
2 12
76
3397
82
7
123
5429
5 -
(I n
com
p Je
te)
Sig
hti
ng
s si
nce
19
22
_sect
2
_lil
~
~
162
1 bull 1
86
1 0
7075
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middot middot
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lt
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Total 152
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Cape Agutka Cape Kum 1 i k Sutwick I~ Kujul ik Bay Cape Kum 1 i un Unavikshak 1
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1199 88 62
1614
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MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970
OBSERVER J Vania J Faro
PILOT George Tibbets Jr
AIRCRAFr Piper Azetex
middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good
Departed King Salmon 1115 AM Returned to King Salmon 745 PM
FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out
We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9
The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down
SEA OTTER SIGHTINGS
Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed
The number counted in each pod is as follows
1071 520 357 68 36 75 30
TOTAL 2157
Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island
171
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SEA OTTER COUNT BARREN ISLANDS and SHUYAK - AFOGNAK ISLANDS
June 9 1970
On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows
Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete
Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good
Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial
Area
BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island
AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island
TOTAL
TOTAL
Number of Sea Otters
0 2 0
76 200
29 0
307
18 6 3
121 0
33 26
207
Complete or Visibility Partial Count
Very good Complete ll If II
It II
II
II IJ
Good Complete II
Fair Partial If Complete II Partial II II
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149
51
SEA OTTER COUNTS - KENAI June J970 to January 1971
Carl Divinyi flew aerial surveys of seals along the outside of the
Kenai Peninsula on a monthly basis He recorded all sea otters sighted
on these surveys The surveys did not cover the entire coast line and
the type of aircraft weather conditions and observers varied from one
survey to another Therefore the seperate counts cannot be compared
However when viewed in total they indicate the areas used by sea otters
and the relative abundance of otters in each area
The attached table presents the counts by broad areas
--
SEA
OTT
ERS
CO
UN
TED
ON
A
ER
IAL
SUR
VEYS
O
F K
EN
AI
PEN
I NS
ULP
Ju
ne
1
97
0-
Jan
uar
y
1971
8i5
8
JUN
E 5
amp 9
JU
LY
15-2
0 A
UG
1l
t oc
r 12
NO
V
12
JAN
12
C
Junk
en
-C
R
esu
rrec
tio
n
5 30
42
27
10
30
Res
urr
ecti
on
Bay
2
2 0
4 2
NS
Ai a
1 i k
Bay
1
20
5 8
0 21
Har
ris
Bay
8
18
7 5
3 25
N
uka
Bay
10
6 56
NS
31
28
27
Po
rt
Dic
k 0
l1
NS
NS
3 23
Roc
ky
Bay
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AERIAL SURVEY Prince William Sound
April 1970
Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey
I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife
April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay
Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals
Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina
Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)
April 16 Weather inclement - no flying
April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island
April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)
Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands
The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed
SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date
PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418
CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52
PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS
ESTHER PASSAGE - VALDEZ ARM NS
GALENA BAY FISH BAY 103 415
FISH BAY - GRAVINA POINT 1 415
GRAVINA POINT - CORDOVA 1 415
HAWKINS ISLfu~D 1 4l7
HINCHINBROOK ISLfu~ 99 417
EGG ISLAND 2 417
MONTAGUE ISLAND 259 417
GREEN AND LITTLE GREEN ISLANDS 102 4J7
KNIGHT ISLAND 136 52
INGOT ISLAND 6 52
ELEANOR ISLAND 3 52
SMITH ISLAND 4 52
SEAL ISLAND 0 52
APPLEGATE ROCK 1 52
PERRY ISLAND NS
(continued) SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date-
LONE ISLAND NS
NAKED ISLAND NS
PEAK ISLAND NS
STOREY ISLAND NS
KAYAK - WINGHAM ISLAND 5 417
TOTAL 892
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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970
On July 17 1970 an aerial survey of sea otters was made in the area
around Green Island and the adjacent coast of Montague Island The purpose
of the survey was to locate concentrations of animals prior to a capturing
operation which began the next day A Dornier twin-engine arrcraft was used
with Schneider Reynolds and Somerville acting as observers The sky was
overcast almost no wind and the sea calm However heavy rain showers intershy
fered with forward visibility and the counting conditions were only fair on
the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay
The count began at 210pm and ended at 345 pm
The numbers and locations of otters are shown on the map In addition
16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy
in brook Island and a pod of 35 were seen three miles east of Johnstone Pt
Most of the area included in the survey was traveled repeatedly in a
skiff over the next four days Durlng this time the weather became calm and
clear for the first time in several days and the bulk of the sea otters shifted
from the coast of Montague out into the shallow areas between there and Green 1
and to Green and Channel Islands It was obvious that many otters had been missed
Fn the aerial sur~ey and that the population in the area is quite dense A
total of 48 sea otters were captured in three days of netting
j
SEA OTTER SURVEY Salisbury Sound to Cape Spencer
August 23-25~ 1970
On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and
middotWalt Cunningham serving as observers
The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good
On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere
On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May
Rough weather and outboard problems restricted tl the search area~
and the effectiveness of the search in that area The following sightings were made
82470 I Adult West of Granite Islands 1 Adult South of Granite Islands
11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands
These sightings represent from four to seven animals
Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull
The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal
The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait
While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island
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Harvest
In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island
and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year
The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial
survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs
Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers
Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same
Transplants
Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent
The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island
Table 1
April 20
middotMay 1
May 2
May 3
May 4
May 5
May 6
May 7
May 8
May 9
May 10
May 11
May 12
May 13
Schedule of the 1970 harvest
1030 pm depart Homer
Midnight arrive Adak MV Active
Refuel and prepare boat
Arrive off Tanaga at noon
Complete skinning pack hides etc in pm
Delarofs
Spent am getting water at Amchitka
Clean up boat take down shelter pack gear
900 am arrive Amchitka fly to Anchorage
Killed 53
138
88
131
143
53
144
43
79
83
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(am)
(pm)
(by mid afternoon)
Table 2 Projected and actual harvest numbers May 1970
Projected Actual Tana~a Island Harvest Harvest
Bumpy Point - Hot Springs Bay 200 191
Hot Springs Bay - Twin Bays 50 50
Twin Bays - Cape Sasmik 50 ) ) 199
Cape Sasmik - Tower 120 )
Tower Tanaga Bay 180 166
TOTAL 600 606
Delarof Islands
Gareloi 20-25 0
Kavalga Ogliuga and Skagul 75-100 125
Ulak and Amatignak 35-55 19
TOTAL 150 144
Amchitka Island saved for
USFWS Counting Units 1 amp 2 100 transplant
3 30 82
4 50 36
5 120 87
TOTAL 300 205
Table 3 Details of sea otters harvested in May 1970
Tanaga Island
Total kill 606
Number of pelts saved 598
Number of small pup pelts discarded 8
Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796
(Seal 3770 void - damaged)
Approximate sex ratio - 220 males 386 females or approximately 64 percent females
Delarof Islands
Total kill 144
Number of pelts saved 138
Number of pups discarded 6
Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934
Sex ratio- 44 males 100 females or approximately 69 percent females
Amchitka Island
Total kill 205
Number of pelts saved 194
Number of pups discarded 11
Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128
Sex ratio- 27 males 178 females or approximately 867 percent females
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A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality
Summary of Harvests and Transplants
Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area
The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5
----
----
----
----
----
----
----
----
----
----
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----
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----
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Tab
le 4
N
umbe
rs
of
sea o
tters
re
mov
ed
fro
m Ala
skan
pop
ulat
ions
~ 1
95
17
01
19
51
5
4
55
56
57
59
60
62
6
3
65
66
67
68
69
70
Am
chit
ka I
U
SFW
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ies
35
22
37
2
6
21
39
2
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ran
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nt
47
6
237
86
A
DFamp
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Har
ves
t 1
80
3
11
2
05
2
3
205
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er
5-
24
1
14
-1
To
tal
35
22
37
26
21
39
1
80
31
1
210
50
0
25
1
292
Tan
aga
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ves
t 6
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aga
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ves
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k I
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ves
t 30
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94
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in Is
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ies
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ak amp
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mak
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ok
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nt
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nta
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e amp
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nsp
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t 39
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reen
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res
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agu
l 1
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lak
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9
I
Inclu
des
all
an
imal
s th
at
wer
e h
arv
est
ed
d
ied
d
uri
ng
st
ud
ies
and
tran
spla
nt
att
em
pts
o
r w
ere
tran
spla
nte
d
to
oth
er
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as
or
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s
Doe
s n
ot
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de n
atu
ral
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rtali
ty o
r il
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al
kil
ls
2
Woo
dlan
d P
ark
Zoo
3
B
att
ell
e M
emo
rial
In
sti
tute
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Tab
le 5
N
umbe
rs
of
sea o
tters
tr
an
spla
nte
d 1
95
5-1
97
0
1955
19
56
1959
19
65
--
~ 1
96
6
1968
19
69
1970
Ale
uti
an
s A
ttu
I
5
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er
I
19
1
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bil
ofs
S
t
Pau
l I
7 S
t
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rge
I
57
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uta
t B
ay
10
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z B
ay
23
20
93
58
(Ch
ich
igo
f L
)
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uth
east
Y
akob
i I
30
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lask
a B
iork
a I
48
Barr
ier
Is
55
Hec
eta
I
51
Cap
e S
pen
cer
25
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tish
C
olum
bia
29
14
291
W
ash
ing
ton
Ore
gon
29
1
Non
e b
eli
ev
ed
to
hav
e su
rviv
ed
II
A
t le
ast
13
die
d s
ho
rtly
aft
er
rele
ase
NO
TE
1955
to
19
59 b
y U
SFW
S
1965
to
19
70 b
y
AD
FampG
In
som
e ca
ses
on
e o
r tw
o o
f th
e
abo
ve
anim
als
die
d n
ear
the
tim
e o
f re
lease
30
SEXUAL SEGREGATION IN SEA OTTER POPULATIONS
by Karl Schneider March 1971
Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males
Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands
Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals
Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas
A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population
In
the
By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled
Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet
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1970
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1970
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54
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ka
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To
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8 18
3
08
6
92
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9
52
6
47
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13
2
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70
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lan
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60
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66
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28
6
72
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7
83
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30
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69
4
TA~AGA
I M
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1970
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dan
t P
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13
7
29
2
71
4
1 8
00
2
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6
64
3
36
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arn
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Po
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2 6
25
0
75
0
2 2
50
0
50
0
83
9
17
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run
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in B
ays
3 6
33
3
66
7
1 1
50
0
50
0
14
0
86
0
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in B
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B
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87
5
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71
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28
6
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85
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3 3
50
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25
4
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68
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31
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lak
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SE
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11
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9
Tan
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55
59
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6
07
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93
3
65
6
35
Sexual Segregation - 2 - March 7 1971
Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples
Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground
Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas
Identification of Hale Areas by Pup Counts
In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable
From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen
Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area
The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on
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Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side
The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed
Kanaga Island
Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$
Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point
Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak
Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass
The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring
Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area
Tab
le
5
Com
pari
son
of
Pup
s C
ou
nte
d w
ith
S
ex R
atio
o
f H
arv
est
Jun
e
1968
A
rea
Su
rvey
O
cto
ber
19
68 H
arv
est
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l KA
NAGA
IS
LAN
D
Ind
ep
Ott
er
Ind
ep
Ott
er
M
M
F
Rou
nd
Hea
d-S
ho
al P
oin
t 0
85
3
45
6
55
4
71
5
29
Nag
a P
oin
t-K
anag
a B
ay
66
1
64
middot
18
5
81
5
20
5
79
5
Cap
e C
hla
nak
-Cap
e T
usi
k
51
1
93
2
40
7
60
3
53
6
47
Edd
y R
ock
-Hiv
e R
ock
61
3
4
35
7
64
3
47
1
52
9
Jun
e
1968
A
rea
Su
rvey
H
ay
1970
Har
ves
t
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l TA
NA
GA
IS
LAN
D
Ind
ep
Ott
er ~ter
M
F
M
F
Cap
e S
ud
ak-P
end
ant
Po
int
0 4
4
60
6
39
4
66
4
33
6
Bar
nes
P
oin
t-T
run
k P
oin
t 7
9
59
3
3
96
7
83
9
17
Tru
nk
Po
int-
Tw
in B
ays
26
4
2
77
9
23
1
40
8
60
D
rin
Bay
s-S
ou
th B
ay
96
1
11
7
3
92
7
14
3
85
7
So
uth
Bay
-Har
em R
ock
14
8
31
6
7
middot93
3
91
9
09
Las
h
Bay
-In
fern
o
Ree
f 1
5
73
0
27
0
74
6
25
4
Har
em R
ock
-Ku
lak
Po
int
0 2
7
64
4
35
6
68
4
31
6
Ku
lak
Po
int-
SE
B
igh
t 6
9
82
1
32
8
68
1
51
8
49
Sexual Segregation - 4 - March~ 1971
Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands
Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area
As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method
Identification of
Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area
The follmving is a description of the sex composition of areas for which no skiff count data was available
Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island
Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill
Sexual Segregation - 5 - March 1971
Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point
The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it
Composition of Female Areas
Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female
The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure
Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters
Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males
Sexual Segregation - 6 - March~ 1971
apparently defending a territory in California but from his observations t
it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change
Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year
A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest
The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas
The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males
There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas
Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak
within Female
In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of
Sexual Segregation - 7 - March 1971
these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June
The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups
Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population
ition of Male Areas
As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles
Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements
le 6
R
epro
du
ctiv
e S
tag
e o
f S
exu
ally
Mat
ure
S
ea O
tters
Har
ves
ted
on
Tan
aga
Isla
nd
-by
Are
a -
May
1
97
0
Un
imp
lan
ted
Im
pla
nte
d
Pre
gn
ant
Pre
gp
ant
Fet
us
Wei
gh
t C
lass
A
rea
No
np
reg
nan
t N
o
No
1
2 3
4 5
Res
orp
tio
n
To
tal
A
Su
dak
-Pen
dan
t P
oin
t 12
7
26
8 30
0
2 0
3 2
2 27
( B
arn
es
Po
int-
Ho
t S
pri
ng
s B
ay
12
7 24
10
36
3
0 1
4 2
2 29
B
( (
Bar
nes
P
oin
t-T
run
k P
oin
t 1
1
6 25
7
29
0 3
1 3
0 2
24
( T
run
k P
oin
t-T
win
Bay
s 7
9 39
7
30
2 1
1 1
2 2
23
c ( (
Haz
ard
P
oin
t-T
win
Bay
s 4
3 27
4
36
0 0
1 1
2 11
( T
win
B
ays-
Her
d
Roc
k 1
2
8 28
9
31
1 1
1 Lf
2
29
D
( ( T
win
B
ays-
So
uth
Bay
2
9 45
9
45
3 0
1 1
Lf
1 20
( S
ou
th B
ay-L
ash
B
ay
6 9
29
16
51
1 0
0 9
6 1
31
E
( ((
So
uth
Bay
-Har
em R
ock
17
14
31
14
31
2 0
1 5
6 45
(
F ( (
L
ash
B
ay-I
nfe
rno
Ree
f 4
1 9
6 55
2
1 1
0 2
11
( G
(
Har
em R
ock
-Ku
lak
Po
int
12
6 29
3
14
1 0
1 1
0 21
H
Ku
lak
Po
int-
SE
B
igh
t 13
9
28
10
31
2 0
1 2
5 32
Bra
cket
s in
dic
ate
o
ver
lap
pin
g are
as
Sexual Segregation - 8 - March 1971
Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area
While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring
The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area
Sex Ratio
With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants
On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL
All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females
There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among
Sexual Segregation - 9 - March 1971
newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population
It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality
The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor
If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve
The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes
REPRODUCTION IN THE FElIALE SEA OTTER
by Karl Schneider Narch 1971
A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle
The follovJing criteria were used in classification
Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation
Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)
Anestrus - Largest follicle under 3 5 rnm no corpus luteum
Proestrus - Follicles over 35 mm
Es - Follicles approximately 10 rrm
Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm
- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~
al scar newly formed corpus albicans with some luteal tissue remaining
The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever
Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies
Tab
le
l Re
p iv
e co
nd
itio
n o
f se
xual
ly m
atur
e se
a o
tters
_
IF
etus
Wei
--------------+
__
_
ln~a
ctive
ov
arie
s 1
Act
ive ovari~
ht C
Jas
s
jmiddot
I P
ost
-I P
roes
tru
s U
nim
pl
lmpl
D
ates
L
ocat
ioQ
1A
nest
rus
Part
um
Res
orpt
ion
l +
E
stru
s P
reo
Pr
eQ
2 3
lJ
S
~
4-8
1970
T
anag
a 1
51
41
0
10
104
17
8 10
33
30
4 (1
68)
(1
35)
(3
3)
(3 3
) (2
89)
(34~
(5
6)
(27
) (3
3)
(11
4)(
10
9)
May
9
19
70
Del
arof
Is
18
14
1
4 13
27
10
0
3 6
8 77
4
) (1
82)
( l
3)
(5
(16
9)
( o
) (1
30
) (3
9)
(78
)(1
04
)
10-1
219
70 A
mch
ltka
I
34
18
3 8
27
48
5 6
2 23
12
13
8
( 0
(24
6)
(13
0)
(22
) (5
8)
(19
6)
(34
8)
(36
) (4
3)
(14
) (1
67)
(8
7)
~lune
23
middotmiddot ~middot~~middot~Amchitka
1
17
2 1
2 4
18
1 2
3 5
7 44
A
ugus
t 13
196
8 (3
86)
(4
5)
(23
) (4
5)
(90
) (4
09)
(2
3)
(45
) (6
8)
(11
3)(
15
8)
July
6-
~dgtAmchitka
1
17
1 0
4 14
10
0
1 1
2 6
46
Aug
ust
819
69
(36
9)
( 2
0 2
) (8
6)
(30
4)
(21
9)
(22
) (2
2)
(43
)(1
32
)
Sep
t 1
0-17
A
dak
72
5 0
16
31
40
6 6
8 9
11
164
1967
(4
39
) ( 3
1)
(98
) ( 1
89)
(
4)
(3 7
) (3
7)
(49
) (5
5)
(6
7)
Sep
t
25
-A
mch
itk
a l
55
6
0 18
19
17
4
0 0
0 3
115
OcL
6
19
67
(4
7~8)
(5
2)
(15
7)
(16
5)
(14
8)
(35
) (8
7)
(26
)
Oct
o 1
2-1
5
Kan
aga
I
58
6 1
13
31
31
4 4
7 11
5
140
1968
(4
13)
(4
3)
(o 7
) (9
3)
(22
2)
(22
2)
(29
) (2
9)
(50
) (7
8)
(36
)
Oct
18
-20
A
dak
l
46
6 0
1 24
13
2
3 0
4 4
100
1968
(L
16 0
) ( 6
0)
(11
0)
(24
0)
(13
0)
(20
) (3
0)
0)
(40
)
Num
bers
in
re
nth
esis
are
pe
rcen
t se
xu
ally
mat
ure
fem
ales
F
etus
wei
ght
clas
s 1
=
0-l
g
2 =
1-l
Og
3
= 1
0-lO
Og
4
= 1
00-lO
OO
g
5 =
Tra
nsp
lan
t m
ort
alit
ies
(all
o
ther
sam
ples
fr
om
har
ves
ts)
Reproduction - 2 - March 1971
There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time
The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable
Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries
Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d
2
~ _s ~
~
c U)
~-1
r
~J
lt
c])
Reproduction - 3 - Narch 1971
to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors
cussion of the Annual
The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases
The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season
The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses
By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7
blastocysts implanting as is shown the increase in Class 1 fetuses
By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses
This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay
At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling
Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses
a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die
Reproduction - 4 - March 1971
There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August
Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)
If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another
The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b
The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short
There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points
At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping
Reproduction - 5 - March 1971
Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever
At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month
These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup
Gestation Period
Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature
A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth
The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)
Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April
It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t
Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo
Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted
Reproduction - 6 - March 1971
to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year
From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months
The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest
Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period
The following is a comparison of the two estimates
Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s
Total Gestation Period 399 d2ys 154 days or about 13 months 5 months
Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period
This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad
Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)
Table 2 Estimated Length of Period
Percent of Fetuses in each Height Class
Time of Collection After January
1 month 31 15 15 38 0
3 months 18 12 22 35 12
5 months 18 8 8 36 30
7 months 3 8 16 29 45
9 months 20 7 10 40 23
10 months
14 18 12 33 23
---~--
r1ean Time in Nonths After January 50 57 53 58 70
Cube Root of Umveighted Mean 063 17 36 7lf lllf
Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days
Table 3 Length of Unimplanted Period
Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted
1 month 35 26 58
3 months 49 49 so
5 months 128 179 42
7 months 22 37 37
9 months 50 57 47
10 months 55 44 56
Unweighted
48
Reproduction - 7 - March 1971
All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples
The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available
This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless
Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent
There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months
Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts
Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of
11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data
The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a
Reproduction - 8 - March 1971
Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters
If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months
The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping
The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve
Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination
There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months
Fetal Rate
the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e
--
3 0
~
shy
----- -shy --middotmiddot~middotmiddot
___ ___1__
_
(middot-)
_ ft -) J ~)___
~-
3
Reproduction - 9 - IYarch 1971
The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)
Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year
Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)
The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based
For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)
A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit
The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined
2
G
X
middot ~ -- (-- ) r~ ~ - - -- ~
yen~ r- ~--~
3
i__middot -~---
~
gt
- cshy
Reproduction - 10 - March 1971
~e of _Sexual Maturity
Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition
Ovulation
From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)
faximum
Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point
Ovulation
Many mustelids are induced ovulatoTs
On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous
ficance of
Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the
season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary
----
a - o lt bull y bull bull l w laquo bull _ ~ ~
Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968
N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A
I
~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy
- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot
1-2
2
3
4 1 I
2 (1)
6
middot5
1 (1)1
2 (l) 17
1 ( 1 )
2 ( 1)
18
l (1) I ( 1 ) 9
10 1 ( 1) L ~j( 2 1 ( 1 )
2 ( l) 1
12
11
1 (1)
I I (I)
1
2 ( 1)
13
1 ( 1 )
2
114
115 I116
17
I 18
I I
I
119 I
Numbers inside parentheses =Number of individuals with corpus luteum
11
Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968
N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I
AGE 84 I I
5 I l 6 7z 3(Years) 1
l rbull ~middot--
--- shy
0-l
J l-2
~ 2
3
4 2 ( 1)
5 3
Ibullbull 6 6 (3)
4 (4)7
8 4
3 9 2 ( 1 )
j iI 10
I 1 (1)11
12
13
14
1 ( 1)
16
15
17
18
Numbers irisi
-
J
(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)
1
1 (1)
5 (1)
10(5)
3
3 1
3 (2)
2 ( 1) 3 (3)
1 ( 1)
2
1 ( 1 ) 5 ( 1)
1 ( 1 ) 3 ( 1)
1
3
2 ( 1 )
1 (l)
1 (1)4 ( 1)
2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )
1 ( 1)
1
2
1 ( 1)
1 ( 1)
1
I I parentheses = Number of individuals with corpus luteum
11
Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968
N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A
-1AGE
~
9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--
- _ Q~L
1-2
(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3
4 2 (1)
l5
16 1 (1)
1 1)7
8
2 (1)9
10
111
12
I13
14
15 1 (1) 16
17
18
1 ( 1)
1 (l)
1
1
1
2
1
1
1
1(1 )
1
1 (1)
1 (1)
1
I
1
1
1
1
-I
Numbers Inside rentheses = Number of individuals with corpus luteum
1
Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968
AGE (Years)
Q-1
t-2
2
3
4
s 6
7
8
9
10
11
12
13
14
15
16
17
18
Numbers
N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A
~L 3 4 slE - 1middotshy
7 1~8 __J~l_11 shy
1-
One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I
One vJi th corpus luteum but no corpora al bicantia
12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3
1 (1) 2 (1)
1 2 (2) l 1
1 (1) 4 (1)
2 (1)
2 1 )
1 ( 1 ) 1
2 (1)
2 (2)
l22 3
3 ( 1)2 21 ( 1)
2 ( 1)
2 ( 1)
l ( 1)1
1 1
1 I ll
I
I I
inside parentheses Nurnber of individuals with corpus luteurn
Reproduction - 11 - March 1971
The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year
Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out
The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve
Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es
It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the
ion and ended around parturi tioD
Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists
Fetal
Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land
Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation
The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first
If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water
Reproduction - 12 - tltIarch 1971
At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water
Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample
Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn
It appears that blastocysts rarely cross from one horn to the other
Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other
When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time
A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active
Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus
Reproductive - 13 - March 1971
Birth iltleigh t
Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g
The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g
One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected
From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high
This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation
This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier
This may have sone effect on information on this
Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females
~
(
~
-s 2 0 C
U)
0
--
ez ()
~) I) middot _t t- (
j -~middot -~
Reproduction - 14 - March 1971
This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition
Sex Ratio at Birth
Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained
In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female
Sex Ratio of the
In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales
The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population
The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be
Lit
The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported
In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8
Table 8 Multiple Corpora Lutea Fetuses
Total Pregnant Females 565
Implanted Pregnancies 316
Unimplanted with 2 CL (corpora lutea) 9
Implanted Pregnancies with ) L CL and 1 Fetus 8
Implanted Pregnancies Vlith 3 CL and 1 Fetus 1
Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5
Implanted Pregnancies middotwith 3 CL and gt Fetuses 1
Total Nultiple Ovulations
I
Percent Nultiple Ovulations 42
Percent of Pregnancies tvith Nultiple Fetuses 19
Corpora Lutea per Pregnancy 105
Fetuses per 102
Reproduction - 15 - lvlarch 1971
From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally
All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo
in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time
With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations
The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus
A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some
Des
Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)
Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)
Resorbed or aborted fetus (implantation at end of horn) 3
Resorbed fetus (umbilicus tvTisted tightly) 1
Resorbed fetuses (three or more fetuses in one horn) 2
Resorbed fetus (cause unknmm) 5
Reproduction - 16 - March 1971
Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents
The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area
While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail
near Parturition
Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition
Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality
after
Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9
It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long
We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life
Table 9 Pups Accompanying Pos artum Females
Sex llleight
M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11
Total Length
47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)
Pups Accompanying Anestrous Females
F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103
Reproduction - 17 - Narch 1971
Frequency of Breeding
Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups
I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases
Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample
Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large
vhich had luteinized but tvere not actually pregnant The remainder had follicles
This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt
Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning
Table 10 Reproductive Condition of Lactating Females
Location
Bay of Is Adak I Sept 1967 9 1 100
Allchitka I Sept-Oct 1967 17 2 105
Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I
Bay of Is Adak I Oct 1968 22 4 154 3
Kanaga I Oct 1968 32 s- 135
Al1chitka I July-Aug 1969 4 2 333
iTanaga I May 1970 56 10 152
Delarof Is May 1970 18 3 143
Amchitka I May 1970 36 3 77
---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy
TOTAL 213 33 134
Anestrous or pregnant with term fetus
]_ Large follicles or corpus luteum present
3 Includes one implanted pregnancy with small fetus
Reproduction - 18 - March 1971
The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy
The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year
Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period
This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup
We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year
Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~
Reproduction - 19 - March 1971
observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less
Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years
This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias
Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years
Reproduction - 20 - March 7 1971
Conclusions
1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population
2 Breeding activity is highest in September October and November
3 More implantations occur during January February and March than at other times of year
4 The birth rate is highest during April Y~y and June
5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity
6 Similarly two and a half to three times as many females pup during peak pupping months
7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed
8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup
9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period
10 Female sea otters become sexually mature ihen 3 to 4 years old
11 Females may ovulate on an average of once a year after sexual maturity is reached
12 Females continue to breed at Cn old age
13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution
14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented
15 Sea otters probably birth both on land and in the water
16 Blastocysts rarely cross from one uterine horn to the other
17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random
Reproduction - 21 - March 1971
18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn
19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth
20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal
21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth
22 Body condition of the female has little effect on the growth rate of the fetus
23 There is a rapid ~Jeight gain shortly after parturition
24 The sex ratio at birth is 44 percent male and 56 percent female
25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term
26 Multiple ovulations occur in 4 percent of the estrus cycles
27 es do not appeErc- to be able to more than two fetuses in a single horn
28 Up to 5 percent of the p may fail dne to in after implantation
29 An unknown but probably significant number of first pregnancies fail before implantation
30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life
31 Sexually matu1middote females normally have one pup every two years
32 Females vrith a pup may ovulate but rarely mate
33 Slightly less than half of the sexually mature females are in any given year
34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived
35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus
Reproduction - 22 - March 1971
CITED
Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp
Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317
Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68
Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657
Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130
Snow H J 1910 In forbidden seas Edward Arnold London 303 pp
Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp
Sea Otter Studies - 2 - ]1archt 1971
Location of Pregnancy Birth Weight Sex Ratio at Birth Sex Ratio of the Population Litter Size In Utero Mortality Mortality at or near Parturition Recovery of the Uterus after Parturition Frequency of Breeding Conclusions Literature Cited
AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OFTHE ALASKA PENINSULA
March 1970
Between March 16 and March 27 1970 an aerial count of sea otters was rnade along the south side of the Alaska Peninsula from Shaw Island to Cape Lazaref on Unlmak Island Sanak Island the Sandman Reefs the Pavlof Islands northern Shumagin Islands and Sutwick Island were included A BLM Aero Commander N6392U was flown by Cal Ward at an altitude averaging 150 feet and an airspeed bullgtf 100 mphmiddot Observers were Karl Schneider next to the pilot and Jim Faro behind the pilot This is the same technique aircraft pilot and observers as were used in the 1969 Aleutian count The count on March 27 was made in a Gruman Goose 1r1ith Robert Wolfe substituting for Faro Pups with their mothers were not counted
Because of conditions of visibility and distribution of animals all of the counts made are considered to be low
In general the variability in factors influencing this type of count is so great that the results are not rei iable for population estimates or for determining short-term fluctuations in dense populations However aerial counts should be useful to determine general distribution and abundance and to follow large changes in population size Such changes are occurring where relatively dense populations are expanding into habitat that is either unshyoccupied or sparsely occupied Aerial counts are also the quickest way to familiarize new personnel with large areas of habitat
The following is a 1ist of definitions of terms used to describe counting conditions Application of these terms is completely subjective and based on the writers experience
Excellent- Surface of water relatively calm Usually overcast with 1ittle surface glare Single animals easy to spot at a distance
Very good - Surface may be slightly choppy but not enough confusing reflection to prevent spotting of animals off shore
Good - Off shore animals may be difficult to spot but single animals near shore and in kelp beds and small pods everywhere are readily spotted
Fair -Usually choppy or surface glare Single animals ~in kelp beds or in the lee of rocks or shore and most groups of animals relatively easy to see Other single animals and some pods in deep bays in the shadow of cliffs or off shore may be missed
Poor -Single animals difficult to spot and many pods may be missed but conditions still good enough to get a rough idea of the distribution of animals
The conditions encountered for each area are listed below
Description of Conditions During March 1970 Aerial Count of Sea Otters
March 20 Aniakchak Bay to Chignik Lagoon (1022 AM- 1240 PM) Conditions were generally fair with a heavy chop The south side of Sutwick Island and the area between the island and mainland were poor with heavy surf Visibility was very good inside Kujulik Bay however many animals were scattered throughout the area On two passes ~cross the bay 12 and 27 were seen therefore many were probably missed Similar scattering occured in Chignik Bay The overall count for the area is probably low
Pavlof Bay to Cold Bay (130PM-200PM) Conditions poor with heavy chop and squalls The count was superficial however a second superfi cia 1 count of this a rea was made on March 21 and only one otter was seen in Cold Bay
Cold Bay to Caee Lazaref (310PM -420PM) Conditions very poor fair in a few spots Heavy chop dense kelp
March 21 Northern Shumagin Islands (943AM- 1137 AM) Poo~ with chop on windward sides Fai_r on leeward sides (westerly wind) however air turbulence forced us away from high cliffs Heavy surf created very poor conditions on the south side of Unga
Pavlof Islands (1145 AM- 125ff PM) Conditions similar to those in the Shumagins
Chignik Lagoon to Pavlof Bay (305 PM- 535 PM) Conditions varying from fair to poor with chop and surface glare Count concentrated on points and better looking areas Most deep bays not completely surveyed
March 22 Sanak Islands (849AM- 1020 AM) Conditions fair to poor with chop on south side and surface glare on north side Animals scattered and very difficult to see Many otters on rocks Count is probably very low
Sandman Reefs ( 1025 AM~ 1115 AM) Conditions similar to Sanak With the exception of one pod of 450+ the animals were widely scattered and very difficult to pick out Only those very close to the aircraft could be seen Bad squalls moved in and the wind increased preventing a complete count of Deer Island and the eastern half of the reefs
March 23 Wide Bay to Aniakchak Bay (1123 AM- 140PM) Conditions generally good at first becoming fair after Cape Providence with a light chop and some surface glare Otters in Amber and Aniakchak Bays were widely scattered and many may have been missed
Port Heiden to Naknek_ Superficial counts were made from Ugashik Bay to Naknek on March 19 and from Port Heiden to Egegik on March 23 These were made under poor conditions by flying outside the surf line The water was choppy and full of silt No sea otters were seen
11arch 27 Shaw Island to Puale Bay (1030 AM - 1205 PM) Conditions very good to excellent High overcast and very little wind down to Katmai Bay After this the conditions deteriorated rapidly to fair and the count had to be stopped because of a combination of bad weather and lack of fuel As shown on the charts the count was concentrated on the points and islands however the good visibility allowed us to see well into the bays We feel certain no concentrations were missed
The numbers of sea otters counted are presented in the Table Specific locations and numbers are shown on the charts Where coverage of an area was not complete the approximate path of the aircraft is shown
Discussion of Results
Surveys along the Alaska Peninsula have been fragmentary and most of those made were done under less than ideal conditionsmiddot There are a number of reasons for this Weather tends to be poor along the south shore where squall5 form along the mountains Areas where an aircraft can refuel are north of the mountains and there are relatively few passes therefore range of the aircraft is a problem Also there is much shallow water off shore in the area from the Shumagin Islands to Sanak Island Coverage of this area is difficult and many animals are undoubtably missed
The present survey covered most of the area however conditions were such that most of the counts are probably quite low Despite these problems and a lack of continuity we have a fairly good picture of the recovery of sea otter populations in this area
Reports from various individuals are available from the 1930s and 1940s 11ajor surveys by the Fish and Wildlife Service in 1951 (Jones) 1957 (Lensink) 1962 and 1965 (Kenyon and Spencer) covered at least portions of the area In 1969 the southern Shumagin Islands were counted by the Alaska Department of Fish and Game and the present survey covered most of the rest of the area
Basically there are four distinct population nuclei in the area These centered around the Sanak Island-Sandman Reef area the Shumagin Island area the Sutwick Island area and the Augustine Island-Cape Douglas area The following discussion is based on information from reports by Lensink and Kenyon and from Alaska Department of Fish and Game surveys middot
~anak Island-Sandman Reefs
Small numbers of sea otters have been reported at Sanak Island since 1922 No reports came from the Sandman Reefs until 1942 In 1948 27 were sighted at
----Cherni Island The 1951 aerial survey showed 65 around Sanak and 97 in the Sandman Reefs In 1957 251 were seen around Sanak and 508 around the Sandman Reefs In addition two were seen in the Pavlof Islands however few were on the mainland In 1962 548 were counted in the Sanak area and 638 in the Sandman Reefs None were reported from the mainland or Pavlof lsland~however
much of the increase was in the northern area around Deer Island The 1962 survey was probably the best being made under excel lent conditions
AERIAL COUNT OF SEA OTTERS MARCH 1970
Partial or Location Count Date Camp 1ete Count Visibility
Cape Lazaref - Cold lsecty_
Cape Lazaref 3 320 Complete Poor
lkatan Peninsula 71 320 Complete Poor
False Pass - Amagat I 66 320 Camp 1ete Poor
Cold Bay 321 Partial Poor
TOTAL 141
Sanak Islands 239 322 Camp 1ete Fair to Poor
Sandman Reefs
Clubbing Rocks 12 322 Complete Fair to Poor
Cherni I amp Rocks 495+ 322 Complete fair to Poor
Goose I 7 322 Complete Fair to Poor
Hay I 18 322 Camp lete Fair to Poor
Hunt I 2 322 Complete Fair to Poor
Deer I 322 Partial Fair to Poor~
TOTAL 568+
Pavlof Islands
Inner 11iasik 2 321 Camp 1ete Fair to Poor
Outer lliasik 16 321 Camp 1ete Fair to Poor
Goloi 2 321 Complete Fair to Poor
Dolgoi 67 321 Complete Fair to Poor
Poperechnoi 29 321 Complete Fair to Poor
Ukolnoi 2 321 Complete Fair to Poor
yenWosnesenski 4 321 Complete Fair to Poor
TOTAL 122
Partial or Location Count Date Comp 1ete Count vis ib j 1i ty
Northern Shumagin Islands
Unga 184 321 Complete Fair to Poor
Popof 52 321 Complete Fair to Poor
Korovin 46 321 Complete Fair to Poor
Karpa __2t 321 Complete Fair to Poor
TOTAL 286
Cold ~ to Beaver ~ 0 320-21 Partial Poor
Beaver Bay to Kupreanof Pen
Beaver Bay 2 321 Partial Fair to Poor
Cape A 1 iaks in 4 321 Partial Fair to Poor
Guillemot I 16 321 Partial Fair to Poor
__1Kupreanof Peninsula 321 Partial Fair to Poor
TOTAL 23
Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor
AntJrrCastle Cape to-1-Ji e ~
Castle Cape-Castle Bay 16 321 Partial Fair to Poor
Chignik Bay 118 320 Complete Fair
Nakchamik I 5 320 Complete Fair
Hook Bay 13 320 Complete Fair
Cape Kum 1i un 88 320 Complete Fair
Kujul ik Bay 1199 320 Complete Very Good
Univikshak I 62 320 Complete Fair
Cape Kuml ik 54 320 Complete Fair to Poor
Sutwick I 14 320 Complete Fair to Poor
Cape Agutka 323 Complete Fair~
TOTAL 1766
Partial or Location Count Date Comp 1ete Count vis i b i 1i ty
Cape Kunmik 13 323 Complete Fair
Naka 1i kok Bay amp offshore islands 16 323 Complete Fair
Chiginagak Bay 5 323 Complete Fair
Agripina amp lmuya Bay 105 323 Complete Good
Wide Bay to Puale Bay N 0 T S U R V E Y E D
Puale Bay to c Kubugakl i __]_ 327 Partial Fair
TOTAL 146
Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent
Cape Chiniak to Shaw _I
Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent
Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent
TOTAL 71
In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring
A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak
With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time
As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population
Shumagin Is 1 ands
This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population
There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion
In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good
There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied
There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas
Sutwi ck Area
Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather
In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved
~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather
The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys
There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years
to come
Augustine Island-Cape Douqlas
For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro
On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group
It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there
The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table
SUM
MAR
Y OF
SE
A OT
TER
SURV
EYS
(Com
pile
d by
p
op
ula
tio
n f
rom
L
ensi
nk
(196
2 T
hes
is
K
enyo
n 0
96
2 amp
196
5 R
epor
ts
amp M
anu
scri
pt)
an
d AD
FampG
Dat
a)
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
AU
GU
STIN
E -
C
DOUG
LAS
Aug
usti
ne
Isla
nd
A
ppro
xim
atel
y 50
(1
948)
Shaw
1
amp
Dou
glas
Are
a S
igh
tin
gs
from
Sh
aw
Is
to
Tux
edn
i B
ay
TOTA
L
40
___
_
41
52
-shy 52
18
_lQ
J
119
130+
-shy
130+
J
71
ALAS
KA
PEN
INSU
LA
c
Kul
iak
-
c
lgva
k
c
lgva
k -
c
Kun
mik
Ani
akch
ak
Bay
amp
Am
ber
Bay
Sutw
i ck
Are
a M
isce
llan
eous
R
epor
ts
Kuj
ul i
k B
ay
amp c
K
uml
ik
Sin
ce
1936
C
Kum
1 i u
n
Uni
viks
hak
Isla
nd
N
akch
amik
Is
lan
d
8
355 12
13
0 19 6
581
103
180
1
22
47
109
684 86
Not
S
urve
yed
7(+
)
139
197 14
1 2
53
101 62
5
I
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
ALAS
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nt
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gnik
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ay
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tle
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ay
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cle
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del
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Few
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19
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388
889 1 2 2 1
149
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338 I105
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118
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12
23
184 52
46 4
75
232 8 27 6
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
SHUM
AGIN
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on
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500
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imat
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ay
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141 4 2 29
67
Lo
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re
1951
19
51
1957
19
59
1962
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65
1969
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70
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r Is
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ther
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13 118
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Castle Bay - Castle Cape 16 Nakcharriik I 5 -- Katmai Reef 0
Total 152
middot
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Cape Agutka Cape Kum 1 i k Sutwick I~ Kujul ik Bay Cape Kum 1 i un Unavikshak 1
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X
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1199 88 62
1614
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MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970
OBSERVER J Vania J Faro
PILOT George Tibbets Jr
AIRCRAFr Piper Azetex
middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good
Departed King Salmon 1115 AM Returned to King Salmon 745 PM
FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out
We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9
The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down
SEA OTTER SIGHTINGS
Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed
The number counted in each pod is as follows
1071 520 357 68 36 75 30
TOTAL 2157
Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island
171
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53
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SEA OTTER COUNT BARREN ISLANDS and SHUYAK - AFOGNAK ISLANDS
June 9 1970
On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows
Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete
Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good
Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial
Area
BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island
AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island
TOTAL
TOTAL
Number of Sea Otters
0 2 0
76 200
29 0
307
18 6 3
121 0
33 26
207
Complete or Visibility Partial Count
Very good Complete ll If II
It II
II
II IJ
Good Complete II
Fair Partial If Complete II Partial II II
II
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SEA OTTER COUNTS - KENAI June J970 to January 1971
Carl Divinyi flew aerial surveys of seals along the outside of the
Kenai Peninsula on a monthly basis He recorded all sea otters sighted
on these surveys The surveys did not cover the entire coast line and
the type of aircraft weather conditions and observers varied from one
survey to another Therefore the seperate counts cannot be compared
However when viewed in total they indicate the areas used by sea otters
and the relative abundance of otters in each area
The attached table presents the counts by broad areas
--
SEA
OTT
ERS
CO
UN
TED
ON
A
ER
IAL
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O
F K
EN
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ne
1
97
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Jan
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1971
8i5
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amp 9
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JAN
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54
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tters
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20
70
AERIAL SURVEY Prince William Sound
April 1970
Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey
I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife
April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay
Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals
Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina
Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)
April 16 Weather inclement - no flying
April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island
April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)
Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands
The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed
SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date
PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418
CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52
PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS
ESTHER PASSAGE - VALDEZ ARM NS
GALENA BAY FISH BAY 103 415
FISH BAY - GRAVINA POINT 1 415
GRAVINA POINT - CORDOVA 1 415
HAWKINS ISLfu~D 1 4l7
HINCHINBROOK ISLfu~ 99 417
EGG ISLAND 2 417
MONTAGUE ISLAND 259 417
GREEN AND LITTLE GREEN ISLANDS 102 4J7
KNIGHT ISLAND 136 52
INGOT ISLAND 6 52
ELEANOR ISLAND 3 52
SMITH ISLAND 4 52
SEAL ISLAND 0 52
APPLEGATE ROCK 1 52
PERRY ISLAND NS
(continued) SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date-
LONE ISLAND NS
NAKED ISLAND NS
PEAK ISLAND NS
STOREY ISLAND NS
KAYAK - WINGHAM ISLAND 5 417
TOTAL 892
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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970
On July 17 1970 an aerial survey of sea otters was made in the area
around Green Island and the adjacent coast of Montague Island The purpose
of the survey was to locate concentrations of animals prior to a capturing
operation which began the next day A Dornier twin-engine arrcraft was used
with Schneider Reynolds and Somerville acting as observers The sky was
overcast almost no wind and the sea calm However heavy rain showers intershy
fered with forward visibility and the counting conditions were only fair on
the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay
The count began at 210pm and ended at 345 pm
The numbers and locations of otters are shown on the map In addition
16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy
in brook Island and a pod of 35 were seen three miles east of Johnstone Pt
Most of the area included in the survey was traveled repeatedly in a
skiff over the next four days Durlng this time the weather became calm and
clear for the first time in several days and the bulk of the sea otters shifted
from the coast of Montague out into the shallow areas between there and Green 1
and to Green and Channel Islands It was obvious that many otters had been missed
Fn the aerial sur~ey and that the population in the area is quite dense A
total of 48 sea otters were captured in three days of netting
j
SEA OTTER SURVEY Salisbury Sound to Cape Spencer
August 23-25~ 1970
On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and
middotWalt Cunningham serving as observers
The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good
On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere
On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May
Rough weather and outboard problems restricted tl the search area~
and the effectiveness of the search in that area The following sightings were made
82470 I Adult West of Granite Islands 1 Adult South of Granite Islands
11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands
These sightings represent from four to seven animals
Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull
The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal
The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait
While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island
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55
HARVEST AND TRANSPLANTS - 1970
Harvest
In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island
and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year
The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial
survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs
Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers
Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same
Transplants
Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent
The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island
Table 1
April 20
middotMay 1
May 2
May 3
May 4
May 5
May 6
May 7
May 8
May 9
May 10
May 11
May 12
May 13
Schedule of the 1970 harvest
1030 pm depart Homer
Midnight arrive Adak MV Active
Refuel and prepare boat
Arrive off Tanaga at noon
Complete skinning pack hides etc in pm
Delarofs
Spent am getting water at Amchitka
Clean up boat take down shelter pack gear
900 am arrive Amchitka fly to Anchorage
Killed 53
138
88
131
143
53
144
43
79
83
otter
(am)
(pm)
(by mid afternoon)
Table 2 Projected and actual harvest numbers May 1970
Projected Actual Tana~a Island Harvest Harvest
Bumpy Point - Hot Springs Bay 200 191
Hot Springs Bay - Twin Bays 50 50
Twin Bays - Cape Sasmik 50 ) ) 199
Cape Sasmik - Tower 120 )
Tower Tanaga Bay 180 166
TOTAL 600 606
Delarof Islands
Gareloi 20-25 0
Kavalga Ogliuga and Skagul 75-100 125
Ulak and Amatignak 35-55 19
TOTAL 150 144
Amchitka Island saved for
USFWS Counting Units 1 amp 2 100 transplant
3 30 82
4 50 36
5 120 87
TOTAL 300 205
Table 3 Details of sea otters harvested in May 1970
Tanaga Island
Total kill 606
Number of pelts saved 598
Number of small pup pelts discarded 8
Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796
(Seal 3770 void - damaged)
Approximate sex ratio - 220 males 386 females or approximately 64 percent females
Delarof Islands
Total kill 144
Number of pelts saved 138
Number of pups discarded 6
Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934
Sex ratio- 44 males 100 females or approximately 69 percent females
Amchitka Island
Total kill 205
Number of pelts saved 194
Number of pups discarded 11
Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128
Sex ratio- 27 males 178 females or approximately 867 percent females
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A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality
Summary of Harvests and Transplants
Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area
The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5
----
----
----
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----
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----
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Tab
le 4
N
umbe
rs
of
sea o
tters
re
mov
ed
fro
m Ala
skan
pop
ulat
ions
~ 1
95
17
01
19
51
5
4
55
56
57
59
60
62
6
3
65
66
67
68
69
70
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chit
ka I
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ies
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22
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2
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ran
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nt
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6
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86
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DFamp
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ves
t 1
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2
3
205
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er
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1
14
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To
tal
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22
37
26
21
39
1
80
31
1
210
50
0
25
1
292
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aga
I
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ves
t 6
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aga
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ves
t 3
18
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k I
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ves
t 30
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94
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in Is
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ies
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ak amp
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mak
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chin
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ok
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ran
spla
nt
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die
s 1
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nta
gu
e amp
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nsp
lan
t 39
46
G
reen
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tud
ies
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aro
f Is
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res
t O
gli
ug
a amp
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agu
l 1
25
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lak
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9
I
Inclu
des
all
an
imal
s th
at
wer
e h
arv
est
ed
d
ied
d
uri
ng
st
ud
ies
and
tran
spla
nt
att
em
pts
o
r w
ere
tran
spla
nte
d
to
oth
er
are
as
or
zoo
s
Doe
s n
ot
inclu
de n
atu
ral
mo
rtali
ty o
r il
leg
al
kil
ls
2
Woo
dlan
d P
ark
Zoo
3
B
att
ell
e M
emo
rial
In
sti
tute
--
Tab
le 5
N
umbe
rs
of
sea o
tters
tr
an
spla
nte
d 1
95
5-1
97
0
1955
19
56
1959
19
65
--
~ 1
96
6
1968
19
69
1970
Ale
uti
an
s A
ttu
I
5
Ott
er
I
19
1
Pri
bil
ofs
S
t
Pau
l I
7 S
t
Geo
rge
I
57
Yak
uta
t B
ay
10
Kha
z B
ay
23
20
93
58
(Ch
ich
igo
f L
)
So
uth
east
Y
akob
i I
30
A
lask
a B
iork
a I
48
Barr
ier
Is
55
Hec
eta
I
51
Cap
e S
pen
cer
25
Bri
tish
C
olum
bia
29
14
291
W
ash
ing
ton
Ore
gon
29
1
Non
e b
eli
ev
ed
to
hav
e su
rviv
ed
II
A
t le
ast
13
die
d s
ho
rtly
aft
er
rele
ase
NO
TE
1955
to
19
59 b
y U
SFW
S
1965
to
19
70 b
y
AD
FampG
In
som
e ca
ses
on
e o
r tw
o o
f th
e
abo
ve
anim
als
die
d n
ear
the
tim
e o
f re
lease
30
SEXUAL SEGREGATION IN SEA OTTER POPULATIONS
by Karl Schneider March 1971
Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males
Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands
Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals
Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas
A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population
In
the
By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled
Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet
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Sexual Segregation - 2 - March 7 1971
Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples
Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground
Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas
Identification of Hale Areas by Pup Counts
In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable
From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen
Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area
The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on
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Sexual Segregation - 3 - March 1 1971
Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side
The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed
Kanaga Island
Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$
Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point
Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak
Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass
The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring
Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area
Tab
le
5
Com
pari
son
of
Pup
s C
ou
nte
d w
ith
S
ex R
atio
o
f H
arv
est
Jun
e
1968
A
rea
Su
rvey
O
cto
ber
19
68 H
arv
est
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l KA
NAGA
IS
LAN
D
Ind
ep
Ott
er
Ind
ep
Ott
er
M
M
F
Rou
nd
Hea
d-S
ho
al P
oin
t 0
85
3
45
6
55
4
71
5
29
Nag
a P
oin
t-K
anag
a B
ay
66
1
64
middot
18
5
81
5
20
5
79
5
Cap
e C
hla
nak
-Cap
e T
usi
k
51
1
93
2
40
7
60
3
53
6
47
Edd
y R
ock
-Hiv
e R
ock
61
3
4
35
7
64
3
47
1
52
9
Jun
e
1968
A
rea
Su
rvey
H
ay
1970
Har
ves
t
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l TA
NA
GA
IS
LAN
D
Ind
ep
Ott
er ~ter
M
F
M
F
Cap
e S
ud
ak-P
end
ant
Po
int
0 4
4
60
6
39
4
66
4
33
6
Bar
nes
P
oin
t-T
run
k P
oin
t 7
9
59
3
3
96
7
83
9
17
Tru
nk
Po
int-
Tw
in B
ays
26
4
2
77
9
23
1
40
8
60
D
rin
Bay
s-S
ou
th B
ay
96
1
11
7
3
92
7
14
3
85
7
So
uth
Bay
-Har
em R
ock
14
8
31
6
7
middot93
3
91
9
09
Las
h
Bay
-In
fern
o
Ree
f 1
5
73
0
27
0
74
6
25
4
Har
em R
ock
-Ku
lak
Po
int
0 2
7
64
4
35
6
68
4
31
6
Ku
lak
Po
int-
SE
B
igh
t 6
9
82
1
32
8
68
1
51
8
49
Sexual Segregation - 4 - March~ 1971
Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands
Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area
As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method
Identification of
Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area
The follmving is a description of the sex composition of areas for which no skiff count data was available
Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island
Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill
Sexual Segregation - 5 - March 1971
Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point
The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it
Composition of Female Areas
Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female
The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure
Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters
Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males
Sexual Segregation - 6 - March~ 1971
apparently defending a territory in California but from his observations t
it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change
Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year
A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest
The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas
The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males
There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas
Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak
within Female
In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of
Sexual Segregation - 7 - March 1971
these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June
The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups
Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population
ition of Male Areas
As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles
Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements
le 6
R
epro
du
ctiv
e S
tag
e o
f S
exu
ally
Mat
ure
S
ea O
tters
Har
ves
ted
on
Tan
aga
Isla
nd
-by
Are
a -
May
1
97
0
Un
imp
lan
ted
Im
pla
nte
d
Pre
gn
ant
Pre
gp
ant
Fet
us
Wei
gh
t C
lass
A
rea
No
np
reg
nan
t N
o
No
1
2 3
4 5
Res
orp
tio
n
To
tal
A
Su
dak
-Pen
dan
t P
oin
t 12
7
26
8 30
0
2 0
3 2
2 27
( B
arn
es
Po
int-
Ho
t S
pri
ng
s B
ay
12
7 24
10
36
3
0 1
4 2
2 29
B
( (
Bar
nes
P
oin
t-T
run
k P
oin
t 1
1
6 25
7
29
0 3
1 3
0 2
24
( T
run
k P
oin
t-T
win
Bay
s 7
9 39
7
30
2 1
1 1
2 2
23
c ( (
Haz
ard
P
oin
t-T
win
Bay
s 4
3 27
4
36
0 0
1 1
2 11
( T
win
B
ays-
Her
d
Roc
k 1
2
8 28
9
31
1 1
1 Lf
2
29
D
( ( T
win
B
ays-
So
uth
Bay
2
9 45
9
45
3 0
1 1
Lf
1 20
( S
ou
th B
ay-L
ash
B
ay
6 9
29
16
51
1 0
0 9
6 1
31
E
( ((
So
uth
Bay
-Har
em R
ock
17
14
31
14
31
2 0
1 5
6 45
(
F ( (
L
ash
B
ay-I
nfe
rno
Ree
f 4
1 9
6 55
2
1 1
0 2
11
( G
(
Har
em R
ock
-Ku
lak
Po
int
12
6 29
3
14
1 0
1 1
0 21
H
Ku
lak
Po
int-
SE
B
igh
t 13
9
28
10
31
2 0
1 2
5 32
Bra
cket
s in
dic
ate
o
ver
lap
pin
g are
as
Sexual Segregation - 8 - March 1971
Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area
While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring
The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area
Sex Ratio
With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants
On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL
All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females
There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among
Sexual Segregation - 9 - March 1971
newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population
It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality
The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor
If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve
The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes
REPRODUCTION IN THE FElIALE SEA OTTER
by Karl Schneider Narch 1971
A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle
The follovJing criteria were used in classification
Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation
Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)
Anestrus - Largest follicle under 3 5 rnm no corpus luteum
Proestrus - Follicles over 35 mm
Es - Follicles approximately 10 rrm
Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm
- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~
al scar newly formed corpus albicans with some luteal tissue remaining
The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever
Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies
Tab
le
l Re
p iv
e co
nd
itio
n o
f se
xual
ly m
atur
e se
a o
tters
_
IF
etus
Wei
--------------+
__
_
ln~a
ctive
ov
arie
s 1
Act
ive ovari~
ht C
Jas
s
jmiddot
I P
ost
-I P
roes
tru
s U
nim
pl
lmpl
D
ates
L
ocat
ioQ
1A
nest
rus
Part
um
Res
orpt
ion
l +
E
stru
s P
reo
Pr
eQ
2 3
lJ
S
~
4-8
1970
T
anag
a 1
51
41
0
10
104
17
8 10
33
30
4 (1
68)
(1
35)
(3
3)
(3 3
) (2
89)
(34~
(5
6)
(27
) (3
3)
(11
4)(
10
9)
May
9
19
70
Del
arof
Is
18
14
1
4 13
27
10
0
3 6
8 77
4
) (1
82)
( l
3)
(5
(16
9)
( o
) (1
30
) (3
9)
(78
)(1
04
)
10-1
219
70 A
mch
ltka
I
34
18
3 8
27
48
5 6
2 23
12
13
8
( 0
(24
6)
(13
0)
(22
) (5
8)
(19
6)
(34
8)
(36
) (4
3)
(14
) (1
67)
(8
7)
~lune
23
middotmiddot ~middot~~middot~Amchitka
1
17
2 1
2 4
18
1 2
3 5
7 44
A
ugus
t 13
196
8 (3
86)
(4
5)
(23
) (4
5)
(90
) (4
09)
(2
3)
(45
) (6
8)
(11
3)(
15
8)
July
6-
~dgtAmchitka
1
17
1 0
4 14
10
0
1 1
2 6
46
Aug
ust
819
69
(36
9)
( 2
0 2
) (8
6)
(30
4)
(21
9)
(22
) (2
2)
(43
)(1
32
)
Sep
t 1
0-17
A
dak
72
5 0
16
31
40
6 6
8 9
11
164
1967
(4
39
) ( 3
1)
(98
) ( 1
89)
(
4)
(3 7
) (3
7)
(49
) (5
5)
(6
7)
Sep
t
25
-A
mch
itk
a l
55
6
0 18
19
17
4
0 0
0 3
115
OcL
6
19
67
(4
7~8)
(5
2)
(15
7)
(16
5)
(14
8)
(35
) (8
7)
(26
)
Oct
o 1
2-1
5
Kan
aga
I
58
6 1
13
31
31
4 4
7 11
5
140
1968
(4
13)
(4
3)
(o 7
) (9
3)
(22
2)
(22
2)
(29
) (2
9)
(50
) (7
8)
(36
)
Oct
18
-20
A
dak
l
46
6 0
1 24
13
2
3 0
4 4
100
1968
(L
16 0
) ( 6
0)
(11
0)
(24
0)
(13
0)
(20
) (3
0)
0)
(40
)
Num
bers
in
re
nth
esis
are
pe
rcen
t se
xu
ally
mat
ure
fem
ales
F
etus
wei
ght
clas
s 1
=
0-l
g
2 =
1-l
Og
3
= 1
0-lO
Og
4
= 1
00-lO
OO
g
5 =
Tra
nsp
lan
t m
ort
alit
ies
(all
o
ther
sam
ples
fr
om
har
ves
ts)
Reproduction - 2 - March 1971
There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time
The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable
Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries
Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d
2
~ _s ~
~
c U)
~-1
r
~J
lt
c])
Reproduction - 3 - Narch 1971
to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors
cussion of the Annual
The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases
The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season
The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses
By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7
blastocysts implanting as is shown the increase in Class 1 fetuses
By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses
This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay
At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling
Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses
a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die
Reproduction - 4 - March 1971
There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August
Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)
If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another
The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b
The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short
There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points
At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping
Reproduction - 5 - March 1971
Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever
At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month
These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup
Gestation Period
Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature
A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth
The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)
Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April
It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t
Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo
Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted
Reproduction - 6 - March 1971
to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year
From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months
The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest
Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period
The following is a comparison of the two estimates
Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s
Total Gestation Period 399 d2ys 154 days or about 13 months 5 months
Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period
This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad
Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)
Table 2 Estimated Length of Period
Percent of Fetuses in each Height Class
Time of Collection After January
1 month 31 15 15 38 0
3 months 18 12 22 35 12
5 months 18 8 8 36 30
7 months 3 8 16 29 45
9 months 20 7 10 40 23
10 months
14 18 12 33 23
---~--
r1ean Time in Nonths After January 50 57 53 58 70
Cube Root of Umveighted Mean 063 17 36 7lf lllf
Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days
Table 3 Length of Unimplanted Period
Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted
1 month 35 26 58
3 months 49 49 so
5 months 128 179 42
7 months 22 37 37
9 months 50 57 47
10 months 55 44 56
Unweighted
48
Reproduction - 7 - March 1971
All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples
The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available
This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless
Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent
There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months
Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts
Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of
11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data
The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a
Reproduction - 8 - March 1971
Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters
If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months
The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping
The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve
Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination
There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months
Fetal Rate
the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e
--
3 0
~
shy
----- -shy --middotmiddot~middotmiddot
___ ___1__
_
(middot-)
_ ft -) J ~)___
~-
3
Reproduction - 9 - IYarch 1971
The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)
Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year
Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)
The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based
For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)
A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit
The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined
2
G
X
middot ~ -- (-- ) r~ ~ - - -- ~
yen~ r- ~--~
3
i__middot -~---
~
gt
- cshy
Reproduction - 10 - March 1971
~e of _Sexual Maturity
Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition
Ovulation
From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)
faximum
Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point
Ovulation
Many mustelids are induced ovulatoTs
On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous
ficance of
Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the
season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary
----
a - o lt bull y bull bull l w laquo bull _ ~ ~
Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968
N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A
I
~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy
- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot
1-2
2
3
4 1 I
2 (1)
6
middot5
1 (1)1
2 (l) 17
1 ( 1 )
2 ( 1)
18
l (1) I ( 1 ) 9
10 1 ( 1) L ~j( 2 1 ( 1 )
2 ( l) 1
12
11
1 (1)
I I (I)
1
2 ( 1)
13
1 ( 1 )
2
114
115 I116
17
I 18
I I
I
119 I
Numbers inside parentheses =Number of individuals with corpus luteum
11
Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968
N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I
AGE 84 I I
5 I l 6 7z 3(Years) 1
l rbull ~middot--
--- shy
0-l
J l-2
~ 2
3
4 2 ( 1)
5 3
Ibullbull 6 6 (3)
4 (4)7
8 4
3 9 2 ( 1 )
j iI 10
I 1 (1)11
12
13
14
1 ( 1)
16
15
17
18
Numbers irisi
-
J
(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)
1
1 (1)
5 (1)
10(5)
3
3 1
3 (2)
2 ( 1) 3 (3)
1 ( 1)
2
1 ( 1 ) 5 ( 1)
1 ( 1 ) 3 ( 1)
1
3
2 ( 1 )
1 (l)
1 (1)4 ( 1)
2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )
1 ( 1)
1
2
1 ( 1)
1 ( 1)
1
I I parentheses = Number of individuals with corpus luteum
11
Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968
N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A
-1AGE
~
9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--
- _ Q~L
1-2
(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3
4 2 (1)
l5
16 1 (1)
1 1)7
8
2 (1)9
10
111
12
I13
14
15 1 (1) 16
17
18
1 ( 1)
1 (l)
1
1
1
2
1
1
1
1(1 )
1
1 (1)
1 (1)
1
I
1
1
1
1
-I
Numbers Inside rentheses = Number of individuals with corpus luteum
1
Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968
AGE (Years)
Q-1
t-2
2
3
4
s 6
7
8
9
10
11
12
13
14
15
16
17
18
Numbers
N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A
~L 3 4 slE - 1middotshy
7 1~8 __J~l_11 shy
1-
One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I
One vJi th corpus luteum but no corpora al bicantia
12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3
1 (1) 2 (1)
1 2 (2) l 1
1 (1) 4 (1)
2 (1)
2 1 )
1 ( 1 ) 1
2 (1)
2 (2)
l22 3
3 ( 1)2 21 ( 1)
2 ( 1)
2 ( 1)
l ( 1)1
1 1
1 I ll
I
I I
inside parentheses Nurnber of individuals with corpus luteurn
Reproduction - 11 - March 1971
The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year
Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out
The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve
Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es
It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the
ion and ended around parturi tioD
Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists
Fetal
Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land
Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation
The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first
If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water
Reproduction - 12 - tltIarch 1971
At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water
Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample
Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn
It appears that blastocysts rarely cross from one horn to the other
Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other
When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time
A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active
Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus
Reproductive - 13 - March 1971
Birth iltleigh t
Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g
The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g
One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected
From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high
This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation
This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier
This may have sone effect on information on this
Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females
~
(
~
-s 2 0 C
U)
0
--
ez ()
~) I) middot _t t- (
j -~middot -~
Reproduction - 14 - March 1971
This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition
Sex Ratio at Birth
Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained
In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female
Sex Ratio of the
In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales
The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population
The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be
Lit
The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported
In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8
Table 8 Multiple Corpora Lutea Fetuses
Total Pregnant Females 565
Implanted Pregnancies 316
Unimplanted with 2 CL (corpora lutea) 9
Implanted Pregnancies with ) L CL and 1 Fetus 8
Implanted Pregnancies Vlith 3 CL and 1 Fetus 1
Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5
Implanted Pregnancies middotwith 3 CL and gt Fetuses 1
Total Nultiple Ovulations
I
Percent Nultiple Ovulations 42
Percent of Pregnancies tvith Nultiple Fetuses 19
Corpora Lutea per Pregnancy 105
Fetuses per 102
Reproduction - 15 - lvlarch 1971
From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally
All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo
in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time
With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations
The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus
A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some
Des
Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)
Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)
Resorbed or aborted fetus (implantation at end of horn) 3
Resorbed fetus (umbilicus tvTisted tightly) 1
Resorbed fetuses (three or more fetuses in one horn) 2
Resorbed fetus (cause unknmm) 5
Reproduction - 16 - March 1971
Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents
The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area
While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail
near Parturition
Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition
Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality
after
Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9
It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long
We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life
Table 9 Pups Accompanying Pos artum Females
Sex llleight
M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11
Total Length
47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)
Pups Accompanying Anestrous Females
F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103
Reproduction - 17 - Narch 1971
Frequency of Breeding
Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups
I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases
Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample
Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large
vhich had luteinized but tvere not actually pregnant The remainder had follicles
This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt
Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning
Table 10 Reproductive Condition of Lactating Females
Location
Bay of Is Adak I Sept 1967 9 1 100
Allchitka I Sept-Oct 1967 17 2 105
Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I
Bay of Is Adak I Oct 1968 22 4 154 3
Kanaga I Oct 1968 32 s- 135
Al1chitka I July-Aug 1969 4 2 333
iTanaga I May 1970 56 10 152
Delarof Is May 1970 18 3 143
Amchitka I May 1970 36 3 77
---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy
TOTAL 213 33 134
Anestrous or pregnant with term fetus
]_ Large follicles or corpus luteum present
3 Includes one implanted pregnancy with small fetus
Reproduction - 18 - March 1971
The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy
The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year
Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period
This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup
We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year
Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~
Reproduction - 19 - March 1971
observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less
Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years
This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias
Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years
Reproduction - 20 - March 7 1971
Conclusions
1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population
2 Breeding activity is highest in September October and November
3 More implantations occur during January February and March than at other times of year
4 The birth rate is highest during April Y~y and June
5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity
6 Similarly two and a half to three times as many females pup during peak pupping months
7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed
8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup
9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period
10 Female sea otters become sexually mature ihen 3 to 4 years old
11 Females may ovulate on an average of once a year after sexual maturity is reached
12 Females continue to breed at Cn old age
13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution
14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented
15 Sea otters probably birth both on land and in the water
16 Blastocysts rarely cross from one uterine horn to the other
17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random
Reproduction - 21 - March 1971
18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn
19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth
20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal
21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth
22 Body condition of the female has little effect on the growth rate of the fetus
23 There is a rapid ~Jeight gain shortly after parturition
24 The sex ratio at birth is 44 percent male and 56 percent female
25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term
26 Multiple ovulations occur in 4 percent of the estrus cycles
27 es do not appeErc- to be able to more than two fetuses in a single horn
28 Up to 5 percent of the p may fail dne to in after implantation
29 An unknown but probably significant number of first pregnancies fail before implantation
30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life
31 Sexually matu1middote females normally have one pup every two years
32 Females vrith a pup may ovulate but rarely mate
33 Slightly less than half of the sexually mature females are in any given year
34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived
35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus
Reproduction - 22 - March 1971
CITED
Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp
Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317
Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68
Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657
Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130
Snow H J 1910 In forbidden seas Edward Arnold London 303 pp
Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp
AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OFTHE ALASKA PENINSULA
March 1970
Between March 16 and March 27 1970 an aerial count of sea otters was rnade along the south side of the Alaska Peninsula from Shaw Island to Cape Lazaref on Unlmak Island Sanak Island the Sandman Reefs the Pavlof Islands northern Shumagin Islands and Sutwick Island were included A BLM Aero Commander N6392U was flown by Cal Ward at an altitude averaging 150 feet and an airspeed bullgtf 100 mphmiddot Observers were Karl Schneider next to the pilot and Jim Faro behind the pilot This is the same technique aircraft pilot and observers as were used in the 1969 Aleutian count The count on March 27 was made in a Gruman Goose 1r1ith Robert Wolfe substituting for Faro Pups with their mothers were not counted
Because of conditions of visibility and distribution of animals all of the counts made are considered to be low
In general the variability in factors influencing this type of count is so great that the results are not rei iable for population estimates or for determining short-term fluctuations in dense populations However aerial counts should be useful to determine general distribution and abundance and to follow large changes in population size Such changes are occurring where relatively dense populations are expanding into habitat that is either unshyoccupied or sparsely occupied Aerial counts are also the quickest way to familiarize new personnel with large areas of habitat
The following is a 1ist of definitions of terms used to describe counting conditions Application of these terms is completely subjective and based on the writers experience
Excellent- Surface of water relatively calm Usually overcast with 1ittle surface glare Single animals easy to spot at a distance
Very good - Surface may be slightly choppy but not enough confusing reflection to prevent spotting of animals off shore
Good - Off shore animals may be difficult to spot but single animals near shore and in kelp beds and small pods everywhere are readily spotted
Fair -Usually choppy or surface glare Single animals ~in kelp beds or in the lee of rocks or shore and most groups of animals relatively easy to see Other single animals and some pods in deep bays in the shadow of cliffs or off shore may be missed
Poor -Single animals difficult to spot and many pods may be missed but conditions still good enough to get a rough idea of the distribution of animals
The conditions encountered for each area are listed below
Description of Conditions During March 1970 Aerial Count of Sea Otters
March 20 Aniakchak Bay to Chignik Lagoon (1022 AM- 1240 PM) Conditions were generally fair with a heavy chop The south side of Sutwick Island and the area between the island and mainland were poor with heavy surf Visibility was very good inside Kujulik Bay however many animals were scattered throughout the area On two passes ~cross the bay 12 and 27 were seen therefore many were probably missed Similar scattering occured in Chignik Bay The overall count for the area is probably low
Pavlof Bay to Cold Bay (130PM-200PM) Conditions poor with heavy chop and squalls The count was superficial however a second superfi cia 1 count of this a rea was made on March 21 and only one otter was seen in Cold Bay
Cold Bay to Caee Lazaref (310PM -420PM) Conditions very poor fair in a few spots Heavy chop dense kelp
March 21 Northern Shumagin Islands (943AM- 1137 AM) Poo~ with chop on windward sides Fai_r on leeward sides (westerly wind) however air turbulence forced us away from high cliffs Heavy surf created very poor conditions on the south side of Unga
Pavlof Islands (1145 AM- 125ff PM) Conditions similar to those in the Shumagins
Chignik Lagoon to Pavlof Bay (305 PM- 535 PM) Conditions varying from fair to poor with chop and surface glare Count concentrated on points and better looking areas Most deep bays not completely surveyed
March 22 Sanak Islands (849AM- 1020 AM) Conditions fair to poor with chop on south side and surface glare on north side Animals scattered and very difficult to see Many otters on rocks Count is probably very low
Sandman Reefs ( 1025 AM~ 1115 AM) Conditions similar to Sanak With the exception of one pod of 450+ the animals were widely scattered and very difficult to pick out Only those very close to the aircraft could be seen Bad squalls moved in and the wind increased preventing a complete count of Deer Island and the eastern half of the reefs
March 23 Wide Bay to Aniakchak Bay (1123 AM- 140PM) Conditions generally good at first becoming fair after Cape Providence with a light chop and some surface glare Otters in Amber and Aniakchak Bays were widely scattered and many may have been missed
Port Heiden to Naknek_ Superficial counts were made from Ugashik Bay to Naknek on March 19 and from Port Heiden to Egegik on March 23 These were made under poor conditions by flying outside the surf line The water was choppy and full of silt No sea otters were seen
11arch 27 Shaw Island to Puale Bay (1030 AM - 1205 PM) Conditions very good to excellent High overcast and very little wind down to Katmai Bay After this the conditions deteriorated rapidly to fair and the count had to be stopped because of a combination of bad weather and lack of fuel As shown on the charts the count was concentrated on the points and islands however the good visibility allowed us to see well into the bays We feel certain no concentrations were missed
The numbers of sea otters counted are presented in the Table Specific locations and numbers are shown on the charts Where coverage of an area was not complete the approximate path of the aircraft is shown
Discussion of Results
Surveys along the Alaska Peninsula have been fragmentary and most of those made were done under less than ideal conditionsmiddot There are a number of reasons for this Weather tends to be poor along the south shore where squall5 form along the mountains Areas where an aircraft can refuel are north of the mountains and there are relatively few passes therefore range of the aircraft is a problem Also there is much shallow water off shore in the area from the Shumagin Islands to Sanak Island Coverage of this area is difficult and many animals are undoubtably missed
The present survey covered most of the area however conditions were such that most of the counts are probably quite low Despite these problems and a lack of continuity we have a fairly good picture of the recovery of sea otter populations in this area
Reports from various individuals are available from the 1930s and 1940s 11ajor surveys by the Fish and Wildlife Service in 1951 (Jones) 1957 (Lensink) 1962 and 1965 (Kenyon and Spencer) covered at least portions of the area In 1969 the southern Shumagin Islands were counted by the Alaska Department of Fish and Game and the present survey covered most of the rest of the area
Basically there are four distinct population nuclei in the area These centered around the Sanak Island-Sandman Reef area the Shumagin Island area the Sutwick Island area and the Augustine Island-Cape Douglas area The following discussion is based on information from reports by Lensink and Kenyon and from Alaska Department of Fish and Game surveys middot
~anak Island-Sandman Reefs
Small numbers of sea otters have been reported at Sanak Island since 1922 No reports came from the Sandman Reefs until 1942 In 1948 27 were sighted at
----Cherni Island The 1951 aerial survey showed 65 around Sanak and 97 in the Sandman Reefs In 1957 251 were seen around Sanak and 508 around the Sandman Reefs In addition two were seen in the Pavlof Islands however few were on the mainland In 1962 548 were counted in the Sanak area and 638 in the Sandman Reefs None were reported from the mainland or Pavlof lsland~however
much of the increase was in the northern area around Deer Island The 1962 survey was probably the best being made under excel lent conditions
AERIAL COUNT OF SEA OTTERS MARCH 1970
Partial or Location Count Date Camp 1ete Count Visibility
Cape Lazaref - Cold lsecty_
Cape Lazaref 3 320 Complete Poor
lkatan Peninsula 71 320 Complete Poor
False Pass - Amagat I 66 320 Camp 1ete Poor
Cold Bay 321 Partial Poor
TOTAL 141
Sanak Islands 239 322 Camp 1ete Fair to Poor
Sandman Reefs
Clubbing Rocks 12 322 Complete Fair to Poor
Cherni I amp Rocks 495+ 322 Complete fair to Poor
Goose I 7 322 Complete Fair to Poor
Hay I 18 322 Camp lete Fair to Poor
Hunt I 2 322 Complete Fair to Poor
Deer I 322 Partial Fair to Poor~
TOTAL 568+
Pavlof Islands
Inner 11iasik 2 321 Camp 1ete Fair to Poor
Outer lliasik 16 321 Camp 1ete Fair to Poor
Goloi 2 321 Complete Fair to Poor
Dolgoi 67 321 Complete Fair to Poor
Poperechnoi 29 321 Complete Fair to Poor
Ukolnoi 2 321 Complete Fair to Poor
yenWosnesenski 4 321 Complete Fair to Poor
TOTAL 122
Partial or Location Count Date Comp 1ete Count vis ib j 1i ty
Northern Shumagin Islands
Unga 184 321 Complete Fair to Poor
Popof 52 321 Complete Fair to Poor
Korovin 46 321 Complete Fair to Poor
Karpa __2t 321 Complete Fair to Poor
TOTAL 286
Cold ~ to Beaver ~ 0 320-21 Partial Poor
Beaver Bay to Kupreanof Pen
Beaver Bay 2 321 Partial Fair to Poor
Cape A 1 iaks in 4 321 Partial Fair to Poor
Guillemot I 16 321 Partial Fair to Poor
__1Kupreanof Peninsula 321 Partial Fair to Poor
TOTAL 23
Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor
AntJrrCastle Cape to-1-Ji e ~
Castle Cape-Castle Bay 16 321 Partial Fair to Poor
Chignik Bay 118 320 Complete Fair
Nakchamik I 5 320 Complete Fair
Hook Bay 13 320 Complete Fair
Cape Kum 1i un 88 320 Complete Fair
Kujul ik Bay 1199 320 Complete Very Good
Univikshak I 62 320 Complete Fair
Cape Kuml ik 54 320 Complete Fair to Poor
Sutwick I 14 320 Complete Fair to Poor
Cape Agutka 323 Complete Fair~
TOTAL 1766
Partial or Location Count Date Comp 1ete Count vis i b i 1i ty
Cape Kunmik 13 323 Complete Fair
Naka 1i kok Bay amp offshore islands 16 323 Complete Fair
Chiginagak Bay 5 323 Complete Fair
Agripina amp lmuya Bay 105 323 Complete Good
Wide Bay to Puale Bay N 0 T S U R V E Y E D
Puale Bay to c Kubugakl i __]_ 327 Partial Fair
TOTAL 146
Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent
Cape Chiniak to Shaw _I
Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent
Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent
TOTAL 71
In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring
A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak
With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time
As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population
Shumagin Is 1 ands
This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population
There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion
In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good
There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied
There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas
Sutwi ck Area
Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather
In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved
~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather
The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys
There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years
to come
Augustine Island-Cape Douqlas
For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro
On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group
It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there
The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table
SUM
MAR
Y OF
SE
A OT
TER
SURV
EYS
(Com
pile
d by
p
op
ula
tio
n f
rom
L
ensi
nk
(196
2 T
hes
is
K
enyo
n 0
96
2 amp
196
5 R
epor
ts
amp M
anu
scri
pt)
an
d AD
FampG
Dat
a)
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
AU
GU
STIN
E -
C
DOUG
LAS
Aug
usti
ne
Isla
nd
A
ppro
xim
atel
y 50
(1
948)
Shaw
1
amp
Dou
glas
Are
a S
igh
tin
gs
from
Sh
aw
Is
to
Tux
edn
i B
ay
TOTA
L
40
___
_
41
52
-shy 52
18
_lQ
J
119
130+
-shy
130+
J
71
ALAS
KA
PEN
INSU
LA
c
Kul
iak
-
c
lgva
k
c
lgva
k -
c
Kun
mik
Ani
akch
ak
Bay
amp
Am
ber
Bay
Sutw
i ck
Are
a M
isce
llan
eous
R
epor
ts
Kuj
ul i
k B
ay
amp c
K
uml
ik
Sin
ce
1936
C
Kum
1 i u
n
Uni
viks
hak
Isla
nd
N
akch
amik
Is
lan
d
8
355 12
13
0 19 6
581
103
180
1
22
47
109
684 86
Not
S
urve
yed
7(+
)
139
197 14
1 2
53
101 62
5
I
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
ALAS
KA
PEN
INSU
LA
(Co
nt
)
Chi
gnik
B
ay
Cas
tle
Bay
amp
Cap
e
TOTA
L
SHUM
AGIN
Mai
nlan
d S
ho
re
Kup
rean
of
Pen
to
P
avlo
f B
ay
Ung
a Is
lan
d
Popo
f Is
lan
d
Kor
ovin
Is
lan
d
Kar
pa
Isla
nd
And
roni
ca
amp t
he
Hay
stac
ks
Nag
ai
Sp
ecta
cle
Ben
del
Tu
rner
Tw
ins
Few
R
epor
ts
Bef
ore
19
40
0
-shy
-shy
388
889 1 2 2 1
149
26
8
~-
7
- 949 4
338 I105
-~-middot
118
_lsect
1 9
12
23
184 52
46 4
75
232 8 27 6
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
SHUM
AGIN
(C
on
t)
Nea
r
Pen
insu
la
Big
Kon
i u
j i
Lit
tle K
oniu
j i
amp A
tkin
s
Sim
eono
f
Bir
d
Che
rnab
ura
TOTA
L
500
Est
imat
e (1
947)
l63
3 (
195 3
)
3 0
220
430
455
160
___L
ll
183
0
14 3
222
255
294 38
_J
l
1 35
2
150 15
296
290
329 76
_6
15
1 0
-shy 309
SANA
K -
SAND
MAN
Mai
nlan
d S
ho
re
Pav
lof
Bay
to
Un
imak
B
ay
Pav
lof
Isla
nd
s
Wos
nese
nski
Is
lan
d
Uko
l noi
Is
lan
d
Pop
erec
hnoi
Is
lan
d
Dol
goi
Isla
nd
2
141 4 2 29
67
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
SAN
AK
-SA
NDM
AN
Pav
lof
Isla
nd
s
Gol
oi
Isla
nd
Out
er
llia
sik
Inne
r ll
iasik
Sand
man
R
eefs
Che
rni
r-s I
and
(Co
nt
) (C
on
t)
Isla
nd
Isla
nd
C1u
bb i
ng
Roc
ks
Goo
se
Isla
nd
Dee
r Is
lan
d
amp O
ther
R
eefs
San
ak
Isla
nd
TOTA
L
2 16 2
27
(194
8)
271
259
495+
No
sig
hti
ng
s b
efo
re
1942
~
2 12
76
3397
82
7
123
5429
5 -
(I n
com
p Je
te)
Sig
hti
ng
s si
nce
19
22
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1199 88 62
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MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970
OBSERVER J Vania J Faro
PILOT George Tibbets Jr
AIRCRAFr Piper Azetex
middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good
Departed King Salmon 1115 AM Returned to King Salmon 745 PM
FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out
We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9
The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down
SEA OTTER SIGHTINGS
Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed
The number counted in each pod is as follows
1071 520 357 68 36 75 30
TOTAL 2157
Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island
171
-amp
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SEA OTTER COUNT BARREN ISLANDS and SHUYAK - AFOGNAK ISLANDS
June 9 1970
On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows
Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete
Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good
Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial
Area
BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island
AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island
TOTAL
TOTAL
Number of Sea Otters
0 2 0
76 200
29 0
307
18 6 3
121 0
33 26
207
Complete or Visibility Partial Count
Very good Complete ll If II
It II
II
II IJ
Good Complete II
Fair Partial If Complete II Partial II II
II
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51
SEA OTTER COUNTS - KENAI June J970 to January 1971
Carl Divinyi flew aerial surveys of seals along the outside of the
Kenai Peninsula on a monthly basis He recorded all sea otters sighted
on these surveys The surveys did not cover the entire coast line and
the type of aircraft weather conditions and observers varied from one
survey to another Therefore the seperate counts cannot be compared
However when viewed in total they indicate the areas used by sea otters
and the relative abundance of otters in each area
The attached table presents the counts by broad areas
--
SEA
OTT
ERS
CO
UN
TED
ON
A
ER
IAL
SUR
VEYS
O
F K
EN
AI
PEN
I NS
ULP
Ju
ne
1
97
0-
Jan
uar
y
1971
8i5
8
JUN
E 5
amp 9
JU
LY
15-2
0 A
UG
1l
t oc
r 12
NO
V
12
JAN
12
C
Junk
en
-C
R
esu
rrec
tio
n
5 30
42
27
10
30
Res
urr
ecti
on
Bay
2
2 0
4 2
NS
Ai a
1 i k
Bay
1
20
5 8
0 21
Har
ris
Bay
8
18
7 5
3 25
N
uka
Bay
10
6 56
NS
31
28
27
Po
rt
Dic
k 0
l1
NS
NS
3 23
Roc
ky
Bay
-
Po
rt C
hath
am
121
125
NS
NS
9 26
Koy
ukto
lik
Bay
-P
ort
Gra
ham
0
0 NS
NS
0
NS-
To
tal
243
262
54
75
55
152
middot3
8 se
a o
tters
cou
nted
fro
m
shor
e an
d sk
iff
11
20
70
AERIAL SURVEY Prince William Sound
April 1970
Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey
I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife
April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay
Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals
Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina
Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)
April 16 Weather inclement - no flying
April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island
April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)
Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands
The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed
SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date
PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418
CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52
PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS
ESTHER PASSAGE - VALDEZ ARM NS
GALENA BAY FISH BAY 103 415
FISH BAY - GRAVINA POINT 1 415
GRAVINA POINT - CORDOVA 1 415
HAWKINS ISLfu~D 1 4l7
HINCHINBROOK ISLfu~ 99 417
EGG ISLAND 2 417
MONTAGUE ISLAND 259 417
GREEN AND LITTLE GREEN ISLANDS 102 4J7
KNIGHT ISLAND 136 52
INGOT ISLAND 6 52
ELEANOR ISLAND 3 52
SMITH ISLAND 4 52
SEAL ISLAND 0 52
APPLEGATE ROCK 1 52
PERRY ISLAND NS
(continued) SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date-
LONE ISLAND NS
NAKED ISLAND NS
PEAK ISLAND NS
STOREY ISLAND NS
KAYAK - WINGHAM ISLAND 5 417
TOTAL 892
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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970
On July 17 1970 an aerial survey of sea otters was made in the area
around Green Island and the adjacent coast of Montague Island The purpose
of the survey was to locate concentrations of animals prior to a capturing
operation which began the next day A Dornier twin-engine arrcraft was used
with Schneider Reynolds and Somerville acting as observers The sky was
overcast almost no wind and the sea calm However heavy rain showers intershy
fered with forward visibility and the counting conditions were only fair on
the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay
The count began at 210pm and ended at 345 pm
The numbers and locations of otters are shown on the map In addition
16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy
in brook Island and a pod of 35 were seen three miles east of Johnstone Pt
Most of the area included in the survey was traveled repeatedly in a
skiff over the next four days Durlng this time the weather became calm and
clear for the first time in several days and the bulk of the sea otters shifted
from the coast of Montague out into the shallow areas between there and Green 1
and to Green and Channel Islands It was obvious that many otters had been missed
Fn the aerial sur~ey and that the population in the area is quite dense A
total of 48 sea otters were captured in three days of netting
j
SEA OTTER SURVEY Salisbury Sound to Cape Spencer
August 23-25~ 1970
On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and
middotWalt Cunningham serving as observers
The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good
On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere
On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May
Rough weather and outboard problems restricted tl the search area~
and the effectiveness of the search in that area The following sightings were made
82470 I Adult West of Granite Islands 1 Adult South of Granite Islands
11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands
These sightings represent from four to seven animals
Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull
The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal
The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait
While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island
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Harvest
In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island
and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year
The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial
survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs
Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers
Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same
Transplants
Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent
The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island
Table 1
April 20
middotMay 1
May 2
May 3
May 4
May 5
May 6
May 7
May 8
May 9
May 10
May 11
May 12
May 13
Schedule of the 1970 harvest
1030 pm depart Homer
Midnight arrive Adak MV Active
Refuel and prepare boat
Arrive off Tanaga at noon
Complete skinning pack hides etc in pm
Delarofs
Spent am getting water at Amchitka
Clean up boat take down shelter pack gear
900 am arrive Amchitka fly to Anchorage
Killed 53
138
88
131
143
53
144
43
79
83
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(am)
(pm)
(by mid afternoon)
Table 2 Projected and actual harvest numbers May 1970
Projected Actual Tana~a Island Harvest Harvest
Bumpy Point - Hot Springs Bay 200 191
Hot Springs Bay - Twin Bays 50 50
Twin Bays - Cape Sasmik 50 ) ) 199
Cape Sasmik - Tower 120 )
Tower Tanaga Bay 180 166
TOTAL 600 606
Delarof Islands
Gareloi 20-25 0
Kavalga Ogliuga and Skagul 75-100 125
Ulak and Amatignak 35-55 19
TOTAL 150 144
Amchitka Island saved for
USFWS Counting Units 1 amp 2 100 transplant
3 30 82
4 50 36
5 120 87
TOTAL 300 205
Table 3 Details of sea otters harvested in May 1970
Tanaga Island
Total kill 606
Number of pelts saved 598
Number of small pup pelts discarded 8
Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796
(Seal 3770 void - damaged)
Approximate sex ratio - 220 males 386 females or approximately 64 percent females
Delarof Islands
Total kill 144
Number of pelts saved 138
Number of pups discarded 6
Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934
Sex ratio- 44 males 100 females or approximately 69 percent females
Amchitka Island
Total kill 205
Number of pelts saved 194
Number of pups discarded 11
Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128
Sex ratio- 27 males 178 females or approximately 867 percent females
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A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality
Summary of Harvests and Transplants
Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area
The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5
----
----
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----
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----
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Tab
le 4
N
umbe
rs
of
sea o
tters
re
mov
ed
fro
m Ala
skan
pop
ulat
ions
~ 1
95
17
01
19
51
5
4
55
56
57
59
60
62
6
3
65
66
67
68
69
70
Am
chit
ka I
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ies
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22
37
2
6
21
39
2
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ran
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nt
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6
237
86
A
DFamp
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Har
ves
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3
11
2
05
2
3
205
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er
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1
14
-1
To
tal
35
22
37
26
21
39
1
80
31
1
210
50
0
25
1
292
Tan
aga
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ves
t 6
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k I
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ves
t 30
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94
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in Is
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ak amp
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mak
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ok
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nt
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nta
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e amp
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reen
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res
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agu
l 1
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lak
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Inclu
des
all
an
imal
s th
at
wer
e h
arv
est
ed
d
ied
d
uri
ng
st
ud
ies
and
tran
spla
nt
att
em
pts
o
r w
ere
tran
spla
nte
d
to
oth
er
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as
or
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s
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s n
ot
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de n
atu
ral
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rtali
ty o
r il
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al
kil
ls
2
Woo
dlan
d P
ark
Zoo
3
B
att
ell
e M
emo
rial
In
sti
tute
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Tab
le 5
N
umbe
rs
of
sea o
tters
tr
an
spla
nte
d 1
95
5-1
97
0
1955
19
56
1959
19
65
--
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96
6
1968
19
69
1970
Ale
uti
an
s A
ttu
I
5
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er
I
19
1
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bil
ofs
S
t
Pau
l I
7 S
t
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rge
I
57
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uta
t B
ay
10
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z B
ay
23
20
93
58
(Ch
ich
igo
f L
)
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uth
east
Y
akob
i I
30
A
lask
a B
iork
a I
48
Barr
ier
Is
55
Hec
eta
I
51
Cap
e S
pen
cer
25
Bri
tish
C
olum
bia
29
14
291
W
ash
ing
ton
Ore
gon
29
1
Non
e b
eli
ev
ed
to
hav
e su
rviv
ed
II
A
t le
ast
13
die
d s
ho
rtly
aft
er
rele
ase
NO
TE
1955
to
19
59 b
y U
SFW
S
1965
to
19
70 b
y
AD
FampG
In
som
e ca
ses
on
e o
r tw
o o
f th
e
abo
ve
anim
als
die
d n
ear
the
tim
e o
f re
lease
30
SEXUAL SEGREGATION IN SEA OTTER POPULATIONS
by Karl Schneider March 1971
Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males
Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands
Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals
Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas
A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population
In
the
By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled
Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet
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1970
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1970
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35
Sexual Segregation - 2 - March 7 1971
Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples
Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground
Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas
Identification of Hale Areas by Pup Counts
In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable
From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen
Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area
The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on
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Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side
The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed
Kanaga Island
Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$
Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point
Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak
Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass
The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring
Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area
Tab
le
5
Com
pari
son
of
Pup
s C
ou
nte
d w
ith
S
ex R
atio
o
f H
arv
est
Jun
e
1968
A
rea
Su
rvey
O
cto
ber
19
68 H
arv
est
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l KA
NAGA
IS
LAN
D
Ind
ep
Ott
er
Ind
ep
Ott
er
M
M
F
Rou
nd
Hea
d-S
ho
al P
oin
t 0
85
3
45
6
55
4
71
5
29
Nag
a P
oin
t-K
anag
a B
ay
66
1
64
middot
18
5
81
5
20
5
79
5
Cap
e C
hla
nak
-Cap
e T
usi
k
51
1
93
2
40
7
60
3
53
6
47
Edd
y R
ock
-Hiv
e R
ock
61
3
4
35
7
64
3
47
1
52
9
Jun
e
1968
A
rea
Su
rvey
H
ay
1970
Har
ves
t
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l TA
NA
GA
IS
LAN
D
Ind
ep
Ott
er ~ter
M
F
M
F
Cap
e S
ud
ak-P
end
ant
Po
int
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4
60
6
39
4
66
4
33
6
Bar
nes
P
oin
t-T
run
k P
oin
t 7
9
59
3
3
96
7
83
9
17
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nk
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int-
Tw
in B
ays
26
4
2
77
9
23
1
40
8
60
D
rin
Bay
s-S
ou
th B
ay
96
1
11
7
3
92
7
14
3
85
7
So
uth
Bay
-Har
em R
ock
14
8
31
6
7
middot93
3
91
9
09
Las
h
Bay
-In
fern
o
Ree
f 1
5
73
0
27
0
74
6
25
4
Har
em R
ock
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lak
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int
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7
64
4
35
6
68
4
31
6
Ku
lak
Po
int-
SE
B
igh
t 6
9
82
1
32
8
68
1
51
8
49
Sexual Segregation - 4 - March~ 1971
Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands
Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area
As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method
Identification of
Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area
The follmving is a description of the sex composition of areas for which no skiff count data was available
Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island
Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill
Sexual Segregation - 5 - March 1971
Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point
The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it
Composition of Female Areas
Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female
The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure
Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters
Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males
Sexual Segregation - 6 - March~ 1971
apparently defending a territory in California but from his observations t
it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change
Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year
A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest
The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas
The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males
There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas
Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak
within Female
In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of
Sexual Segregation - 7 - March 1971
these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June
The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups
Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population
ition of Male Areas
As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles
Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements
le 6
R
epro
du
ctiv
e S
tag
e o
f S
exu
ally
Mat
ure
S
ea O
tters
Har
ves
ted
on
Tan
aga
Isla
nd
-by
Are
a -
May
1
97
0
Un
imp
lan
ted
Im
pla
nte
d
Pre
gn
ant
Pre
gp
ant
Fet
us
Wei
gh
t C
lass
A
rea
No
np
reg
nan
t N
o
No
1
2 3
4 5
Res
orp
tio
n
To
tal
A
Su
dak
-Pen
dan
t P
oin
t 12
7
26
8 30
0
2 0
3 2
2 27
( B
arn
es
Po
int-
Ho
t S
pri
ng
s B
ay
12
7 24
10
36
3
0 1
4 2
2 29
B
( (
Bar
nes
P
oin
t-T
run
k P
oin
t 1
1
6 25
7
29
0 3
1 3
0 2
24
( T
run
k P
oin
t-T
win
Bay
s 7
9 39
7
30
2 1
1 1
2 2
23
c ( (
Haz
ard
P
oin
t-T
win
Bay
s 4
3 27
4
36
0 0
1 1
2 11
( T
win
B
ays-
Her
d
Roc
k 1
2
8 28
9
31
1 1
1 Lf
2
29
D
( ( T
win
B
ays-
So
uth
Bay
2
9 45
9
45
3 0
1 1
Lf
1 20
( S
ou
th B
ay-L
ash
B
ay
6 9
29
16
51
1 0
0 9
6 1
31
E
( ((
So
uth
Bay
-Har
em R
ock
17
14
31
14
31
2 0
1 5
6 45
(
F ( (
L
ash
B
ay-I
nfe
rno
Ree
f 4
1 9
6 55
2
1 1
0 2
11
( G
(
Har
em R
ock
-Ku
lak
Po
int
12
6 29
3
14
1 0
1 1
0 21
H
Ku
lak
Po
int-
SE
B
igh
t 13
9
28
10
31
2 0
1 2
5 32
Bra
cket
s in
dic
ate
o
ver
lap
pin
g are
as
Sexual Segregation - 8 - March 1971
Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area
While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring
The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area
Sex Ratio
With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants
On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL
All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females
There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among
Sexual Segregation - 9 - March 1971
newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population
It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality
The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor
If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve
The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes
REPRODUCTION IN THE FElIALE SEA OTTER
by Karl Schneider Narch 1971
A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle
The follovJing criteria were used in classification
Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation
Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)
Anestrus - Largest follicle under 3 5 rnm no corpus luteum
Proestrus - Follicles over 35 mm
Es - Follicles approximately 10 rrm
Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm
- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~
al scar newly formed corpus albicans with some luteal tissue remaining
The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever
Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies
Tab
le
l Re
p iv
e co
nd
itio
n o
f se
xual
ly m
atur
e se
a o
tters
_
IF
etus
Wei
--------------+
__
_
ln~a
ctive
ov
arie
s 1
Act
ive ovari~
ht C
Jas
s
jmiddot
I P
ost
-I P
roes
tru
s U
nim
pl
lmpl
D
ates
L
ocat
ioQ
1A
nest
rus
Part
um
Res
orpt
ion
l +
E
stru
s P
reo
Pr
eQ
2 3
lJ
S
~
4-8
1970
T
anag
a 1
51
41
0
10
104
17
8 10
33
30
4 (1
68)
(1
35)
(3
3)
(3 3
) (2
89)
(34~
(5
6)
(27
) (3
3)
(11
4)(
10
9)
May
9
19
70
Del
arof
Is
18
14
1
4 13
27
10
0
3 6
8 77
4
) (1
82)
( l
3)
(5
(16
9)
( o
) (1
30
) (3
9)
(78
)(1
04
)
10-1
219
70 A
mch
ltka
I
34
18
3 8
27
48
5 6
2 23
12
13
8
( 0
(24
6)
(13
0)
(22
) (5
8)
(19
6)
(34
8)
(36
) (4
3)
(14
) (1
67)
(8
7)
~lune
23
middotmiddot ~middot~~middot~Amchitka
1
17
2 1
2 4
18
1 2
3 5
7 44
A
ugus
t 13
196
8 (3
86)
(4
5)
(23
) (4
5)
(90
) (4
09)
(2
3)
(45
) (6
8)
(11
3)(
15
8)
July
6-
~dgtAmchitka
1
17
1 0
4 14
10
0
1 1
2 6
46
Aug
ust
819
69
(36
9)
( 2
0 2
) (8
6)
(30
4)
(21
9)
(22
) (2
2)
(43
)(1
32
)
Sep
t 1
0-17
A
dak
72
5 0
16
31
40
6 6
8 9
11
164
1967
(4
39
) ( 3
1)
(98
) ( 1
89)
(
4)
(3 7
) (3
7)
(49
) (5
5)
(6
7)
Sep
t
25
-A
mch
itk
a l
55
6
0 18
19
17
4
0 0
0 3
115
OcL
6
19
67
(4
7~8)
(5
2)
(15
7)
(16
5)
(14
8)
(35
) (8
7)
(26
)
Oct
o 1
2-1
5
Kan
aga
I
58
6 1
13
31
31
4 4
7 11
5
140
1968
(4
13)
(4
3)
(o 7
) (9
3)
(22
2)
(22
2)
(29
) (2
9)
(50
) (7
8)
(36
)
Oct
18
-20
A
dak
l
46
6 0
1 24
13
2
3 0
4 4
100
1968
(L
16 0
) ( 6
0)
(11
0)
(24
0)
(13
0)
(20
) (3
0)
0)
(40
)
Num
bers
in
re
nth
esis
are
pe
rcen
t se
xu
ally
mat
ure
fem
ales
F
etus
wei
ght
clas
s 1
=
0-l
g
2 =
1-l
Og
3
= 1
0-lO
Og
4
= 1
00-lO
OO
g
5 =
Tra
nsp
lan
t m
ort
alit
ies
(all
o
ther
sam
ples
fr
om
har
ves
ts)
Reproduction - 2 - March 1971
There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time
The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable
Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries
Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d
2
~ _s ~
~
c U)
~-1
r
~J
lt
c])
Reproduction - 3 - Narch 1971
to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors
cussion of the Annual
The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases
The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season
The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses
By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7
blastocysts implanting as is shown the increase in Class 1 fetuses
By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses
This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay
At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling
Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses
a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die
Reproduction - 4 - March 1971
There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August
Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)
If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another
The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b
The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short
There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points
At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping
Reproduction - 5 - March 1971
Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever
At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month
These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup
Gestation Period
Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature
A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth
The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)
Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April
It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t
Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo
Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted
Reproduction - 6 - March 1971
to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year
From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months
The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest
Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period
The following is a comparison of the two estimates
Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s
Total Gestation Period 399 d2ys 154 days or about 13 months 5 months
Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period
This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad
Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)
Table 2 Estimated Length of Period
Percent of Fetuses in each Height Class
Time of Collection After January
1 month 31 15 15 38 0
3 months 18 12 22 35 12
5 months 18 8 8 36 30
7 months 3 8 16 29 45
9 months 20 7 10 40 23
10 months
14 18 12 33 23
---~--
r1ean Time in Nonths After January 50 57 53 58 70
Cube Root of Umveighted Mean 063 17 36 7lf lllf
Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days
Table 3 Length of Unimplanted Period
Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted
1 month 35 26 58
3 months 49 49 so
5 months 128 179 42
7 months 22 37 37
9 months 50 57 47
10 months 55 44 56
Unweighted
48
Reproduction - 7 - March 1971
All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples
The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available
This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless
Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent
There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months
Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts
Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of
11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data
The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a
Reproduction - 8 - March 1971
Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters
If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months
The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping
The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve
Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination
There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months
Fetal Rate
the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e
--
3 0
~
shy
----- -shy --middotmiddot~middotmiddot
___ ___1__
_
(middot-)
_ ft -) J ~)___
~-
3
Reproduction - 9 - IYarch 1971
The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)
Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year
Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)
The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based
For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)
A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit
The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined
2
G
X
middot ~ -- (-- ) r~ ~ - - -- ~
yen~ r- ~--~
3
i__middot -~---
~
gt
- cshy
Reproduction - 10 - March 1971
~e of _Sexual Maturity
Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition
Ovulation
From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)
faximum
Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point
Ovulation
Many mustelids are induced ovulatoTs
On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous
ficance of
Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the
season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary
----
a - o lt bull y bull bull l w laquo bull _ ~ ~
Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968
N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A
I
~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy
- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot
1-2
2
3
4 1 I
2 (1)
6
middot5
1 (1)1
2 (l) 17
1 ( 1 )
2 ( 1)
18
l (1) I ( 1 ) 9
10 1 ( 1) L ~j( 2 1 ( 1 )
2 ( l) 1
12
11
1 (1)
I I (I)
1
2 ( 1)
13
1 ( 1 )
2
114
115 I116
17
I 18
I I
I
119 I
Numbers inside parentheses =Number of individuals with corpus luteum
11
Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968
N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I
AGE 84 I I
5 I l 6 7z 3(Years) 1
l rbull ~middot--
--- shy
0-l
J l-2
~ 2
3
4 2 ( 1)
5 3
Ibullbull 6 6 (3)
4 (4)7
8 4
3 9 2 ( 1 )
j iI 10
I 1 (1)11
12
13
14
1 ( 1)
16
15
17
18
Numbers irisi
-
J
(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)
1
1 (1)
5 (1)
10(5)
3
3 1
3 (2)
2 ( 1) 3 (3)
1 ( 1)
2
1 ( 1 ) 5 ( 1)
1 ( 1 ) 3 ( 1)
1
3
2 ( 1 )
1 (l)
1 (1)4 ( 1)
2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )
1 ( 1)
1
2
1 ( 1)
1 ( 1)
1
I I parentheses = Number of individuals with corpus luteum
11
Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968
N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A
-1AGE
~
9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--
- _ Q~L
1-2
(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3
4 2 (1)
l5
16 1 (1)
1 1)7
8
2 (1)9
10
111
12
I13
14
15 1 (1) 16
17
18
1 ( 1)
1 (l)
1
1
1
2
1
1
1
1(1 )
1
1 (1)
1 (1)
1
I
1
1
1
1
-I
Numbers Inside rentheses = Number of individuals with corpus luteum
1
Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968
AGE (Years)
Q-1
t-2
2
3
4
s 6
7
8
9
10
11
12
13
14
15
16
17
18
Numbers
N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A
~L 3 4 slE - 1middotshy
7 1~8 __J~l_11 shy
1-
One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I
One vJi th corpus luteum but no corpora al bicantia
12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3
1 (1) 2 (1)
1 2 (2) l 1
1 (1) 4 (1)
2 (1)
2 1 )
1 ( 1 ) 1
2 (1)
2 (2)
l22 3
3 ( 1)2 21 ( 1)
2 ( 1)
2 ( 1)
l ( 1)1
1 1
1 I ll
I
I I
inside parentheses Nurnber of individuals with corpus luteurn
Reproduction - 11 - March 1971
The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year
Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out
The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve
Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es
It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the
ion and ended around parturi tioD
Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists
Fetal
Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land
Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation
The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first
If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water
Reproduction - 12 - tltIarch 1971
At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water
Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample
Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn
It appears that blastocysts rarely cross from one horn to the other
Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other
When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time
A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active
Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus
Reproductive - 13 - March 1971
Birth iltleigh t
Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g
The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g
One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected
From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high
This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation
This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier
This may have sone effect on information on this
Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females
~
(
~
-s 2 0 C
U)
0
--
ez ()
~) I) middot _t t- (
j -~middot -~
Reproduction - 14 - March 1971
This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition
Sex Ratio at Birth
Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained
In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female
Sex Ratio of the
In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales
The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population
The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be
Lit
The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported
In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8
Table 8 Multiple Corpora Lutea Fetuses
Total Pregnant Females 565
Implanted Pregnancies 316
Unimplanted with 2 CL (corpora lutea) 9
Implanted Pregnancies with ) L CL and 1 Fetus 8
Implanted Pregnancies Vlith 3 CL and 1 Fetus 1
Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5
Implanted Pregnancies middotwith 3 CL and gt Fetuses 1
Total Nultiple Ovulations
I
Percent Nultiple Ovulations 42
Percent of Pregnancies tvith Nultiple Fetuses 19
Corpora Lutea per Pregnancy 105
Fetuses per 102
Reproduction - 15 - lvlarch 1971
From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally
All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo
in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time
With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations
The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus
A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some
Des
Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)
Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)
Resorbed or aborted fetus (implantation at end of horn) 3
Resorbed fetus (umbilicus tvTisted tightly) 1
Resorbed fetuses (three or more fetuses in one horn) 2
Resorbed fetus (cause unknmm) 5
Reproduction - 16 - March 1971
Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents
The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area
While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail
near Parturition
Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition
Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality
after
Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9
It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long
We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life
Table 9 Pups Accompanying Pos artum Females
Sex llleight
M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11
Total Length
47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)
Pups Accompanying Anestrous Females
F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103
Reproduction - 17 - Narch 1971
Frequency of Breeding
Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups
I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases
Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample
Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large
vhich had luteinized but tvere not actually pregnant The remainder had follicles
This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt
Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning
Table 10 Reproductive Condition of Lactating Females
Location
Bay of Is Adak I Sept 1967 9 1 100
Allchitka I Sept-Oct 1967 17 2 105
Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I
Bay of Is Adak I Oct 1968 22 4 154 3
Kanaga I Oct 1968 32 s- 135
Al1chitka I July-Aug 1969 4 2 333
iTanaga I May 1970 56 10 152
Delarof Is May 1970 18 3 143
Amchitka I May 1970 36 3 77
---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy
TOTAL 213 33 134
Anestrous or pregnant with term fetus
]_ Large follicles or corpus luteum present
3 Includes one implanted pregnancy with small fetus
Reproduction - 18 - March 1971
The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy
The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year
Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period
This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup
We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year
Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~
Reproduction - 19 - March 1971
observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less
Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years
This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias
Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years
Reproduction - 20 - March 7 1971
Conclusions
1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population
2 Breeding activity is highest in September October and November
3 More implantations occur during January February and March than at other times of year
4 The birth rate is highest during April Y~y and June
5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity
6 Similarly two and a half to three times as many females pup during peak pupping months
7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed
8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup
9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period
10 Female sea otters become sexually mature ihen 3 to 4 years old
11 Females may ovulate on an average of once a year after sexual maturity is reached
12 Females continue to breed at Cn old age
13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution
14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented
15 Sea otters probably birth both on land and in the water
16 Blastocysts rarely cross from one uterine horn to the other
17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random
Reproduction - 21 - March 1971
18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn
19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth
20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal
21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth
22 Body condition of the female has little effect on the growth rate of the fetus
23 There is a rapid ~Jeight gain shortly after parturition
24 The sex ratio at birth is 44 percent male and 56 percent female
25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term
26 Multiple ovulations occur in 4 percent of the estrus cycles
27 es do not appeErc- to be able to more than two fetuses in a single horn
28 Up to 5 percent of the p may fail dne to in after implantation
29 An unknown but probably significant number of first pregnancies fail before implantation
30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life
31 Sexually matu1middote females normally have one pup every two years
32 Females vrith a pup may ovulate but rarely mate
33 Slightly less than half of the sexually mature females are in any given year
34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived
35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus
Reproduction - 22 - March 1971
CITED
Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp
Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317
Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68
Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657
Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130
Snow H J 1910 In forbidden seas Edward Arnold London 303 pp
Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp
Description of Conditions During March 1970 Aerial Count of Sea Otters
March 20 Aniakchak Bay to Chignik Lagoon (1022 AM- 1240 PM) Conditions were generally fair with a heavy chop The south side of Sutwick Island and the area between the island and mainland were poor with heavy surf Visibility was very good inside Kujulik Bay however many animals were scattered throughout the area On two passes ~cross the bay 12 and 27 were seen therefore many were probably missed Similar scattering occured in Chignik Bay The overall count for the area is probably low
Pavlof Bay to Cold Bay (130PM-200PM) Conditions poor with heavy chop and squalls The count was superficial however a second superfi cia 1 count of this a rea was made on March 21 and only one otter was seen in Cold Bay
Cold Bay to Caee Lazaref (310PM -420PM) Conditions very poor fair in a few spots Heavy chop dense kelp
March 21 Northern Shumagin Islands (943AM- 1137 AM) Poo~ with chop on windward sides Fai_r on leeward sides (westerly wind) however air turbulence forced us away from high cliffs Heavy surf created very poor conditions on the south side of Unga
Pavlof Islands (1145 AM- 125ff PM) Conditions similar to those in the Shumagins
Chignik Lagoon to Pavlof Bay (305 PM- 535 PM) Conditions varying from fair to poor with chop and surface glare Count concentrated on points and better looking areas Most deep bays not completely surveyed
March 22 Sanak Islands (849AM- 1020 AM) Conditions fair to poor with chop on south side and surface glare on north side Animals scattered and very difficult to see Many otters on rocks Count is probably very low
Sandman Reefs ( 1025 AM~ 1115 AM) Conditions similar to Sanak With the exception of one pod of 450+ the animals were widely scattered and very difficult to pick out Only those very close to the aircraft could be seen Bad squalls moved in and the wind increased preventing a complete count of Deer Island and the eastern half of the reefs
March 23 Wide Bay to Aniakchak Bay (1123 AM- 140PM) Conditions generally good at first becoming fair after Cape Providence with a light chop and some surface glare Otters in Amber and Aniakchak Bays were widely scattered and many may have been missed
Port Heiden to Naknek_ Superficial counts were made from Ugashik Bay to Naknek on March 19 and from Port Heiden to Egegik on March 23 These were made under poor conditions by flying outside the surf line The water was choppy and full of silt No sea otters were seen
11arch 27 Shaw Island to Puale Bay (1030 AM - 1205 PM) Conditions very good to excellent High overcast and very little wind down to Katmai Bay After this the conditions deteriorated rapidly to fair and the count had to be stopped because of a combination of bad weather and lack of fuel As shown on the charts the count was concentrated on the points and islands however the good visibility allowed us to see well into the bays We feel certain no concentrations were missed
The numbers of sea otters counted are presented in the Table Specific locations and numbers are shown on the charts Where coverage of an area was not complete the approximate path of the aircraft is shown
Discussion of Results
Surveys along the Alaska Peninsula have been fragmentary and most of those made were done under less than ideal conditionsmiddot There are a number of reasons for this Weather tends to be poor along the south shore where squall5 form along the mountains Areas where an aircraft can refuel are north of the mountains and there are relatively few passes therefore range of the aircraft is a problem Also there is much shallow water off shore in the area from the Shumagin Islands to Sanak Island Coverage of this area is difficult and many animals are undoubtably missed
The present survey covered most of the area however conditions were such that most of the counts are probably quite low Despite these problems and a lack of continuity we have a fairly good picture of the recovery of sea otter populations in this area
Reports from various individuals are available from the 1930s and 1940s 11ajor surveys by the Fish and Wildlife Service in 1951 (Jones) 1957 (Lensink) 1962 and 1965 (Kenyon and Spencer) covered at least portions of the area In 1969 the southern Shumagin Islands were counted by the Alaska Department of Fish and Game and the present survey covered most of the rest of the area
Basically there are four distinct population nuclei in the area These centered around the Sanak Island-Sandman Reef area the Shumagin Island area the Sutwick Island area and the Augustine Island-Cape Douglas area The following discussion is based on information from reports by Lensink and Kenyon and from Alaska Department of Fish and Game surveys middot
~anak Island-Sandman Reefs
Small numbers of sea otters have been reported at Sanak Island since 1922 No reports came from the Sandman Reefs until 1942 In 1948 27 were sighted at
----Cherni Island The 1951 aerial survey showed 65 around Sanak and 97 in the Sandman Reefs In 1957 251 were seen around Sanak and 508 around the Sandman Reefs In addition two were seen in the Pavlof Islands however few were on the mainland In 1962 548 were counted in the Sanak area and 638 in the Sandman Reefs None were reported from the mainland or Pavlof lsland~however
much of the increase was in the northern area around Deer Island The 1962 survey was probably the best being made under excel lent conditions
AERIAL COUNT OF SEA OTTERS MARCH 1970
Partial or Location Count Date Camp 1ete Count Visibility
Cape Lazaref - Cold lsecty_
Cape Lazaref 3 320 Complete Poor
lkatan Peninsula 71 320 Complete Poor
False Pass - Amagat I 66 320 Camp 1ete Poor
Cold Bay 321 Partial Poor
TOTAL 141
Sanak Islands 239 322 Camp 1ete Fair to Poor
Sandman Reefs
Clubbing Rocks 12 322 Complete Fair to Poor
Cherni I amp Rocks 495+ 322 Complete fair to Poor
Goose I 7 322 Complete Fair to Poor
Hay I 18 322 Camp lete Fair to Poor
Hunt I 2 322 Complete Fair to Poor
Deer I 322 Partial Fair to Poor~
TOTAL 568+
Pavlof Islands
Inner 11iasik 2 321 Camp 1ete Fair to Poor
Outer lliasik 16 321 Camp 1ete Fair to Poor
Goloi 2 321 Complete Fair to Poor
Dolgoi 67 321 Complete Fair to Poor
Poperechnoi 29 321 Complete Fair to Poor
Ukolnoi 2 321 Complete Fair to Poor
yenWosnesenski 4 321 Complete Fair to Poor
TOTAL 122
Partial or Location Count Date Comp 1ete Count vis ib j 1i ty
Northern Shumagin Islands
Unga 184 321 Complete Fair to Poor
Popof 52 321 Complete Fair to Poor
Korovin 46 321 Complete Fair to Poor
Karpa __2t 321 Complete Fair to Poor
TOTAL 286
Cold ~ to Beaver ~ 0 320-21 Partial Poor
Beaver Bay to Kupreanof Pen
Beaver Bay 2 321 Partial Fair to Poor
Cape A 1 iaks in 4 321 Partial Fair to Poor
Guillemot I 16 321 Partial Fair to Poor
__1Kupreanof Peninsula 321 Partial Fair to Poor
TOTAL 23
Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor
AntJrrCastle Cape to-1-Ji e ~
Castle Cape-Castle Bay 16 321 Partial Fair to Poor
Chignik Bay 118 320 Complete Fair
Nakchamik I 5 320 Complete Fair
Hook Bay 13 320 Complete Fair
Cape Kum 1i un 88 320 Complete Fair
Kujul ik Bay 1199 320 Complete Very Good
Univikshak I 62 320 Complete Fair
Cape Kuml ik 54 320 Complete Fair to Poor
Sutwick I 14 320 Complete Fair to Poor
Cape Agutka 323 Complete Fair~
TOTAL 1766
Partial or Location Count Date Comp 1ete Count vis i b i 1i ty
Cape Kunmik 13 323 Complete Fair
Naka 1i kok Bay amp offshore islands 16 323 Complete Fair
Chiginagak Bay 5 323 Complete Fair
Agripina amp lmuya Bay 105 323 Complete Good
Wide Bay to Puale Bay N 0 T S U R V E Y E D
Puale Bay to c Kubugakl i __]_ 327 Partial Fair
TOTAL 146
Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent
Cape Chiniak to Shaw _I
Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent
Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent
TOTAL 71
In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring
A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak
With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time
As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population
Shumagin Is 1 ands
This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population
There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion
In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good
There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied
There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas
Sutwi ck Area
Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather
In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved
~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather
The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys
There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years
to come
Augustine Island-Cape Douqlas
For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro
On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group
It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there
The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table
SUM
MAR
Y OF
SE
A OT
TER
SURV
EYS
(Com
pile
d by
p
op
ula
tio
n f
rom
L
ensi
nk
(196
2 T
hes
is
K
enyo
n 0
96
2 amp
196
5 R
epor
ts
amp M
anu
scri
pt)
an
d AD
FampG
Dat
a)
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
AU
GU
STIN
E -
C
DOUG
LAS
Aug
usti
ne
Isla
nd
A
ppro
xim
atel
y 50
(1
948)
Shaw
1
amp
Dou
glas
Are
a S
igh
tin
gs
from
Sh
aw
Is
to
Tux
edn
i B
ay
TOTA
L
40
___
_
41
52
-shy 52
18
_lQ
J
119
130+
-shy
130+
J
71
ALAS
KA
PEN
INSU
LA
c
Kul
iak
-
c
lgva
k
c
lgva
k -
c
Kun
mik
Ani
akch
ak
Bay
amp
Am
ber
Bay
Sutw
i ck
Are
a M
isce
llan
eous
R
epor
ts
Kuj
ul i
k B
ay
amp c
K
uml
ik
Sin
ce
1936
C
Kum
1 i u
n
Uni
viks
hak
Isla
nd
N
akch
amik
Is
lan
d
8
355 12
13
0 19 6
581
103
180
1
22
47
109
684 86
Not
S
urve
yed
7(+
)
139
197 14
1 2
53
101 62
5
I
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
ALAS
KA
PEN
INSU
LA
(Co
nt
)
Chi
gnik
B
ay
Cas
tle
Bay
amp
Cap
e
TOTA
L
SHUM
AGIN
Mai
nlan
d S
ho
re
Kup
rean
of
Pen
to
P
avlo
f B
ay
Ung
a Is
lan
d
Popo
f Is
lan
d
Kor
ovin
Is
lan
d
Kar
pa
Isla
nd
And
roni
ca
amp t
he
Hay
stac
ks
Nag
ai
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t
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Hook Bay Chignik Bay
13 118
-1 shy
shy
shy
bull
I +
Castle Bay - Castle Cape 16 Nakcharriik I 5 -- Katmai Reef 0
Total 152
middot
~ middot-
bull -middot
bull
-middot
Cape Agutka Cape Kum 1 i k Sutwick I~ Kujul ik Bay Cape Kum 1 i un Unavikshak 1
Total
X
-------8
197 54 14
1199 88 62
1614
X
E c 1 ~
(l) a 10
u
0 ---_ +-
Q) u
gt-sect OJ al middotshy
gt 10 0 c 1
middot- a a middot- (l)1 a Olfil lt(U
bull c
)
r r -middot
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middot i 1~ bull middot- i ~ shy
i
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- _ ~~---Q middot- ---- ~ - ~ ~ --~ G lt-~ bulllt00 middot __ ~-middot middot-middot +~ yen_-_____ _ ---
_-y _ -
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bull ~
v
4gt
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rd ltgt
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9
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lt
bull
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I I
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smiddotmiddot I
l bullt
middot
99
10
i
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9
75
~-
MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970
OBSERVER J Vania J Faro
PILOT George Tibbets Jr
AIRCRAFr Piper Azetex
middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good
Departed King Salmon 1115 AM Returned to King Salmon 745 PM
FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out
We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9
The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down
SEA OTTER SIGHTINGS
Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed
The number counted in each pod is as follows
1071 520 357 68 36 75 30
TOTAL 2157
Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island
171
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54
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53
51
52
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48
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54
58
54
53
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s 43
36 ~
44
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39
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53
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SEA OTTER COUNT BARREN ISLANDS and SHUYAK - AFOGNAK ISLANDS
June 9 1970
On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows
Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete
Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good
Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial
Area
BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island
AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island
TOTAL
TOTAL
Number of Sea Otters
0 2 0
76 200
29 0
307
18 6 3
121 0
33 26
207
Complete or Visibility Partial Count
Very good Complete ll If II
It II
II
II IJ
Good Complete II
Fair Partial If Complete II Partial II II
II
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lt6gt iltj- cgtcgt Jgt
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9
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64
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675
6
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SEA OTTER COUNTS - KENAI June J970 to January 1971
Carl Divinyi flew aerial surveys of seals along the outside of the
Kenai Peninsula on a monthly basis He recorded all sea otters sighted
on these surveys The surveys did not cover the entire coast line and
the type of aircraft weather conditions and observers varied from one
survey to another Therefore the seperate counts cannot be compared
However when viewed in total they indicate the areas used by sea otters
and the relative abundance of otters in each area
The attached table presents the counts by broad areas
--
SEA
OTT
ERS
CO
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TED
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ER
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AERIAL SURVEY Prince William Sound
April 1970
Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey
I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife
April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay
Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals
Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina
Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)
April 16 Weather inclement - no flying
April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island
April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)
Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands
The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed
SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date
PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418
CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52
PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS
ESTHER PASSAGE - VALDEZ ARM NS
GALENA BAY FISH BAY 103 415
FISH BAY - GRAVINA POINT 1 415
GRAVINA POINT - CORDOVA 1 415
HAWKINS ISLfu~D 1 4l7
HINCHINBROOK ISLfu~ 99 417
EGG ISLAND 2 417
MONTAGUE ISLAND 259 417
GREEN AND LITTLE GREEN ISLANDS 102 4J7
KNIGHT ISLAND 136 52
INGOT ISLAND 6 52
ELEANOR ISLAND 3 52
SMITH ISLAND 4 52
SEAL ISLAND 0 52
APPLEGATE ROCK 1 52
PERRY ISLAND NS
(continued) SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date-
LONE ISLAND NS
NAKED ISLAND NS
PEAK ISLAND NS
STOREY ISLAND NS
KAYAK - WINGHAM ISLAND 5 417
TOTAL 892
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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970
On July 17 1970 an aerial survey of sea otters was made in the area
around Green Island and the adjacent coast of Montague Island The purpose
of the survey was to locate concentrations of animals prior to a capturing
operation which began the next day A Dornier twin-engine arrcraft was used
with Schneider Reynolds and Somerville acting as observers The sky was
overcast almost no wind and the sea calm However heavy rain showers intershy
fered with forward visibility and the counting conditions were only fair on
the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay
The count began at 210pm and ended at 345 pm
The numbers and locations of otters are shown on the map In addition
16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy
in brook Island and a pod of 35 were seen three miles east of Johnstone Pt
Most of the area included in the survey was traveled repeatedly in a
skiff over the next four days Durlng this time the weather became calm and
clear for the first time in several days and the bulk of the sea otters shifted
from the coast of Montague out into the shallow areas between there and Green 1
and to Green and Channel Islands It was obvious that many otters had been missed
Fn the aerial sur~ey and that the population in the area is quite dense A
total of 48 sea otters were captured in three days of netting
j
SEA OTTER SURVEY Salisbury Sound to Cape Spencer
August 23-25~ 1970
On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and
middotWalt Cunningham serving as observers
The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good
On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere
On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May
Rough weather and outboard problems restricted tl the search area~
and the effectiveness of the search in that area The following sightings were made
82470 I Adult West of Granite Islands 1 Adult South of Granite Islands
11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands
These sightings represent from four to seven animals
Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull
The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal
The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait
While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island
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HARVEST AND TRANSPLANTS - 1970
Harvest
In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island
and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year
The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial
survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs
Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers
Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same
Transplants
Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent
The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island
Table 1
April 20
middotMay 1
May 2
May 3
May 4
May 5
May 6
May 7
May 8
May 9
May 10
May 11
May 12
May 13
Schedule of the 1970 harvest
1030 pm depart Homer
Midnight arrive Adak MV Active
Refuel and prepare boat
Arrive off Tanaga at noon
Complete skinning pack hides etc in pm
Delarofs
Spent am getting water at Amchitka
Clean up boat take down shelter pack gear
900 am arrive Amchitka fly to Anchorage
Killed 53
138
88
131
143
53
144
43
79
83
otter
(am)
(pm)
(by mid afternoon)
Table 2 Projected and actual harvest numbers May 1970
Projected Actual Tana~a Island Harvest Harvest
Bumpy Point - Hot Springs Bay 200 191
Hot Springs Bay - Twin Bays 50 50
Twin Bays - Cape Sasmik 50 ) ) 199
Cape Sasmik - Tower 120 )
Tower Tanaga Bay 180 166
TOTAL 600 606
Delarof Islands
Gareloi 20-25 0
Kavalga Ogliuga and Skagul 75-100 125
Ulak and Amatignak 35-55 19
TOTAL 150 144
Amchitka Island saved for
USFWS Counting Units 1 amp 2 100 transplant
3 30 82
4 50 36
5 120 87
TOTAL 300 205
Table 3 Details of sea otters harvested in May 1970
Tanaga Island
Total kill 606
Number of pelts saved 598
Number of small pup pelts discarded 8
Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796
(Seal 3770 void - damaged)
Approximate sex ratio - 220 males 386 females or approximately 64 percent females
Delarof Islands
Total kill 144
Number of pelts saved 138
Number of pups discarded 6
Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934
Sex ratio- 44 males 100 females or approximately 69 percent females
Amchitka Island
Total kill 205
Number of pelts saved 194
Number of pups discarded 11
Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128
Sex ratio- 27 males 178 females or approximately 867 percent females
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A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality
Summary of Harvests and Transplants
Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area
The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5
----
----
----
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----
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Tab
le 4
N
umbe
rs
of
sea o
tters
re
mov
ed
fro
m Ala
skan
pop
ulat
ions
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95
17
01
19
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4
55
56
57
59
60
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6
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65
66
67
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69
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chit
ka I
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nt
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er
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tal
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1
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ves
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in Is
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ak amp
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mak
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nt
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nta
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reen
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res
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Inclu
des
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imal
s th
at
wer
e h
arv
est
ed
d
ied
d
uri
ng
st
ud
ies
and
tran
spla
nt
att
em
pts
o
r w
ere
tran
spla
nte
d
to
oth
er
are
as
or
zoo
s
Doe
s n
ot
inclu
de n
atu
ral
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rtali
ty o
r il
leg
al
kil
ls
2
Woo
dlan
d P
ark
Zoo
3
B
att
ell
e M
emo
rial
In
sti
tute
--
Tab
le 5
N
umbe
rs
of
sea o
tters
tr
an
spla
nte
d 1
95
5-1
97
0
1955
19
56
1959
19
65
--
~ 1
96
6
1968
19
69
1970
Ale
uti
an
s A
ttu
I
5
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er
I
19
1
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bil
ofs
S
t
Pau
l I
7 S
t
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rge
I
57
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uta
t B
ay
10
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z B
ay
23
20
93
58
(Ch
ich
igo
f L
)
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uth
east
Y
akob
i I
30
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lask
a B
iork
a I
48
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ier
Is
55
Hec
eta
I
51
Cap
e S
pen
cer
25
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tish
C
olum
bia
29
14
291
W
ash
ing
ton
Ore
gon
29
1
Non
e b
eli
ev
ed
to
hav
e su
rviv
ed
II
A
t le
ast
13
die
d s
ho
rtly
aft
er
rele
ase
NO
TE
1955
to
19
59 b
y U
SFW
S
1965
to
19
70 b
y
AD
FampG
In
som
e ca
ses
on
e o
r tw
o o
f th
e
abo
ve
anim
als
die
d n
ear
the
tim
e o
f re
lease
30
SEXUAL SEGREGATION IN SEA OTTER POPULATIONS
by Karl Schneider March 1971
Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males
Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands
Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals
Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas
A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population
In
the
By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled
Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet
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Sexual Segregation - 2 - March 7 1971
Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples
Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground
Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas
Identification of Hale Areas by Pup Counts
In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable
From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen
Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area
The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on
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Gl
~ middot
4
~
middot
(r
l
117
214
54
2
l
so
O
U--
~ 4
u~lt
27
~ ~5
9 -
139
r9
rk_
y_ 14
7
Tb
n9
middota l ~
---~poundiir
r) 3-~-
4~i
~ ~o-
~ 68
a
71Ja
cttt
~7~-
~15
9 1
159
122
l 1~J
J ~
1 4
0
rky
I )
ct
bull
t o
il
7
0 8
4
88
r 42
J
66
1
43
131
7
0 3
if
ll~
ts c~4
~
--middot~
0
5 2
65
Ill
12
7 2
33
3
78
4
42
f
ty
fj
t1
jbull
8
0
16
1
37
15
2 6
3
108
45
592
I
2 C
) P
s
2
I
2
02
25
2 I
I J
flI
s 41
52
39
3
57
5
58
5
28
16
1 3
30
3
62
1 35
7 2
58
2
47
2
52
29
5 3
00
424
44
56
(-
Y v
n 7
-
A
59
64
3
5
59
v
-s
4
52
I
n 7
-
1-
J _
fs3~
_
) 7
32
7
-7
7A
Sexual Segregation - 3 - March 1 1971
Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side
The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed
Kanaga Island
Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$
Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point
Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak
Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass
The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring
Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area
Tab
le
5
Com
pari
son
of
Pup
s C
ou
nte
d w
ith
S
ex R
atio
o
f H
arv
est
Jun
e
1968
A
rea
Su
rvey
O
cto
ber
19
68 H
arv
est
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l KA
NAGA
IS
LAN
D
Ind
ep
Ott
er
Ind
ep
Ott
er
M
M
F
Rou
nd
Hea
d-S
ho
al P
oin
t 0
85
3
45
6
55
4
71
5
29
Nag
a P
oin
t-K
anag
a B
ay
66
1
64
middot
18
5
81
5
20
5
79
5
Cap
e C
hla
nak
-Cap
e T
usi
k
51
1
93
2
40
7
60
3
53
6
47
Edd
y R
ock
-Hiv
e R
ock
61
3
4
35
7
64
3
47
1
52
9
Jun
e
1968
A
rea
Su
rvey
H
ay
1970
Har
ves
t
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l TA
NA
GA
IS
LAN
D
Ind
ep
Ott
er ~ter
M
F
M
F
Cap
e S
ud
ak-P
end
ant
Po
int
0 4
4
60
6
39
4
66
4
33
6
Bar
nes
P
oin
t-T
run
k P
oin
t 7
9
59
3
3
96
7
83
9
17
Tru
nk
Po
int-
Tw
in B
ays
26
4
2
77
9
23
1
40
8
60
D
rin
Bay
s-S
ou
th B
ay
96
1
11
7
3
92
7
14
3
85
7
So
uth
Bay
-Har
em R
ock
14
8
31
6
7
middot93
3
91
9
09
Las
h
Bay
-In
fern
o
Ree
f 1
5
73
0
27
0
74
6
25
4
Har
em R
ock
-Ku
lak
Po
int
0 2
7
64
4
35
6
68
4
31
6
Ku
lak
Po
int-
SE
B
igh
t 6
9
82
1
32
8
68
1
51
8
49
Sexual Segregation - 4 - March~ 1971
Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands
Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area
As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method
Identification of
Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area
The follmving is a description of the sex composition of areas for which no skiff count data was available
Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island
Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill
Sexual Segregation - 5 - March 1971
Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point
The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it
Composition of Female Areas
Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female
The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure
Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters
Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males
Sexual Segregation - 6 - March~ 1971
apparently defending a territory in California but from his observations t
it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change
Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year
A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest
The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas
The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males
There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas
Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak
within Female
In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of
Sexual Segregation - 7 - March 1971
these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June
The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups
Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population
ition of Male Areas
As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles
Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements
le 6
R
epro
du
ctiv
e S
tag
e o
f S
exu
ally
Mat
ure
S
ea O
tters
Har
ves
ted
on
Tan
aga
Isla
nd
-by
Are
a -
May
1
97
0
Un
imp
lan
ted
Im
pla
nte
d
Pre
gn
ant
Pre
gp
ant
Fet
us
Wei
gh
t C
lass
A
rea
No
np
reg
nan
t N
o
No
1
2 3
4 5
Res
orp
tio
n
To
tal
A
Su
dak
-Pen
dan
t P
oin
t 12
7
26
8 30
0
2 0
3 2
2 27
( B
arn
es
Po
int-
Ho
t S
pri
ng
s B
ay
12
7 24
10
36
3
0 1
4 2
2 29
B
( (
Bar
nes
P
oin
t-T
run
k P
oin
t 1
1
6 25
7
29
0 3
1 3
0 2
24
( T
run
k P
oin
t-T
win
Bay
s 7
9 39
7
30
2 1
1 1
2 2
23
c ( (
Haz
ard
P
oin
t-T
win
Bay
s 4
3 27
4
36
0 0
1 1
2 11
( T
win
B
ays-
Her
d
Roc
k 1
2
8 28
9
31
1 1
1 Lf
2
29
D
( ( T
win
B
ays-
So
uth
Bay
2
9 45
9
45
3 0
1 1
Lf
1 20
( S
ou
th B
ay-L
ash
B
ay
6 9
29
16
51
1 0
0 9
6 1
31
E
( ((
So
uth
Bay
-Har
em R
ock
17
14
31
14
31
2 0
1 5
6 45
(
F ( (
L
ash
B
ay-I
nfe
rno
Ree
f 4
1 9
6 55
2
1 1
0 2
11
( G
(
Har
em R
ock
-Ku
lak
Po
int
12
6 29
3
14
1 0
1 1
0 21
H
Ku
lak
Po
int-
SE
B
igh
t 13
9
28
10
31
2 0
1 2
5 32
Bra
cket
s in
dic
ate
o
ver
lap
pin
g are
as
Sexual Segregation - 8 - March 1971
Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area
While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring
The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area
Sex Ratio
With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants
On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL
All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females
There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among
Sexual Segregation - 9 - March 1971
newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population
It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality
The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor
If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve
The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes
REPRODUCTION IN THE FElIALE SEA OTTER
by Karl Schneider Narch 1971
A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle
The follovJing criteria were used in classification
Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation
Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)
Anestrus - Largest follicle under 3 5 rnm no corpus luteum
Proestrus - Follicles over 35 mm
Es - Follicles approximately 10 rrm
Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm
- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~
al scar newly formed corpus albicans with some luteal tissue remaining
The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever
Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies
Tab
le
l Re
p iv
e co
nd
itio
n o
f se
xual
ly m
atur
e se
a o
tters
_
IF
etus
Wei
--------------+
__
_
ln~a
ctive
ov
arie
s 1
Act
ive ovari~
ht C
Jas
s
jmiddot
I P
ost
-I P
roes
tru
s U
nim
pl
lmpl
D
ates
L
ocat
ioQ
1A
nest
rus
Part
um
Res
orpt
ion
l +
E
stru
s P
reo
Pr
eQ
2 3
lJ
S
~
4-8
1970
T
anag
a 1
51
41
0
10
104
17
8 10
33
30
4 (1
68)
(1
35)
(3
3)
(3 3
) (2
89)
(34~
(5
6)
(27
) (3
3)
(11
4)(
10
9)
May
9
19
70
Del
arof
Is
18
14
1
4 13
27
10
0
3 6
8 77
4
) (1
82)
( l
3)
(5
(16
9)
( o
) (1
30
) (3
9)
(78
)(1
04
)
10-1
219
70 A
mch
ltka
I
34
18
3 8
27
48
5 6
2 23
12
13
8
( 0
(24
6)
(13
0)
(22
) (5
8)
(19
6)
(34
8)
(36
) (4
3)
(14
) (1
67)
(8
7)
~lune
23
middotmiddot ~middot~~middot~Amchitka
1
17
2 1
2 4
18
1 2
3 5
7 44
A
ugus
t 13
196
8 (3
86)
(4
5)
(23
) (4
5)
(90
) (4
09)
(2
3)
(45
) (6
8)
(11
3)(
15
8)
July
6-
~dgtAmchitka
1
17
1 0
4 14
10
0
1 1
2 6
46
Aug
ust
819
69
(36
9)
( 2
0 2
) (8
6)
(30
4)
(21
9)
(22
) (2
2)
(43
)(1
32
)
Sep
t 1
0-17
A
dak
72
5 0
16
31
40
6 6
8 9
11
164
1967
(4
39
) ( 3
1)
(98
) ( 1
89)
(
4)
(3 7
) (3
7)
(49
) (5
5)
(6
7)
Sep
t
25
-A
mch
itk
a l
55
6
0 18
19
17
4
0 0
0 3
115
OcL
6
19
67
(4
7~8)
(5
2)
(15
7)
(16
5)
(14
8)
(35
) (8
7)
(26
)
Oct
o 1
2-1
5
Kan
aga
I
58
6 1
13
31
31
4 4
7 11
5
140
1968
(4
13)
(4
3)
(o 7
) (9
3)
(22
2)
(22
2)
(29
) (2
9)
(50
) (7
8)
(36
)
Oct
18
-20
A
dak
l
46
6 0
1 24
13
2
3 0
4 4
100
1968
(L
16 0
) ( 6
0)
(11
0)
(24
0)
(13
0)
(20
) (3
0)
0)
(40
)
Num
bers
in
re
nth
esis
are
pe
rcen
t se
xu
ally
mat
ure
fem
ales
F
etus
wei
ght
clas
s 1
=
0-l
g
2 =
1-l
Og
3
= 1
0-lO
Og
4
= 1
00-lO
OO
g
5 =
Tra
nsp
lan
t m
ort
alit
ies
(all
o
ther
sam
ples
fr
om
har
ves
ts)
Reproduction - 2 - March 1971
There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time
The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable
Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries
Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d
2
~ _s ~
~
c U)
~-1
r
~J
lt
c])
Reproduction - 3 - Narch 1971
to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors
cussion of the Annual
The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases
The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season
The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses
By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7
blastocysts implanting as is shown the increase in Class 1 fetuses
By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses
This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay
At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling
Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses
a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die
Reproduction - 4 - March 1971
There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August
Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)
If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another
The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b
The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short
There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points
At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping
Reproduction - 5 - March 1971
Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever
At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month
These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup
Gestation Period
Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature
A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth
The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)
Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April
It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t
Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo
Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted
Reproduction - 6 - March 1971
to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year
From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months
The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest
Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period
The following is a comparison of the two estimates
Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s
Total Gestation Period 399 d2ys 154 days or about 13 months 5 months
Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period
This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad
Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)
Table 2 Estimated Length of Period
Percent of Fetuses in each Height Class
Time of Collection After January
1 month 31 15 15 38 0
3 months 18 12 22 35 12
5 months 18 8 8 36 30
7 months 3 8 16 29 45
9 months 20 7 10 40 23
10 months
14 18 12 33 23
---~--
r1ean Time in Nonths After January 50 57 53 58 70
Cube Root of Umveighted Mean 063 17 36 7lf lllf
Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days
Table 3 Length of Unimplanted Period
Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted
1 month 35 26 58
3 months 49 49 so
5 months 128 179 42
7 months 22 37 37
9 months 50 57 47
10 months 55 44 56
Unweighted
48
Reproduction - 7 - March 1971
All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples
The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available
This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless
Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent
There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months
Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts
Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of
11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data
The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a
Reproduction - 8 - March 1971
Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters
If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months
The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping
The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve
Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination
There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months
Fetal Rate
the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e
--
3 0
~
shy
----- -shy --middotmiddot~middotmiddot
___ ___1__
_
(middot-)
_ ft -) J ~)___
~-
3
Reproduction - 9 - IYarch 1971
The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)
Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year
Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)
The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based
For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)
A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit
The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined
2
G
X
middot ~ -- (-- ) r~ ~ - - -- ~
yen~ r- ~--~
3
i__middot -~---
~
gt
- cshy
Reproduction - 10 - March 1971
~e of _Sexual Maturity
Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition
Ovulation
From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)
faximum
Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point
Ovulation
Many mustelids are induced ovulatoTs
On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous
ficance of
Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the
season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary
----
a - o lt bull y bull bull l w laquo bull _ ~ ~
Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968
N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A
I
~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy
- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot
1-2
2
3
4 1 I
2 (1)
6
middot5
1 (1)1
2 (l) 17
1 ( 1 )
2 ( 1)
18
l (1) I ( 1 ) 9
10 1 ( 1) L ~j( 2 1 ( 1 )
2 ( l) 1
12
11
1 (1)
I I (I)
1
2 ( 1)
13
1 ( 1 )
2
114
115 I116
17
I 18
I I
I
119 I
Numbers inside parentheses =Number of individuals with corpus luteum
11
Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968
N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I
AGE 84 I I
5 I l 6 7z 3(Years) 1
l rbull ~middot--
--- shy
0-l
J l-2
~ 2
3
4 2 ( 1)
5 3
Ibullbull 6 6 (3)
4 (4)7
8 4
3 9 2 ( 1 )
j iI 10
I 1 (1)11
12
13
14
1 ( 1)
16
15
17
18
Numbers irisi
-
J
(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)
1
1 (1)
5 (1)
10(5)
3
3 1
3 (2)
2 ( 1) 3 (3)
1 ( 1)
2
1 ( 1 ) 5 ( 1)
1 ( 1 ) 3 ( 1)
1
3
2 ( 1 )
1 (l)
1 (1)4 ( 1)
2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )
1 ( 1)
1
2
1 ( 1)
1 ( 1)
1
I I parentheses = Number of individuals with corpus luteum
11
Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968
N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A
-1AGE
~
9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--
- _ Q~L
1-2
(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3
4 2 (1)
l5
16 1 (1)
1 1)7
8
2 (1)9
10
111
12
I13
14
15 1 (1) 16
17
18
1 ( 1)
1 (l)
1
1
1
2
1
1
1
1(1 )
1
1 (1)
1 (1)
1
I
1
1
1
1
-I
Numbers Inside rentheses = Number of individuals with corpus luteum
1
Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968
AGE (Years)
Q-1
t-2
2
3
4
s 6
7
8
9
10
11
12
13
14
15
16
17
18
Numbers
N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A
~L 3 4 slE - 1middotshy
7 1~8 __J~l_11 shy
1-
One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I
One vJi th corpus luteum but no corpora al bicantia
12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3
1 (1) 2 (1)
1 2 (2) l 1
1 (1) 4 (1)
2 (1)
2 1 )
1 ( 1 ) 1
2 (1)
2 (2)
l22 3
3 ( 1)2 21 ( 1)
2 ( 1)
2 ( 1)
l ( 1)1
1 1
1 I ll
I
I I
inside parentheses Nurnber of individuals with corpus luteurn
Reproduction - 11 - March 1971
The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year
Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out
The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve
Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es
It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the
ion and ended around parturi tioD
Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists
Fetal
Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land
Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation
The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first
If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water
Reproduction - 12 - tltIarch 1971
At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water
Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample
Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn
It appears that blastocysts rarely cross from one horn to the other
Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other
When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time
A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active
Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus
Reproductive - 13 - March 1971
Birth iltleigh t
Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g
The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g
One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected
From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high
This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation
This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier
This may have sone effect on information on this
Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females
~
(
~
-s 2 0 C
U)
0
--
ez ()
~) I) middot _t t- (
j -~middot -~
Reproduction - 14 - March 1971
This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition
Sex Ratio at Birth
Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained
In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female
Sex Ratio of the
In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales
The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population
The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be
Lit
The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported
In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8
Table 8 Multiple Corpora Lutea Fetuses
Total Pregnant Females 565
Implanted Pregnancies 316
Unimplanted with 2 CL (corpora lutea) 9
Implanted Pregnancies with ) L CL and 1 Fetus 8
Implanted Pregnancies Vlith 3 CL and 1 Fetus 1
Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5
Implanted Pregnancies middotwith 3 CL and gt Fetuses 1
Total Nultiple Ovulations
I
Percent Nultiple Ovulations 42
Percent of Pregnancies tvith Nultiple Fetuses 19
Corpora Lutea per Pregnancy 105
Fetuses per 102
Reproduction - 15 - lvlarch 1971
From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally
All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo
in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time
With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations
The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus
A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some
Des
Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)
Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)
Resorbed or aborted fetus (implantation at end of horn) 3
Resorbed fetus (umbilicus tvTisted tightly) 1
Resorbed fetuses (three or more fetuses in one horn) 2
Resorbed fetus (cause unknmm) 5
Reproduction - 16 - March 1971
Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents
The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area
While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail
near Parturition
Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition
Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality
after
Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9
It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long
We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life
Table 9 Pups Accompanying Pos artum Females
Sex llleight
M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11
Total Length
47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)
Pups Accompanying Anestrous Females
F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103
Reproduction - 17 - Narch 1971
Frequency of Breeding
Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups
I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases
Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample
Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large
vhich had luteinized but tvere not actually pregnant The remainder had follicles
This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt
Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning
Table 10 Reproductive Condition of Lactating Females
Location
Bay of Is Adak I Sept 1967 9 1 100
Allchitka I Sept-Oct 1967 17 2 105
Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I
Bay of Is Adak I Oct 1968 22 4 154 3
Kanaga I Oct 1968 32 s- 135
Al1chitka I July-Aug 1969 4 2 333
iTanaga I May 1970 56 10 152
Delarof Is May 1970 18 3 143
Amchitka I May 1970 36 3 77
---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy
TOTAL 213 33 134
Anestrous or pregnant with term fetus
]_ Large follicles or corpus luteum present
3 Includes one implanted pregnancy with small fetus
Reproduction - 18 - March 1971
The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy
The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year
Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period
This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup
We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year
Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~
Reproduction - 19 - March 1971
observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less
Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years
This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias
Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years
Reproduction - 20 - March 7 1971
Conclusions
1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population
2 Breeding activity is highest in September October and November
3 More implantations occur during January February and March than at other times of year
4 The birth rate is highest during April Y~y and June
5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity
6 Similarly two and a half to three times as many females pup during peak pupping months
7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed
8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup
9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period
10 Female sea otters become sexually mature ihen 3 to 4 years old
11 Females may ovulate on an average of once a year after sexual maturity is reached
12 Females continue to breed at Cn old age
13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution
14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented
15 Sea otters probably birth both on land and in the water
16 Blastocysts rarely cross from one uterine horn to the other
17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random
Reproduction - 21 - March 1971
18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn
19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth
20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal
21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth
22 Body condition of the female has little effect on the growth rate of the fetus
23 There is a rapid ~Jeight gain shortly after parturition
24 The sex ratio at birth is 44 percent male and 56 percent female
25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term
26 Multiple ovulations occur in 4 percent of the estrus cycles
27 es do not appeErc- to be able to more than two fetuses in a single horn
28 Up to 5 percent of the p may fail dne to in after implantation
29 An unknown but probably significant number of first pregnancies fail before implantation
30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life
31 Sexually matu1middote females normally have one pup every two years
32 Females vrith a pup may ovulate but rarely mate
33 Slightly less than half of the sexually mature females are in any given year
34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived
35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus
Reproduction - 22 - March 1971
CITED
Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp
Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317
Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68
Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657
Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130
Snow H J 1910 In forbidden seas Edward Arnold London 303 pp
Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp
11arch 27 Shaw Island to Puale Bay (1030 AM - 1205 PM) Conditions very good to excellent High overcast and very little wind down to Katmai Bay After this the conditions deteriorated rapidly to fair and the count had to be stopped because of a combination of bad weather and lack of fuel As shown on the charts the count was concentrated on the points and islands however the good visibility allowed us to see well into the bays We feel certain no concentrations were missed
The numbers of sea otters counted are presented in the Table Specific locations and numbers are shown on the charts Where coverage of an area was not complete the approximate path of the aircraft is shown
Discussion of Results
Surveys along the Alaska Peninsula have been fragmentary and most of those made were done under less than ideal conditionsmiddot There are a number of reasons for this Weather tends to be poor along the south shore where squall5 form along the mountains Areas where an aircraft can refuel are north of the mountains and there are relatively few passes therefore range of the aircraft is a problem Also there is much shallow water off shore in the area from the Shumagin Islands to Sanak Island Coverage of this area is difficult and many animals are undoubtably missed
The present survey covered most of the area however conditions were such that most of the counts are probably quite low Despite these problems and a lack of continuity we have a fairly good picture of the recovery of sea otter populations in this area
Reports from various individuals are available from the 1930s and 1940s 11ajor surveys by the Fish and Wildlife Service in 1951 (Jones) 1957 (Lensink) 1962 and 1965 (Kenyon and Spencer) covered at least portions of the area In 1969 the southern Shumagin Islands were counted by the Alaska Department of Fish and Game and the present survey covered most of the rest of the area
Basically there are four distinct population nuclei in the area These centered around the Sanak Island-Sandman Reef area the Shumagin Island area the Sutwick Island area and the Augustine Island-Cape Douglas area The following discussion is based on information from reports by Lensink and Kenyon and from Alaska Department of Fish and Game surveys middot
~anak Island-Sandman Reefs
Small numbers of sea otters have been reported at Sanak Island since 1922 No reports came from the Sandman Reefs until 1942 In 1948 27 were sighted at
----Cherni Island The 1951 aerial survey showed 65 around Sanak and 97 in the Sandman Reefs In 1957 251 were seen around Sanak and 508 around the Sandman Reefs In addition two were seen in the Pavlof Islands however few were on the mainland In 1962 548 were counted in the Sanak area and 638 in the Sandman Reefs None were reported from the mainland or Pavlof lsland~however
much of the increase was in the northern area around Deer Island The 1962 survey was probably the best being made under excel lent conditions
AERIAL COUNT OF SEA OTTERS MARCH 1970
Partial or Location Count Date Camp 1ete Count Visibility
Cape Lazaref - Cold lsecty_
Cape Lazaref 3 320 Complete Poor
lkatan Peninsula 71 320 Complete Poor
False Pass - Amagat I 66 320 Camp 1ete Poor
Cold Bay 321 Partial Poor
TOTAL 141
Sanak Islands 239 322 Camp 1ete Fair to Poor
Sandman Reefs
Clubbing Rocks 12 322 Complete Fair to Poor
Cherni I amp Rocks 495+ 322 Complete fair to Poor
Goose I 7 322 Complete Fair to Poor
Hay I 18 322 Camp lete Fair to Poor
Hunt I 2 322 Complete Fair to Poor
Deer I 322 Partial Fair to Poor~
TOTAL 568+
Pavlof Islands
Inner 11iasik 2 321 Camp 1ete Fair to Poor
Outer lliasik 16 321 Camp 1ete Fair to Poor
Goloi 2 321 Complete Fair to Poor
Dolgoi 67 321 Complete Fair to Poor
Poperechnoi 29 321 Complete Fair to Poor
Ukolnoi 2 321 Complete Fair to Poor
yenWosnesenski 4 321 Complete Fair to Poor
TOTAL 122
Partial or Location Count Date Comp 1ete Count vis ib j 1i ty
Northern Shumagin Islands
Unga 184 321 Complete Fair to Poor
Popof 52 321 Complete Fair to Poor
Korovin 46 321 Complete Fair to Poor
Karpa __2t 321 Complete Fair to Poor
TOTAL 286
Cold ~ to Beaver ~ 0 320-21 Partial Poor
Beaver Bay to Kupreanof Pen
Beaver Bay 2 321 Partial Fair to Poor
Cape A 1 iaks in 4 321 Partial Fair to Poor
Guillemot I 16 321 Partial Fair to Poor
__1Kupreanof Peninsula 321 Partial Fair to Poor
TOTAL 23
Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor
AntJrrCastle Cape to-1-Ji e ~
Castle Cape-Castle Bay 16 321 Partial Fair to Poor
Chignik Bay 118 320 Complete Fair
Nakchamik I 5 320 Complete Fair
Hook Bay 13 320 Complete Fair
Cape Kum 1i un 88 320 Complete Fair
Kujul ik Bay 1199 320 Complete Very Good
Univikshak I 62 320 Complete Fair
Cape Kuml ik 54 320 Complete Fair to Poor
Sutwick I 14 320 Complete Fair to Poor
Cape Agutka 323 Complete Fair~
TOTAL 1766
Partial or Location Count Date Comp 1ete Count vis i b i 1i ty
Cape Kunmik 13 323 Complete Fair
Naka 1i kok Bay amp offshore islands 16 323 Complete Fair
Chiginagak Bay 5 323 Complete Fair
Agripina amp lmuya Bay 105 323 Complete Good
Wide Bay to Puale Bay N 0 T S U R V E Y E D
Puale Bay to c Kubugakl i __]_ 327 Partial Fair
TOTAL 146
Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent
Cape Chiniak to Shaw _I
Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent
Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent
TOTAL 71
In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring
A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak
With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time
As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population
Shumagin Is 1 ands
This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population
There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion
In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good
There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied
There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas
Sutwi ck Area
Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather
In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved
~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather
The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys
There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years
to come
Augustine Island-Cape Douqlas
For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro
On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group
It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there
The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table
SUM
MAR
Y OF
SE
A OT
TER
SURV
EYS
(Com
pile
d by
p
op
ula
tio
n f
rom
L
ensi
nk
(196
2 T
hes
is
K
enyo
n 0
96
2 amp
196
5 R
epor
ts
amp M
anu
scri
pt)
an
d AD
FampG
Dat
a)
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
AU
GU
STIN
E -
C
DOUG
LAS
Aug
usti
ne
Isla
nd
A
ppro
xim
atel
y 50
(1
948)
Shaw
1
amp
Dou
glas
Are
a S
igh
tin
gs
from
Sh
aw
Is
to
Tux
edn
i B
ay
TOTA
L
40
___
_
41
52
-shy 52
18
_lQ
J
119
130+
-shy
130+
J
71
ALAS
KA
PEN
INSU
LA
c
Kul
iak
-
c
lgva
k
c
lgva
k -
c
Kun
mik
Ani
akch
ak
Bay
amp
Am
ber
Bay
Sutw
i ck
Are
a M
isce
llan
eous
R
epor
ts
Kuj
ul i
k B
ay
amp c
K
uml
ik
Sin
ce
1936
C
Kum
1 i u
n
Uni
viks
hak
Isla
nd
N
akch
amik
Is
lan
d
8
355 12
13
0 19 6
581
103
180
1
22
47
109
684 86
Not
S
urve
yed
7(+
)
139
197 14
1 2
53
101 62
5
I
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
ALAS
KA
PEN
INSU
LA
(Co
nt
)
Chi
gnik
B
ay
Cas
tle
Bay
amp
Cap
e
TOTA
L
SHUM
AGIN
Mai
nlan
d S
ho
re
Kup
rean
of
Pen
to
P
avlo
f B
ay
Ung
a Is
lan
d
Popo
f Is
lan
d
Kor
ovin
Is
lan
d
Kar
pa
Isla
nd
And
roni
ca
amp t
he
Hay
stac
ks
Nag
ai
Sp
ecta
cle
Ben
del
Tu
rner
Tw
ins
Few
R
epor
ts
Bef
ore
19
40
0
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-shy
388
889 1 2 2 1
149
26
8
~-
7
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338 I105
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118
_lsect
1 9
12
23
184 52
46 4
75
232 8 27 6
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
SHUM
AGIN
(C
on
t)
Nea
r
Pen
insu
la
Big
Kon
i u
j i
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tle K
oniu
j i
amp A
tkin
s
Sim
eono
f
Bir
d
Che
rnab
ura
TOTA
L
500
Est
imat
e (1
947)
l63
3 (
195 3
)
3 0
220
430
455
160
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ll
183
0
14 3
222
255
294 38
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2
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296
290
329 76
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SANA
K -
SAND
MAN
Mai
nlan
d S
ho
re
Pav
lof
Bay
to
Un
imak
B
ay
Pav
lof
Isla
nd
s
Wos
nese
nski
Is
lan
d
Uko
l noi
Is
lan
d
Pop
erec
hnoi
Is
lan
d
Dol
goi
Isla
nd
2
141 4 2 29
67
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
SAN
AK
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NDM
AN
Pav
lof
Isla
nd
s
Gol
oi
Isla
nd
Out
er
llia
sik
Inne
r ll
iasik
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man
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eefs
Che
rni
r-s I
and
(Co
nt
) (C
on
t)
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nd
Isla
nd
C1u
bb i
ng
Roc
ks
Goo
se
Isla
nd
Dee
r Is
lan
d
amp O
ther
R
eefs
San
ak
Isla
nd
TOTA
L
2 16 2
27
(194
8)
271
259
495+
No
sig
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ng
s b
efo
re
1942
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76
3397
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p Je
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Hook Bay Chignik Bay
13 118
-1 shy
shy
shy
bull
I +
Castle Bay - Castle Cape 16 Nakcharriik I 5 -- Katmai Reef 0
Total 152
middot
~ middot-
bull -middot
bull
-middot
Cape Agutka Cape Kum 1 i k Sutwick I~ Kujul ik Bay Cape Kum 1 i un Unavikshak 1
Total
X
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197 54 14
1199 88 62
1614
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10
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75
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MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970
OBSERVER J Vania J Faro
PILOT George Tibbets Jr
AIRCRAFr Piper Azetex
middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good
Departed King Salmon 1115 AM Returned to King Salmon 745 PM
FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out
We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9
The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down
SEA OTTER SIGHTINGS
Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed
The number counted in each pod is as follows
1071 520 357 68 36 75 30
TOTAL 2157
Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island
171
-amp
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53
51
52
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54
58
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36 ~
44
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SEA OTTER COUNT BARREN ISLANDS and SHUYAK - AFOGNAK ISLANDS
June 9 1970
On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows
Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete
Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good
Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial
Area
BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island
AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island
TOTAL
TOTAL
Number of Sea Otters
0 2 0
76 200
29 0
307
18 6 3
121 0
33 26
207
Complete or Visibility Partial Count
Very good Complete ll If II
It II
II
II IJ
Good Complete II
Fair Partial If Complete II Partial II II
II
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64
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675
6
57
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SEA OTTER COUNTS - KENAI June J970 to January 1971
Carl Divinyi flew aerial surveys of seals along the outside of the
Kenai Peninsula on a monthly basis He recorded all sea otters sighted
on these surveys The surveys did not cover the entire coast line and
the type of aircraft weather conditions and observers varied from one
survey to another Therefore the seperate counts cannot be compared
However when viewed in total they indicate the areas used by sea otters
and the relative abundance of otters in each area
The attached table presents the counts by broad areas
--
SEA
OTT
ERS
CO
UN
TED
ON
A
ER
IAL
SUR
VEYS
O
F K
EN
AI
PEN
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Ju
ne
1
97
0-
Jan
uar
y
1971
8i5
8
JUN
E 5
amp 9
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t oc
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NO
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12
JAN
12
C
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n
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42
27
10
30
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ecti
on
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NS
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ris
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31
28
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rt
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NS
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rt C
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125
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Gra
ham
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0
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To
tal
243
262
54
75
55
152
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tters
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shor
e an
d sk
iff
11
20
70
AERIAL SURVEY Prince William Sound
April 1970
Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey
I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife
April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay
Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals
Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina
Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)
April 16 Weather inclement - no flying
April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island
April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)
Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands
The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed
SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date
PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418
CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52
PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS
ESTHER PASSAGE - VALDEZ ARM NS
GALENA BAY FISH BAY 103 415
FISH BAY - GRAVINA POINT 1 415
GRAVINA POINT - CORDOVA 1 415
HAWKINS ISLfu~D 1 4l7
HINCHINBROOK ISLfu~ 99 417
EGG ISLAND 2 417
MONTAGUE ISLAND 259 417
GREEN AND LITTLE GREEN ISLANDS 102 4J7
KNIGHT ISLAND 136 52
INGOT ISLAND 6 52
ELEANOR ISLAND 3 52
SMITH ISLAND 4 52
SEAL ISLAND 0 52
APPLEGATE ROCK 1 52
PERRY ISLAND NS
(continued) SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date-
LONE ISLAND NS
NAKED ISLAND NS
PEAK ISLAND NS
STOREY ISLAND NS
KAYAK - WINGHAM ISLAND 5 417
TOTAL 892
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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970
On July 17 1970 an aerial survey of sea otters was made in the area
around Green Island and the adjacent coast of Montague Island The purpose
of the survey was to locate concentrations of animals prior to a capturing
operation which began the next day A Dornier twin-engine arrcraft was used
with Schneider Reynolds and Somerville acting as observers The sky was
overcast almost no wind and the sea calm However heavy rain showers intershy
fered with forward visibility and the counting conditions were only fair on
the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay
The count began at 210pm and ended at 345 pm
The numbers and locations of otters are shown on the map In addition
16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy
in brook Island and a pod of 35 were seen three miles east of Johnstone Pt
Most of the area included in the survey was traveled repeatedly in a
skiff over the next four days Durlng this time the weather became calm and
clear for the first time in several days and the bulk of the sea otters shifted
from the coast of Montague out into the shallow areas between there and Green 1
and to Green and Channel Islands It was obvious that many otters had been missed
Fn the aerial sur~ey and that the population in the area is quite dense A
total of 48 sea otters were captured in three days of netting
j
SEA OTTER SURVEY Salisbury Sound to Cape Spencer
August 23-25~ 1970
On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and
middotWalt Cunningham serving as observers
The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good
On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere
On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May
Rough weather and outboard problems restricted tl the search area~
and the effectiveness of the search in that area The following sightings were made
82470 I Adult West of Granite Islands 1 Adult South of Granite Islands
11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands
These sightings represent from four to seven animals
Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull
The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal
The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait
While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island
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55
HARVEST AND TRANSPLANTS - 1970
Harvest
In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island
and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year
The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial
survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs
Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers
Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same
Transplants
Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent
The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island
Table 1
April 20
middotMay 1
May 2
May 3
May 4
May 5
May 6
May 7
May 8
May 9
May 10
May 11
May 12
May 13
Schedule of the 1970 harvest
1030 pm depart Homer
Midnight arrive Adak MV Active
Refuel and prepare boat
Arrive off Tanaga at noon
Complete skinning pack hides etc in pm
Delarofs
Spent am getting water at Amchitka
Clean up boat take down shelter pack gear
900 am arrive Amchitka fly to Anchorage
Killed 53
138
88
131
143
53
144
43
79
83
otter
(am)
(pm)
(by mid afternoon)
Table 2 Projected and actual harvest numbers May 1970
Projected Actual Tana~a Island Harvest Harvest
Bumpy Point - Hot Springs Bay 200 191
Hot Springs Bay - Twin Bays 50 50
Twin Bays - Cape Sasmik 50 ) ) 199
Cape Sasmik - Tower 120 )
Tower Tanaga Bay 180 166
TOTAL 600 606
Delarof Islands
Gareloi 20-25 0
Kavalga Ogliuga and Skagul 75-100 125
Ulak and Amatignak 35-55 19
TOTAL 150 144
Amchitka Island saved for
USFWS Counting Units 1 amp 2 100 transplant
3 30 82
4 50 36
5 120 87
TOTAL 300 205
Table 3 Details of sea otters harvested in May 1970
Tanaga Island
Total kill 606
Number of pelts saved 598
Number of small pup pelts discarded 8
Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796
(Seal 3770 void - damaged)
Approximate sex ratio - 220 males 386 females or approximately 64 percent females
Delarof Islands
Total kill 144
Number of pelts saved 138
Number of pups discarded 6
Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934
Sex ratio- 44 males 100 females or approximately 69 percent females
Amchitka Island
Total kill 205
Number of pelts saved 194
Number of pups discarded 11
Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128
Sex ratio- 27 males 178 females or approximately 867 percent females
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bull
A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality
Summary of Harvests and Transplants
Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area
The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
----
---
Tab
le 4
N
umbe
rs
of
sea o
tters
re
mov
ed
fro
m Ala
skan
pop
ulat
ions
~ 1
95
17
01
19
51
5
4
55
56
57
59
60
62
6
3
65
66
67
68
69
70
Am
chit
ka I
U
SFW
S S
tud
ies
35
22
37
2
6
21
39
2
AD
FampG
T
ran
spla
nt
47
6
237
86
A
DFamp
G
Har
ves
t 1
80
3
11
2
05
2
3
205
Oth
er
5-
24
1
14
-1
To
tal
35
22
37
26
21
39
1
80
31
1
210
50
0
25
1
292
Tan
aga
I
AD
FampG
Har
ves
t 6
06
Kan
aga
I
AD
FampG
Har
ves
t 3
18
~
Ada
k I
A
DFamp
G
Har
ves
t 30
0 1
94
Shu
mag
in Is
U
SFW
S S
tud
ies
14
2
San
ak amp
Uni
mak
Is
Hin
chin
b ro
ok
A
DFamp
G T
ran
spla
nt
41
I
A
DFamp
G
Stu
die
s 1
Ho
nta
gu
e amp
A
DFamp
G
Tra
nsp
lan
t 39
46
G
reen
Is
AD
FampG
S
tud
ies
1
Del
aro
f Is
A
DFamp
G
Han
res
t O
gli
ug
a amp
Sk
agu
l 1
25
U
lak
1
9
I
Inclu
des
all
an
imal
s th
at
wer
e h
arv
est
ed
d
ied
d
uri
ng
st
ud
ies
and
tran
spla
nt
att
em
pts
o
r w
ere
tran
spla
nte
d
to
oth
er
are
as
or
zoo
s
Doe
s n
ot
inclu
de n
atu
ral
mo
rtali
ty o
r il
leg
al
kil
ls
2
Woo
dlan
d P
ark
Zoo
3
B
att
ell
e M
emo
rial
In
sti
tute
--
Tab
le 5
N
umbe
rs
of
sea o
tters
tr
an
spla
nte
d 1
95
5-1
97
0
1955
19
56
1959
19
65
--
~ 1
96
6
1968
19
69
1970
Ale
uti
an
s A
ttu
I
5
Ott
er
I
19
1
Pri
bil
ofs
S
t
Pau
l I
7 S
t
Geo
rge
I
57
Yak
uta
t B
ay
10
Kha
z B
ay
23
20
93
58
(Ch
ich
igo
f L
)
So
uth
east
Y
akob
i I
30
A
lask
a B
iork
a I
48
Barr
ier
Is
55
Hec
eta
I
51
Cap
e S
pen
cer
25
Bri
tish
C
olum
bia
29
14
291
W
ash
ing
ton
Ore
gon
29
1
Non
e b
eli
ev
ed
to
hav
e su
rviv
ed
II
A
t le
ast
13
die
d s
ho
rtly
aft
er
rele
ase
NO
TE
1955
to
19
59 b
y U
SFW
S
1965
to
19
70 b
y
AD
FampG
In
som
e ca
ses
on
e o
r tw
o o
f th
e
abo
ve
anim
als
die
d n
ear
the
tim
e o
f re
lease
30
SEXUAL SEGREGATION IN SEA OTTER POPULATIONS
by Karl Schneider March 1971
Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males
Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands
Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals
Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas
A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population
In
the
By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled
Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet
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y R
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Hiv
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30
54
35
7
64
3
57
64
47
1
52
9
Kan
aga
Isla
nd
To
tal
63
161
28
1
71
9
121
197
38
1
61
9
Tab
le
2
Sex
C
om
po
siti
on
of
Har
ves
ted
S
ea O
tters
-
by
Are
a -
Sp
rin
g
Sam
ple
s
AM
CHIT
KA
I
May
19
70
Ad
ult
s T
ota
l (a
ll a
ges
) M
F
M
F
11
F M
F
A
Cro
wn
Ree
fer
Po
int-
Ch
itk
a C
ove
3 32
8
6
91
4
6 37
1
40
8
60
B
W
est
of
Ch
itk
a C
ove
1 32
3
0
97
0
5 39
1
14
8
86
c
8
-10
M
i
SE
o
f A
leu
t P
oin
t 1
28
34
9
66
4
32
11
1
88
9
D
10
-15
Mi
S
E
of
Ale
ut
Po
int
4 59
6
3
93
7
12
70
14
6
85
4
Am
chit
ka
Isla
nd
To
tal
9 15
1 5
6
94
4
27
178
13
2
86
8
DEL
ARO
F IS
M
ay
1970
A
Og
liu
ga
Isla
nd
amp
Sk
agu
l Is
lan
d
21
67
23
9
76
1
41
84
32
8
67
2
B
U1a
k Is
lan
d
2 10
1
67
8
3 3
3
16
15
8
84
2
Del
aro
f Is
lan
ds
To
tal
23
77
23
0
77 o
44
10
0 3
06
6
94
TAN
AG
A
I M
ay
1970
A
Cap
e S
ud
ak-P
end
ant
Po
int
40
26
60
6
39
4
79
40
66
4
33
6
B
Bar
nes
P
oin
t-T
run
k P
oin
t 2
58
33
9
67
6
66
83
9
17
c
T
run
k P
oin
t-T
win
Bay
s 3
36
77
9
23
7
43
14
0
86
0
D
Tw
in
Bay
s-S
ou
th
Bay
4
51
73
9
27
10
60
1
43
8
57
E
S
ou
th
Bay
-Har
em R
ock
6 84
6
7
93
3
9 90
9
1
90
9
F
Las
h B
aymiddot-
Infe
rno
R
eef
27
10
73
0
27
0
50
17
74
6
25
4
G
Har
em
Roc
k-K
ulak
Po
int
38
21
6t
4
35
6
52
24
68
L
31
6
H
Ku
lak
Po
int-
SE
5
33
13
2
86
8
8 45
1
51
8
49
Tan
aga
Isla
nd
To
tal
125
319
28
2
71
8
221
385
36
5
63
5
Tab
le
3
Sex
Co
mp
osi
tio
n
of
Pup
and
Su
bad
ult
S
ea O
tters
Har
ves
ted
~
by A
rea
-F
all
S
ampl
e
ADAK
I
Sep
tem
ber
196
7
Su
bad
ult
s PU
ES
All
Age
s M
F
M
F M
F
M
F
M
F
A
Bay
o
f Is
lan
ds
3
20
13
0
87
0
middot 9
4 6
92
3
08
2
67
7
33
ADAK
I
Oct
ob
er 1
968
A
Bay
o
f Is
lan
ds
0
15
0
10
00
8
3 7
27
2
73
2
40
7
60
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Tl1re~
Arm
Bay
0
1 0
10
00
1
1 5
00
5
00
2
00
8
00
c
M
id
Po
int-
Bli
nd
Po
int
5 11
3
12
6
88
3
2 6
00
4
00
3
37
6
63
Ada
k Is
lan
d T
ota
l 5
27
L5
6
8L~
4
12
6 6
66
3
33
2
78
7
22
KANA
GA
I O
cto
ber
19
68
A
Rou
nd
Hea
d-S
ho
al P
oin
t 1
1
7 6
11
3
89
3
1 7
50
2
50
4
71
5
29
B
N
aga
Po
int-
Kan
aga
Bay
0
2
0 1
00
0
4 7
36
4
63
6
20
5
79
5
c
Cap
e C
h1an
ak-C
ape
Tu
sik
3
8 2
73
7
27
10
1
90
9
91
3
53
6
47
a
D
Edd
y R
ock-
Hiv
e R
ock
24
72
7
27
3
3 1
75
0
25
0
47
1
52
9J
Kan
aga
Isla
nd
To
tal
38
26
59
4
40
6
20
10
66
6
33
3
38
1
61
9
Tab
le
4
Sex
C
om
po
siti
on
of
Pup
an
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ub
adu
lt
Sea
Ott
ers
H
arv
este
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-by
A
rea
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pri
ng
S
amp
le
AM
CHIT
KA
I
May
19
70
Su
bad
ult
s PU
J2S
All
A
ges
M
F M
F M
F
M
F
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F
A
Cro
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Reefe
r P
oin
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ka
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e 1
3 2
50
7
50
2
2 5
00
5
00
1
40
8
60
B
W
est
of
Ch
itk
a
Cov
e 1
3 2
50
7
50
3
4 5
71
4
29
1
14
8
86
c
8
-10
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S
E
of
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ut
Po
int
2 2
50
0
50
0
1
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33
6
66
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11
8
89
D
1
0-1
5 M
i
SE
o
f A
leu
t P
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t 4
10
28
6
711
+
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1 8
00
2
00
1
46
8
54
Am
ehit
ka
Isla
nd
To
tal
8 18
3
08
6
92
10
9
52
6
47
4
13
2
86
8
DEL
AR
OF
IS
1-fa
y 19
70
A
Og
liu
ga
Isla
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amp
Sk
agu
l Is
lan
d
16
9 6
40
3
60
4
8 3
33
6
66
3
28
6
72
B
U
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lan
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1 5
16
7
83
3
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00
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15
8
84
2
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rof
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ota
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llf
54
8
45
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30
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69
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30
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69
4
TA~AGA
I M
ay
1970
A
Cap
e S
ud
ak-middot
Pen
dan
t P
oin
t 35
13
7
29
2
71
4
1 8
00
2
00
6
64
3
36
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B
arn
es
Po
int-
Tru
nk
Po
int
2 6
25
0
75
0
2 2
50
0
50
0
83
9
17
c
T
run
k
Po
int-
Tw
in B
ays
3 6
33
3
66
7
1 1
50
0
50
0
14
0
86
0
D
Tw
in B
ays-
So
uth
B
ay
1 7
12
5
87
5
5 2
71
4
28
6
14
3
85
7
E
So
uth
Bay
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em R
ock
3 3
50
0
50
0
0 3
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00
0
91
9
09
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L
ash
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ay
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fern
o
Ree
f 22
7
75
9
24
1
1 0
10
00
0
74
6
25
4
G
Har
em R
ock
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lak
Po
int
13
2
86
7
13
3
1 1
50
0
50
0
68
4
31
6
H
Ku
lak
Po
int-
SE
B
igh
t 0
11
0 1
00
0
3 1
75
0
25
0
15
1
84
9
Tan
aga
Isla
nd
T
ota
l 79
55
59
o
l1l
O
17
11
6
07
3
93
3
65
6
35
Sexual Segregation - 2 - March 7 1971
Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples
Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground
Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas
Identification of Hale Areas by Pup Counts
In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable
From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen
Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area
The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on
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Sexual Segregation - 3 - March 1 1971
Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side
The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed
Kanaga Island
Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$
Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point
Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak
Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass
The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring
Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area
Tab
le
5
Com
pari
son
of
Pup
s C
ou
nte
d w
ith
S
ex R
atio
o
f H
arv
est
Jun
e
1968
A
rea
Su
rvey
O
cto
ber
19
68 H
arv
est
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l KA
NAGA
IS
LAN
D
Ind
ep
Ott
er
Ind
ep
Ott
er
M
M
F
Rou
nd
Hea
d-S
ho
al P
oin
t 0
85
3
45
6
55
4
71
5
29
Nag
a P
oin
t-K
anag
a B
ay
66
1
64
middot
18
5
81
5
20
5
79
5
Cap
e C
hla
nak
-Cap
e T
usi
k
51
1
93
2
40
7
60
3
53
6
47
Edd
y R
ock
-Hiv
e R
ock
61
3
4
35
7
64
3
47
1
52
9
Jun
e
1968
A
rea
Su
rvey
H
ay
1970
Har
ves
t
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l TA
NA
GA
IS
LAN
D
Ind
ep
Ott
er ~ter
M
F
M
F
Cap
e S
ud
ak-P
end
ant
Po
int
0 4
4
60
6
39
4
66
4
33
6
Bar
nes
P
oin
t-T
run
k P
oin
t 7
9
59
3
3
96
7
83
9
17
Tru
nk
Po
int-
Tw
in B
ays
26
4
2
77
9
23
1
40
8
60
D
rin
Bay
s-S
ou
th B
ay
96
1
11
7
3
92
7
14
3
85
7
So
uth
Bay
-Har
em R
ock
14
8
31
6
7
middot93
3
91
9
09
Las
h
Bay
-In
fern
o
Ree
f 1
5
73
0
27
0
74
6
25
4
Har
em R
ock
-Ku
lak
Po
int
0 2
7
64
4
35
6
68
4
31
6
Ku
lak
Po
int-
SE
B
igh
t 6
9
82
1
32
8
68
1
51
8
49
Sexual Segregation - 4 - March~ 1971
Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands
Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area
As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method
Identification of
Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area
The follmving is a description of the sex composition of areas for which no skiff count data was available
Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island
Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill
Sexual Segregation - 5 - March 1971
Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point
The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it
Composition of Female Areas
Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female
The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure
Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters
Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males
Sexual Segregation - 6 - March~ 1971
apparently defending a territory in California but from his observations t
it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change
Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year
A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest
The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas
The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males
There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas
Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak
within Female
In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of
Sexual Segregation - 7 - March 1971
these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June
The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups
Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population
ition of Male Areas
As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles
Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements
le 6
R
epro
du
ctiv
e S
tag
e o
f S
exu
ally
Mat
ure
S
ea O
tters
Har
ves
ted
on
Tan
aga
Isla
nd
-by
Are
a -
May
1
97
0
Un
imp
lan
ted
Im
pla
nte
d
Pre
gn
ant
Pre
gp
ant
Fet
us
Wei
gh
t C
lass
A
rea
No
np
reg
nan
t N
o
No
1
2 3
4 5
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orp
tio
n
To
tal
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dak
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dan
t P
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t 12
7
26
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0
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arn
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nes
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t-T
run
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t 1
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7
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0 3
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24
( T
run
k P
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t-T
win
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s 7
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30
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c ( (
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ard
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9
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ock
-Ku
lak
Po
int
12
6 29
3
14
1 0
1 1
0 21
H
Ku
lak
Po
int-
SE
B
igh
t 13
9
28
10
31
2 0
1 2
5 32
Bra
cket
s in
dic
ate
o
ver
lap
pin
g are
as
Sexual Segregation - 8 - March 1971
Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area
While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring
The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area
Sex Ratio
With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants
On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL
All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females
There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among
Sexual Segregation - 9 - March 1971
newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population
It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality
The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor
If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve
The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes
REPRODUCTION IN THE FElIALE SEA OTTER
by Karl Schneider Narch 1971
A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle
The follovJing criteria were used in classification
Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation
Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)
Anestrus - Largest follicle under 3 5 rnm no corpus luteum
Proestrus - Follicles over 35 mm
Es - Follicles approximately 10 rrm
Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm
- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~
al scar newly formed corpus albicans with some luteal tissue remaining
The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever
Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies
Tab
le
l Re
p iv
e co
nd
itio
n o
f se
xual
ly m
atur
e se
a o
tters
_
IF
etus
Wei
--------------+
__
_
ln~a
ctive
ov
arie
s 1
Act
ive ovari~
ht C
Jas
s
jmiddot
I P
ost
-I P
roes
tru
s U
nim
pl
lmpl
D
ates
L
ocat
ioQ
1A
nest
rus
Part
um
Res
orpt
ion
l +
E
stru
s P
reo
Pr
eQ
2 3
lJ
S
~
4-8
1970
T
anag
a 1
51
41
0
10
104
17
8 10
33
30
4 (1
68)
(1
35)
(3
3)
(3 3
) (2
89)
(34~
(5
6)
(27
) (3
3)
(11
4)(
10
9)
May
9
19
70
Del
arof
Is
18
14
1
4 13
27
10
0
3 6
8 77
4
) (1
82)
( l
3)
(5
(16
9)
( o
) (1
30
) (3
9)
(78
)(1
04
)
10-1
219
70 A
mch
ltka
I
34
18
3 8
27
48
5 6
2 23
12
13
8
( 0
(24
6)
(13
0)
(22
) (5
8)
(19
6)
(34
8)
(36
) (4
3)
(14
) (1
67)
(8
7)
~lune
23
middotmiddot ~middot~~middot~Amchitka
1
17
2 1
2 4
18
1 2
3 5
7 44
A
ugus
t 13
196
8 (3
86)
(4
5)
(23
) (4
5)
(90
) (4
09)
(2
3)
(45
) (6
8)
(11
3)(
15
8)
July
6-
~dgtAmchitka
1
17
1 0
4 14
10
0
1 1
2 6
46
Aug
ust
819
69
(36
9)
( 2
0 2
) (8
6)
(30
4)
(21
9)
(22
) (2
2)
(43
)(1
32
)
Sep
t 1
0-17
A
dak
72
5 0
16
31
40
6 6
8 9
11
164
1967
(4
39
) ( 3
1)
(98
) ( 1
89)
(
4)
(3 7
) (3
7)
(49
) (5
5)
(6
7)
Sep
t
25
-A
mch
itk
a l
55
6
0 18
19
17
4
0 0
0 3
115
OcL
6
19
67
(4
7~8)
(5
2)
(15
7)
(16
5)
(14
8)
(35
) (8
7)
(26
)
Oct
o 1
2-1
5
Kan
aga
I
58
6 1
13
31
31
4 4
7 11
5
140
1968
(4
13)
(4
3)
(o 7
) (9
3)
(22
2)
(22
2)
(29
) (2
9)
(50
) (7
8)
(36
)
Oct
18
-20
A
dak
l
46
6 0
1 24
13
2
3 0
4 4
100
1968
(L
16 0
) ( 6
0)
(11
0)
(24
0)
(13
0)
(20
) (3
0)
0)
(40
)
Num
bers
in
re
nth
esis
are
pe
rcen
t se
xu
ally
mat
ure
fem
ales
F
etus
wei
ght
clas
s 1
=
0-l
g
2 =
1-l
Og
3
= 1
0-lO
Og
4
= 1
00-lO
OO
g
5 =
Tra
nsp
lan
t m
ort
alit
ies
(all
o
ther
sam
ples
fr
om
har
ves
ts)
Reproduction - 2 - March 1971
There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time
The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable
Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries
Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d
2
~ _s ~
~
c U)
~-1
r
~J
lt
c])
Reproduction - 3 - Narch 1971
to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors
cussion of the Annual
The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases
The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season
The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses
By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7
blastocysts implanting as is shown the increase in Class 1 fetuses
By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses
This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay
At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling
Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses
a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die
Reproduction - 4 - March 1971
There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August
Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)
If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another
The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b
The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short
There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points
At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping
Reproduction - 5 - March 1971
Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever
At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month
These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup
Gestation Period
Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature
A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth
The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)
Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April
It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t
Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo
Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted
Reproduction - 6 - March 1971
to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year
From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months
The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest
Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period
The following is a comparison of the two estimates
Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s
Total Gestation Period 399 d2ys 154 days or about 13 months 5 months
Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period
This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad
Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)
Table 2 Estimated Length of Period
Percent of Fetuses in each Height Class
Time of Collection After January
1 month 31 15 15 38 0
3 months 18 12 22 35 12
5 months 18 8 8 36 30
7 months 3 8 16 29 45
9 months 20 7 10 40 23
10 months
14 18 12 33 23
---~--
r1ean Time in Nonths After January 50 57 53 58 70
Cube Root of Umveighted Mean 063 17 36 7lf lllf
Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days
Table 3 Length of Unimplanted Period
Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted
1 month 35 26 58
3 months 49 49 so
5 months 128 179 42
7 months 22 37 37
9 months 50 57 47
10 months 55 44 56
Unweighted
48
Reproduction - 7 - March 1971
All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples
The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available
This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless
Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent
There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months
Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts
Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of
11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data
The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a
Reproduction - 8 - March 1971
Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters
If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months
The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping
The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve
Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination
There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months
Fetal Rate
the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e
--
3 0
~
shy
----- -shy --middotmiddot~middotmiddot
___ ___1__
_
(middot-)
_ ft -) J ~)___
~-
3
Reproduction - 9 - IYarch 1971
The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)
Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year
Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)
The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based
For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)
A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit
The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined
2
G
X
middot ~ -- (-- ) r~ ~ - - -- ~
yen~ r- ~--~
3
i__middot -~---
~
gt
- cshy
Reproduction - 10 - March 1971
~e of _Sexual Maturity
Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition
Ovulation
From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)
faximum
Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point
Ovulation
Many mustelids are induced ovulatoTs
On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous
ficance of
Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the
season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary
----
a - o lt bull y bull bull l w laquo bull _ ~ ~
Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968
N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A
I
~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy
- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot
1-2
2
3
4 1 I
2 (1)
6
middot5
1 (1)1
2 (l) 17
1 ( 1 )
2 ( 1)
18
l (1) I ( 1 ) 9
10 1 ( 1) L ~j( 2 1 ( 1 )
2 ( l) 1
12
11
1 (1)
I I (I)
1
2 ( 1)
13
1 ( 1 )
2
114
115 I116
17
I 18
I I
I
119 I
Numbers inside parentheses =Number of individuals with corpus luteum
11
Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968
N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I
AGE 84 I I
5 I l 6 7z 3(Years) 1
l rbull ~middot--
--- shy
0-l
J l-2
~ 2
3
4 2 ( 1)
5 3
Ibullbull 6 6 (3)
4 (4)7
8 4
3 9 2 ( 1 )
j iI 10
I 1 (1)11
12
13
14
1 ( 1)
16
15
17
18
Numbers irisi
-
J
(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)
1
1 (1)
5 (1)
10(5)
3
3 1
3 (2)
2 ( 1) 3 (3)
1 ( 1)
2
1 ( 1 ) 5 ( 1)
1 ( 1 ) 3 ( 1)
1
3
2 ( 1 )
1 (l)
1 (1)4 ( 1)
2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )
1 ( 1)
1
2
1 ( 1)
1 ( 1)
1
I I parentheses = Number of individuals with corpus luteum
11
Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968
N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A
-1AGE
~
9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--
- _ Q~L
1-2
(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3
4 2 (1)
l5
16 1 (1)
1 1)7
8
2 (1)9
10
111
12
I13
14
15 1 (1) 16
17
18
1 ( 1)
1 (l)
1
1
1
2
1
1
1
1(1 )
1
1 (1)
1 (1)
1
I
1
1
1
1
-I
Numbers Inside rentheses = Number of individuals with corpus luteum
1
Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968
AGE (Years)
Q-1
t-2
2
3
4
s 6
7
8
9
10
11
12
13
14
15
16
17
18
Numbers
N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A
~L 3 4 slE - 1middotshy
7 1~8 __J~l_11 shy
1-
One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I
One vJi th corpus luteum but no corpora al bicantia
12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3
1 (1) 2 (1)
1 2 (2) l 1
1 (1) 4 (1)
2 (1)
2 1 )
1 ( 1 ) 1
2 (1)
2 (2)
l22 3
3 ( 1)2 21 ( 1)
2 ( 1)
2 ( 1)
l ( 1)1
1 1
1 I ll
I
I I
inside parentheses Nurnber of individuals with corpus luteurn
Reproduction - 11 - March 1971
The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year
Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out
The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve
Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es
It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the
ion and ended around parturi tioD
Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists
Fetal
Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land
Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation
The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first
If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water
Reproduction - 12 - tltIarch 1971
At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water
Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample
Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn
It appears that blastocysts rarely cross from one horn to the other
Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other
When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time
A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active
Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus
Reproductive - 13 - March 1971
Birth iltleigh t
Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g
The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g
One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected
From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high
This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation
This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier
This may have sone effect on information on this
Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females
~
(
~
-s 2 0 C
U)
0
--
ez ()
~) I) middot _t t- (
j -~middot -~
Reproduction - 14 - March 1971
This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition
Sex Ratio at Birth
Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained
In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female
Sex Ratio of the
In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales
The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population
The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be
Lit
The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported
In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8
Table 8 Multiple Corpora Lutea Fetuses
Total Pregnant Females 565
Implanted Pregnancies 316
Unimplanted with 2 CL (corpora lutea) 9
Implanted Pregnancies with ) L CL and 1 Fetus 8
Implanted Pregnancies Vlith 3 CL and 1 Fetus 1
Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5
Implanted Pregnancies middotwith 3 CL and gt Fetuses 1
Total Nultiple Ovulations
I
Percent Nultiple Ovulations 42
Percent of Pregnancies tvith Nultiple Fetuses 19
Corpora Lutea per Pregnancy 105
Fetuses per 102
Reproduction - 15 - lvlarch 1971
From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally
All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo
in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time
With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations
The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus
A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some
Des
Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)
Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)
Resorbed or aborted fetus (implantation at end of horn) 3
Resorbed fetus (umbilicus tvTisted tightly) 1
Resorbed fetuses (three or more fetuses in one horn) 2
Resorbed fetus (cause unknmm) 5
Reproduction - 16 - March 1971
Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents
The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area
While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail
near Parturition
Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition
Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality
after
Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9
It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long
We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life
Table 9 Pups Accompanying Pos artum Females
Sex llleight
M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11
Total Length
47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)
Pups Accompanying Anestrous Females
F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103
Reproduction - 17 - Narch 1971
Frequency of Breeding
Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups
I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases
Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample
Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large
vhich had luteinized but tvere not actually pregnant The remainder had follicles
This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt
Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning
Table 10 Reproductive Condition of Lactating Females
Location
Bay of Is Adak I Sept 1967 9 1 100
Allchitka I Sept-Oct 1967 17 2 105
Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I
Bay of Is Adak I Oct 1968 22 4 154 3
Kanaga I Oct 1968 32 s- 135
Al1chitka I July-Aug 1969 4 2 333
iTanaga I May 1970 56 10 152
Delarof Is May 1970 18 3 143
Amchitka I May 1970 36 3 77
---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy
TOTAL 213 33 134
Anestrous or pregnant with term fetus
]_ Large follicles or corpus luteum present
3 Includes one implanted pregnancy with small fetus
Reproduction - 18 - March 1971
The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy
The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year
Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period
This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup
We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year
Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~
Reproduction - 19 - March 1971
observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less
Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years
This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias
Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years
Reproduction - 20 - March 7 1971
Conclusions
1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population
2 Breeding activity is highest in September October and November
3 More implantations occur during January February and March than at other times of year
4 The birth rate is highest during April Y~y and June
5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity
6 Similarly two and a half to three times as many females pup during peak pupping months
7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed
8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup
9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period
10 Female sea otters become sexually mature ihen 3 to 4 years old
11 Females may ovulate on an average of once a year after sexual maturity is reached
12 Females continue to breed at Cn old age
13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution
14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented
15 Sea otters probably birth both on land and in the water
16 Blastocysts rarely cross from one uterine horn to the other
17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random
Reproduction - 21 - March 1971
18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn
19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth
20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal
21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth
22 Body condition of the female has little effect on the growth rate of the fetus
23 There is a rapid ~Jeight gain shortly after parturition
24 The sex ratio at birth is 44 percent male and 56 percent female
25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term
26 Multiple ovulations occur in 4 percent of the estrus cycles
27 es do not appeErc- to be able to more than two fetuses in a single horn
28 Up to 5 percent of the p may fail dne to in after implantation
29 An unknown but probably significant number of first pregnancies fail before implantation
30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life
31 Sexually matu1middote females normally have one pup every two years
32 Females vrith a pup may ovulate but rarely mate
33 Slightly less than half of the sexually mature females are in any given year
34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived
35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus
Reproduction - 22 - March 1971
CITED
Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp
Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317
Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68
Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657
Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130
Snow H J 1910 In forbidden seas Edward Arnold London 303 pp
Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp
AERIAL COUNT OF SEA OTTERS MARCH 1970
Partial or Location Count Date Camp 1ete Count Visibility
Cape Lazaref - Cold lsecty_
Cape Lazaref 3 320 Complete Poor
lkatan Peninsula 71 320 Complete Poor
False Pass - Amagat I 66 320 Camp 1ete Poor
Cold Bay 321 Partial Poor
TOTAL 141
Sanak Islands 239 322 Camp 1ete Fair to Poor
Sandman Reefs
Clubbing Rocks 12 322 Complete Fair to Poor
Cherni I amp Rocks 495+ 322 Complete fair to Poor
Goose I 7 322 Complete Fair to Poor
Hay I 18 322 Camp lete Fair to Poor
Hunt I 2 322 Complete Fair to Poor
Deer I 322 Partial Fair to Poor~
TOTAL 568+
Pavlof Islands
Inner 11iasik 2 321 Camp 1ete Fair to Poor
Outer lliasik 16 321 Camp 1ete Fair to Poor
Goloi 2 321 Complete Fair to Poor
Dolgoi 67 321 Complete Fair to Poor
Poperechnoi 29 321 Complete Fair to Poor
Ukolnoi 2 321 Complete Fair to Poor
yenWosnesenski 4 321 Complete Fair to Poor
TOTAL 122
Partial or Location Count Date Comp 1ete Count vis ib j 1i ty
Northern Shumagin Islands
Unga 184 321 Complete Fair to Poor
Popof 52 321 Complete Fair to Poor
Korovin 46 321 Complete Fair to Poor
Karpa __2t 321 Complete Fair to Poor
TOTAL 286
Cold ~ to Beaver ~ 0 320-21 Partial Poor
Beaver Bay to Kupreanof Pen
Beaver Bay 2 321 Partial Fair to Poor
Cape A 1 iaks in 4 321 Partial Fair to Poor
Guillemot I 16 321 Partial Fair to Poor
__1Kupreanof Peninsula 321 Partial Fair to Poor
TOTAL 23
Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor
AntJrrCastle Cape to-1-Ji e ~
Castle Cape-Castle Bay 16 321 Partial Fair to Poor
Chignik Bay 118 320 Complete Fair
Nakchamik I 5 320 Complete Fair
Hook Bay 13 320 Complete Fair
Cape Kum 1i un 88 320 Complete Fair
Kujul ik Bay 1199 320 Complete Very Good
Univikshak I 62 320 Complete Fair
Cape Kuml ik 54 320 Complete Fair to Poor
Sutwick I 14 320 Complete Fair to Poor
Cape Agutka 323 Complete Fair~
TOTAL 1766
Partial or Location Count Date Comp 1ete Count vis i b i 1i ty
Cape Kunmik 13 323 Complete Fair
Naka 1i kok Bay amp offshore islands 16 323 Complete Fair
Chiginagak Bay 5 323 Complete Fair
Agripina amp lmuya Bay 105 323 Complete Good
Wide Bay to Puale Bay N 0 T S U R V E Y E D
Puale Bay to c Kubugakl i __]_ 327 Partial Fair
TOTAL 146
Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent
Cape Chiniak to Shaw _I
Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent
Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent
TOTAL 71
In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring
A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak
With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time
As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population
Shumagin Is 1 ands
This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population
There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion
In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good
There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied
There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas
Sutwi ck Area
Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather
In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved
~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather
The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys
There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years
to come
Augustine Island-Cape Douqlas
For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro
On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group
It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there
The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table
SUM
MAR
Y OF
SE
A OT
TER
SURV
EYS
(Com
pile
d by
p
op
ula
tio
n f
rom
L
ensi
nk
(196
2 T
hes
is
K
enyo
n 0
96
2 amp
196
5 R
epor
ts
amp M
anu
scri
pt)
an
d AD
FampG
Dat
a)
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
AU
GU
STIN
E -
C
DOUG
LAS
Aug
usti
ne
Isla
nd
A
ppro
xim
atel
y 50
(1
948)
Shaw
1
amp
Dou
glas
Are
a S
igh
tin
gs
from
Sh
aw
Is
to
Tux
edn
i B
ay
TOTA
L
40
___
_
41
52
-shy 52
18
_lQ
J
119
130+
-shy
130+
J
71
ALAS
KA
PEN
INSU
LA
c
Kul
iak
-
c
lgva
k
c
lgva
k -
c
Kun
mik
Ani
akch
ak
Bay
amp
Am
ber
Bay
Sutw
i ck
Are
a M
isce
llan
eous
R
epor
ts
Kuj
ul i
k B
ay
amp c
K
uml
ik
Sin
ce
1936
C
Kum
1 i u
n
Uni
viks
hak
Isla
nd
N
akch
amik
Is
lan
d
8
355 12
13
0 19 6
581
103
180
1
22
47
109
684 86
Not
S
urve
yed
7(+
)
139
197 14
1 2
53
101 62
5
I
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
ALAS
KA
PEN
INSU
LA
(Co
nt
)
Chi
gnik
B
ay
Cas
tle
Bay
amp
Cap
e
TOTA
L
SHUM
AGIN
Mai
nlan
d S
ho
re
Kup
rean
of
Pen
to
P
avlo
f B
ay
Ung
a Is
lan
d
Popo
f Is
lan
d
Kor
ovin
Is
lan
d
Kar
pa
Isla
nd
And
roni
ca
amp t
he
Hay
stac
ks
Nag
ai
Sp
ecta
cle
Ben
del
Tu
rner
Tw
ins
Few
R
epor
ts
Bef
ore
19
40
0
-shy
-shy
388
889 1 2 2 1
149
26
8
~-
7
- 949 4
338 I105
-~-middot
118
_lsect
1 9
12
23
184 52
46 4
75
232 8 27 6
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
SHUM
AGIN
(C
on
t)
Nea
r
Pen
insu
la
Big
Kon
i u
j i
Lit
tle K
oniu
j i
amp A
tkin
s
Sim
eono
f
Bir
d
Che
rnab
ura
TOTA
L
500
Est
imat
e (1
947)
l63
3 (
195 3
)
3 0
220
430
455
160
___L
ll
183
0
14 3
222
255
294 38
_J
l
1 35
2
150 15
296
290
329 76
_6
15
1 0
-shy 309
SANA
K -
SAND
MAN
Mai
nlan
d S
ho
re
Pav
lof
Bay
to
Un
imak
B
ay
Pav
lof
Isla
nd
s
Wos
nese
nski
Is
lan
d
Uko
l noi
Is
lan
d
Pop
erec
hnoi
Is
lan
d
Dol
goi
Isla
nd
2
141 4 2 29
67
Lo
cati
on
P
re
1951
19
51
1957
19
59
1962
19
65
1969
19
70
SAN
AK
-SA
NDM
AN
Pav
lof
Isla
nd
s
Gol
oi
Isla
nd
Out
er
llia
sik
Inne
r ll
iasik
Sand
man
R
eefs
Che
rni
r-s I
and
(Co
nt
) (C
on
t)
Isla
nd
Isla
nd
C1u
bb i
ng
Roc
ks
Goo
se
Isla
nd
Dee
r Is
lan
d
amp O
ther
R
eefs
San
ak
Isla
nd
TOTA
L
2 16 2
27
(194
8)
271
259
495+
No
sig
hti
ng
s b
efo
re
1942
~
2 12
76
3397
82
7
123
5429
5 -
(I n
com
p Je
te)
Sig
hti
ng
s si
nce
19
22
_sect
2
_lil
~
~
162
1 bull 1
86
1 0
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MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970
OBSERVER J Vania J Faro
PILOT George Tibbets Jr
AIRCRAFr Piper Azetex
middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good
Departed King Salmon 1115 AM Returned to King Salmon 745 PM
FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out
We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9
The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down
SEA OTTER SIGHTINGS
Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed
The number counted in each pod is as follows
1071 520 357 68 36 75 30
TOTAL 2157
Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island
171
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SEA OTTER COUNT BARREN ISLANDS and SHUYAK - AFOGNAK ISLANDS
June 9 1970
On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows
Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete
Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good
Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial
Area
BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island
AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island
TOTAL
TOTAL
Number of Sea Otters
0 2 0
76 200
29 0
307
18 6 3
121 0
33 26
207
Complete or Visibility Partial Count
Very good Complete ll If II
It II
II
II IJ
Good Complete II
Fair Partial If Complete II Partial II II
II
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51
SEA OTTER COUNTS - KENAI June J970 to January 1971
Carl Divinyi flew aerial surveys of seals along the outside of the
Kenai Peninsula on a monthly basis He recorded all sea otters sighted
on these surveys The surveys did not cover the entire coast line and
the type of aircraft weather conditions and observers varied from one
survey to another Therefore the seperate counts cannot be compared
However when viewed in total they indicate the areas used by sea otters
and the relative abundance of otters in each area
The attached table presents the counts by broad areas
--
SEA
OTT
ERS
CO
UN
TED
ON
A
ER
IAL
SUR
VEYS
O
F K
EN
AI
PEN
I NS
ULP
Ju
ne
1
97
0-
Jan
uar
y
1971
8i5
8
JUN
E 5
amp 9
JU
LY
15-2
0 A
UG
1l
t oc
r 12
NO
V
12
JAN
12
C
Junk
en
-C
R
esu
rrec
tio
n
5 30
42
27
10
30
Res
urr
ecti
on
Bay
2
2 0
4 2
NS
Ai a
1 i k
Bay
1
20
5 8
0 21
Har
ris
Bay
8
18
7 5
3 25
N
uka
Bay
10
6 56
NS
31
28
27
Po
rt
Dic
k 0
l1
NS
NS
3 23
Roc
ky
Bay
-
Po
rt C
hath
am
121
125
NS
NS
9 26
Koy
ukto
lik
Bay
-P
ort
Gra
ham
0
0 NS
NS
0
NS-
To
tal
243
262
54
75
55
152
middot3
8 se
a o
tters
cou
nted
fro
m
shor
e an
d sk
iff
11
20
70
AERIAL SURVEY Prince William Sound
April 1970
Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey
I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife
April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay
Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals
Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina
Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)
April 16 Weather inclement - no flying
April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island
April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)
Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands
The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed
SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date
PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418
CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52
PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS
ESTHER PASSAGE - VALDEZ ARM NS
GALENA BAY FISH BAY 103 415
FISH BAY - GRAVINA POINT 1 415
GRAVINA POINT - CORDOVA 1 415
HAWKINS ISLfu~D 1 4l7
HINCHINBROOK ISLfu~ 99 417
EGG ISLAND 2 417
MONTAGUE ISLAND 259 417
GREEN AND LITTLE GREEN ISLANDS 102 4J7
KNIGHT ISLAND 136 52
INGOT ISLAND 6 52
ELEANOR ISLAND 3 52
SMITH ISLAND 4 52
SEAL ISLAND 0 52
APPLEGATE ROCK 1 52
PERRY ISLAND NS
(continued) SEA OTTER COUNTED Prince William Sound
April 1970
Number of Area Sea Otter Date-
LONE ISLAND NS
NAKED ISLAND NS
PEAK ISLAND NS
STOREY ISLAND NS
KAYAK - WINGHAM ISLAND 5 417
TOTAL 892
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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970
On July 17 1970 an aerial survey of sea otters was made in the area
around Green Island and the adjacent coast of Montague Island The purpose
of the survey was to locate concentrations of animals prior to a capturing
operation which began the next day A Dornier twin-engine arrcraft was used
with Schneider Reynolds and Somerville acting as observers The sky was
overcast almost no wind and the sea calm However heavy rain showers intershy
fered with forward visibility and the counting conditions were only fair on
the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay
The count began at 210pm and ended at 345 pm
The numbers and locations of otters are shown on the map In addition
16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy
in brook Island and a pod of 35 were seen three miles east of Johnstone Pt
Most of the area included in the survey was traveled repeatedly in a
skiff over the next four days Durlng this time the weather became calm and
clear for the first time in several days and the bulk of the sea otters shifted
from the coast of Montague out into the shallow areas between there and Green 1
and to Green and Channel Islands It was obvious that many otters had been missed
Fn the aerial sur~ey and that the population in the area is quite dense A
total of 48 sea otters were captured in three days of netting
j
SEA OTTER SURVEY Salisbury Sound to Cape Spencer
August 23-25~ 1970
On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and
middotWalt Cunningham serving as observers
The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good
On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere
On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May
Rough weather and outboard problems restricted tl the search area~
and the effectiveness of the search in that area The following sightings were made
82470 I Adult West of Granite Islands 1 Adult South of Granite Islands
11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands
These sightings represent from four to seven animals
Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull
The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal
The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait
While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island
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HARVEST AND TRANSPLANTS - 1970
Harvest
In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island
and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year
The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial
survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs
Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers
Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same
Transplants
Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent
The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island
Table 1
April 20
middotMay 1
May 2
May 3
May 4
May 5
May 6
May 7
May 8
May 9
May 10
May 11
May 12
May 13
Schedule of the 1970 harvest
1030 pm depart Homer
Midnight arrive Adak MV Active
Refuel and prepare boat
Arrive off Tanaga at noon
Complete skinning pack hides etc in pm
Delarofs
Spent am getting water at Amchitka
Clean up boat take down shelter pack gear
900 am arrive Amchitka fly to Anchorage
Killed 53
138
88
131
143
53
144
43
79
83
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(am)
(pm)
(by mid afternoon)
Table 2 Projected and actual harvest numbers May 1970
Projected Actual Tana~a Island Harvest Harvest
Bumpy Point - Hot Springs Bay 200 191
Hot Springs Bay - Twin Bays 50 50
Twin Bays - Cape Sasmik 50 ) ) 199
Cape Sasmik - Tower 120 )
Tower Tanaga Bay 180 166
TOTAL 600 606
Delarof Islands
Gareloi 20-25 0
Kavalga Ogliuga and Skagul 75-100 125
Ulak and Amatignak 35-55 19
TOTAL 150 144
Amchitka Island saved for
USFWS Counting Units 1 amp 2 100 transplant
3 30 82
4 50 36
5 120 87
TOTAL 300 205
Table 3 Details of sea otters harvested in May 1970
Tanaga Island
Total kill 606
Number of pelts saved 598
Number of small pup pelts discarded 8
Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796
(Seal 3770 void - damaged)
Approximate sex ratio - 220 males 386 females or approximately 64 percent females
Delarof Islands
Total kill 144
Number of pelts saved 138
Number of pups discarded 6
Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934
Sex ratio- 44 males 100 females or approximately 69 percent females
Amchitka Island
Total kill 205
Number of pelts saved 194
Number of pups discarded 11
Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128
Sex ratio- 27 males 178 females or approximately 867 percent females
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A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality
Summary of Harvests and Transplants
Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area
The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5
----
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Tab
le 4
N
umbe
rs
of
sea o
tters
re
mov
ed
fro
m Ala
skan
pop
ulat
ions
~ 1
95
17
01
19
51
5
4
55
56
57
59
60
62
6
3
65
66
67
68
69
70
Am
chit
ka I
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SFW
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ies
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22
37
2
6
21
39
2
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ran
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nt
47
6
237
86
A
DFamp
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Har
ves
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80
3
11
2
05
2
3
205
Oth
er
5-
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1
14
-1
To
tal
35
22
37
26
21
39
1
80
31
1
210
50
0
25
1
292
Tan
aga
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ves
t 6
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ves
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94
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in Is
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ak amp
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nt
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nta
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reen
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res
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l 1
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Inclu
des
all
an
imal
s th
at
wer
e h
arv
est
ed
d
ied
d
uri
ng
st
ud
ies
and
tran
spla
nt
att
em
pts
o
r w
ere
tran
spla
nte
d
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oth
er
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as
or
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s
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s n
ot
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de n
atu
ral
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rtali
ty o
r il
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al
kil
ls
2
Woo
dlan
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ark
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3
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att
ell
e M
emo
rial
In
sti
tute
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le 5
N
umbe
rs
of
sea o
tters
tr
an
spla
nte
d 1
95
5-1
97
0
1955
19
56
1959
19
65
--
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96
6
1968
19
69
1970
Ale
uti
an
s A
ttu
I
5
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er
I
19
1
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bil
ofs
S
t
Pau
l I
7 S
t
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rge
I
57
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uta
t B
ay
10
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z B
ay
23
20
93
58
(Ch
ich
igo
f L
)
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uth
east
Y
akob
i I
30
A
lask
a B
iork
a I
48
Barr
ier
Is
55
Hec
eta
I
51
Cap
e S
pen
cer
25
Bri
tish
C
olum
bia
29
14
291
W
ash
ing
ton
Ore
gon
29
1
Non
e b
eli
ev
ed
to
hav
e su
rviv
ed
II
A
t le
ast
13
die
d s
ho
rtly
aft
er
rele
ase
NO
TE
1955
to
19
59 b
y U
SFW
S
1965
to
19
70 b
y
AD
FampG
In
som
e ca
ses
on
e o
r tw
o o
f th
e
abo
ve
anim
als
die
d n
ear
the
tim
e o
f re
lease
30
SEXUAL SEGREGATION IN SEA OTTER POPULATIONS
by Karl Schneider March 1971
Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males
Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands
Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals
Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas
A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population
In
the
By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled
Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet
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1970
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1970
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I M
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1970
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29
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64
3
36
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75
0
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50
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9
17
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50
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55
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07
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6
35
Sexual Segregation - 2 - March 7 1971
Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples
Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground
Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas
Identification of Hale Areas by Pup Counts
In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable
From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen
Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area
The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on
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Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side
The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed
Kanaga Island
Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$
Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point
Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak
Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass
The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring
Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area
Tab
le
5
Com
pari
son
of
Pup
s C
ou
nte
d w
ith
S
ex R
atio
o
f H
arv
est
Jun
e
1968
A
rea
Su
rvey
O
cto
ber
19
68 H
arv
est
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l KA
NAGA
IS
LAN
D
Ind
ep
Ott
er
Ind
ep
Ott
er
M
M
F
Rou
nd
Hea
d-S
ho
al P
oin
t 0
85
3
45
6
55
4
71
5
29
Nag
a P
oin
t-K
anag
a B
ay
66
1
64
middot
18
5
81
5
20
5
79
5
Cap
e C
hla
nak
-Cap
e T
usi
k
51
1
93
2
40
7
60
3
53
6
47
Edd
y R
ock
-Hiv
e R
ock
61
3
4
35
7
64
3
47
1
52
9
Jun
e
1968
A
rea
Su
rvey
H
ay
1970
Har
ves
t
Pu
ps
10
0
Pu
ps
10
0
Ad
ult
s T
ota
l TA
NA
GA
IS
LAN
D
Ind
ep
Ott
er ~ter
M
F
M
F
Cap
e S
ud
ak-P
end
ant
Po
int
0 4
4
60
6
39
4
66
4
33
6
Bar
nes
P
oin
t-T
run
k P
oin
t 7
9
59
3
3
96
7
83
9
17
Tru
nk
Po
int-
Tw
in B
ays
26
4
2
77
9
23
1
40
8
60
D
rin
Bay
s-S
ou
th B
ay
96
1
11
7
3
92
7
14
3
85
7
So
uth
Bay
-Har
em R
ock
14
8
31
6
7
middot93
3
91
9
09
Las
h
Bay
-In
fern
o
Ree
f 1
5
73
0
27
0
74
6
25
4
Har
em R
ock
-Ku
lak
Po
int
0 2
7
64
4
35
6
68
4
31
6
Ku
lak
Po
int-
SE
B
igh
t 6
9
82
1
32
8
68
1
51
8
49
Sexual Segregation - 4 - March~ 1971
Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands
Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area
As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method
Identification of
Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area
The follmving is a description of the sex composition of areas for which no skiff count data was available
Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island
Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill
Sexual Segregation - 5 - March 1971
Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point
The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it
Composition of Female Areas
Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female
The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure
Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters
Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males
Sexual Segregation - 6 - March~ 1971
apparently defending a territory in California but from his observations t
it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change
Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year
A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest
The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas
The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males
There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas
Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak
within Female
In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of
Sexual Segregation - 7 - March 1971
these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June
The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups
Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population
ition of Male Areas
As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles
Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements
le 6
R
epro
du
ctiv
e S
tag
e o
f S
exu
ally
Mat
ure
S
ea O
tters
Har
ves
ted
on
Tan
aga
Isla
nd
-by
Are
a -
May
1
97
0
Un
imp
lan
ted
Im
pla
nte
d
Pre
gn
ant
Pre
gp
ant
Fet
us
Wei
gh
t C
lass
A
rea
No
np
reg
nan
t N
o
No
1
2 3
4 5
Res
orp
tio
n
To
tal
A
Su
dak
-Pen
dan
t P
oin
t 12
7
26
8 30
0
2 0
3 2
2 27
( B
arn
es
Po
int-
Ho
t S
pri
ng
s B
ay
12
7 24
10
36
3
0 1
4 2
2 29
B
( (
Bar
nes
P
oin
t-T
run
k P
oin
t 1
1
6 25
7
29
0 3
1 3
0 2
24
( T
run
k P
oin
t-T
win
Bay
s 7
9 39
7
30
2 1
1 1
2 2
23
c ( (
Haz
ard
P
oin
t-T
win
Bay
s 4
3 27
4
36
0 0
1 1
2 11
( T
win
B
ays-
Her
d
Roc
k 1
2
8 28
9
31
1 1
1 Lf
2
29
D
( ( T
win
B
ays-
So
uth
Bay
2
9 45
9
45
3 0
1 1
Lf
1 20
( S
ou
th B
ay-L
ash
B
ay
6 9
29
16
51
1 0
0 9
6 1
31
E
( ((
So
uth
Bay
-Har
em R
ock
17
14
31
14
31
2 0
1 5
6 45
(
F ( (
L
ash
B
ay-I
nfe
rno
Ree
f 4
1 9
6 55
2
1 1
0 2
11
( G
(
Har
em R
ock
-Ku
lak
Po
int
12
6 29
3
14
1 0
1 1
0 21
H
Ku
lak
Po
int-
SE
B
igh
t 13
9
28
10
31
2 0
1 2
5 32
Bra
cket
s in
dic
ate
o
ver
lap
pin
g are
as
Sexual Segregation - 8 - March 1971
Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area
While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring
The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area
Sex Ratio
With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants
On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL
All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females
There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among
Sexual Segregation - 9 - March 1971
newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population
It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality
The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor
If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve
The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes
REPRODUCTION IN THE FElIALE SEA OTTER
by Karl Schneider Narch 1971
A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle
The follovJing criteria were used in classification
Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation
Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)
Anestrus - Largest follicle under 3 5 rnm no corpus luteum
Proestrus - Follicles over 35 mm
Es - Follicles approximately 10 rrm
Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm
- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~
al scar newly formed corpus albicans with some luteal tissue remaining
The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever
Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies
Tab
le
l Re
p iv
e co
nd
itio
n o
f se
xual
ly m
atur
e se
a o
tters
_
IF
etus
Wei
--------------+
__
_
ln~a
ctive
ov
arie
s 1
Act
ive ovari~
ht C
Jas
s
jmiddot
I P
ost
-I P
roes
tru
s U
nim
pl
lmpl
D
ates
L
ocat
ioQ
1A
nest
rus
Part
um
Res
orpt
ion
l +
E
stru
s P
reo
Pr
eQ
2 3
lJ
S
~
4-8
1970
T
anag
a 1
51
41
0
10
104
17
8 10
33
30
4 (1
68)
(1
35)
(3
3)
(3 3
) (2
89)
(34~
(5
6)
(27
) (3
3)
(11
4)(
10
9)
May
9
19
70
Del
arof
Is
18
14
1
4 13
27
10
0
3 6
8 77
4
) (1
82)
( l
3)
(5
(16
9)
( o
) (1
30
) (3
9)
(78
)(1
04
)
10-1
219
70 A
mch
ltka
I
34
18
3 8
27
48
5 6
2 23
12
13
8
( 0
(24
6)
(13
0)
(22
) (5
8)
(19
6)
(34
8)
(36
) (4
3)
(14
) (1
67)
(8
7)
~lune
23
middotmiddot ~middot~~middot~Amchitka
1
17
2 1
2 4
18
1 2
3 5
7 44
A
ugus
t 13
196
8 (3
86)
(4
5)
(23
) (4
5)
(90
) (4
09)
(2
3)
(45
) (6
8)
(11
3)(
15
8)
July
6-
~dgtAmchitka
1
17
1 0
4 14
10
0
1 1
2 6
46
Aug
ust
819
69
(36
9)
( 2
0 2
) (8
6)
(30
4)
(21
9)
(22
) (2
2)
(43
)(1
32
)
Sep
t 1
0-17
A
dak
72
5 0
16
31
40
6 6
8 9
11
164
1967
(4
39
) ( 3
1)
(98
) ( 1
89)
(
4)
(3 7
) (3
7)
(49
) (5
5)
(6
7)
Sep
t
25
-A
mch
itk
a l
55
6
0 18
19
17
4
0 0
0 3
115
OcL
6
19
67
(4
7~8)
(5
2)
(15
7)
(16
5)
(14
8)
(35
) (8
7)
(26
)
Oct
o 1
2-1
5
Kan
aga
I
58
6 1
13
31
31
4 4
7 11
5
140
1968
(4
13)
(4
3)
(o 7
) (9
3)
(22
2)
(22
2)
(29
) (2
9)
(50
) (7
8)
(36
)
Oct
18
-20
A
dak
l
46
6 0
1 24
13
2
3 0
4 4
100
1968
(L
16 0
) ( 6
0)
(11
0)
(24
0)
(13
0)
(20
) (3
0)
0)
(40
)
Num
bers
in
re
nth
esis
are
pe
rcen
t se
xu
ally
mat
ure
fem
ales
F
etus
wei
ght
clas
s 1
=
0-l
g
2 =
1-l
Og
3
= 1
0-lO
Og
4
= 1
00-lO
OO
g
5 =
Tra
nsp
lan
t m
ort
alit
ies
(all
o
ther
sam
ples
fr
om
har
ves
ts)
Reproduction - 2 - March 1971
There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time
The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable
Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries
Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d
2
~ _s ~
~
c U)
~-1
r
~J
lt
c])
Reproduction - 3 - Narch 1971
to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors
cussion of the Annual
The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases
The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season
The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses
By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7
blastocysts implanting as is shown the increase in Class 1 fetuses
By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses
This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay
At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling
Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses
a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die
Reproduction - 4 - March 1971
There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August
Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)
If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another
The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b
The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short
There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points
At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping
Reproduction - 5 - March 1971
Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever
At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month
These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup
Gestation Period
Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature
A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth
The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)
Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April
It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t
Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo
Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted
Reproduction - 6 - March 1971
to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year
From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months
The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest
Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period
The following is a comparison of the two estimates
Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s
Total Gestation Period 399 d2ys 154 days or about 13 months 5 months
Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period
This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad
Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)
Table 2 Estimated Length of Period
Percent of Fetuses in each Height Class
Time of Collection After January
1 month 31 15 15 38 0
3 months 18 12 22 35 12
5 months 18 8 8 36 30
7 months 3 8 16 29 45
9 months 20 7 10 40 23
10 months
14 18 12 33 23
---~--
r1ean Time in Nonths After January 50 57 53 58 70
Cube Root of Umveighted Mean 063 17 36 7lf lllf
Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days
Table 3 Length of Unimplanted Period
Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted
1 month 35 26 58
3 months 49 49 so
5 months 128 179 42
7 months 22 37 37
9 months 50 57 47
10 months 55 44 56
Unweighted
48
Reproduction - 7 - March 1971
All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples
The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available
This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless
Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent
There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months
Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts
Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of
11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data
The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a
Reproduction - 8 - March 1971
Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters
If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months
The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping
The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve
Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination
There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months
Fetal Rate
the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e
--
3 0
~
shy
----- -shy --middotmiddot~middotmiddot
___ ___1__
_
(middot-)
_ ft -) J ~)___
~-
3
Reproduction - 9 - IYarch 1971
The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)
Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year
Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)
The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based
For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)
A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit
The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined
2
G
X
middot ~ -- (-- ) r~ ~ - - -- ~
yen~ r- ~--~
3
i__middot -~---
~
gt
- cshy
Reproduction - 10 - March 1971
~e of _Sexual Maturity
Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition
Ovulation
From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)
faximum
Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point
Ovulation
Many mustelids are induced ovulatoTs
On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous
ficance of
Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the
season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary
----
a - o lt bull y bull bull l w laquo bull _ ~ ~
Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968
N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A
I
~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy
- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot
1-2
2
3
4 1 I
2 (1)
6
middot5
1 (1)1
2 (l) 17
1 ( 1 )
2 ( 1)
18
l (1) I ( 1 ) 9
10 1 ( 1) L ~j( 2 1 ( 1 )
2 ( l) 1
12
11
1 (1)
I I (I)
1
2 ( 1)
13
1 ( 1 )
2
114
115 I116
17
I 18
I I
I
119 I
Numbers inside parentheses =Number of individuals with corpus luteum
11
Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968
N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I
AGE 84 I I
5 I l 6 7z 3(Years) 1
l rbull ~middot--
--- shy
0-l
J l-2
~ 2
3
4 2 ( 1)
5 3
Ibullbull 6 6 (3)
4 (4)7
8 4
3 9 2 ( 1 )
j iI 10
I 1 (1)11
12
13
14
1 ( 1)
16
15
17
18
Numbers irisi
-
J
(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)
1
1 (1)
5 (1)
10(5)
3
3 1
3 (2)
2 ( 1) 3 (3)
1 ( 1)
2
1 ( 1 ) 5 ( 1)
1 ( 1 ) 3 ( 1)
1
3
2 ( 1 )
1 (l)
1 (1)4 ( 1)
2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )
1 ( 1)
1
2
1 ( 1)
1 ( 1)
1
I I parentheses = Number of individuals with corpus luteum
11
Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968
N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A
-1AGE
~
9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--
- _ Q~L
1-2
(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3
4 2 (1)
l5
16 1 (1)
1 1)7
8
2 (1)9
10
111
12
I13
14
15 1 (1) 16
17
18
1 ( 1)
1 (l)
1
1
1
2
1
1
1
1(1 )
1
1 (1)
1 (1)
1
I
1
1
1
1
-I
Numbers Inside rentheses = Number of individuals with corpus luteum
1
Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968
AGE (Years)
Q-1
t-2
2
3
4
s 6
7
8
9
10
11
12
13
14
15
16
17
18
Numbers
N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A
~L 3 4 slE - 1middotshy
7 1~8 __J~l_11 shy
1-
One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I
One vJi th corpus luteum but no corpora al bicantia
12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3
1 (1) 2 (1)
1 2 (2) l 1
1 (1) 4 (1)
2 (1)
2 1 )
1 ( 1 ) 1
2 (1)
2 (2)
l22 3
3 ( 1)2 21 ( 1)
2 ( 1)
2 ( 1)
l ( 1)1
1 1
1 I ll
I
I I
inside parentheses Nurnber of individuals with corpus luteurn
Reproduction - 11 - March 1971
The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year
Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out
The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve
Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es
It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the
ion and ended around parturi tioD
Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists
Fetal
Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land
Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation
The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first
If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water
Reproduction - 12 - tltIarch 1971
At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water
Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample
Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn
It appears that blastocysts rarely cross from one horn to the other
Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other
When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time
A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active
Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus
Reproductive - 13 - March 1971
Birth iltleigh t
Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g
The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g
One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected
From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high
This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation
This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier
This may have sone effect on information on this
Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females
~
(
~
-s 2 0 C
U)
0
--
ez ()
~) I) middot _t t- (
j -~middot -~
Reproduction - 14 - March 1971
This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition
Sex Ratio at Birth
Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained
In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female
Sex Ratio of the
In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales
The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population
The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be
Lit
The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported
In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8
Table 8 Multiple Corpora Lutea Fetuses
Total Pregnant Females 565
Implanted Pregnancies 316
Unimplanted with 2 CL (corpora lutea) 9
Implanted Pregnancies with ) L CL and 1 Fetus 8
Implanted Pregnancies Vlith 3 CL and 1 Fetus 1
Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5
Implanted Pregnancies middotwith 3 CL and gt Fetuses 1
Total Nultiple Ovulations
I
Percent Nultiple Ovulations 42
Percent of Pregnancies tvith Nultiple Fetuses 19
Corpora Lutea per Pregnancy 105
Fetuses per 102
Reproduction - 15 - lvlarch 1971
From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally
All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo
in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time
With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations
The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus
A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some
Des
Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)
Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)
Resorbed or aborted fetus (implantation at end of horn) 3
Resorbed fetus (umbilicus tvTisted tightly) 1
Resorbed fetuses (three or more fetuses in one horn) 2
Resorbed fetus (cause unknmm) 5
Reproduction - 16 - March 1971
Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents
The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area
While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail
near Parturition
Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition
Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality
after
Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9
It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long
We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life
Table 9 Pups Accompanying Pos artum Females
Sex llleight
M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11
Total Length
47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)
Pups Accompanying Anestrous Females
F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103
Reproduction - 17 - Narch 1971
Frequency of Breeding
Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups
I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases
Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample
Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large
vhich had luteinized but tvere not actually pregnant The remainder had follicles
This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt
Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning
Table 10 Reproductive Condition of Lactating Females
Location
Bay of Is Adak I Sept 1967 9 1 100
Allchitka I Sept-Oct 1967 17 2 105
Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I
Bay of Is Adak I Oct 1968 22 4 154 3
Kanaga I Oct 1968 32 s- 135
Al1chitka I July-Aug 1969 4 2 333
iTanaga I May 1970 56 10 152
Delarof Is May 1970 18 3 143
Amchitka I May 1970 36 3 77
---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy
TOTAL 213 33 134
Anestrous or pregnant with term fetus
]_ Large follicles or corpus luteum present
3 Includes one implanted pregnancy with small fetus
Reproduction - 18 - March 1971
The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy
The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year
Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period
This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup
We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year
Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~
Reproduction - 19 - March 1971
observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less
Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years
This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias
Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years
Reproduction - 20 - March 7 1971
Conclusions
1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population
2 Breeding activity is highest in September October and November
3 More implantations occur during January February and March than at other times of year
4 The birth rate is highest during April Y~y and June
5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity
6 Similarly two and a half to three times as many females pup during peak pupping months
7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed
8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup
9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period
10 Female sea otters become sexually mature ihen 3 to 4 years old
11 Females may ovulate on an average of once a year after sexual maturity is reached
12 Females continue to breed at Cn old age
13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution
14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented
15 Sea otters probably birth both on land and in the water
16 Blastocysts rarely cross from one uterine horn to the other
17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random
Reproduction - 21 - March 1971
18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn
19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth
20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal
21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth
22 Body condition of the female has little effect on the growth rate of the fetus
23 There is a rapid ~Jeight gain shortly after parturition
24 The sex ratio at birth is 44 percent male and 56 percent female
25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term
26 Multiple ovulations occur in 4 percent of the estrus cycles
27 es do not appeErc- to be able to more than two fetuses in a single horn
28 Up to 5 percent of the p may fail dne to in after implantation
29 An unknown but probably significant number of first pregnancies fail before implantation
30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life
31 Sexually matu1middote females normally have one pup every two years
32 Females vrith a pup may ovulate but rarely mate
33 Slightly less than half of the sexually mature females are in any given year
34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived
35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus
Reproduction - 22 - March 1971
CITED
Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp
Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317
Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68
Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657
Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130
Snow H J 1910 In forbidden seas Edward Arnold London 303 pp
Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp