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BULLETIN OF MARINE SCIENCE 45(2) 316-337 ]989
COMPARISON OF MYODOCOPID OSTRACODES IN TWO ZONESOF THE BELIZE BARRIER REEF NEAR CARRIE BOW CAY
WITH CHANGES IN DISTRIBUTION 1978-1981
Anne Constant Cohen
ABSTRACTThe number of myodocopid species (51) in the vicinity of Carrie Bow Cay Belize is the
highest yet reported for any geographic area of similar size Replicate sampling in two differentsand-bottomed zones a shallow (15 m) lagoon site and a deeper (18-30 m) fore-reef sitefrom 1978-1981 shows that myodocopid species and families differed between the zonesand that species and family composition varied from year to year possibly due to disturbanceby and recovery from a hurricane Myodocopids were less abundant but species diversitygreater and yearly changes much smaller in the deeper less disturbed fore-reef site than inthe shallower lagoon site which is subject to greater storm disturbance Some changes inspecies and family abundance following a storm may be related to differences in life-historystrategies
With one exception (Kornicker 1958) previous collections ofmyodocopids oncoral reefs have consisted of descriptions of a few species andor have been fromsingle samples made in each of a relatively few locations (Poulsen 1962 1965Hartmann 1964 Kornicker and King 1965 Hall 1985) Kornicker (1958) dis-cussed the systematics and distribution of myodocopids (21 species) on the Ba-hama Bank near Bimini an island in the northern part of the range of Caribbeancoral reefs and with reefs less developed than on the barrier reef off Belize
My study is part of a larger research project begun in 1972 by the SmithsonianInstitution at Carrie Bow Cay in the central province of the 100-mile-Iong coralbarrier reef lying 10-12 miles off the coast of Belize (Riitz1er and Macintyre1982) The central province has the highest coral species abundance and sea levelreef development on the barrier reef (Burke 1982) In 1978 Belize was hit byHurricane Greta providing an opportunity to study the effects of a major dis-turbance on the stability and persistence of the myodocopid fauna Field studieson abundance of two or more species in any community covering a number ofgenerations are rare but important (Connell and Sousa 1983) The finding ofnonstable conditions on a local scale andor over the course of a few turnovershas important ecological and evolutionary implications Disturbances causinglocal nonstable or persistent conditions are frequently essential for the stabilityor persistence of species on larger scales (Connell and Sousa 1983) This studyis the first survey of the myodocopid fauna on a well developed coral reef It isalso the first analysis of distribution of the myodocopid fauna in relation to reefzonation temporal fluctuations and life-history
METHODS
Collection Site -Specimens were collected from a variety of habitats in the vicinity of Carrie BowCay (16deg48N 88deg05W) mostly from 1976 to 1981 (Fig I) Samples used for numerical comparisonswere taken during 1978-1981 in the following two reef zones near a fixed transect line crossing thebarrier reef just north of Carrie Bow Cay the sand and rubble zone of the lagoon (Figs 2 3A) andthe sand trough of the outer fore-reef (Figs 2 3B) These two sites were chosen for three reasons (I)the fixed reef transect line had been previously photographed mapped and divided into units definedand described on the basis of dominant biological and geological features (Riitzler and Macintyre1982) (2) although the two zones differed in reef zonation and depth each had broad sandy bottoms
316
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 317
II]Shoals
~ Sand Bores ampPatch Reefs
o km
oQ)
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Tobacco Range ~gt0
Figure I Map of bamer reef complex in the vicinity of Came Bow Cay area of larger map locatedon inset by rectangle [From Riitzler and Macintyre 1982 used with permission of Riitzler]
which could be easily sampled by the same method and (3) in preliminary samples many myodocopidostracodes were found in these two sites The sand and rubble sampling zone is about 15 m deepwith a bottom of poorly sorted sediment ranging in size from silt to gravel and 10-14 m thick overa hard bottom (Riitzler and Macintyre 1982) The area sampled in the sand trough is about 18-30m deep with substrate that is poorly sorted silt size to very coarse sand sediment plain This sedimentis mainly very fine to fine sand but coarser material consisting of Halimeda plates mollusks benthicforaminiferans and echinoids is scattered throughout Probings indicated that sediment is morethan 12 m in the axis of the trough (Riitzler and Macintyre 1982)
To investigate local ranges of myodocopids additional nonquantitative samples were taken from(I) other zones along the fixed reef transect line (2) the following areas in the vicinity of Came BowCay lagoon and back-reef around Came Bow Cay (sand cay) and South Water Cay (mangrove cay)south shores and a channel through Twin Cays (mangroves) South Water Cut (channel through the
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318 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 319
Figure 3 A Photo of bottom in sand and rubble zone ofIagoon depth 15 m B Photo of bottomin sand trough of outer fore-reef depth 30 m showing part of movable transect line
barrier reef) Curlew Bank (submerged cay) lagoon patch reefs and adjacent sea grass (in deep lagoonin barrier reef complex) Ragged Cay and adjacent bank (western or inner edge of barrier reef complex)and (3) on the mainland subtidal shores of northern (sand beach) and just north of (mangroves andsea grass) Dandriga Nonquantitative collections were made by the author in May 1976 April 1977January 1978 June and October 1979 and June 1981 Other ostracodes collected at the study siteand deposited in the National Museum of Natural History (USNM) were also used to complete dataon indi vidual species ranges (listed in Cohen 1987) Additional data on the local range of Skogsbergialerneri were obtained from baited traps Traps were left overnight (about 1700-0900) in the lagoon
320 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
and back-reef(one in June 1979 two in June 1981) during the day (0900-1700) in the back reef (onein June 1981) and for about 24 h (two in the lagoon and back-reef one in the spur and groove inJanuary 1978 one in the lagoon and one in the spur and groove in June 1979 one each in the lagoonback-reef spur and groove sand trough and outer ridge in October 1979 one in the back-reef in1981)
Sampling Methods -Samples were made by diving usually with SCUBA except in very shallow waterwhere collections were made while wading In depths accessible by diving this method is superior todredging because the area sampled can be chosen and the sampling monitored visually Coring wastried but found to yield very few ostracodes and therefore not used Quantitative samples in the twostudy areas were made by dragging a rectangular hand net (16 cm high 22 cm wide mesh opening02 mm) along the surface of the sediment sequentially along both sides of a movable 3-m transectline The net lightly scraped the surface sediment so that the net sampled the upper 1-2 em of sedimentand 14-15 em of water above it thus sampling the strata occupied by ostracods observed undisturbedand disturbed in dishes of sediment and seawater The net was immediately twisted to trap materialat the distal end and enclosed in a plastic bag Quantitative comparisons are based only on samplesmade with the movable transect first used in 1978 Single quantitative samples were taken in eachof the two zones in 1978 At least three quantitative samples were taken in each zone during eachsubsequent field trip (June 1979 October 1979 and June 1981) and three per zone used in analysisof those dates (see below) An aquarium net was used in 1978 but replaced in 1979 with a net of thesame dimensions as the aquarium net but having a deeper bag Net samples were decanted as describedin Cohen (1983 1987)
Additional nonquantitative ostracode samples were made in surface sediment rubble and by trapsbut these collections were used only for life-history studies and in determining range limits of speciesSampling of rubble and by traps is described in detail in Cohen (1983 1987)
Sorting Preparation and Identification of Ostracodes - The preserved samples were stirred vigorouslydrawing the lightweight ostracodes to the surface and allowing them to be decanted This procedurewas repeated until no more ostracodes were found The thoroughness of the method was checked byexamining the remaining residue of two samples not more than two to four additional ostracodeswere found in these residues Other residues were spot-checked from time to time to insure that allostracodes were being extracted Because one 1981 lagoon sample contained almost 2000 myodocopidostracodes a plankton splitter was used to divide another 1981 lagoon sample in half The numberof ostracodes extracted from that half was doubled for comparison with other samples
From the three field trips 1979-1981 only three ofthe quantitative samples taken each time in thesand and rubble zone were fully sorted because of time constraints and large sample sizes All of thesand trough transect samples (18 samples) were sorted but only three quantitative samples from eachdate (1979-1981) were chosen for quantitative comparisons on the basis of matching collecting periodsand thoroughness of sorting and identifying Samples not used were those which had originally beenlive-sorted in Belize and the residue then discarded Subsequent checking of preserved residues ofother samples 1 had believed were thoroughly live-sorted showed that ostracodes remained in theresidue
The abundance of juveniles collected throughout this study complicated species identification butalso presented an opportunity to investigate developmental aspects such as clutch and embryo sizeinstar number juvenile taxonomic characters and to some extent development time and life historypatterns Juveniles representing all myodocopid families and many species were found in the samplesLive specimens were maintained and reared as described in Cohen (1983 1987) Skogsbergia lerneriwere reared from egg to adult Rutiderma new species A were reared from egg to second instar partialmolt series were obtained from several other species
Deposition of Specimens - Material is deposited in the Los Angeles County Museum of Natural History(LACM) When taxonomic descriptions are published holotypes and paratypes will be deposited inthe LACM and additional paratypes in the Smithsonian Institution (USNM)
RESULTS
Comparison of Two Ecological Zones-LAGOON SAND AND RUBBLE ZONE (l5mdepth) Twenty-nine species and 3281 individual myodocopid ostracodes weretaken from the 10 quantitative samples (Table 1) Two measures of species di-versity for the lagoon myodocopids were calculated using one pooled sample (10samples combined) species richness H = 090 (Shannon-Weaver diversity index)and species evenness (equitability) E = 061 The most abundant species wasRutiderma dinochelatum Komicker 1958 (1045 specimens) (Fig 4 Tables 12)
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 321
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322 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 323
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324 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 2 Comparison of most abundant species in 10 combined lagoon and 10 combined sand troughsamples (1978-1981)
Lagoon Sand trough
Species (No) () (No) ()
S lerneri 2 006 190 164V new species ST 0 0 95 82V new species SA 0 0 26 22P sex 2 006 36 31P new species N 791 245 0 0P muelleri 124 38 12 10S setisparsa 28 087 31 27A americana 3 009 34 29A monambon 30 093 8 069H paucichelatus 172 53 187 161R new species A 158 49 48 41R dinochelatum 1045 324 25 22R hartmanni 88 27 I 009R new species B 104 32 21 18E new species KE 479 149 0 0E new species KG 112 35 143 123E new species R 19 059 9 077E new species P 22 068 21 18E new species B 0 0 141 121E new species MJ 0 0 31 27E new species MR 0 0 32 28E donabbotti 32 099 57 49E new species J 12 037 14 12Total 3223 100 1162 100
Also abundant were Parasterope new species N (791 specimens) and Eusarsiellanew species KE (479 specimens) (Fig 4) Two species R dinochelatum andParasterope new species N accounted for 56 of the myodocopids collected inthat zone The three most abundant species accounted for 71 of the myodocopidscollected The most abundant families in the sand and rubble zone were Ruti-dermatidae (5 species 1396 specimens) and Cylindroleberididae (8 species 983specimens all but 14 of these belonging to the Cylindroleberidinae) The Philo-medidae were represented by only 172 specimens all belonging to Harbansuspaucichelatus (Kornicker 1958) Species belonging to the Cypridinidae were veryrare in the samples (two species four specimens)
Almost half of the specimens collected in the lagoon zone belong to specieseither not present (eight species) or represented by fewer than 10 specimens inthe fore-reef samples Two additional species Rutiderma new species Land Eu-sarsiella new species MO were collected in nonquantitative sand samples fromthis zone (Table 3) but they were not found in the 10 samples used for numericalcomparisons (Tables I 2)
The largest sample (973 myodocopids June 1981) represents a density of about737 m-2 of surface sediment (973 divided by 022 6 m)
OUTERFORE-REEFSANDTROUGH(18-30 m depth) Thirty species and 1188individual myodocopid specimens were found in the 10 samples analyzed Speciesdiversity values were H = 116 E = 078 The most abundant species wasSkogsbergia lerneri (Kornicker 1958) (190 specimens) (Fig 4 Tables I 2) Alsoabundant were Harbansus paucichelatus (187) Eusarsiella new species KO (143)and E new species B (141) which together with S lerneri accounted for 55 of
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 325
11001000900800700600500400300200100
aRd PN EKE
MAJOR LAGOON SPECIES
1100~ 1000~ 9008 800Q
CIl 700~ 0gt 600
eJ ~ 500~ at bullbull00~ 300t 2002 100
a
MAJOR SAND TROUGH SPECIES
SI Hp EKD LB VST Ed RA
Figure 4 Comparison often combined samples in lagoon and ten combined samples in sand troughhistogram shows abundance of most common species in each zone (those species contributing 71-72 of the total ostracodes collected in each zone) Species shown in lagoon histogram arc R d =Rutiderma dinochelatum P N = Parasterope new species N E KE = Eusarsiella new species KESpecies shown in sand trough histogram arc S I = Skogsbergia lerneri H p = Harbansus pauci-cheallls E KG = Eusarsiella new species KO E B = E new species B V ST = Vargula n spST E d = Eusarsiella donabbotti R A = Rutiderma new species A Bar key black = Rutidermatidaeright diagonal = Cylindroleberididae left diagonal = Sarsiellidae cross-hatch = Cypridinidae open= Philomedidae
the myodocopids collected in the samples These four species along with threeothers Vargula new species ST (95 specimens) Eusarsiella donabbotti new species(57) and Rutiderma new species A (48) accounted for 72 of the myodocopidscollected in the analyzed samples The five most abundant species belong to thefamilies Cypridinidae (4 species 347 specimens) Sarsiellidae (13 species 459specimens) and Philomedidae (3 species 191 specimens) the families most abun-dant in the sand trough Specimens belonging to the other two myodocopid fam-ilies were relatively rare Rutidermatidae (6 species 98 specimens) and Cylin-droleberididae (5 species 93 specimens with only 43 of these belonging to theCylindroleberidinae)
About half of the specimens collected belong to species either not present (10)or not represented by more than 10 specimens in the lagoon samples Threeadditional species Vargula new species M Parasterope new species E and Ruti-derma new species K were found in nonquantitative samples from sand troughrubble (Table 3)
Temporal Changes in Myodocopid Fauna-LAGOON SAND AND RUBBLE ZONEThere was no change in species representation (except that rarer species did notoccur in all samples) but there was a shift in relative species family and totalabundance in collections made from January 1978 to June 1981 (Table 1Fig 5)
The Rutidermatidae was the most abundant family at the beginning (1978263specimens) and end (1981 946 specimens) of the study although the most abun-dant species in the family changed from Rutiderma new species A in January1978 to R dinochelatum in all subsequent samples Rutiderma hartmanni andR new species B increased in abundance following each sampling period
The largest sample was the single 1978 sample of 387 ostracodes Ostracodeabundance in single replicate samples declined in 1979 and increased in 1981The single 1978 sample was larger than any of the six replicate samples collectedin 1979 (Table 1) but smaller than each of the replicate samples of 1981 Fewcylindroleberids were collected in 1978 (four species five specimens in the singlesample) The Cylindroleberididae (particularly Parasterope new species N 184specimens) was the most abundant family in the June 1979 samples (8 species
326 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 3 List of myodocopid species collected only in nonquantitative samples from Belize (alsolisted other published localities)
Taxa
CypridinidaeVargula new species M
CylindroleberididaeCylindroleberidinae
Parasterope new species E
Postasterope abacoPostasterope new species BPostasterope new species X
Bruuniella new species A
Diaslerope new species D
AsteropteroninaeAsteropella maclaughlinae
Actinoseta hummelincki
RutidermatidaeAlternochelata polychelataRutiderma new species L
Rutiderma new species KE
SarsiellidaeEusarsiella new species MO
Locality (and year)
Belize only sand trough (trap and rubble 1979) lagoon patchreefs (1976) South Water Cut (1978) spur and groove(1976) outer reef slope (1979)
Belize only sand trough (rubble 1978) plankton (1976 5males) reef crest (1978)
Belize plankton (1975 I male) BahamasBelize only sand and rubble W Carrie Bow Cay (1978)Belize only South Water Cut (1977) W side Carrie Bow Cay
( 1978)Belize back-reef (1974) inner slope of outer ridge (1976) spur
and groove (1978) BermudaBelize only W side Carrie Bow Cay (1978 A-I male)
Belize Carrie Bow Cay (1978) Blue Ground (1981) W Flori-da Texas
Belize back-reef (1974 1976) spur and groove (1976 1978)Twin Cays (1977) South Water Cut (1978) Florida Vene-zuela W Panama
Belize Avicennia mangroves (I male) BahamasBelize only sand and rubble zone (1979) lagoon plankton
( 1978)Belize only sand trough (rubble 1978) outer fore-reef slope
(1979 1982) spur and groove (1979) back-reef (I 974)
Belize only sand and rubble zone (I 979) South Water Cut(I 977)
Source of localities located outside Belize POitaslerope abaca Kornicker 1986 Bruumela new species A Komicker 1981 Acrinosetamaclallghtinae Komicker 1981 (Kornicker 1986) Aclmoseta hllmmelillcki Komicker 1981 (Kornicker 1986)
253 specimens) and continued to increase in abundance subsequently thoughnot as rapidly as rutidermatids and sarsiellids
Representatives of the Sarsiellidae were most abundant in June 1981 samples(l0 species 376 specimens) and least abundant in June 1979 (8 species 70 spec-imens) The most abundant sarsiellid species in January 1978 was Eusarsiellanew species P (17) Eusarsiella new species KE and E new species KO increasedin number in subsequent samplings Three species of Eusarsiella were absent fromthe January 1978 sample but present in later samples Their absence in the 1978sample may be related to smaller sample size (single replicate) as most of thesespecies were rare in later samples Six Eusarsiella species were absent in eitherthe June or October 1979 collections but present in the June 1981 collections
The Philomedidae are represented in this zone only by Harbansus paucichelatuswhich declined in number in 1979 and increased in 1981 The Cypridinidae wererare in all samples
FORE-REEFSANDTROUGHThere was also no change in species compositionin this sample area (except that some less abundant species did not occur in allsamples) but there were shifts in relative species and family abundances between
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
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JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
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336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 317
II]Shoals
~ Sand Bores ampPatch Reefs
o km
oQ)
ltJ)
coQ)
DD 0~ 17Nu
lt~t~middotmiddotmiddot
Tobacco Range ~gt0
Figure I Map of bamer reef complex in the vicinity of Came Bow Cay area of larger map locatedon inset by rectangle [From Riitzler and Macintyre 1982 used with permission of Riitzler]
which could be easily sampled by the same method and (3) in preliminary samples many myodocopidostracodes were found in these two sites The sand and rubble sampling zone is about 15 m deepwith a bottom of poorly sorted sediment ranging in size from silt to gravel and 10-14 m thick overa hard bottom (Riitzler and Macintyre 1982) The area sampled in the sand trough is about 18-30m deep with substrate that is poorly sorted silt size to very coarse sand sediment plain This sedimentis mainly very fine to fine sand but coarser material consisting of Halimeda plates mollusks benthicforaminiferans and echinoids is scattered throughout Probings indicated that sediment is morethan 12 m in the axis of the trough (Riitzler and Macintyre 1982)
To investigate local ranges of myodocopids additional nonquantitative samples were taken from(I) other zones along the fixed reef transect line (2) the following areas in the vicinity of Came BowCay lagoon and back-reef around Came Bow Cay (sand cay) and South Water Cay (mangrove cay)south shores and a channel through Twin Cays (mangroves) South Water Cut (channel through the
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318 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 319
Figure 3 A Photo of bottom in sand and rubble zone ofIagoon depth 15 m B Photo of bottomin sand trough of outer fore-reef depth 30 m showing part of movable transect line
barrier reef) Curlew Bank (submerged cay) lagoon patch reefs and adjacent sea grass (in deep lagoonin barrier reef complex) Ragged Cay and adjacent bank (western or inner edge of barrier reef complex)and (3) on the mainland subtidal shores of northern (sand beach) and just north of (mangroves andsea grass) Dandriga Nonquantitative collections were made by the author in May 1976 April 1977January 1978 June and October 1979 and June 1981 Other ostracodes collected at the study siteand deposited in the National Museum of Natural History (USNM) were also used to complete dataon indi vidual species ranges (listed in Cohen 1987) Additional data on the local range of Skogsbergialerneri were obtained from baited traps Traps were left overnight (about 1700-0900) in the lagoon
320 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
and back-reef(one in June 1979 two in June 1981) during the day (0900-1700) in the back reef (onein June 1981) and for about 24 h (two in the lagoon and back-reef one in the spur and groove inJanuary 1978 one in the lagoon and one in the spur and groove in June 1979 one each in the lagoonback-reef spur and groove sand trough and outer ridge in October 1979 one in the back-reef in1981)
Sampling Methods -Samples were made by diving usually with SCUBA except in very shallow waterwhere collections were made while wading In depths accessible by diving this method is superior todredging because the area sampled can be chosen and the sampling monitored visually Coring wastried but found to yield very few ostracodes and therefore not used Quantitative samples in the twostudy areas were made by dragging a rectangular hand net (16 cm high 22 cm wide mesh opening02 mm) along the surface of the sediment sequentially along both sides of a movable 3-m transectline The net lightly scraped the surface sediment so that the net sampled the upper 1-2 em of sedimentand 14-15 em of water above it thus sampling the strata occupied by ostracods observed undisturbedand disturbed in dishes of sediment and seawater The net was immediately twisted to trap materialat the distal end and enclosed in a plastic bag Quantitative comparisons are based only on samplesmade with the movable transect first used in 1978 Single quantitative samples were taken in eachof the two zones in 1978 At least three quantitative samples were taken in each zone during eachsubsequent field trip (June 1979 October 1979 and June 1981) and three per zone used in analysisof those dates (see below) An aquarium net was used in 1978 but replaced in 1979 with a net of thesame dimensions as the aquarium net but having a deeper bag Net samples were decanted as describedin Cohen (1983 1987)
Additional nonquantitative ostracode samples were made in surface sediment rubble and by trapsbut these collections were used only for life-history studies and in determining range limits of speciesSampling of rubble and by traps is described in detail in Cohen (1983 1987)
Sorting Preparation and Identification of Ostracodes - The preserved samples were stirred vigorouslydrawing the lightweight ostracodes to the surface and allowing them to be decanted This procedurewas repeated until no more ostracodes were found The thoroughness of the method was checked byexamining the remaining residue of two samples not more than two to four additional ostracodeswere found in these residues Other residues were spot-checked from time to time to insure that allostracodes were being extracted Because one 1981 lagoon sample contained almost 2000 myodocopidostracodes a plankton splitter was used to divide another 1981 lagoon sample in half The numberof ostracodes extracted from that half was doubled for comparison with other samples
From the three field trips 1979-1981 only three ofthe quantitative samples taken each time in thesand and rubble zone were fully sorted because of time constraints and large sample sizes All of thesand trough transect samples (18 samples) were sorted but only three quantitative samples from eachdate (1979-1981) were chosen for quantitative comparisons on the basis of matching collecting periodsand thoroughness of sorting and identifying Samples not used were those which had originally beenlive-sorted in Belize and the residue then discarded Subsequent checking of preserved residues ofother samples 1 had believed were thoroughly live-sorted showed that ostracodes remained in theresidue
The abundance of juveniles collected throughout this study complicated species identification butalso presented an opportunity to investigate developmental aspects such as clutch and embryo sizeinstar number juvenile taxonomic characters and to some extent development time and life historypatterns Juveniles representing all myodocopid families and many species were found in the samplesLive specimens were maintained and reared as described in Cohen (1983 1987) Skogsbergia lerneriwere reared from egg to adult Rutiderma new species A were reared from egg to second instar partialmolt series were obtained from several other species
Deposition of Specimens - Material is deposited in the Los Angeles County Museum of Natural History(LACM) When taxonomic descriptions are published holotypes and paratypes will be deposited inthe LACM and additional paratypes in the Smithsonian Institution (USNM)
RESULTS
Comparison of Two Ecological Zones-LAGOON SAND AND RUBBLE ZONE (l5mdepth) Twenty-nine species and 3281 individual myodocopid ostracodes weretaken from the 10 quantitative samples (Table 1) Two measures of species di-versity for the lagoon myodocopids were calculated using one pooled sample (10samples combined) species richness H = 090 (Shannon-Weaver diversity index)and species evenness (equitability) E = 061 The most abundant species wasRutiderma dinochelatum Komicker 1958 (1045 specimens) (Fig 4 Tables 12)
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 321
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QC
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NQbull
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322 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
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0000 QQ bullbullbullbull QQ bullbullbullbullbullbull NN -- 00 00 00 00 00 QIQI 0011 bullbullbull
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 323
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324 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 2 Comparison of most abundant species in 10 combined lagoon and 10 combined sand troughsamples (1978-1981)
Lagoon Sand trough
Species (No) () (No) ()
S lerneri 2 006 190 164V new species ST 0 0 95 82V new species SA 0 0 26 22P sex 2 006 36 31P new species N 791 245 0 0P muelleri 124 38 12 10S setisparsa 28 087 31 27A americana 3 009 34 29A monambon 30 093 8 069H paucichelatus 172 53 187 161R new species A 158 49 48 41R dinochelatum 1045 324 25 22R hartmanni 88 27 I 009R new species B 104 32 21 18E new species KE 479 149 0 0E new species KG 112 35 143 123E new species R 19 059 9 077E new species P 22 068 21 18E new species B 0 0 141 121E new species MJ 0 0 31 27E new species MR 0 0 32 28E donabbotti 32 099 57 49E new species J 12 037 14 12Total 3223 100 1162 100
Also abundant were Parasterope new species N (791 specimens) and Eusarsiellanew species KE (479 specimens) (Fig 4) Two species R dinochelatum andParasterope new species N accounted for 56 of the myodocopids collected inthat zone The three most abundant species accounted for 71 of the myodocopidscollected The most abundant families in the sand and rubble zone were Ruti-dermatidae (5 species 1396 specimens) and Cylindroleberididae (8 species 983specimens all but 14 of these belonging to the Cylindroleberidinae) The Philo-medidae were represented by only 172 specimens all belonging to Harbansuspaucichelatus (Kornicker 1958) Species belonging to the Cypridinidae were veryrare in the samples (two species four specimens)
Almost half of the specimens collected in the lagoon zone belong to specieseither not present (eight species) or represented by fewer than 10 specimens inthe fore-reef samples Two additional species Rutiderma new species Land Eu-sarsiella new species MO were collected in nonquantitative sand samples fromthis zone (Table 3) but they were not found in the 10 samples used for numericalcomparisons (Tables I 2)
The largest sample (973 myodocopids June 1981) represents a density of about737 m-2 of surface sediment (973 divided by 022 6 m)
OUTERFORE-REEFSANDTROUGH(18-30 m depth) Thirty species and 1188individual myodocopid specimens were found in the 10 samples analyzed Speciesdiversity values were H = 116 E = 078 The most abundant species wasSkogsbergia lerneri (Kornicker 1958) (190 specimens) (Fig 4 Tables I 2) Alsoabundant were Harbansus paucichelatus (187) Eusarsiella new species KO (143)and E new species B (141) which together with S lerneri accounted for 55 of
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 325
11001000900800700600500400300200100
aRd PN EKE
MAJOR LAGOON SPECIES
1100~ 1000~ 9008 800Q
CIl 700~ 0gt 600
eJ ~ 500~ at bullbull00~ 300t 2002 100
a
MAJOR SAND TROUGH SPECIES
SI Hp EKD LB VST Ed RA
Figure 4 Comparison often combined samples in lagoon and ten combined samples in sand troughhistogram shows abundance of most common species in each zone (those species contributing 71-72 of the total ostracodes collected in each zone) Species shown in lagoon histogram arc R d =Rutiderma dinochelatum P N = Parasterope new species N E KE = Eusarsiella new species KESpecies shown in sand trough histogram arc S I = Skogsbergia lerneri H p = Harbansus pauci-cheallls E KG = Eusarsiella new species KO E B = E new species B V ST = Vargula n spST E d = Eusarsiella donabbotti R A = Rutiderma new species A Bar key black = Rutidermatidaeright diagonal = Cylindroleberididae left diagonal = Sarsiellidae cross-hatch = Cypridinidae open= Philomedidae
the myodocopids collected in the samples These four species along with threeothers Vargula new species ST (95 specimens) Eusarsiella donabbotti new species(57) and Rutiderma new species A (48) accounted for 72 of the myodocopidscollected in the analyzed samples The five most abundant species belong to thefamilies Cypridinidae (4 species 347 specimens) Sarsiellidae (13 species 459specimens) and Philomedidae (3 species 191 specimens) the families most abun-dant in the sand trough Specimens belonging to the other two myodocopid fam-ilies were relatively rare Rutidermatidae (6 species 98 specimens) and Cylin-droleberididae (5 species 93 specimens with only 43 of these belonging to theCylindroleberidinae)
About half of the specimens collected belong to species either not present (10)or not represented by more than 10 specimens in the lagoon samples Threeadditional species Vargula new species M Parasterope new species E and Ruti-derma new species K were found in nonquantitative samples from sand troughrubble (Table 3)
Temporal Changes in Myodocopid Fauna-LAGOON SAND AND RUBBLE ZONEThere was no change in species representation (except that rarer species did notoccur in all samples) but there was a shift in relative species family and totalabundance in collections made from January 1978 to June 1981 (Table 1Fig 5)
The Rutidermatidae was the most abundant family at the beginning (1978263specimens) and end (1981 946 specimens) of the study although the most abun-dant species in the family changed from Rutiderma new species A in January1978 to R dinochelatum in all subsequent samples Rutiderma hartmanni andR new species B increased in abundance following each sampling period
The largest sample was the single 1978 sample of 387 ostracodes Ostracodeabundance in single replicate samples declined in 1979 and increased in 1981The single 1978 sample was larger than any of the six replicate samples collectedin 1979 (Table 1) but smaller than each of the replicate samples of 1981 Fewcylindroleberids were collected in 1978 (four species five specimens in the singlesample) The Cylindroleberididae (particularly Parasterope new species N 184specimens) was the most abundant family in the June 1979 samples (8 species
326 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 3 List of myodocopid species collected only in nonquantitative samples from Belize (alsolisted other published localities)
Taxa
CypridinidaeVargula new species M
CylindroleberididaeCylindroleberidinae
Parasterope new species E
Postasterope abacoPostasterope new species BPostasterope new species X
Bruuniella new species A
Diaslerope new species D
AsteropteroninaeAsteropella maclaughlinae
Actinoseta hummelincki
RutidermatidaeAlternochelata polychelataRutiderma new species L
Rutiderma new species KE
SarsiellidaeEusarsiella new species MO
Locality (and year)
Belize only sand trough (trap and rubble 1979) lagoon patchreefs (1976) South Water Cut (1978) spur and groove(1976) outer reef slope (1979)
Belize only sand trough (rubble 1978) plankton (1976 5males) reef crest (1978)
Belize plankton (1975 I male) BahamasBelize only sand and rubble W Carrie Bow Cay (1978)Belize only South Water Cut (1977) W side Carrie Bow Cay
( 1978)Belize back-reef (1974) inner slope of outer ridge (1976) spur
and groove (1978) BermudaBelize only W side Carrie Bow Cay (1978 A-I male)
Belize Carrie Bow Cay (1978) Blue Ground (1981) W Flori-da Texas
Belize back-reef (1974 1976) spur and groove (1976 1978)Twin Cays (1977) South Water Cut (1978) Florida Vene-zuela W Panama
Belize Avicennia mangroves (I male) BahamasBelize only sand and rubble zone (1979) lagoon plankton
( 1978)Belize only sand trough (rubble 1978) outer fore-reef slope
(1979 1982) spur and groove (1979) back-reef (I 974)
Belize only sand and rubble zone (I 979) South Water Cut(I 977)
Source of localities located outside Belize POitaslerope abaca Kornicker 1986 Bruumela new species A Komicker 1981 Acrinosetamaclallghtinae Komicker 1981 (Kornicker 1986) Aclmoseta hllmmelillcki Komicker 1981 (Kornicker 1986)
253 specimens) and continued to increase in abundance subsequently thoughnot as rapidly as rutidermatids and sarsiellids
Representatives of the Sarsiellidae were most abundant in June 1981 samples(l0 species 376 specimens) and least abundant in June 1979 (8 species 70 spec-imens) The most abundant sarsiellid species in January 1978 was Eusarsiellanew species P (17) Eusarsiella new species KE and E new species KO increasedin number in subsequent samplings Three species of Eusarsiella were absent fromthe January 1978 sample but present in later samples Their absence in the 1978sample may be related to smaller sample size (single replicate) as most of thesespecies were rare in later samples Six Eusarsiella species were absent in eitherthe June or October 1979 collections but present in the June 1981 collections
The Philomedidae are represented in this zone only by Harbansus paucichelatuswhich declined in number in 1979 and increased in 1981 The Cypridinidae wererare in all samples
FORE-REEFSANDTROUGHThere was also no change in species compositionin this sample area (except that some less abundant species did not occur in allsamples) but there were shifts in relative species and family abundances between
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
ramllv
JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
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336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
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--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 319
Figure 3 A Photo of bottom in sand and rubble zone ofIagoon depth 15 m B Photo of bottomin sand trough of outer fore-reef depth 30 m showing part of movable transect line
barrier reef) Curlew Bank (submerged cay) lagoon patch reefs and adjacent sea grass (in deep lagoonin barrier reef complex) Ragged Cay and adjacent bank (western or inner edge of barrier reef complex)and (3) on the mainland subtidal shores of northern (sand beach) and just north of (mangroves andsea grass) Dandriga Nonquantitative collections were made by the author in May 1976 April 1977January 1978 June and October 1979 and June 1981 Other ostracodes collected at the study siteand deposited in the National Museum of Natural History (USNM) were also used to complete dataon indi vidual species ranges (listed in Cohen 1987) Additional data on the local range of Skogsbergialerneri were obtained from baited traps Traps were left overnight (about 1700-0900) in the lagoon
320 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
and back-reef(one in June 1979 two in June 1981) during the day (0900-1700) in the back reef (onein June 1981) and for about 24 h (two in the lagoon and back-reef one in the spur and groove inJanuary 1978 one in the lagoon and one in the spur and groove in June 1979 one each in the lagoonback-reef spur and groove sand trough and outer ridge in October 1979 one in the back-reef in1981)
Sampling Methods -Samples were made by diving usually with SCUBA except in very shallow waterwhere collections were made while wading In depths accessible by diving this method is superior todredging because the area sampled can be chosen and the sampling monitored visually Coring wastried but found to yield very few ostracodes and therefore not used Quantitative samples in the twostudy areas were made by dragging a rectangular hand net (16 cm high 22 cm wide mesh opening02 mm) along the surface of the sediment sequentially along both sides of a movable 3-m transectline The net lightly scraped the surface sediment so that the net sampled the upper 1-2 em of sedimentand 14-15 em of water above it thus sampling the strata occupied by ostracods observed undisturbedand disturbed in dishes of sediment and seawater The net was immediately twisted to trap materialat the distal end and enclosed in a plastic bag Quantitative comparisons are based only on samplesmade with the movable transect first used in 1978 Single quantitative samples were taken in eachof the two zones in 1978 At least three quantitative samples were taken in each zone during eachsubsequent field trip (June 1979 October 1979 and June 1981) and three per zone used in analysisof those dates (see below) An aquarium net was used in 1978 but replaced in 1979 with a net of thesame dimensions as the aquarium net but having a deeper bag Net samples were decanted as describedin Cohen (1983 1987)
Additional nonquantitative ostracode samples were made in surface sediment rubble and by trapsbut these collections were used only for life-history studies and in determining range limits of speciesSampling of rubble and by traps is described in detail in Cohen (1983 1987)
Sorting Preparation and Identification of Ostracodes - The preserved samples were stirred vigorouslydrawing the lightweight ostracodes to the surface and allowing them to be decanted This procedurewas repeated until no more ostracodes were found The thoroughness of the method was checked byexamining the remaining residue of two samples not more than two to four additional ostracodeswere found in these residues Other residues were spot-checked from time to time to insure that allostracodes were being extracted Because one 1981 lagoon sample contained almost 2000 myodocopidostracodes a plankton splitter was used to divide another 1981 lagoon sample in half The numberof ostracodes extracted from that half was doubled for comparison with other samples
From the three field trips 1979-1981 only three ofthe quantitative samples taken each time in thesand and rubble zone were fully sorted because of time constraints and large sample sizes All of thesand trough transect samples (18 samples) were sorted but only three quantitative samples from eachdate (1979-1981) were chosen for quantitative comparisons on the basis of matching collecting periodsand thoroughness of sorting and identifying Samples not used were those which had originally beenlive-sorted in Belize and the residue then discarded Subsequent checking of preserved residues ofother samples 1 had believed were thoroughly live-sorted showed that ostracodes remained in theresidue
The abundance of juveniles collected throughout this study complicated species identification butalso presented an opportunity to investigate developmental aspects such as clutch and embryo sizeinstar number juvenile taxonomic characters and to some extent development time and life historypatterns Juveniles representing all myodocopid families and many species were found in the samplesLive specimens were maintained and reared as described in Cohen (1983 1987) Skogsbergia lerneriwere reared from egg to adult Rutiderma new species A were reared from egg to second instar partialmolt series were obtained from several other species
Deposition of Specimens - Material is deposited in the Los Angeles County Museum of Natural History(LACM) When taxonomic descriptions are published holotypes and paratypes will be deposited inthe LACM and additional paratypes in the Smithsonian Institution (USNM)
RESULTS
Comparison of Two Ecological Zones-LAGOON SAND AND RUBBLE ZONE (l5mdepth) Twenty-nine species and 3281 individual myodocopid ostracodes weretaken from the 10 quantitative samples (Table 1) Two measures of species di-versity for the lagoon myodocopids were calculated using one pooled sample (10samples combined) species richness H = 090 (Shannon-Weaver diversity index)and species evenness (equitability) E = 061 The most abundant species wasRutiderma dinochelatum Komicker 1958 (1045 specimens) (Fig 4 Tables 12)
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 321
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322 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
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324 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 2 Comparison of most abundant species in 10 combined lagoon and 10 combined sand troughsamples (1978-1981)
Lagoon Sand trough
Species (No) () (No) ()
S lerneri 2 006 190 164V new species ST 0 0 95 82V new species SA 0 0 26 22P sex 2 006 36 31P new species N 791 245 0 0P muelleri 124 38 12 10S setisparsa 28 087 31 27A americana 3 009 34 29A monambon 30 093 8 069H paucichelatus 172 53 187 161R new species A 158 49 48 41R dinochelatum 1045 324 25 22R hartmanni 88 27 I 009R new species B 104 32 21 18E new species KE 479 149 0 0E new species KG 112 35 143 123E new species R 19 059 9 077E new species P 22 068 21 18E new species B 0 0 141 121E new species MJ 0 0 31 27E new species MR 0 0 32 28E donabbotti 32 099 57 49E new species J 12 037 14 12Total 3223 100 1162 100
Also abundant were Parasterope new species N (791 specimens) and Eusarsiellanew species KE (479 specimens) (Fig 4) Two species R dinochelatum andParasterope new species N accounted for 56 of the myodocopids collected inthat zone The three most abundant species accounted for 71 of the myodocopidscollected The most abundant families in the sand and rubble zone were Ruti-dermatidae (5 species 1396 specimens) and Cylindroleberididae (8 species 983specimens all but 14 of these belonging to the Cylindroleberidinae) The Philo-medidae were represented by only 172 specimens all belonging to Harbansuspaucichelatus (Kornicker 1958) Species belonging to the Cypridinidae were veryrare in the samples (two species four specimens)
Almost half of the specimens collected in the lagoon zone belong to specieseither not present (eight species) or represented by fewer than 10 specimens inthe fore-reef samples Two additional species Rutiderma new species Land Eu-sarsiella new species MO were collected in nonquantitative sand samples fromthis zone (Table 3) but they were not found in the 10 samples used for numericalcomparisons (Tables I 2)
The largest sample (973 myodocopids June 1981) represents a density of about737 m-2 of surface sediment (973 divided by 022 6 m)
OUTERFORE-REEFSANDTROUGH(18-30 m depth) Thirty species and 1188individual myodocopid specimens were found in the 10 samples analyzed Speciesdiversity values were H = 116 E = 078 The most abundant species wasSkogsbergia lerneri (Kornicker 1958) (190 specimens) (Fig 4 Tables I 2) Alsoabundant were Harbansus paucichelatus (187) Eusarsiella new species KO (143)and E new species B (141) which together with S lerneri accounted for 55 of
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 325
11001000900800700600500400300200100
aRd PN EKE
MAJOR LAGOON SPECIES
1100~ 1000~ 9008 800Q
CIl 700~ 0gt 600
eJ ~ 500~ at bullbull00~ 300t 2002 100
a
MAJOR SAND TROUGH SPECIES
SI Hp EKD LB VST Ed RA
Figure 4 Comparison often combined samples in lagoon and ten combined samples in sand troughhistogram shows abundance of most common species in each zone (those species contributing 71-72 of the total ostracodes collected in each zone) Species shown in lagoon histogram arc R d =Rutiderma dinochelatum P N = Parasterope new species N E KE = Eusarsiella new species KESpecies shown in sand trough histogram arc S I = Skogsbergia lerneri H p = Harbansus pauci-cheallls E KG = Eusarsiella new species KO E B = E new species B V ST = Vargula n spST E d = Eusarsiella donabbotti R A = Rutiderma new species A Bar key black = Rutidermatidaeright diagonal = Cylindroleberididae left diagonal = Sarsiellidae cross-hatch = Cypridinidae open= Philomedidae
the myodocopids collected in the samples These four species along with threeothers Vargula new species ST (95 specimens) Eusarsiella donabbotti new species(57) and Rutiderma new species A (48) accounted for 72 of the myodocopidscollected in the analyzed samples The five most abundant species belong to thefamilies Cypridinidae (4 species 347 specimens) Sarsiellidae (13 species 459specimens) and Philomedidae (3 species 191 specimens) the families most abun-dant in the sand trough Specimens belonging to the other two myodocopid fam-ilies were relatively rare Rutidermatidae (6 species 98 specimens) and Cylin-droleberididae (5 species 93 specimens with only 43 of these belonging to theCylindroleberidinae)
About half of the specimens collected belong to species either not present (10)or not represented by more than 10 specimens in the lagoon samples Threeadditional species Vargula new species M Parasterope new species E and Ruti-derma new species K were found in nonquantitative samples from sand troughrubble (Table 3)
Temporal Changes in Myodocopid Fauna-LAGOON SAND AND RUBBLE ZONEThere was no change in species representation (except that rarer species did notoccur in all samples) but there was a shift in relative species family and totalabundance in collections made from January 1978 to June 1981 (Table 1Fig 5)
The Rutidermatidae was the most abundant family at the beginning (1978263specimens) and end (1981 946 specimens) of the study although the most abun-dant species in the family changed from Rutiderma new species A in January1978 to R dinochelatum in all subsequent samples Rutiderma hartmanni andR new species B increased in abundance following each sampling period
The largest sample was the single 1978 sample of 387 ostracodes Ostracodeabundance in single replicate samples declined in 1979 and increased in 1981The single 1978 sample was larger than any of the six replicate samples collectedin 1979 (Table 1) but smaller than each of the replicate samples of 1981 Fewcylindroleberids were collected in 1978 (four species five specimens in the singlesample) The Cylindroleberididae (particularly Parasterope new species N 184specimens) was the most abundant family in the June 1979 samples (8 species
326 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 3 List of myodocopid species collected only in nonquantitative samples from Belize (alsolisted other published localities)
Taxa
CypridinidaeVargula new species M
CylindroleberididaeCylindroleberidinae
Parasterope new species E
Postasterope abacoPostasterope new species BPostasterope new species X
Bruuniella new species A
Diaslerope new species D
AsteropteroninaeAsteropella maclaughlinae
Actinoseta hummelincki
RutidermatidaeAlternochelata polychelataRutiderma new species L
Rutiderma new species KE
SarsiellidaeEusarsiella new species MO
Locality (and year)
Belize only sand trough (trap and rubble 1979) lagoon patchreefs (1976) South Water Cut (1978) spur and groove(1976) outer reef slope (1979)
Belize only sand trough (rubble 1978) plankton (1976 5males) reef crest (1978)
Belize plankton (1975 I male) BahamasBelize only sand and rubble W Carrie Bow Cay (1978)Belize only South Water Cut (1977) W side Carrie Bow Cay
( 1978)Belize back-reef (1974) inner slope of outer ridge (1976) spur
and groove (1978) BermudaBelize only W side Carrie Bow Cay (1978 A-I male)
Belize Carrie Bow Cay (1978) Blue Ground (1981) W Flori-da Texas
Belize back-reef (1974 1976) spur and groove (1976 1978)Twin Cays (1977) South Water Cut (1978) Florida Vene-zuela W Panama
Belize Avicennia mangroves (I male) BahamasBelize only sand and rubble zone (1979) lagoon plankton
( 1978)Belize only sand trough (rubble 1978) outer fore-reef slope
(1979 1982) spur and groove (1979) back-reef (I 974)
Belize only sand and rubble zone (I 979) South Water Cut(I 977)
Source of localities located outside Belize POitaslerope abaca Kornicker 1986 Bruumela new species A Komicker 1981 Acrinosetamaclallghtinae Komicker 1981 (Kornicker 1986) Aclmoseta hllmmelillcki Komicker 1981 (Kornicker 1986)
253 specimens) and continued to increase in abundance subsequently thoughnot as rapidly as rutidermatids and sarsiellids
Representatives of the Sarsiellidae were most abundant in June 1981 samples(l0 species 376 specimens) and least abundant in June 1979 (8 species 70 spec-imens) The most abundant sarsiellid species in January 1978 was Eusarsiellanew species P (17) Eusarsiella new species KE and E new species KO increasedin number in subsequent samplings Three species of Eusarsiella were absent fromthe January 1978 sample but present in later samples Their absence in the 1978sample may be related to smaller sample size (single replicate) as most of thesespecies were rare in later samples Six Eusarsiella species were absent in eitherthe June or October 1979 collections but present in the June 1981 collections
The Philomedidae are represented in this zone only by Harbansus paucichelatuswhich declined in number in 1979 and increased in 1981 The Cypridinidae wererare in all samples
FORE-REEFSANDTROUGHThere was also no change in species compositionin this sample area (except that some less abundant species did not occur in allsamples) but there were shifts in relative species and family abundances between
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
ramllv
JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
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Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 319
Figure 3 A Photo of bottom in sand and rubble zone ofIagoon depth 15 m B Photo of bottomin sand trough of outer fore-reef depth 30 m showing part of movable transect line
barrier reef) Curlew Bank (submerged cay) lagoon patch reefs and adjacent sea grass (in deep lagoonin barrier reef complex) Ragged Cay and adjacent bank (western or inner edge of barrier reef complex)and (3) on the mainland subtidal shores of northern (sand beach) and just north of (mangroves andsea grass) Dandriga Nonquantitative collections were made by the author in May 1976 April 1977January 1978 June and October 1979 and June 1981 Other ostracodes collected at the study siteand deposited in the National Museum of Natural History (USNM) were also used to complete dataon indi vidual species ranges (listed in Cohen 1987) Additional data on the local range of Skogsbergialerneri were obtained from baited traps Traps were left overnight (about 1700-0900) in the lagoon
320 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
and back-reef(one in June 1979 two in June 1981) during the day (0900-1700) in the back reef (onein June 1981) and for about 24 h (two in the lagoon and back-reef one in the spur and groove inJanuary 1978 one in the lagoon and one in the spur and groove in June 1979 one each in the lagoonback-reef spur and groove sand trough and outer ridge in October 1979 one in the back-reef in1981)
Sampling Methods -Samples were made by diving usually with SCUBA except in very shallow waterwhere collections were made while wading In depths accessible by diving this method is superior todredging because the area sampled can be chosen and the sampling monitored visually Coring wastried but found to yield very few ostracodes and therefore not used Quantitative samples in the twostudy areas were made by dragging a rectangular hand net (16 cm high 22 cm wide mesh opening02 mm) along the surface of the sediment sequentially along both sides of a movable 3-m transectline The net lightly scraped the surface sediment so that the net sampled the upper 1-2 em of sedimentand 14-15 em of water above it thus sampling the strata occupied by ostracods observed undisturbedand disturbed in dishes of sediment and seawater The net was immediately twisted to trap materialat the distal end and enclosed in a plastic bag Quantitative comparisons are based only on samplesmade with the movable transect first used in 1978 Single quantitative samples were taken in eachof the two zones in 1978 At least three quantitative samples were taken in each zone during eachsubsequent field trip (June 1979 October 1979 and June 1981) and three per zone used in analysisof those dates (see below) An aquarium net was used in 1978 but replaced in 1979 with a net of thesame dimensions as the aquarium net but having a deeper bag Net samples were decanted as describedin Cohen (1983 1987)
Additional nonquantitative ostracode samples were made in surface sediment rubble and by trapsbut these collections were used only for life-history studies and in determining range limits of speciesSampling of rubble and by traps is described in detail in Cohen (1983 1987)
Sorting Preparation and Identification of Ostracodes - The preserved samples were stirred vigorouslydrawing the lightweight ostracodes to the surface and allowing them to be decanted This procedurewas repeated until no more ostracodes were found The thoroughness of the method was checked byexamining the remaining residue of two samples not more than two to four additional ostracodeswere found in these residues Other residues were spot-checked from time to time to insure that allostracodes were being extracted Because one 1981 lagoon sample contained almost 2000 myodocopidostracodes a plankton splitter was used to divide another 1981 lagoon sample in half The numberof ostracodes extracted from that half was doubled for comparison with other samples
From the three field trips 1979-1981 only three ofthe quantitative samples taken each time in thesand and rubble zone were fully sorted because of time constraints and large sample sizes All of thesand trough transect samples (18 samples) were sorted but only three quantitative samples from eachdate (1979-1981) were chosen for quantitative comparisons on the basis of matching collecting periodsand thoroughness of sorting and identifying Samples not used were those which had originally beenlive-sorted in Belize and the residue then discarded Subsequent checking of preserved residues ofother samples 1 had believed were thoroughly live-sorted showed that ostracodes remained in theresidue
The abundance of juveniles collected throughout this study complicated species identification butalso presented an opportunity to investigate developmental aspects such as clutch and embryo sizeinstar number juvenile taxonomic characters and to some extent development time and life historypatterns Juveniles representing all myodocopid families and many species were found in the samplesLive specimens were maintained and reared as described in Cohen (1983 1987) Skogsbergia lerneriwere reared from egg to adult Rutiderma new species A were reared from egg to second instar partialmolt series were obtained from several other species
Deposition of Specimens - Material is deposited in the Los Angeles County Museum of Natural History(LACM) When taxonomic descriptions are published holotypes and paratypes will be deposited inthe LACM and additional paratypes in the Smithsonian Institution (USNM)
RESULTS
Comparison of Two Ecological Zones-LAGOON SAND AND RUBBLE ZONE (l5mdepth) Twenty-nine species and 3281 individual myodocopid ostracodes weretaken from the 10 quantitative samples (Table 1) Two measures of species di-versity for the lagoon myodocopids were calculated using one pooled sample (10samples combined) species richness H = 090 (Shannon-Weaver diversity index)and species evenness (equitability) E = 061 The most abundant species wasRutiderma dinochelatum Komicker 1958 (1045 specimens) (Fig 4 Tables 12)
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 321
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00lt11 00+ +00 000000 CIO 0-bullbullbullbullbull fI) -(I)
bull
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NQbull
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bullbullbullbullbullbull 00 VoVo 00 00 NN NN 00 -- -- 00 00 00 00 00NN
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322 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
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+ bullbullbullI-lIl
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+
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bullbull
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o+N
+
5o
0000 QQ bullbullbullbull QQ bullbullbullbullbullbull NN -- 00 00 00 00 00 QIQI 0011 bullbullbull
o+0o+o
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+o
+IIl
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t-
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+ltt
+
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+ +o
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Nbullbullbull
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 323
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+ +o
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+
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00
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324 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 2 Comparison of most abundant species in 10 combined lagoon and 10 combined sand troughsamples (1978-1981)
Lagoon Sand trough
Species (No) () (No) ()
S lerneri 2 006 190 164V new species ST 0 0 95 82V new species SA 0 0 26 22P sex 2 006 36 31P new species N 791 245 0 0P muelleri 124 38 12 10S setisparsa 28 087 31 27A americana 3 009 34 29A monambon 30 093 8 069H paucichelatus 172 53 187 161R new species A 158 49 48 41R dinochelatum 1045 324 25 22R hartmanni 88 27 I 009R new species B 104 32 21 18E new species KE 479 149 0 0E new species KG 112 35 143 123E new species R 19 059 9 077E new species P 22 068 21 18E new species B 0 0 141 121E new species MJ 0 0 31 27E new species MR 0 0 32 28E donabbotti 32 099 57 49E new species J 12 037 14 12Total 3223 100 1162 100
Also abundant were Parasterope new species N (791 specimens) and Eusarsiellanew species KE (479 specimens) (Fig 4) Two species R dinochelatum andParasterope new species N accounted for 56 of the myodocopids collected inthat zone The three most abundant species accounted for 71 of the myodocopidscollected The most abundant families in the sand and rubble zone were Ruti-dermatidae (5 species 1396 specimens) and Cylindroleberididae (8 species 983specimens all but 14 of these belonging to the Cylindroleberidinae) The Philo-medidae were represented by only 172 specimens all belonging to Harbansuspaucichelatus (Kornicker 1958) Species belonging to the Cypridinidae were veryrare in the samples (two species four specimens)
Almost half of the specimens collected in the lagoon zone belong to specieseither not present (eight species) or represented by fewer than 10 specimens inthe fore-reef samples Two additional species Rutiderma new species Land Eu-sarsiella new species MO were collected in nonquantitative sand samples fromthis zone (Table 3) but they were not found in the 10 samples used for numericalcomparisons (Tables I 2)
The largest sample (973 myodocopids June 1981) represents a density of about737 m-2 of surface sediment (973 divided by 022 6 m)
OUTERFORE-REEFSANDTROUGH(18-30 m depth) Thirty species and 1188individual myodocopid specimens were found in the 10 samples analyzed Speciesdiversity values were H = 116 E = 078 The most abundant species wasSkogsbergia lerneri (Kornicker 1958) (190 specimens) (Fig 4 Tables I 2) Alsoabundant were Harbansus paucichelatus (187) Eusarsiella new species KO (143)and E new species B (141) which together with S lerneri accounted for 55 of
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 325
11001000900800700600500400300200100
aRd PN EKE
MAJOR LAGOON SPECIES
1100~ 1000~ 9008 800Q
CIl 700~ 0gt 600
eJ ~ 500~ at bullbull00~ 300t 2002 100
a
MAJOR SAND TROUGH SPECIES
SI Hp EKD LB VST Ed RA
Figure 4 Comparison often combined samples in lagoon and ten combined samples in sand troughhistogram shows abundance of most common species in each zone (those species contributing 71-72 of the total ostracodes collected in each zone) Species shown in lagoon histogram arc R d =Rutiderma dinochelatum P N = Parasterope new species N E KE = Eusarsiella new species KESpecies shown in sand trough histogram arc S I = Skogsbergia lerneri H p = Harbansus pauci-cheallls E KG = Eusarsiella new species KO E B = E new species B V ST = Vargula n spST E d = Eusarsiella donabbotti R A = Rutiderma new species A Bar key black = Rutidermatidaeright diagonal = Cylindroleberididae left diagonal = Sarsiellidae cross-hatch = Cypridinidae open= Philomedidae
the myodocopids collected in the samples These four species along with threeothers Vargula new species ST (95 specimens) Eusarsiella donabbotti new species(57) and Rutiderma new species A (48) accounted for 72 of the myodocopidscollected in the analyzed samples The five most abundant species belong to thefamilies Cypridinidae (4 species 347 specimens) Sarsiellidae (13 species 459specimens) and Philomedidae (3 species 191 specimens) the families most abun-dant in the sand trough Specimens belonging to the other two myodocopid fam-ilies were relatively rare Rutidermatidae (6 species 98 specimens) and Cylin-droleberididae (5 species 93 specimens with only 43 of these belonging to theCylindroleberidinae)
About half of the specimens collected belong to species either not present (10)or not represented by more than 10 specimens in the lagoon samples Threeadditional species Vargula new species M Parasterope new species E and Ruti-derma new species K were found in nonquantitative samples from sand troughrubble (Table 3)
Temporal Changes in Myodocopid Fauna-LAGOON SAND AND RUBBLE ZONEThere was no change in species representation (except that rarer species did notoccur in all samples) but there was a shift in relative species family and totalabundance in collections made from January 1978 to June 1981 (Table 1Fig 5)
The Rutidermatidae was the most abundant family at the beginning (1978263specimens) and end (1981 946 specimens) of the study although the most abun-dant species in the family changed from Rutiderma new species A in January1978 to R dinochelatum in all subsequent samples Rutiderma hartmanni andR new species B increased in abundance following each sampling period
The largest sample was the single 1978 sample of 387 ostracodes Ostracodeabundance in single replicate samples declined in 1979 and increased in 1981The single 1978 sample was larger than any of the six replicate samples collectedin 1979 (Table 1) but smaller than each of the replicate samples of 1981 Fewcylindroleberids were collected in 1978 (four species five specimens in the singlesample) The Cylindroleberididae (particularly Parasterope new species N 184specimens) was the most abundant family in the June 1979 samples (8 species
326 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 3 List of myodocopid species collected only in nonquantitative samples from Belize (alsolisted other published localities)
Taxa
CypridinidaeVargula new species M
CylindroleberididaeCylindroleberidinae
Parasterope new species E
Postasterope abacoPostasterope new species BPostasterope new species X
Bruuniella new species A
Diaslerope new species D
AsteropteroninaeAsteropella maclaughlinae
Actinoseta hummelincki
RutidermatidaeAlternochelata polychelataRutiderma new species L
Rutiderma new species KE
SarsiellidaeEusarsiella new species MO
Locality (and year)
Belize only sand trough (trap and rubble 1979) lagoon patchreefs (1976) South Water Cut (1978) spur and groove(1976) outer reef slope (1979)
Belize only sand trough (rubble 1978) plankton (1976 5males) reef crest (1978)
Belize plankton (1975 I male) BahamasBelize only sand and rubble W Carrie Bow Cay (1978)Belize only South Water Cut (1977) W side Carrie Bow Cay
( 1978)Belize back-reef (1974) inner slope of outer ridge (1976) spur
and groove (1978) BermudaBelize only W side Carrie Bow Cay (1978 A-I male)
Belize Carrie Bow Cay (1978) Blue Ground (1981) W Flori-da Texas
Belize back-reef (1974 1976) spur and groove (1976 1978)Twin Cays (1977) South Water Cut (1978) Florida Vene-zuela W Panama
Belize Avicennia mangroves (I male) BahamasBelize only sand and rubble zone (1979) lagoon plankton
( 1978)Belize only sand trough (rubble 1978) outer fore-reef slope
(1979 1982) spur and groove (1979) back-reef (I 974)
Belize only sand and rubble zone (I 979) South Water Cut(I 977)
Source of localities located outside Belize POitaslerope abaca Kornicker 1986 Bruumela new species A Komicker 1981 Acrinosetamaclallghtinae Komicker 1981 (Kornicker 1986) Aclmoseta hllmmelillcki Komicker 1981 (Kornicker 1986)
253 specimens) and continued to increase in abundance subsequently thoughnot as rapidly as rutidermatids and sarsiellids
Representatives of the Sarsiellidae were most abundant in June 1981 samples(l0 species 376 specimens) and least abundant in June 1979 (8 species 70 spec-imens) The most abundant sarsiellid species in January 1978 was Eusarsiellanew species P (17) Eusarsiella new species KE and E new species KO increasedin number in subsequent samplings Three species of Eusarsiella were absent fromthe January 1978 sample but present in later samples Their absence in the 1978sample may be related to smaller sample size (single replicate) as most of thesespecies were rare in later samples Six Eusarsiella species were absent in eitherthe June or October 1979 collections but present in the June 1981 collections
The Philomedidae are represented in this zone only by Harbansus paucichelatuswhich declined in number in 1979 and increased in 1981 The Cypridinidae wererare in all samples
FORE-REEFSANDTROUGHThere was also no change in species compositionin this sample area (except that some less abundant species did not occur in allsamples) but there were shifts in relative species and family abundances between
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
ramllv
JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
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Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
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336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
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(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
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Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
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--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
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-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
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Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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320 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
and back-reef(one in June 1979 two in June 1981) during the day (0900-1700) in the back reef (onein June 1981) and for about 24 h (two in the lagoon and back-reef one in the spur and groove inJanuary 1978 one in the lagoon and one in the spur and groove in June 1979 one each in the lagoonback-reef spur and groove sand trough and outer ridge in October 1979 one in the back-reef in1981)
Sampling Methods -Samples were made by diving usually with SCUBA except in very shallow waterwhere collections were made while wading In depths accessible by diving this method is superior todredging because the area sampled can be chosen and the sampling monitored visually Coring wastried but found to yield very few ostracodes and therefore not used Quantitative samples in the twostudy areas were made by dragging a rectangular hand net (16 cm high 22 cm wide mesh opening02 mm) along the surface of the sediment sequentially along both sides of a movable 3-m transectline The net lightly scraped the surface sediment so that the net sampled the upper 1-2 em of sedimentand 14-15 em of water above it thus sampling the strata occupied by ostracods observed undisturbedand disturbed in dishes of sediment and seawater The net was immediately twisted to trap materialat the distal end and enclosed in a plastic bag Quantitative comparisons are based only on samplesmade with the movable transect first used in 1978 Single quantitative samples were taken in eachof the two zones in 1978 At least three quantitative samples were taken in each zone during eachsubsequent field trip (June 1979 October 1979 and June 1981) and three per zone used in analysisof those dates (see below) An aquarium net was used in 1978 but replaced in 1979 with a net of thesame dimensions as the aquarium net but having a deeper bag Net samples were decanted as describedin Cohen (1983 1987)
Additional nonquantitative ostracode samples were made in surface sediment rubble and by trapsbut these collections were used only for life-history studies and in determining range limits of speciesSampling of rubble and by traps is described in detail in Cohen (1983 1987)
Sorting Preparation and Identification of Ostracodes - The preserved samples were stirred vigorouslydrawing the lightweight ostracodes to the surface and allowing them to be decanted This procedurewas repeated until no more ostracodes were found The thoroughness of the method was checked byexamining the remaining residue of two samples not more than two to four additional ostracodeswere found in these residues Other residues were spot-checked from time to time to insure that allostracodes were being extracted Because one 1981 lagoon sample contained almost 2000 myodocopidostracodes a plankton splitter was used to divide another 1981 lagoon sample in half The numberof ostracodes extracted from that half was doubled for comparison with other samples
From the three field trips 1979-1981 only three ofthe quantitative samples taken each time in thesand and rubble zone were fully sorted because of time constraints and large sample sizes All of thesand trough transect samples (18 samples) were sorted but only three quantitative samples from eachdate (1979-1981) were chosen for quantitative comparisons on the basis of matching collecting periodsand thoroughness of sorting and identifying Samples not used were those which had originally beenlive-sorted in Belize and the residue then discarded Subsequent checking of preserved residues ofother samples 1 had believed were thoroughly live-sorted showed that ostracodes remained in theresidue
The abundance of juveniles collected throughout this study complicated species identification butalso presented an opportunity to investigate developmental aspects such as clutch and embryo sizeinstar number juvenile taxonomic characters and to some extent development time and life historypatterns Juveniles representing all myodocopid families and many species were found in the samplesLive specimens were maintained and reared as described in Cohen (1983 1987) Skogsbergia lerneriwere reared from egg to adult Rutiderma new species A were reared from egg to second instar partialmolt series were obtained from several other species
Deposition of Specimens - Material is deposited in the Los Angeles County Museum of Natural History(LACM) When taxonomic descriptions are published holotypes and paratypes will be deposited inthe LACM and additional paratypes in the Smithsonian Institution (USNM)
RESULTS
Comparison of Two Ecological Zones-LAGOON SAND AND RUBBLE ZONE (l5mdepth) Twenty-nine species and 3281 individual myodocopid ostracodes weretaken from the 10 quantitative samples (Table 1) Two measures of species di-versity for the lagoon myodocopids were calculated using one pooled sample (10samples combined) species richness H = 090 (Shannon-Weaver diversity index)and species evenness (equitability) E = 061 The most abundant species wasRutiderma dinochelatum Komicker 1958 (1045 specimens) (Fig 4 Tables 12)
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 321
Q
+Q
+
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N
+NQ
+Q
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degC ~ltIIbullbull oscQ~-abullbull ltIIVi bullbull-lt
bull
bull
bull+QC+
QC
+lt11N
+
+
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+o
+
+
+
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IDbullbull0--~ -M
o+0o+o
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+ +o
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o+0o+o+
0gt
+0
o+N
+
+
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+o+
o
+lt1
N
+lt1_ltI
+o
lt1
+lt100+o
o
+ lt1+0 _ltI
o+0o+o
N
+O~+N
+00gt 0ltI
+0
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o
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+~~-N+o
+ IIIQC bullbullbull+IC
00lt11 00+ +00 000000 CIO 0-bullbullbullbullbull fI) -(I)
bull
bull
bull
+
+ICIIlltIIN
+QIII
+bull
+NbullbullbullN
bull
+Ill bullbullbullbull +
NQbull
IIIbull
Q bullbullbullN
+QCQbull
N
+0+r--
o+0o+o
o+0o+o
N Q
+ +ltI+IIl+Nr-- N 00 _ N bullbullbull
+ + + +
o+0o+o
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+00
o+0o+o
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+ +o Volt1
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bullbullbullbullbullbull 00 VoVo 00 00 NN NN 00 -- -- 00 00 00 00 00NN
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bullbull
ICN
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322 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
-1--Co-~-+lttN
+
N
+No+o
c
+QIQ
+bullbullbull
Q
+Q
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+Q
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+ bullbullbullI-lIl
+N
+t-N-+ltt
o+0o+o
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o+0o+o
Q+c1Ilbullbullbullbull
t-
+ bullbull+1Il~ 0
+ +I- 0IIl
+
0+ +0o 0
+ +o 0
o+0o+o
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+o+o
o+0o+o
o+0o+oo
+ +o
+ltgt
+N~
+bullbullbull
t-
+cbullbullbullN
Q
+Q
Q
+ bullbullN
+ltgt
Q
+Q
+QIQIbullbullbull 1-
+
QI
bullbull
+o+o
+N
+N
+o+o
o+N
+
5o
0000 QQ bullbullbullbull QQ bullbullbullbullbullbull NN -- 00 00 00 00 00 QIQI 0011 bullbullbull
o+0o+o
o+0o+o
+o
+IIl
o+ +~~+
t-
O ++0o+o
o+0o+o
o+0o+o
+ltt
+
o
+o
o+0o+o
0
+N_11
+
+N
o++11 0
+ +o
o+0o+o
o+0o+o
o+0o+o
t-11
+0lttoo+N
11
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QC)bullbullbull+bullbullbullbull1IlC bullbullbullbull+c
Nbullbullbull
+QC)cbull IIlQl
+cQI
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QC) QC)
Q Q+ + + +QI QI
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 323
+
o+0o+o
N
+No+o
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ONN
t-
O ++0o+o
o+0o+o
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+o+
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+--00~-
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+
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+
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ZG_ til bull_ 0U bullbull
0 bullbull til~~ 0t
+
+
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+
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+
o+0o+o
o 0
+0 + +o -0- 0
+ + +o 0
o+0o+o
+o
OOOO--OONN--
o+0o+o
o+0o+o
+l-N
o+0o+o
o+0o+o
+
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o+0o+o
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N
+ +o
+ +N 0
+o+o
+o
o+0o+o
o
+o+
+
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+N
+o
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00
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o+0o+o
+
+00
+000 -
00
+N0gt N
+
+00_
o+0o+o
o+0o+o
+
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+o+o
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o+0o+o
o+0o+o
o+0o+o
+o
+t-ltl
1414
ot-
+ltlaobull
o+ +~~+
+bullltl
+t-bull
bullt-
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bullbull
+o+o
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324 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 2 Comparison of most abundant species in 10 combined lagoon and 10 combined sand troughsamples (1978-1981)
Lagoon Sand trough
Species (No) () (No) ()
S lerneri 2 006 190 164V new species ST 0 0 95 82V new species SA 0 0 26 22P sex 2 006 36 31P new species N 791 245 0 0P muelleri 124 38 12 10S setisparsa 28 087 31 27A americana 3 009 34 29A monambon 30 093 8 069H paucichelatus 172 53 187 161R new species A 158 49 48 41R dinochelatum 1045 324 25 22R hartmanni 88 27 I 009R new species B 104 32 21 18E new species KE 479 149 0 0E new species KG 112 35 143 123E new species R 19 059 9 077E new species P 22 068 21 18E new species B 0 0 141 121E new species MJ 0 0 31 27E new species MR 0 0 32 28E donabbotti 32 099 57 49E new species J 12 037 14 12Total 3223 100 1162 100
Also abundant were Parasterope new species N (791 specimens) and Eusarsiellanew species KE (479 specimens) (Fig 4) Two species R dinochelatum andParasterope new species N accounted for 56 of the myodocopids collected inthat zone The three most abundant species accounted for 71 of the myodocopidscollected The most abundant families in the sand and rubble zone were Ruti-dermatidae (5 species 1396 specimens) and Cylindroleberididae (8 species 983specimens all but 14 of these belonging to the Cylindroleberidinae) The Philo-medidae were represented by only 172 specimens all belonging to Harbansuspaucichelatus (Kornicker 1958) Species belonging to the Cypridinidae were veryrare in the samples (two species four specimens)
Almost half of the specimens collected in the lagoon zone belong to specieseither not present (eight species) or represented by fewer than 10 specimens inthe fore-reef samples Two additional species Rutiderma new species Land Eu-sarsiella new species MO were collected in nonquantitative sand samples fromthis zone (Table 3) but they were not found in the 10 samples used for numericalcomparisons (Tables I 2)
The largest sample (973 myodocopids June 1981) represents a density of about737 m-2 of surface sediment (973 divided by 022 6 m)
OUTERFORE-REEFSANDTROUGH(18-30 m depth) Thirty species and 1188individual myodocopid specimens were found in the 10 samples analyzed Speciesdiversity values were H = 116 E = 078 The most abundant species wasSkogsbergia lerneri (Kornicker 1958) (190 specimens) (Fig 4 Tables I 2) Alsoabundant were Harbansus paucichelatus (187) Eusarsiella new species KO (143)and E new species B (141) which together with S lerneri accounted for 55 of
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 325
11001000900800700600500400300200100
aRd PN EKE
MAJOR LAGOON SPECIES
1100~ 1000~ 9008 800Q
CIl 700~ 0gt 600
eJ ~ 500~ at bullbull00~ 300t 2002 100
a
MAJOR SAND TROUGH SPECIES
SI Hp EKD LB VST Ed RA
Figure 4 Comparison often combined samples in lagoon and ten combined samples in sand troughhistogram shows abundance of most common species in each zone (those species contributing 71-72 of the total ostracodes collected in each zone) Species shown in lagoon histogram arc R d =Rutiderma dinochelatum P N = Parasterope new species N E KE = Eusarsiella new species KESpecies shown in sand trough histogram arc S I = Skogsbergia lerneri H p = Harbansus pauci-cheallls E KG = Eusarsiella new species KO E B = E new species B V ST = Vargula n spST E d = Eusarsiella donabbotti R A = Rutiderma new species A Bar key black = Rutidermatidaeright diagonal = Cylindroleberididae left diagonal = Sarsiellidae cross-hatch = Cypridinidae open= Philomedidae
the myodocopids collected in the samples These four species along with threeothers Vargula new species ST (95 specimens) Eusarsiella donabbotti new species(57) and Rutiderma new species A (48) accounted for 72 of the myodocopidscollected in the analyzed samples The five most abundant species belong to thefamilies Cypridinidae (4 species 347 specimens) Sarsiellidae (13 species 459specimens) and Philomedidae (3 species 191 specimens) the families most abun-dant in the sand trough Specimens belonging to the other two myodocopid fam-ilies were relatively rare Rutidermatidae (6 species 98 specimens) and Cylin-droleberididae (5 species 93 specimens with only 43 of these belonging to theCylindroleberidinae)
About half of the specimens collected belong to species either not present (10)or not represented by more than 10 specimens in the lagoon samples Threeadditional species Vargula new species M Parasterope new species E and Ruti-derma new species K were found in nonquantitative samples from sand troughrubble (Table 3)
Temporal Changes in Myodocopid Fauna-LAGOON SAND AND RUBBLE ZONEThere was no change in species representation (except that rarer species did notoccur in all samples) but there was a shift in relative species family and totalabundance in collections made from January 1978 to June 1981 (Table 1Fig 5)
The Rutidermatidae was the most abundant family at the beginning (1978263specimens) and end (1981 946 specimens) of the study although the most abun-dant species in the family changed from Rutiderma new species A in January1978 to R dinochelatum in all subsequent samples Rutiderma hartmanni andR new species B increased in abundance following each sampling period
The largest sample was the single 1978 sample of 387 ostracodes Ostracodeabundance in single replicate samples declined in 1979 and increased in 1981The single 1978 sample was larger than any of the six replicate samples collectedin 1979 (Table 1) but smaller than each of the replicate samples of 1981 Fewcylindroleberids were collected in 1978 (four species five specimens in the singlesample) The Cylindroleberididae (particularly Parasterope new species N 184specimens) was the most abundant family in the June 1979 samples (8 species
326 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 3 List of myodocopid species collected only in nonquantitative samples from Belize (alsolisted other published localities)
Taxa
CypridinidaeVargula new species M
CylindroleberididaeCylindroleberidinae
Parasterope new species E
Postasterope abacoPostasterope new species BPostasterope new species X
Bruuniella new species A
Diaslerope new species D
AsteropteroninaeAsteropella maclaughlinae
Actinoseta hummelincki
RutidermatidaeAlternochelata polychelataRutiderma new species L
Rutiderma new species KE
SarsiellidaeEusarsiella new species MO
Locality (and year)
Belize only sand trough (trap and rubble 1979) lagoon patchreefs (1976) South Water Cut (1978) spur and groove(1976) outer reef slope (1979)
Belize only sand trough (rubble 1978) plankton (1976 5males) reef crest (1978)
Belize plankton (1975 I male) BahamasBelize only sand and rubble W Carrie Bow Cay (1978)Belize only South Water Cut (1977) W side Carrie Bow Cay
( 1978)Belize back-reef (1974) inner slope of outer ridge (1976) spur
and groove (1978) BermudaBelize only W side Carrie Bow Cay (1978 A-I male)
Belize Carrie Bow Cay (1978) Blue Ground (1981) W Flori-da Texas
Belize back-reef (1974 1976) spur and groove (1976 1978)Twin Cays (1977) South Water Cut (1978) Florida Vene-zuela W Panama
Belize Avicennia mangroves (I male) BahamasBelize only sand and rubble zone (1979) lagoon plankton
( 1978)Belize only sand trough (rubble 1978) outer fore-reef slope
(1979 1982) spur and groove (1979) back-reef (I 974)
Belize only sand and rubble zone (I 979) South Water Cut(I 977)
Source of localities located outside Belize POitaslerope abaca Kornicker 1986 Bruumela new species A Komicker 1981 Acrinosetamaclallghtinae Komicker 1981 (Kornicker 1986) Aclmoseta hllmmelillcki Komicker 1981 (Kornicker 1986)
253 specimens) and continued to increase in abundance subsequently thoughnot as rapidly as rutidermatids and sarsiellids
Representatives of the Sarsiellidae were most abundant in June 1981 samples(l0 species 376 specimens) and least abundant in June 1979 (8 species 70 spec-imens) The most abundant sarsiellid species in January 1978 was Eusarsiellanew species P (17) Eusarsiella new species KE and E new species KO increasedin number in subsequent samplings Three species of Eusarsiella were absent fromthe January 1978 sample but present in later samples Their absence in the 1978sample may be related to smaller sample size (single replicate) as most of thesespecies were rare in later samples Six Eusarsiella species were absent in eitherthe June or October 1979 collections but present in the June 1981 collections
The Philomedidae are represented in this zone only by Harbansus paucichelatuswhich declined in number in 1979 and increased in 1981 The Cypridinidae wererare in all samples
FORE-REEFSANDTROUGHThere was also no change in species compositionin this sample area (except that some less abundant species did not occur in allsamples) but there were shifts in relative species and family abundances between
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
ramllv
JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
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Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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0000 QQ bullbullbullbull QQ bullbullbullbullbullbull NN -- 00 00 00 00 00 QIQI 0011 bullbullbull
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 323
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324 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 2 Comparison of most abundant species in 10 combined lagoon and 10 combined sand troughsamples (1978-1981)
Lagoon Sand trough
Species (No) () (No) ()
S lerneri 2 006 190 164V new species ST 0 0 95 82V new species SA 0 0 26 22P sex 2 006 36 31P new species N 791 245 0 0P muelleri 124 38 12 10S setisparsa 28 087 31 27A americana 3 009 34 29A monambon 30 093 8 069H paucichelatus 172 53 187 161R new species A 158 49 48 41R dinochelatum 1045 324 25 22R hartmanni 88 27 I 009R new species B 104 32 21 18E new species KE 479 149 0 0E new species KG 112 35 143 123E new species R 19 059 9 077E new species P 22 068 21 18E new species B 0 0 141 121E new species MJ 0 0 31 27E new species MR 0 0 32 28E donabbotti 32 099 57 49E new species J 12 037 14 12Total 3223 100 1162 100
Also abundant were Parasterope new species N (791 specimens) and Eusarsiellanew species KE (479 specimens) (Fig 4) Two species R dinochelatum andParasterope new species N accounted for 56 of the myodocopids collected inthat zone The three most abundant species accounted for 71 of the myodocopidscollected The most abundant families in the sand and rubble zone were Ruti-dermatidae (5 species 1396 specimens) and Cylindroleberididae (8 species 983specimens all but 14 of these belonging to the Cylindroleberidinae) The Philo-medidae were represented by only 172 specimens all belonging to Harbansuspaucichelatus (Kornicker 1958) Species belonging to the Cypridinidae were veryrare in the samples (two species four specimens)
Almost half of the specimens collected in the lagoon zone belong to specieseither not present (eight species) or represented by fewer than 10 specimens inthe fore-reef samples Two additional species Rutiderma new species Land Eu-sarsiella new species MO were collected in nonquantitative sand samples fromthis zone (Table 3) but they were not found in the 10 samples used for numericalcomparisons (Tables I 2)
The largest sample (973 myodocopids June 1981) represents a density of about737 m-2 of surface sediment (973 divided by 022 6 m)
OUTERFORE-REEFSANDTROUGH(18-30 m depth) Thirty species and 1188individual myodocopid specimens were found in the 10 samples analyzed Speciesdiversity values were H = 116 E = 078 The most abundant species wasSkogsbergia lerneri (Kornicker 1958) (190 specimens) (Fig 4 Tables I 2) Alsoabundant were Harbansus paucichelatus (187) Eusarsiella new species KO (143)and E new species B (141) which together with S lerneri accounted for 55 of
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 325
11001000900800700600500400300200100
aRd PN EKE
MAJOR LAGOON SPECIES
1100~ 1000~ 9008 800Q
CIl 700~ 0gt 600
eJ ~ 500~ at bullbull00~ 300t 2002 100
a
MAJOR SAND TROUGH SPECIES
SI Hp EKD LB VST Ed RA
Figure 4 Comparison often combined samples in lagoon and ten combined samples in sand troughhistogram shows abundance of most common species in each zone (those species contributing 71-72 of the total ostracodes collected in each zone) Species shown in lagoon histogram arc R d =Rutiderma dinochelatum P N = Parasterope new species N E KE = Eusarsiella new species KESpecies shown in sand trough histogram arc S I = Skogsbergia lerneri H p = Harbansus pauci-cheallls E KG = Eusarsiella new species KO E B = E new species B V ST = Vargula n spST E d = Eusarsiella donabbotti R A = Rutiderma new species A Bar key black = Rutidermatidaeright diagonal = Cylindroleberididae left diagonal = Sarsiellidae cross-hatch = Cypridinidae open= Philomedidae
the myodocopids collected in the samples These four species along with threeothers Vargula new species ST (95 specimens) Eusarsiella donabbotti new species(57) and Rutiderma new species A (48) accounted for 72 of the myodocopidscollected in the analyzed samples The five most abundant species belong to thefamilies Cypridinidae (4 species 347 specimens) Sarsiellidae (13 species 459specimens) and Philomedidae (3 species 191 specimens) the families most abun-dant in the sand trough Specimens belonging to the other two myodocopid fam-ilies were relatively rare Rutidermatidae (6 species 98 specimens) and Cylin-droleberididae (5 species 93 specimens with only 43 of these belonging to theCylindroleberidinae)
About half of the specimens collected belong to species either not present (10)or not represented by more than 10 specimens in the lagoon samples Threeadditional species Vargula new species M Parasterope new species E and Ruti-derma new species K were found in nonquantitative samples from sand troughrubble (Table 3)
Temporal Changes in Myodocopid Fauna-LAGOON SAND AND RUBBLE ZONEThere was no change in species representation (except that rarer species did notoccur in all samples) but there was a shift in relative species family and totalabundance in collections made from January 1978 to June 1981 (Table 1Fig 5)
The Rutidermatidae was the most abundant family at the beginning (1978263specimens) and end (1981 946 specimens) of the study although the most abun-dant species in the family changed from Rutiderma new species A in January1978 to R dinochelatum in all subsequent samples Rutiderma hartmanni andR new species B increased in abundance following each sampling period
The largest sample was the single 1978 sample of 387 ostracodes Ostracodeabundance in single replicate samples declined in 1979 and increased in 1981The single 1978 sample was larger than any of the six replicate samples collectedin 1979 (Table 1) but smaller than each of the replicate samples of 1981 Fewcylindroleberids were collected in 1978 (four species five specimens in the singlesample) The Cylindroleberididae (particularly Parasterope new species N 184specimens) was the most abundant family in the June 1979 samples (8 species
326 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 3 List of myodocopid species collected only in nonquantitative samples from Belize (alsolisted other published localities)
Taxa
CypridinidaeVargula new species M
CylindroleberididaeCylindroleberidinae
Parasterope new species E
Postasterope abacoPostasterope new species BPostasterope new species X
Bruuniella new species A
Diaslerope new species D
AsteropteroninaeAsteropella maclaughlinae
Actinoseta hummelincki
RutidermatidaeAlternochelata polychelataRutiderma new species L
Rutiderma new species KE
SarsiellidaeEusarsiella new species MO
Locality (and year)
Belize only sand trough (trap and rubble 1979) lagoon patchreefs (1976) South Water Cut (1978) spur and groove(1976) outer reef slope (1979)
Belize only sand trough (rubble 1978) plankton (1976 5males) reef crest (1978)
Belize plankton (1975 I male) BahamasBelize only sand and rubble W Carrie Bow Cay (1978)Belize only South Water Cut (1977) W side Carrie Bow Cay
( 1978)Belize back-reef (1974) inner slope of outer ridge (1976) spur
and groove (1978) BermudaBelize only W side Carrie Bow Cay (1978 A-I male)
Belize Carrie Bow Cay (1978) Blue Ground (1981) W Flori-da Texas
Belize back-reef (1974 1976) spur and groove (1976 1978)Twin Cays (1977) South Water Cut (1978) Florida Vene-zuela W Panama
Belize Avicennia mangroves (I male) BahamasBelize only sand and rubble zone (1979) lagoon plankton
( 1978)Belize only sand trough (rubble 1978) outer fore-reef slope
(1979 1982) spur and groove (1979) back-reef (I 974)
Belize only sand and rubble zone (I 979) South Water Cut(I 977)
Source of localities located outside Belize POitaslerope abaca Kornicker 1986 Bruumela new species A Komicker 1981 Acrinosetamaclallghtinae Komicker 1981 (Kornicker 1986) Aclmoseta hllmmelillcki Komicker 1981 (Kornicker 1986)
253 specimens) and continued to increase in abundance subsequently thoughnot as rapidly as rutidermatids and sarsiellids
Representatives of the Sarsiellidae were most abundant in June 1981 samples(l0 species 376 specimens) and least abundant in June 1979 (8 species 70 spec-imens) The most abundant sarsiellid species in January 1978 was Eusarsiellanew species P (17) Eusarsiella new species KE and E new species KO increasedin number in subsequent samplings Three species of Eusarsiella were absent fromthe January 1978 sample but present in later samples Their absence in the 1978sample may be related to smaller sample size (single replicate) as most of thesespecies were rare in later samples Six Eusarsiella species were absent in eitherthe June or October 1979 collections but present in the June 1981 collections
The Philomedidae are represented in this zone only by Harbansus paucichelatuswhich declined in number in 1979 and increased in 1981 The Cypridinidae wererare in all samples
FORE-REEFSANDTROUGHThere was also no change in species compositionin this sample area (except that some less abundant species did not occur in allsamples) but there were shifts in relative species and family abundances between
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
ramllv
JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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322 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
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324 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 2 Comparison of most abundant species in 10 combined lagoon and 10 combined sand troughsamples (1978-1981)
Lagoon Sand trough
Species (No) () (No) ()
S lerneri 2 006 190 164V new species ST 0 0 95 82V new species SA 0 0 26 22P sex 2 006 36 31P new species N 791 245 0 0P muelleri 124 38 12 10S setisparsa 28 087 31 27A americana 3 009 34 29A monambon 30 093 8 069H paucichelatus 172 53 187 161R new species A 158 49 48 41R dinochelatum 1045 324 25 22R hartmanni 88 27 I 009R new species B 104 32 21 18E new species KE 479 149 0 0E new species KG 112 35 143 123E new species R 19 059 9 077E new species P 22 068 21 18E new species B 0 0 141 121E new species MJ 0 0 31 27E new species MR 0 0 32 28E donabbotti 32 099 57 49E new species J 12 037 14 12Total 3223 100 1162 100
Also abundant were Parasterope new species N (791 specimens) and Eusarsiellanew species KE (479 specimens) (Fig 4) Two species R dinochelatum andParasterope new species N accounted for 56 of the myodocopids collected inthat zone The three most abundant species accounted for 71 of the myodocopidscollected The most abundant families in the sand and rubble zone were Ruti-dermatidae (5 species 1396 specimens) and Cylindroleberididae (8 species 983specimens all but 14 of these belonging to the Cylindroleberidinae) The Philo-medidae were represented by only 172 specimens all belonging to Harbansuspaucichelatus (Kornicker 1958) Species belonging to the Cypridinidae were veryrare in the samples (two species four specimens)
Almost half of the specimens collected in the lagoon zone belong to specieseither not present (eight species) or represented by fewer than 10 specimens inthe fore-reef samples Two additional species Rutiderma new species Land Eu-sarsiella new species MO were collected in nonquantitative sand samples fromthis zone (Table 3) but they were not found in the 10 samples used for numericalcomparisons (Tables I 2)
The largest sample (973 myodocopids June 1981) represents a density of about737 m-2 of surface sediment (973 divided by 022 6 m)
OUTERFORE-REEFSANDTROUGH(18-30 m depth) Thirty species and 1188individual myodocopid specimens were found in the 10 samples analyzed Speciesdiversity values were H = 116 E = 078 The most abundant species wasSkogsbergia lerneri (Kornicker 1958) (190 specimens) (Fig 4 Tables I 2) Alsoabundant were Harbansus paucichelatus (187) Eusarsiella new species KO (143)and E new species B (141) which together with S lerneri accounted for 55 of
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 325
11001000900800700600500400300200100
aRd PN EKE
MAJOR LAGOON SPECIES
1100~ 1000~ 9008 800Q
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MAJOR SAND TROUGH SPECIES
SI Hp EKD LB VST Ed RA
Figure 4 Comparison often combined samples in lagoon and ten combined samples in sand troughhistogram shows abundance of most common species in each zone (those species contributing 71-72 of the total ostracodes collected in each zone) Species shown in lagoon histogram arc R d =Rutiderma dinochelatum P N = Parasterope new species N E KE = Eusarsiella new species KESpecies shown in sand trough histogram arc S I = Skogsbergia lerneri H p = Harbansus pauci-cheallls E KG = Eusarsiella new species KO E B = E new species B V ST = Vargula n spST E d = Eusarsiella donabbotti R A = Rutiderma new species A Bar key black = Rutidermatidaeright diagonal = Cylindroleberididae left diagonal = Sarsiellidae cross-hatch = Cypridinidae open= Philomedidae
the myodocopids collected in the samples These four species along with threeothers Vargula new species ST (95 specimens) Eusarsiella donabbotti new species(57) and Rutiderma new species A (48) accounted for 72 of the myodocopidscollected in the analyzed samples The five most abundant species belong to thefamilies Cypridinidae (4 species 347 specimens) Sarsiellidae (13 species 459specimens) and Philomedidae (3 species 191 specimens) the families most abun-dant in the sand trough Specimens belonging to the other two myodocopid fam-ilies were relatively rare Rutidermatidae (6 species 98 specimens) and Cylin-droleberididae (5 species 93 specimens with only 43 of these belonging to theCylindroleberidinae)
About half of the specimens collected belong to species either not present (10)or not represented by more than 10 specimens in the lagoon samples Threeadditional species Vargula new species M Parasterope new species E and Ruti-derma new species K were found in nonquantitative samples from sand troughrubble (Table 3)
Temporal Changes in Myodocopid Fauna-LAGOON SAND AND RUBBLE ZONEThere was no change in species representation (except that rarer species did notoccur in all samples) but there was a shift in relative species family and totalabundance in collections made from January 1978 to June 1981 (Table 1Fig 5)
The Rutidermatidae was the most abundant family at the beginning (1978263specimens) and end (1981 946 specimens) of the study although the most abun-dant species in the family changed from Rutiderma new species A in January1978 to R dinochelatum in all subsequent samples Rutiderma hartmanni andR new species B increased in abundance following each sampling period
The largest sample was the single 1978 sample of 387 ostracodes Ostracodeabundance in single replicate samples declined in 1979 and increased in 1981The single 1978 sample was larger than any of the six replicate samples collectedin 1979 (Table 1) but smaller than each of the replicate samples of 1981 Fewcylindroleberids were collected in 1978 (four species five specimens in the singlesample) The Cylindroleberididae (particularly Parasterope new species N 184specimens) was the most abundant family in the June 1979 samples (8 species
326 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 3 List of myodocopid species collected only in nonquantitative samples from Belize (alsolisted other published localities)
Taxa
CypridinidaeVargula new species M
CylindroleberididaeCylindroleberidinae
Parasterope new species E
Postasterope abacoPostasterope new species BPostasterope new species X
Bruuniella new species A
Diaslerope new species D
AsteropteroninaeAsteropella maclaughlinae
Actinoseta hummelincki
RutidermatidaeAlternochelata polychelataRutiderma new species L
Rutiderma new species KE
SarsiellidaeEusarsiella new species MO
Locality (and year)
Belize only sand trough (trap and rubble 1979) lagoon patchreefs (1976) South Water Cut (1978) spur and groove(1976) outer reef slope (1979)
Belize only sand trough (rubble 1978) plankton (1976 5males) reef crest (1978)
Belize plankton (1975 I male) BahamasBelize only sand and rubble W Carrie Bow Cay (1978)Belize only South Water Cut (1977) W side Carrie Bow Cay
( 1978)Belize back-reef (1974) inner slope of outer ridge (1976) spur
and groove (1978) BermudaBelize only W side Carrie Bow Cay (1978 A-I male)
Belize Carrie Bow Cay (1978) Blue Ground (1981) W Flori-da Texas
Belize back-reef (1974 1976) spur and groove (1976 1978)Twin Cays (1977) South Water Cut (1978) Florida Vene-zuela W Panama
Belize Avicennia mangroves (I male) BahamasBelize only sand and rubble zone (1979) lagoon plankton
( 1978)Belize only sand trough (rubble 1978) outer fore-reef slope
(1979 1982) spur and groove (1979) back-reef (I 974)
Belize only sand and rubble zone (I 979) South Water Cut(I 977)
Source of localities located outside Belize POitaslerope abaca Kornicker 1986 Bruumela new species A Komicker 1981 Acrinosetamaclallghtinae Komicker 1981 (Kornicker 1986) Aclmoseta hllmmelillcki Komicker 1981 (Kornicker 1986)
253 specimens) and continued to increase in abundance subsequently thoughnot as rapidly as rutidermatids and sarsiellids
Representatives of the Sarsiellidae were most abundant in June 1981 samples(l0 species 376 specimens) and least abundant in June 1979 (8 species 70 spec-imens) The most abundant sarsiellid species in January 1978 was Eusarsiellanew species P (17) Eusarsiella new species KE and E new species KO increasedin number in subsequent samplings Three species of Eusarsiella were absent fromthe January 1978 sample but present in later samples Their absence in the 1978sample may be related to smaller sample size (single replicate) as most of thesespecies were rare in later samples Six Eusarsiella species were absent in eitherthe June or October 1979 collections but present in the June 1981 collections
The Philomedidae are represented in this zone only by Harbansus paucichelatuswhich declined in number in 1979 and increased in 1981 The Cypridinidae wererare in all samples
FORE-REEFSANDTROUGHThere was also no change in species compositionin this sample area (except that some less abundant species did not occur in allsamples) but there were shifts in relative species and family abundances between
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
ramllv
JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 323
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324 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 2 Comparison of most abundant species in 10 combined lagoon and 10 combined sand troughsamples (1978-1981)
Lagoon Sand trough
Species (No) () (No) ()
S lerneri 2 006 190 164V new species ST 0 0 95 82V new species SA 0 0 26 22P sex 2 006 36 31P new species N 791 245 0 0P muelleri 124 38 12 10S setisparsa 28 087 31 27A americana 3 009 34 29A monambon 30 093 8 069H paucichelatus 172 53 187 161R new species A 158 49 48 41R dinochelatum 1045 324 25 22R hartmanni 88 27 I 009R new species B 104 32 21 18E new species KE 479 149 0 0E new species KG 112 35 143 123E new species R 19 059 9 077E new species P 22 068 21 18E new species B 0 0 141 121E new species MJ 0 0 31 27E new species MR 0 0 32 28E donabbotti 32 099 57 49E new species J 12 037 14 12Total 3223 100 1162 100
Also abundant were Parasterope new species N (791 specimens) and Eusarsiellanew species KE (479 specimens) (Fig 4) Two species R dinochelatum andParasterope new species N accounted for 56 of the myodocopids collected inthat zone The three most abundant species accounted for 71 of the myodocopidscollected The most abundant families in the sand and rubble zone were Ruti-dermatidae (5 species 1396 specimens) and Cylindroleberididae (8 species 983specimens all but 14 of these belonging to the Cylindroleberidinae) The Philo-medidae were represented by only 172 specimens all belonging to Harbansuspaucichelatus (Kornicker 1958) Species belonging to the Cypridinidae were veryrare in the samples (two species four specimens)
Almost half of the specimens collected in the lagoon zone belong to specieseither not present (eight species) or represented by fewer than 10 specimens inthe fore-reef samples Two additional species Rutiderma new species Land Eu-sarsiella new species MO were collected in nonquantitative sand samples fromthis zone (Table 3) but they were not found in the 10 samples used for numericalcomparisons (Tables I 2)
The largest sample (973 myodocopids June 1981) represents a density of about737 m-2 of surface sediment (973 divided by 022 6 m)
OUTERFORE-REEFSANDTROUGH(18-30 m depth) Thirty species and 1188individual myodocopid specimens were found in the 10 samples analyzed Speciesdiversity values were H = 116 E = 078 The most abundant species wasSkogsbergia lerneri (Kornicker 1958) (190 specimens) (Fig 4 Tables I 2) Alsoabundant were Harbansus paucichelatus (187) Eusarsiella new species KO (143)and E new species B (141) which together with S lerneri accounted for 55 of
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 325
11001000900800700600500400300200100
aRd PN EKE
MAJOR LAGOON SPECIES
1100~ 1000~ 9008 800Q
CIl 700~ 0gt 600
eJ ~ 500~ at bullbull00~ 300t 2002 100
a
MAJOR SAND TROUGH SPECIES
SI Hp EKD LB VST Ed RA
Figure 4 Comparison often combined samples in lagoon and ten combined samples in sand troughhistogram shows abundance of most common species in each zone (those species contributing 71-72 of the total ostracodes collected in each zone) Species shown in lagoon histogram arc R d =Rutiderma dinochelatum P N = Parasterope new species N E KE = Eusarsiella new species KESpecies shown in sand trough histogram arc S I = Skogsbergia lerneri H p = Harbansus pauci-cheallls E KG = Eusarsiella new species KO E B = E new species B V ST = Vargula n spST E d = Eusarsiella donabbotti R A = Rutiderma new species A Bar key black = Rutidermatidaeright diagonal = Cylindroleberididae left diagonal = Sarsiellidae cross-hatch = Cypridinidae open= Philomedidae
the myodocopids collected in the samples These four species along with threeothers Vargula new species ST (95 specimens) Eusarsiella donabbotti new species(57) and Rutiderma new species A (48) accounted for 72 of the myodocopidscollected in the analyzed samples The five most abundant species belong to thefamilies Cypridinidae (4 species 347 specimens) Sarsiellidae (13 species 459specimens) and Philomedidae (3 species 191 specimens) the families most abun-dant in the sand trough Specimens belonging to the other two myodocopid fam-ilies were relatively rare Rutidermatidae (6 species 98 specimens) and Cylin-droleberididae (5 species 93 specimens with only 43 of these belonging to theCylindroleberidinae)
About half of the specimens collected belong to species either not present (10)or not represented by more than 10 specimens in the lagoon samples Threeadditional species Vargula new species M Parasterope new species E and Ruti-derma new species K were found in nonquantitative samples from sand troughrubble (Table 3)
Temporal Changes in Myodocopid Fauna-LAGOON SAND AND RUBBLE ZONEThere was no change in species representation (except that rarer species did notoccur in all samples) but there was a shift in relative species family and totalabundance in collections made from January 1978 to June 1981 (Table 1Fig 5)
The Rutidermatidae was the most abundant family at the beginning (1978263specimens) and end (1981 946 specimens) of the study although the most abun-dant species in the family changed from Rutiderma new species A in January1978 to R dinochelatum in all subsequent samples Rutiderma hartmanni andR new species B increased in abundance following each sampling period
The largest sample was the single 1978 sample of 387 ostracodes Ostracodeabundance in single replicate samples declined in 1979 and increased in 1981The single 1978 sample was larger than any of the six replicate samples collectedin 1979 (Table 1) but smaller than each of the replicate samples of 1981 Fewcylindroleberids were collected in 1978 (four species five specimens in the singlesample) The Cylindroleberididae (particularly Parasterope new species N 184specimens) was the most abundant family in the June 1979 samples (8 species
326 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 3 List of myodocopid species collected only in nonquantitative samples from Belize (alsolisted other published localities)
Taxa
CypridinidaeVargula new species M
CylindroleberididaeCylindroleberidinae
Parasterope new species E
Postasterope abacoPostasterope new species BPostasterope new species X
Bruuniella new species A
Diaslerope new species D
AsteropteroninaeAsteropella maclaughlinae
Actinoseta hummelincki
RutidermatidaeAlternochelata polychelataRutiderma new species L
Rutiderma new species KE
SarsiellidaeEusarsiella new species MO
Locality (and year)
Belize only sand trough (trap and rubble 1979) lagoon patchreefs (1976) South Water Cut (1978) spur and groove(1976) outer reef slope (1979)
Belize only sand trough (rubble 1978) plankton (1976 5males) reef crest (1978)
Belize plankton (1975 I male) BahamasBelize only sand and rubble W Carrie Bow Cay (1978)Belize only South Water Cut (1977) W side Carrie Bow Cay
( 1978)Belize back-reef (1974) inner slope of outer ridge (1976) spur
and groove (1978) BermudaBelize only W side Carrie Bow Cay (1978 A-I male)
Belize Carrie Bow Cay (1978) Blue Ground (1981) W Flori-da Texas
Belize back-reef (1974 1976) spur and groove (1976 1978)Twin Cays (1977) South Water Cut (1978) Florida Vene-zuela W Panama
Belize Avicennia mangroves (I male) BahamasBelize only sand and rubble zone (1979) lagoon plankton
( 1978)Belize only sand trough (rubble 1978) outer fore-reef slope
(1979 1982) spur and groove (1979) back-reef (I 974)
Belize only sand and rubble zone (I 979) South Water Cut(I 977)
Source of localities located outside Belize POitaslerope abaca Kornicker 1986 Bruumela new species A Komicker 1981 Acrinosetamaclallghtinae Komicker 1981 (Kornicker 1986) Aclmoseta hllmmelillcki Komicker 1981 (Kornicker 1986)
253 specimens) and continued to increase in abundance subsequently thoughnot as rapidly as rutidermatids and sarsiellids
Representatives of the Sarsiellidae were most abundant in June 1981 samples(l0 species 376 specimens) and least abundant in June 1979 (8 species 70 spec-imens) The most abundant sarsiellid species in January 1978 was Eusarsiellanew species P (17) Eusarsiella new species KE and E new species KO increasedin number in subsequent samplings Three species of Eusarsiella were absent fromthe January 1978 sample but present in later samples Their absence in the 1978sample may be related to smaller sample size (single replicate) as most of thesespecies were rare in later samples Six Eusarsiella species were absent in eitherthe June or October 1979 collections but present in the June 1981 collections
The Philomedidae are represented in this zone only by Harbansus paucichelatuswhich declined in number in 1979 and increased in 1981 The Cypridinidae wererare in all samples
FORE-REEFSANDTROUGHThere was also no change in species compositionin this sample area (except that some less abundant species did not occur in allsamples) but there were shifts in relative species and family abundances between
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
ramllv
JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
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-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
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des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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324 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 2 Comparison of most abundant species in 10 combined lagoon and 10 combined sand troughsamples (1978-1981)
Lagoon Sand trough
Species (No) () (No) ()
S lerneri 2 006 190 164V new species ST 0 0 95 82V new species SA 0 0 26 22P sex 2 006 36 31P new species N 791 245 0 0P muelleri 124 38 12 10S setisparsa 28 087 31 27A americana 3 009 34 29A monambon 30 093 8 069H paucichelatus 172 53 187 161R new species A 158 49 48 41R dinochelatum 1045 324 25 22R hartmanni 88 27 I 009R new species B 104 32 21 18E new species KE 479 149 0 0E new species KG 112 35 143 123E new species R 19 059 9 077E new species P 22 068 21 18E new species B 0 0 141 121E new species MJ 0 0 31 27E new species MR 0 0 32 28E donabbotti 32 099 57 49E new species J 12 037 14 12Total 3223 100 1162 100
Also abundant were Parasterope new species N (791 specimens) and Eusarsiellanew species KE (479 specimens) (Fig 4) Two species R dinochelatum andParasterope new species N accounted for 56 of the myodocopids collected inthat zone The three most abundant species accounted for 71 of the myodocopidscollected The most abundant families in the sand and rubble zone were Ruti-dermatidae (5 species 1396 specimens) and Cylindroleberididae (8 species 983specimens all but 14 of these belonging to the Cylindroleberidinae) The Philo-medidae were represented by only 172 specimens all belonging to Harbansuspaucichelatus (Kornicker 1958) Species belonging to the Cypridinidae were veryrare in the samples (two species four specimens)
Almost half of the specimens collected in the lagoon zone belong to specieseither not present (eight species) or represented by fewer than 10 specimens inthe fore-reef samples Two additional species Rutiderma new species Land Eu-sarsiella new species MO were collected in nonquantitative sand samples fromthis zone (Table 3) but they were not found in the 10 samples used for numericalcomparisons (Tables I 2)
The largest sample (973 myodocopids June 1981) represents a density of about737 m-2 of surface sediment (973 divided by 022 6 m)
OUTERFORE-REEFSANDTROUGH(18-30 m depth) Thirty species and 1188individual myodocopid specimens were found in the 10 samples analyzed Speciesdiversity values were H = 116 E = 078 The most abundant species wasSkogsbergia lerneri (Kornicker 1958) (190 specimens) (Fig 4 Tables I 2) Alsoabundant were Harbansus paucichelatus (187) Eusarsiella new species KO (143)and E new species B (141) which together with S lerneri accounted for 55 of
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 325
11001000900800700600500400300200100
aRd PN EKE
MAJOR LAGOON SPECIES
1100~ 1000~ 9008 800Q
CIl 700~ 0gt 600
eJ ~ 500~ at bullbull00~ 300t 2002 100
a
MAJOR SAND TROUGH SPECIES
SI Hp EKD LB VST Ed RA
Figure 4 Comparison often combined samples in lagoon and ten combined samples in sand troughhistogram shows abundance of most common species in each zone (those species contributing 71-72 of the total ostracodes collected in each zone) Species shown in lagoon histogram arc R d =Rutiderma dinochelatum P N = Parasterope new species N E KE = Eusarsiella new species KESpecies shown in sand trough histogram arc S I = Skogsbergia lerneri H p = Harbansus pauci-cheallls E KG = Eusarsiella new species KO E B = E new species B V ST = Vargula n spST E d = Eusarsiella donabbotti R A = Rutiderma new species A Bar key black = Rutidermatidaeright diagonal = Cylindroleberididae left diagonal = Sarsiellidae cross-hatch = Cypridinidae open= Philomedidae
the myodocopids collected in the samples These four species along with threeothers Vargula new species ST (95 specimens) Eusarsiella donabbotti new species(57) and Rutiderma new species A (48) accounted for 72 of the myodocopidscollected in the analyzed samples The five most abundant species belong to thefamilies Cypridinidae (4 species 347 specimens) Sarsiellidae (13 species 459specimens) and Philomedidae (3 species 191 specimens) the families most abun-dant in the sand trough Specimens belonging to the other two myodocopid fam-ilies were relatively rare Rutidermatidae (6 species 98 specimens) and Cylin-droleberididae (5 species 93 specimens with only 43 of these belonging to theCylindroleberidinae)
About half of the specimens collected belong to species either not present (10)or not represented by more than 10 specimens in the lagoon samples Threeadditional species Vargula new species M Parasterope new species E and Ruti-derma new species K were found in nonquantitative samples from sand troughrubble (Table 3)
Temporal Changes in Myodocopid Fauna-LAGOON SAND AND RUBBLE ZONEThere was no change in species representation (except that rarer species did notoccur in all samples) but there was a shift in relative species family and totalabundance in collections made from January 1978 to June 1981 (Table 1Fig 5)
The Rutidermatidae was the most abundant family at the beginning (1978263specimens) and end (1981 946 specimens) of the study although the most abun-dant species in the family changed from Rutiderma new species A in January1978 to R dinochelatum in all subsequent samples Rutiderma hartmanni andR new species B increased in abundance following each sampling period
The largest sample was the single 1978 sample of 387 ostracodes Ostracodeabundance in single replicate samples declined in 1979 and increased in 1981The single 1978 sample was larger than any of the six replicate samples collectedin 1979 (Table 1) but smaller than each of the replicate samples of 1981 Fewcylindroleberids were collected in 1978 (four species five specimens in the singlesample) The Cylindroleberididae (particularly Parasterope new species N 184specimens) was the most abundant family in the June 1979 samples (8 species
326 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 3 List of myodocopid species collected only in nonquantitative samples from Belize (alsolisted other published localities)
Taxa
CypridinidaeVargula new species M
CylindroleberididaeCylindroleberidinae
Parasterope new species E
Postasterope abacoPostasterope new species BPostasterope new species X
Bruuniella new species A
Diaslerope new species D
AsteropteroninaeAsteropella maclaughlinae
Actinoseta hummelincki
RutidermatidaeAlternochelata polychelataRutiderma new species L
Rutiderma new species KE
SarsiellidaeEusarsiella new species MO
Locality (and year)
Belize only sand trough (trap and rubble 1979) lagoon patchreefs (1976) South Water Cut (1978) spur and groove(1976) outer reef slope (1979)
Belize only sand trough (rubble 1978) plankton (1976 5males) reef crest (1978)
Belize plankton (1975 I male) BahamasBelize only sand and rubble W Carrie Bow Cay (1978)Belize only South Water Cut (1977) W side Carrie Bow Cay
( 1978)Belize back-reef (1974) inner slope of outer ridge (1976) spur
and groove (1978) BermudaBelize only W side Carrie Bow Cay (1978 A-I male)
Belize Carrie Bow Cay (1978) Blue Ground (1981) W Flori-da Texas
Belize back-reef (1974 1976) spur and groove (1976 1978)Twin Cays (1977) South Water Cut (1978) Florida Vene-zuela W Panama
Belize Avicennia mangroves (I male) BahamasBelize only sand and rubble zone (1979) lagoon plankton
( 1978)Belize only sand trough (rubble 1978) outer fore-reef slope
(1979 1982) spur and groove (1979) back-reef (I 974)
Belize only sand and rubble zone (I 979) South Water Cut(I 977)
Source of localities located outside Belize POitaslerope abaca Kornicker 1986 Bruumela new species A Komicker 1981 Acrinosetamaclallghtinae Komicker 1981 (Kornicker 1986) Aclmoseta hllmmelillcki Komicker 1981 (Kornicker 1986)
253 specimens) and continued to increase in abundance subsequently thoughnot as rapidly as rutidermatids and sarsiellids
Representatives of the Sarsiellidae were most abundant in June 1981 samples(l0 species 376 specimens) and least abundant in June 1979 (8 species 70 spec-imens) The most abundant sarsiellid species in January 1978 was Eusarsiellanew species P (17) Eusarsiella new species KE and E new species KO increasedin number in subsequent samplings Three species of Eusarsiella were absent fromthe January 1978 sample but present in later samples Their absence in the 1978sample may be related to smaller sample size (single replicate) as most of thesespecies were rare in later samples Six Eusarsiella species were absent in eitherthe June or October 1979 collections but present in the June 1981 collections
The Philomedidae are represented in this zone only by Harbansus paucichelatuswhich declined in number in 1979 and increased in 1981 The Cypridinidae wererare in all samples
FORE-REEFSANDTROUGHThere was also no change in species compositionin this sample area (except that some less abundant species did not occur in allsamples) but there were shifts in relative species and family abundances between
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
ramllv
JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 325
11001000900800700600500400300200100
aRd PN EKE
MAJOR LAGOON SPECIES
1100~ 1000~ 9008 800Q
CIl 700~ 0gt 600
eJ ~ 500~ at bullbull00~ 300t 2002 100
a
MAJOR SAND TROUGH SPECIES
SI Hp EKD LB VST Ed RA
Figure 4 Comparison often combined samples in lagoon and ten combined samples in sand troughhistogram shows abundance of most common species in each zone (those species contributing 71-72 of the total ostracodes collected in each zone) Species shown in lagoon histogram arc R d =Rutiderma dinochelatum P N = Parasterope new species N E KE = Eusarsiella new species KESpecies shown in sand trough histogram arc S I = Skogsbergia lerneri H p = Harbansus pauci-cheallls E KG = Eusarsiella new species KO E B = E new species B V ST = Vargula n spST E d = Eusarsiella donabbotti R A = Rutiderma new species A Bar key black = Rutidermatidaeright diagonal = Cylindroleberididae left diagonal = Sarsiellidae cross-hatch = Cypridinidae open= Philomedidae
the myodocopids collected in the samples These four species along with threeothers Vargula new species ST (95 specimens) Eusarsiella donabbotti new species(57) and Rutiderma new species A (48) accounted for 72 of the myodocopidscollected in the analyzed samples The five most abundant species belong to thefamilies Cypridinidae (4 species 347 specimens) Sarsiellidae (13 species 459specimens) and Philomedidae (3 species 191 specimens) the families most abun-dant in the sand trough Specimens belonging to the other two myodocopid fam-ilies were relatively rare Rutidermatidae (6 species 98 specimens) and Cylin-droleberididae (5 species 93 specimens with only 43 of these belonging to theCylindroleberidinae)
About half of the specimens collected belong to species either not present (10)or not represented by more than 10 specimens in the lagoon samples Threeadditional species Vargula new species M Parasterope new species E and Ruti-derma new species K were found in nonquantitative samples from sand troughrubble (Table 3)
Temporal Changes in Myodocopid Fauna-LAGOON SAND AND RUBBLE ZONEThere was no change in species representation (except that rarer species did notoccur in all samples) but there was a shift in relative species family and totalabundance in collections made from January 1978 to June 1981 (Table 1Fig 5)
The Rutidermatidae was the most abundant family at the beginning (1978263specimens) and end (1981 946 specimens) of the study although the most abun-dant species in the family changed from Rutiderma new species A in January1978 to R dinochelatum in all subsequent samples Rutiderma hartmanni andR new species B increased in abundance following each sampling period
The largest sample was the single 1978 sample of 387 ostracodes Ostracodeabundance in single replicate samples declined in 1979 and increased in 1981The single 1978 sample was larger than any of the six replicate samples collectedin 1979 (Table 1) but smaller than each of the replicate samples of 1981 Fewcylindroleberids were collected in 1978 (four species five specimens in the singlesample) The Cylindroleberididae (particularly Parasterope new species N 184specimens) was the most abundant family in the June 1979 samples (8 species
326 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 3 List of myodocopid species collected only in nonquantitative samples from Belize (alsolisted other published localities)
Taxa
CypridinidaeVargula new species M
CylindroleberididaeCylindroleberidinae
Parasterope new species E
Postasterope abacoPostasterope new species BPostasterope new species X
Bruuniella new species A
Diaslerope new species D
AsteropteroninaeAsteropella maclaughlinae
Actinoseta hummelincki
RutidermatidaeAlternochelata polychelataRutiderma new species L
Rutiderma new species KE
SarsiellidaeEusarsiella new species MO
Locality (and year)
Belize only sand trough (trap and rubble 1979) lagoon patchreefs (1976) South Water Cut (1978) spur and groove(1976) outer reef slope (1979)
Belize only sand trough (rubble 1978) plankton (1976 5males) reef crest (1978)
Belize plankton (1975 I male) BahamasBelize only sand and rubble W Carrie Bow Cay (1978)Belize only South Water Cut (1977) W side Carrie Bow Cay
( 1978)Belize back-reef (1974) inner slope of outer ridge (1976) spur
and groove (1978) BermudaBelize only W side Carrie Bow Cay (1978 A-I male)
Belize Carrie Bow Cay (1978) Blue Ground (1981) W Flori-da Texas
Belize back-reef (1974 1976) spur and groove (1976 1978)Twin Cays (1977) South Water Cut (1978) Florida Vene-zuela W Panama
Belize Avicennia mangroves (I male) BahamasBelize only sand and rubble zone (1979) lagoon plankton
( 1978)Belize only sand trough (rubble 1978) outer fore-reef slope
(1979 1982) spur and groove (1979) back-reef (I 974)
Belize only sand and rubble zone (I 979) South Water Cut(I 977)
Source of localities located outside Belize POitaslerope abaca Kornicker 1986 Bruumela new species A Komicker 1981 Acrinosetamaclallghtinae Komicker 1981 (Kornicker 1986) Aclmoseta hllmmelillcki Komicker 1981 (Kornicker 1986)
253 specimens) and continued to increase in abundance subsequently thoughnot as rapidly as rutidermatids and sarsiellids
Representatives of the Sarsiellidae were most abundant in June 1981 samples(l0 species 376 specimens) and least abundant in June 1979 (8 species 70 spec-imens) The most abundant sarsiellid species in January 1978 was Eusarsiellanew species P (17) Eusarsiella new species KE and E new species KO increasedin number in subsequent samplings Three species of Eusarsiella were absent fromthe January 1978 sample but present in later samples Their absence in the 1978sample may be related to smaller sample size (single replicate) as most of thesespecies were rare in later samples Six Eusarsiella species were absent in eitherthe June or October 1979 collections but present in the June 1981 collections
The Philomedidae are represented in this zone only by Harbansus paucichelatuswhich declined in number in 1979 and increased in 1981 The Cypridinidae wererare in all samples
FORE-REEFSANDTROUGHThere was also no change in species compositionin this sample area (except that some less abundant species did not occur in allsamples) but there were shifts in relative species and family abundances between
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
ramllv
JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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326 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 3 List of myodocopid species collected only in nonquantitative samples from Belize (alsolisted other published localities)
Taxa
CypridinidaeVargula new species M
CylindroleberididaeCylindroleberidinae
Parasterope new species E
Postasterope abacoPostasterope new species BPostasterope new species X
Bruuniella new species A
Diaslerope new species D
AsteropteroninaeAsteropella maclaughlinae
Actinoseta hummelincki
RutidermatidaeAlternochelata polychelataRutiderma new species L
Rutiderma new species KE
SarsiellidaeEusarsiella new species MO
Locality (and year)
Belize only sand trough (trap and rubble 1979) lagoon patchreefs (1976) South Water Cut (1978) spur and groove(1976) outer reef slope (1979)
Belize only sand trough (rubble 1978) plankton (1976 5males) reef crest (1978)
Belize plankton (1975 I male) BahamasBelize only sand and rubble W Carrie Bow Cay (1978)Belize only South Water Cut (1977) W side Carrie Bow Cay
( 1978)Belize back-reef (1974) inner slope of outer ridge (1976) spur
and groove (1978) BermudaBelize only W side Carrie Bow Cay (1978 A-I male)
Belize Carrie Bow Cay (1978) Blue Ground (1981) W Flori-da Texas
Belize back-reef (1974 1976) spur and groove (1976 1978)Twin Cays (1977) South Water Cut (1978) Florida Vene-zuela W Panama
Belize Avicennia mangroves (I male) BahamasBelize only sand and rubble zone (1979) lagoon plankton
( 1978)Belize only sand trough (rubble 1978) outer fore-reef slope
(1979 1982) spur and groove (1979) back-reef (I 974)
Belize only sand and rubble zone (I 979) South Water Cut(I 977)
Source of localities located outside Belize POitaslerope abaca Kornicker 1986 Bruumela new species A Komicker 1981 Acrinosetamaclallghtinae Komicker 1981 (Kornicker 1986) Aclmoseta hllmmelillcki Komicker 1981 (Kornicker 1986)
253 specimens) and continued to increase in abundance subsequently thoughnot as rapidly as rutidermatids and sarsiellids
Representatives of the Sarsiellidae were most abundant in June 1981 samples(l0 species 376 specimens) and least abundant in June 1979 (8 species 70 spec-imens) The most abundant sarsiellid species in January 1978 was Eusarsiellanew species P (17) Eusarsiella new species KE and E new species KO increasedin number in subsequent samplings Three species of Eusarsiella were absent fromthe January 1978 sample but present in later samples Their absence in the 1978sample may be related to smaller sample size (single replicate) as most of thesespecies were rare in later samples Six Eusarsiella species were absent in eitherthe June or October 1979 collections but present in the June 1981 collections
The Philomedidae are represented in this zone only by Harbansus paucichelatuswhich declined in number in 1979 and increased in 1981 The Cypridinidae wererare in all samples
FORE-REEFSANDTROUGHThere was also no change in species compositionin this sample area (except that some less abundant species did not occur in allsamples) but there were shifts in relative species and family abundances between
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
ramllv
JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
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336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
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Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 327
lOG
LAGOON
FORE REEF
CCdPRS
Fomlly
JAN 1978
co
LAGOON
JUNE 1979
LAGOON
OCTOBER 1979
LAGOON
FORE REEF
CI Cd P R
ramllv
JUNE 1981
Figure 5 Comparison of relative abundance of myodocopid families in lagoon and sand trough atfour sampling dates (1978-] 981) Histograms for January 1978 show number of individuals collectedfor each family in a single sample in each ofthc two zones Histograms for 1979-1981 show meanand standard error of three samples from each zone at each date Samples are same shown in Tab]eI CI = Cylindro]eberididae Cd = Cypridinidae P = Philomedidae R = Rutidermatidae S = Sar-siellidae
January 1978 and June 1981 Initially there was an increase then a decline intotal specimen numbers (Fig 5 Table 1) The smallest sample was 46 ostracodescollected in the single 1978 sample fewer ostracodes than any of the subsequentnine samples The highest number collected was in the combined three October1979 samples (total 473) while about equal numbers were collected in the com-bined three samples in June 1979 (total 356) and combined three samples in June1981 (total 313) The largest single sample was also collected in October 1979
The most abundant species in the sand trough Skogsbergia lerneri as well asthe other three sand trough species that belong to the family Cypridinidae wasmost abundant in the June 1979 collection and declined in subsequent collectionsEach of the three June 1979 samples was larger than the single 1978 sample
Harbansus paucichelatus the only abundant species of Philomedidae was al-most as abundant as S lerneri increasing slightly during the study period untilJune 1981 when it declined considerably
The Rutidermatidae and the most abundant species of that family in the sandtrough Rutiderma new species A both showed numerical increase over most ofthe study period and a slight decline in October 1979 Like all other rutidermatidsin the sand trough Rutiderma dinochelatum was rare but was represented by 19specimens in the June 1981 collection
The Sarsiellidae (and five sarsiellid species) were most abundant in the October1979 collection and two of those species were only present then Six species(including the two only present in October 1979) declined in June 1981 and wereless abundant than in June 1979 as well The most abundant species of theSarsiellidae in the sand trough Eusarsiella new species KO as well as two otherspecies showed an increase in numbers over the entire study period Eight speciesof sarsiellids were not collected in all four sampling periods
COMPARISONOF ABUNDANCEIN THETwo ZoNES About three times as manymyodocopids were collected in the 10 lagoon samples (3281) as in the 10 fore-reef samples (1188) Total abundance of ostracodes in each of the 10 samples(1978-1981) from each zone was significantly different between the two sites [ranksum test (Ambrose and Ambrose 1981) T = 69 P lt 005] and also significantlydifferent between zones in each of the nine replicate samples collected at each site
328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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328 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Table 4 Chi-square comparison of myodocopid abundance in three lagoon and three sand troughsamples collected at each of three dates
Lagoon Sand trough
Date Observed Expected Observed Expected
June 1979 91 137 100 54197 257 162 102181 197 94 78
Oct 1979 238 278 150 110294 295 118 1]7109 225 205 89
June 1981 392 386 147 152973 759 85 300418 358 81 141
(1979-1981) (Table 4) [chi-square test of independence (Ambrose and Ambrose1981) chi-square = 592 eight degrees of freedom P lt 005] Total abundanceof individual species (Table 2) was significantly different between the two zones[chi-square test of independence (Ambrose and Ambrose 1981) chi-square =277622 degrees of freedom P lt 0005]
The total number of individuals and individualsspecies varied widely not onlybetween sample areas and periods but also between samples collected during thesame period and the same sample area (Tables 1 5) Sample distributions wereboth skewed and showed kurtosis even after square root and log transformations(tested by methods in Remington and Schork 1970 219) making parametricstatistical analysis inadvisable Inequality of sample variance is probably due inpart to failure to make collecting and sorting efforts equal but the large differencessuggest that the ostracodes are not distributed evenly in the sandy bottom withineach zone
DISCUSSION
Diversity - The small area sampled on the Belize Barrier Reef in the vicinity ofCarrie Bow Cay was characterized by a very high number (51) of myodocopidspecies relative to previously published totals Twenty-six of the Belize myodo-copid ostracodes were undescribed (Cohen 1987) Only 32 Caribbean myodo-copid species seven from Belize had been reported previously (Poulsen 19621965 Wilson 1913 Kornicker and King 1965 Kornicker and Cohen 1978Kornicker 1978 1981 1983a 1984a 1984b 1986a 1986b Cohen 1983 1987Cohen and Morin 1986) One hundred five species of marine podocopid ostra-codes have also been reported from Belize (Teeter 1975)
Thirty-three (65) of the 51 species were found in the sand trough of the fore-reef in an area only about 300 m long and 50 m wide and 29 species (57) inthe sand and rubble zone of the lagoon in an area only about 50 m long and 50m wide By comparison only two species were reported by Komicker (1974) frommultiple quadrant grab samples in Cape Cod Bay Massachusetts Five species ofmyodocopids were reported by Elofson (1941 1969) and Komicker (1987) fromrepeated dredging in the Skagerak and adjacent areas off Sweden Lie (1968)reported only four myodocopid species (two very common) in repeated samplesat 8 stations in Puget Sound Washington USA Baker (1974 1975) listed 25myodocopid species (two very common) from 491 stations on the continental
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
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Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
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-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 329
Table 5 Mean number of specimens standard deviation and variance of three samples collected ineach of two reef zones at each date
Lagoon Sand trough
Date x SD Variance SD Variance
June 1979 156 57 3249 118 38 1444Oct 1979 214 94 8836 158 44 1936June 1981 595 329 108241 104 37 1369
shelf of southern California Only about 25 myodocopid species have been re-ported off the whole coast of Japan and adjacent seas (Hanai et aI 1977) butthere are more undescribed species there (based on collections by me and byHiruta personal communication) Kornicker (1986b) reported 45 myodocopidspecies in the northern Gulf of Mexico On the Bahama Bank off Bimini Kornicker(1958) collected 21 myodocopid species by dredging over a 10 by 12 km areaThere are no data regarding myodocopid diversity on other well developed coralreefs except 13 species of Myodocopida (11 Sarsiellidae) have been reported fromLizard Island on the Great Barrier Reef Australia (Komicker 1981 1982 1983bHall 1987) Sixteen of the 51 myodocopid species at Carrie Bow Cay belong tothe Sarsiellidae
Comparison of Ecological Zones - While many species have overlapping distri-butions in the reef area studied a comparison of repeated benthic samples in thetwo different sandy reef zones shows that the zones differ in myodocopid speciescomposition particularly in relative abundance (ie number of individuals) ofmany species Dominant macro-organisms also differ between the two zones(Rutzler and Macintyre 1982) and Spracklin (1982) found little overlap of hy-droid species between the back-reef (adjacent to my lagoon study area) and fore-reef sand trough at Carrie Bow Cay
The two areas I sampled differ not only in species composition but in relativeabundance of individuals belonging to each of the myodocopid families as wellComb-feeder cylindroleberid and predatory rutidermatid myodocopids wereabundant in the shallow lagoon site but uncommon at the deeper fore-reef sitewhile the scavenging cypridinids and predatory sarsiellids were more common inthe fore-reef site Cylindroleberidinae have been observed to burrow just underthe surface ofthe sediment and form mucous-like tubes around themselves (Mul-Ier 1893 and personal observation) Perhaps there is less food available forstationary feeders in the sand trough of the fore-reef than in the shallow lagoonarea However while the most abundant fore-reef cypridinid Skogsbergia lerneriwas almost absent from my lagoon sediment samples it was abundant in baitedtraps set there at night and other reef zones (Cohen 1987) (Table 1 6) It wasalso abundant in a rubble sample collected in daytime from the adjacent channel(South Water Cut) cutting through the barrier reef about 400 m from the lagoonsampling area more sparsely present in sediment from a variety of ecological sitesin the area (eg lagoon patch reefs and the mangrove-covered Twin Cays) andhas been found from Panama to the Bahamas and Gulf of Mexico in the northwestAtlantic Ocean at depths of 1-130 m All of the sediment samples were madeduring daytime while trap samples were made day and night but mostly at nightOnly two Skogsbergia lerneri were collected in the back-reef in the single day trapsample but night trap samples each contained about 100 to several thousand
330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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330 BULLETIN OF MARINE SCIENCE VOL 45 NO 2 1989
Apparently S lerneri a good swimmer (personal observation) rarely spends day-light hours in sediment of the sand and rubble zone in the lagoon but at nightswims some distance possibly as much as 400 m to feed there
Species abundant in both zones have distributions extending beyond Belizewhile many species collected in specific reef zones have been found only in BelizeSpecies abundant in both zones (though relatively more abundant in one of thezones) are Harbansus paucichelatus Rutiderma new species A Eusarsiella newspecies KO and to a lesser extent E donabbotti Synasterope setisparsa R din-ochelatum and Parasterope muelleri All of these species occur in a number ofother habitats in the vicinity of Carrie Bow Cay five of them have been collectedin other NW Atlantic localities (Table 6) and therefore may be considered rela-tively widespread species On the other hand of the five species restricted tocollections in the back-reef and lagoon Parasterope new species N Postasteropenew species (1 specimen) E new species KE E new species R and Chelicopiaarostrata only the last-mentioned species is known outside Belize Five specieswere restricted to collections in the fore-reef and channel through the reef Vargulanew species SA V new species ST Harbansus new species A Euphilomedesspecies and Eusarsiella new species MO None of these species have been collectedelsewhere and the last three are rare in Belize
Temporal Changes-I hypothesize that Hurricane Greta (September 1978) af-fected the shallow lagoon site more strongly than the deeper site I also hypothesizethat differences between myodocopids of the lagoon and fore-reef sites in partic-ular and that the high number of myodocopid species occurring in the reef areaat Carrie Bow Cay in general are related to storm disturbance Although speciescomposition remained fairly stable in the two zones relative abundance of in-dividuals of species and families changed during the 3 years particularly in thelagoon samples (Table 1 Fig 5) While total abundance increased considerablyin the lagoon site in each collection period following the hurricane total abundancein the fore-reef site first increased slightly and then declined slightly in 1981 Thesmaller fore-reef fluctuations may be a reflection of lesser disturbance or theymay be insignificant The fore-reef site has a smaller number of ostracodes ahigher species diversity and smaller changes in relative abundance than the shallowlagoon site
A comparison of the effects of storm disturbance on ecological stability in thetwo study areas requires an estimation of relative force in the areas (Connell andSousa 1983) Evidence that the sand trough is less disturbed comes from thenature of its sediment In the sand trough very fine sediment reaches a depth of12 m at the axis of the trough while sediment in the sand and rubble zone ispoorly sorted and ranges in size from silt to gravel (Riitzler and Macintyre 1982)Storm disturbance in the sand trough is probably considerably less than in thelagoon because of the greater depth of the sand trough and its protected positionbetween and below the inner and outer fore-reef ridges I observed damage tocorals in the shallow lagoon following Hurricane Greta Kjerfve and Dinnel (1983)found that substantial waves from Hurricane Greta in 1978 approached CarrieBow Cay from the SW strongly affecting the lagoon Riitzler and Macintyre (1982)found considerable damage and movement of the Acropora cervicornis coral usu-ally scattered about in the sand and rubble zone Many lagoon ostracodes musthave been washed away by the hurricane In analysis of the sandy zone in theback-reef of Tobacco Reef 3 km N of and similar to the lagoon site at CarrieBow Cay Macintyre et al (1987) calculated that sand transport takes place duringbrief intervals of storm activity Sand there was suspended and transported during
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
![Page 16: RRI 1SRVWERW 1SLIR€¦ · il j_lmcmn_h]_ i` mj_]c_m ih f[la_l m][f_m! %?ihh_ff [h^ Oiom[( .650&+ Pbcm mno^s cm nb_ `clmn molp_s i` nb_ gsi^i]ijc^ `[oh[ ih [ q_ff ^_p_fij_^ ]il[f](https://reader035.pdfslide.net/reader035/viewer/2022070919/5fb8926bc6b2b641d4548d53/html5/thumbnails/16.jpg)
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 331
Table 6 Myodocopid species collected in quantitative samples and also collected in nonquantitativesamples from other Belize localities or reported from localities outside of Belize
Taxon
CypridinidaeSkogsbergia lerneri
Vargula new species STV new species SAPlerocypridina sex
CylindroleberididaeParasterope new species NP muelleri
Synaslerope setisparsaS new species AAmboleberis americana
Asteropella monambonAclinosela chelisparsa
PhilomedidaeHarbansus paucichelalus
RutidermatidaeRutiderma new species AR dinochelatumR darbyi
R harmanniR cohenae
RUliderma new species B
R new species KO
SarsiellidaeDantya magnificaChelicopia arostralaEusarsiela uncusE new species KEE new species KO
E new species RUE new species CE new species PE new species MJE new species MRE donabbottiE new species KNE new species JE new species CL
Nonquantitative sample locality (and published source of non-Belize localities)
Belize traps all major zones rubble and algae in back-reef andWater Cut Twin Cays lagoon patch reefs Bahamas E PanamaGulf of Mexico (Kornicker 1984a)
Belize only patch reefs South Water Cut rubbleBelize only edge of outer shelf of fore-reefN and S Carolina E and W Florida (Komicker 1984a)
Belize only lagoon Thalassia back-reef South Water CutBelize back-reef spur and groove reef slope Twin Cays patch
reefs plankton English channel West Indies Mauritania Medi-terranean Florida Bahamas Bermuda (Kornicker 1986b)
Bahamas (Kornicker 1986b)Belize only patch reef South Water CutBelize spur and groove reef slope South Water Cay Texas S
Carolina-Florida Bahamas Brazil W Costa Rica W Panama(Kornicker198I)
Belize back-reef Bahamas Puerto Rico Cuba (Kornicker 1981)Belize spur and groove outer ridge W Carrie Bow Cay South
Water Cut Bahamas W Florida Bonaire Venezuela PanamaCura~o (Kornicker 1981)
Belize patch reef South Water Cut N Carolina Florida Baha-mas Gulf of Mexico (Kornicker 1984b)
Belize back-reef Thalassia plankton Texas (Komicker 1983a)Bahamas (Kornicker 1983a)Belize back-reef spur and groove lagoon patch reefs N Carolina-
W Florida Bahamas (Kornicker 1983a)Belize South Water Cay W Panama (Kornicker ( 985)Belize spur and groove outer fore-reef slope Bahamas Florida
Keys (Kornicker 1983a)Belize only lagoon patch reefs back-reef spur and groove outer
ridge South Water Cut lagoon planktonBelize only spur and groove lagoon plankton
Belize only outer reef slopeBelize back-reef Florida Bahamas (Kornicker 1986a)Florida Bahamas (Kornicker 1986a)Belize only back-reefBelize only lagoon patch reefs Thalassia spur and groove reef
slope South Water CutBelize only back-reefBelize only inner outer ridgeBelize only spur and grooveBelize only sand and rubble zoneBelize only Ragged Cay planktonBelize only lagoon patch reef South Water Cut DangrigaBelize only back-reef plankton Ragged CayBelize only Twin CaysBelize only Twin Cays
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
![Page 17: RRI 1SRVWERW 1SLIR€¦ · il j_lmcmn_h]_ i` mj_]c_m ih f[la_l m][f_m! %?ihh_ff [h^ Oiom[( .650&+ Pbcm mno^s cm nb_ `clmn molp_s i` nb_ gsi^i]ijc^ `[oh[ ih [ q_ff ^_p_fij_^ ]il[f](https://reader035.pdfslide.net/reader035/viewer/2022070919/5fb8926bc6b2b641d4548d53/html5/thumbnails/17.jpg)
332 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
Hurricane Greta disturbed to the point of ripple formation during average stormsand undisturbed by tradewinds Highsmith et a1 (1980) found that HurricaneGreta in 1978 caused fragmentation of coral but increased numbers and redis-tribution of colonies (smaller) of Acropora palmata in the vicinity of Carrie BowCay Belize demonstrating that long-term reef calcification and growth rates maybe highest on reefs periodically disturbed by storms of intermediate intensityThey found that Hurricane Greta caused breakage and scouring of corals in allzones of the reef in the vicinity of Carrie Bow Cay to a depth of approximately25 m An analysis of ecological stability should use appropriate temporal andareal scales to examine the degree of constancy in (I) number of organisms andor (2) presence or absence of taxa (Connell and Sousa 1983) Minimum temporalscale is one turnover or lifetime of the organism (Connell and Sousa 1983) Thestudy period from 1978-1981 greatly exceeds the lifetime of the most abundantspecies in the study Skogsbergia lerneri as this species was reared from egg toadult in a coral reef aquarium within 2 months and survived no longer than 2months as an adult (adult females had two to three successive clutches) (Cohen1983) Some specimens of Rutiderma hartmanni A partially reared under similarconditions were brooded as embryos for 53 days and developed from first tosecond instar in another 53 days (Table 7 Cohen 1987) If duration of all fourjuvenile instars in Rutiderma is similar as it is in Skogsbergia complete devel-opment time from egg to adult would be about 9 months in Rutiderma Arealscale in the two study sites probably exceeds the minimum space needed forgenerational replacement of the ostracodes All instars of the most abundantspecies were usually found in single replicates and during each sampling period
Shifts in abundance of dominant taxa occurred not only at the species levelbut at higher taxonomic levels ie generic and familial These alterations maybe correlated with characters (perhaps synapomorphies) shared by all of the speciesbelonging to a higher taxon Possible characters include those affecting dispersaland life history tactics and feeding strategies When one of these characters hasrelatively more importance in natural selection selection could act at the highertaxonomic level as a byproduct of selection at the individual level (all includedindividuals possessing that same character state)
Following the hurricane the most abundant ostracode species included onlymyodocopid taxa with swimming juveniles Parasterope new species N in thelagoon and the cypridinid Skogsbergia lerneri in the fore-reef site Both speciesare better swimmers even as adults (personal observation) than the rutidermatidsand to a lesser extent than sarsiellids the two families which were more abundantbefore and 3 years after the hurricane In the Rutidermatidae only adults haveswimming hairs on the exopod of the second antenna (Kornicker 1985 Cohen1987) the limb used for swimming In the other three families all instars haveswimming hairs on that limb After emerging from the brood chamber instarsof Rutiderma were not observed to swim for 70 days and then swam for only afew millimeters (Cohen 1987) Rutidermatidjuveniles displaced by a storm couldonly crawl and swim slowly while swimming juveniles of other myodocopid taxacould regain a favorable habitat more rapidly by swifter swimming HoweverRutiderma new species A was the dominant species in the lagoon site (and Eu-sarsiella new species P the dominant sarsiellid) only in the year before the hur-ricane The dominant species 3 years after the hurricane was a different rutider-matid R dinochelatum (and sarsiellid E new species KE)
Predatory species were more abundant before and 3 years after Hurricane Gretawhile just after the hurricane comb-feeders and scavengers were relatively moreimportant Specimens of Rutiderma and Eusarsiella often have whole copepods
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
![Page 18: RRI 1SRVWERW 1SLIR€¦ · il j_lmcmn_h]_ i` mj_]c_m ih f[la_l m][f_m! %?ihh_ff [h^ Oiom[( .650&+ Pbcm mno^s cm nb_ `clmn molp_s i` nb_ gsi^i]ijc^ `[oh[ ih [ q_ff ^_p_fij_^ ]il[f](https://reader035.pdfslide.net/reader035/viewer/2022070919/5fb8926bc6b2b641d4548d53/html5/thumbnails/18.jpg)
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 333
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334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
![Page 19: RRI 1SRVWERW 1SLIR€¦ · il j_lmcmn_h]_ i` mj_]c_m ih f[la_l m][f_m! %?ihh_ff [h^ Oiom[( .650&+ Pbcm mno^s cm nb_ `clmn molp_s i` nb_ gsi^i]ijc^ `[oh[ ih [ q_ff ^_p_fij_^ ]il[f](https://reader035.pdfslide.net/reader035/viewer/2022070919/5fb8926bc6b2b641d4548d53/html5/thumbnails/19.jpg)
334 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
nematodes or podocopid ostracodes in their guts (Komicker 1975 Cohen 1987)Possibly their decline in number just after the hurricane was related to a corre-sponding decline in their prey in the lagoon On the other hand finding suitablefood may have been relatively less difficult then for the comb-feeder Parasteropeand scavenger Skogsbergia Hurricane Greta may have increased temporarily theamount of dead animals available to S lerneri a species caught by the thousandsin traps baited with dead fish and which also ate dead amphipods shrimp andconch (Cohen 1983)
Little is known about comparative life histories of myodocopid ostracodes buta literature review suggests that there may be evolutionary constraints by lineageon life history traits for families or genera of these ostracodes (Cohen 1987)(Table 7) Higher myodocopid taxa have a specific number of juvenile instarsWhile clutch size appears to be related to carapace size in many myodocopids(Komicker 1981 1986a) clutch size for all species ofRutidermatidae is restrictedto three to six and for Rutiderma is three to four (probably four as embryos aresometimes lost in handling) (Cohen and Komicker 1987) Clutch size in theCylindroleberidinae is about 1-30 (Komicker 1981) However both Parasteropemuelleri and P new species N had clutch sizes (seven-nine) higher than that ofthe Belize and other rutidermatids with carapaces similar in length to these twocylindroleberids The uniformity of clutch size unrelated to adult size or habitatsuggests that clutch size is ancestrally determined and a systematic character ofthe genus Hines (1986) lists similar constraints for families of crabs and manyother crustacean taxa Reproduction appears to occur year-round in Belize (Cohen1987) Skogsbergia lerneri has a faster development time a larger clutch size andone more instar than Rutiderma new species A and R hartmanni but one moreinstar than they have (Cohen 1987) (Table 7) Although S lerneri has an addi-tional instar it still developed more quickly than the Rutiderma species underthe aquarium conditions Developmental time is unknown for the Belize sarsiellidand cylindroleberid species Parasterope new species N has a larger clutch sizethan Rutiderma species but two additional instars (Cohen 1987) Sarsiellids haveonly four juvenile instars (Hiruta 1977 1978 1980 1983 Komicker 1969Cohen 1987) and some species of Eusarsiella have double clutches one in theovary and one brooded in the marsupium (Kornicker 1986a Cohen 1987) astrategy that would increase reproductive rate (Table 7) Eusarsiella new speciesKE the only Belize species found to have double clutches was also the third mostabundant species and recovered fairly rapidly after the hurricane (Cohen 1987)
CONCLUSIONS
Myodocopid species were diverse in this small area predominant species andfamilies differed between the sand trough of the fore-reef and the sand and rubblezone of the lagoon and species and family composition varied from year to yearFurther myodocopids were less abundant overall but displayed greater speciesrichness and smaller yearly changes in the samples from the deeper less disturbedfore-reef site than in the shallower lagoon site I hypothesize that this temporaland zonal variation was correlated both with zonal differences in relative physicaldisturbance by Hurricane Greta and with taxonomic differences (at the familynot only the specific level) in life history and functional morphology Taxa col-lected in greatest abundance 9 months after Hurricane Greta include the fast-developing Skogsbergia lerneri (a cypridinid) characterized by large clutch sizes
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
![Page 20: RRI 1SRVWERW 1SLIR€¦ · il j_lmcmn_h]_ i` mj_]c_m ih f[la_l m][f_m! %?ihh_ff [h^ Oiom[( .650&+ Pbcm mno^s cm nb_ `clmn molp_s i` nb_ gsi^i]ijc^ `[oh[ ih [ q_ff ^_p_fij_^ ]il[f](https://reader035.pdfslide.net/reader035/viewer/2022070919/5fb8926bc6b2b641d4548d53/html5/thumbnails/20.jpg)
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 335
juveniles with swimming hairs and scavenging diet Less abundant in collections9 months after the hurricane but very abundant 3 years later were slower-de-veloping rutidermatids with smaller clutch sizes juveniles lacking swimminghairs and predatory diet Also more abundant and recovering more quickly thanits congeners was the only sarsiellid species with double clutches The data fromthis study are unfortunately limited but seem to agree with the theories thatdiversity is related to interaction of physical disturbance with biological factors(Huston 1979 Sousa 1984) and highest diversity to intermediate levels of dis-turbance (Connell 1978)
ACKNOWLEDGMENTS
This paper is dedicated to the memory of Donald P Abbott who launched and encouraged me inthe study of invertebrate zoology
I am grateful also to L S Kornicker for encouragement training and advice in the study of myo-docopid ostracodes in general and in this research in particular I thank R Brusca and an anonymousreviewer for many helpful comments on the manuscript and G Hendler and A Jahn for particularideas but responsibility for final content is mine alone For providing laboratory space advice andencouragement I thank K Riitzler and the Division of Crustacea Smithsonian Institution particularlyT E Bowman B Kensley and R B Manning and at the Allan Hancock Foundation University ofSouthern California R C Brusca (now at the San Diego Museum of Natural History)
This is contribution number 239 Caribbean Coral Reef Ecosystems (CCRE) Smithsonian Insti-tution partly supported by a grant from the Exxon Corporation Field work in 1981 was also supportedby a grant from the Lerner Fund American Museum of Natural History I thank the Mead Foundationfor a grant providing funds for a word processor and a trip to study collections at the SmithsonianInstitution Many provided collecting assistance I particularly thank M Carpenter B Kensley RLarson K Riitzler J D Thomas C A Child G Bretchko S Lewis T Rath and J Ferraris TStebbins instructed me in computer graphics For invaluable help in moving collections from Wash-ington D C to California I thank C S Cohen and for general encouragement I thank C S and DM Cohen and C Cohen Leech This paper represents work that is in partial fulfillment of therequirements of the PhD degree at George Washington University
LITERATURE CITED
Ambrose H W III and K P Ambrose 1981 A handbook of biological investigation HunterPublishing Co Winston-Salem North Carolina 170 pp
Baker J H 1974 Distribution ecology and life history of selected myodocopid ostracods from thesouthern California mainland shelf Texas J Science 25 131-132
--- 1975 Distributions ecology and life histories of selected Cypridinacea (MyodocopidaOstracoda) from the southern California mainland shelf PhD Dissertation University of Hous-ton Texas 184 pp
Burke R B 1982 Reconnaissance study of the geomorphology and benthic communities of theouter barrier reef platform Belize Pages 509-526 in K Riitzler and I Macintyre eds The AtlanticBarrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communities SmithsonianContr Mar Sci 12
Cohen A C 1983 Rearing and postembryonic development of the myodocopid ostracode Skogs-bergia lerneri from coral reefs of Belize and the Bahamas J Crust BioI 3 235-256
--- 1987 Systematics life history and distribution of myodocopid ostracodes on the barrierreef at Carrie Bow Cay Belize PhD Dissertation George Washington University WashingtonDC 620 pp
-- and L S Komicker 1987 Catalog of the Rutidermatidae (Crustacea Ostracoda) Smithso-nian Contr Zool 449 1-11
--- and J G Morin 1986 Three new luminescent ostracodes of the genus Vargula (MyodocopidaCypridinidae) from the San Bias region of Panama Contr Science Nat Hist Mus Los AngelesCo 373 1-23
Connell J H 1978 Diversity in tropical rain forests and coral reefs Science 199 1302-1310--- and W P Sousa 1983 On the evidence needed to judge ecological stability or persistence
Am Nat 121 789-824Elofson O 1941 Zur Kenntnis der marinen Ostracoden Schwedens mit besonderer Beriicksichtigung
des Skagerraks Zoologiska Bidrag fran Uppsala 19 215-534
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
![Page 21: RRI 1SRVWERW 1SLIR€¦ · il j_lmcmn_h]_ i` mj_]c_m ih f[la_l m][f_m! %?ihh_ff [h^ Oiom[( .650&+ Pbcm mno^s cm nb_ `clmn molp_s i` nb_ gsi^i]ijc^ `[oh[ ih [ q_ff ^_p_fij_^ ]il[f](https://reader035.pdfslide.net/reader035/viewer/2022070919/5fb8926bc6b2b641d4548d53/html5/thumbnails/21.jpg)
336 BULLETIN OF MARINE SCIENCE VOL 45 NO2 1989
1969 Marine Ostracoda of Sweden with special consideration of the Skagerrak IsraelProgram Sci Trans Ltd 286 pp Translation from German of 1941 publication
Hall S J 1985 Four new species of myodocopine ostracodes (Sarsiellidae) from Lizard IslandNorth Queensland J Crust BioI 5 500-522
--- 1987 New species of Sarsiella and Anscottiella (Ostracoda Myodocopina) from LizardIsland North Queensland J Crust BioI 7 738-763
Hanai T N Ikeya K Ishizaki Y Sekiguchi and M Yajima 1977 Checklist of Ostracoda fromJapan and its adjacent seas Univ Mus Univ Tokyo Bull 12 1-110
Hartmann G 1964 Zur Kenntnis der Ostracoden des Roten Meeres Kieler Meeresforsch 20 35-127
Highsmith R c A C Riggs and C M DAntonio 1980 Survival of hurricane-generated coralfragments and a disturbancc model of reef calcificationgrowth rate Oecologia 46 322-329
Hines A H 1986 Larval patterns in the life histories of brachyuran crabs (Crustacea DecapodaBrachyura) Bull Mar Sci 39 444-466
Hiruta S 1977 A new species of the genus Sarsiella Norman from Hokkaido with reference to thelarva] stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 44-60
--- 1978 Redescription of Sarsiella misakiensis Kajiyama from Hokkaido with reference tothe larval stages (Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 21 262-278
--- 1979 A new species of the genus Bathyleberis Kornicker from Hokkaido with reference tothe larval stages Ostracoda Myodocopina) J Fac Sci Hokkaido Univ Ser VI Zool 22 99-121
--- 1980 Notes on the life history of Sarsiellajaponica Hiruta (Ostracoda Myodocopina) JHokkaido Univ Educ (Sect lIB) 3](1) 41-45
-- ]983 Post-embryonic development of myodocopid Ostracoda Pages 667-677 in R Mad-docks ed Applications of Ostracoda Proc Eighth International Symposium on Ostracoda July26-29 1982 Univ Houston
Hulings N ]969 The ecology of the marine Ostracoda of Hadley Harbor Massachusetts withspecial reference to the life history of Paraslerope pollex Kornicker ]967 Pages 412-422 in JW Neale ed The taxonomy morphology and ecology of Recent Ostracoda Oliver and BoydEdinburgh
Huston M 1979 A general hypothesis of species diversity Am Nat 113 81-10 IInternational code of zoological nomenclature 1985 Third edition W Ride C Sabrosky G
Barnardi and R Melville eds University of California Press Berkeley 338 ppKjerfve B and S Dinnel 1983 Hindcast hurricane characteristics on the Belize barrier reef Coral
Reefs I 203-207Kornicker L S 1958 Ecology and taxonomy ofrecent marine ostracodes in the Bimini area Great
Bahama Bank Publ Inst Mar Sci Univ Texas 5 194-300--- 1969 Morphology ontogeny and intraspecific variation of Spinacopia a new genus of
myodocopid ostracod (Sarsiellidae) Smithsonian Contr Zool 8 I-55--- 1974 Ostracoda (Myodocopina) of Cape Cod Bay Massachusetts Smithsonian Contr Zool
173 1-20-- 1975 Antarctic Ostracoda (Myodocopina) Smithsonian Contr Zool 163 1-110--- 1978 Harbansus a new genus of marine Ostracoda and a revision of the Philomedidae
(Myodocopina) Smithsonian Contr Zool 260 1-78--- 1981 Revision distribution ecology and ontogeny of the ostracode subfamily Cyclastero-
pinae (Myodocopina Cylindroleberididae) Smithsonian Contr Zool 319 1-548--- 1982 Allernochelala Iizardensis a new species of myodocopine ostracode from the Great
Barrier Reef of Australia (Rutidermatidae) Proc BioI Soc Washington 95 793-806--- 1983a Rutidermatidae of the continental shelf of southeastern North America and the Gulf
of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 371 1-86--- 1983b Harbansus slatteryi a new species of myodocopine ostracode from the Great Barrier
Reef of Australia (Philomedidae) Proc BioI Soc Washington 96 181-188--- 1984a Cypridinidae of the continental shelves of southeastern North America the northern
Gulf of Mexico and the West Indies (Ostracoda Myodocopina) Smithsonian Contr Zool 40 I1-37
--- 1984b Philomedidae of the continental shelf of eastern North America and the northernGulf of Mexico (Ostracoda Myodocopina) Smithsonian Contr Zool 393 1-77
--- 1985 Sexual dimorphism ontogeny and functional morphology of Ruliderma harlmanniPoulsen 1965 (Crustacea Ostracoda) Smithsonian Contr Zool 408 1-28
--- 1986a Sarsiellidae of the western Atlantic and northern Gulf of Mexico and revision ofthe Sarsiellinae (Ostracoda Myodocopina) Smithsonian Contr Zool 415 1-217
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
![Page 22: RRI 1SRVWERW 1SLIR€¦ · il j_lmcmn_h]_ i` mj_]c_m ih f[la_l m][f_m! %?ihh_ff [h^ Oiom[( .650&+ Pbcm mno^s cm nb_ `clmn molp_s i` nb_ gsi^i]ijc^ `[oh[ ih [ q_ff ^_p_fij_^ ]il[f](https://reader035.pdfslide.net/reader035/viewer/2022070919/5fb8926bc6b2b641d4548d53/html5/thumbnails/22.jpg)
COHEN OSTRACOD DISTRIBUTIONS OF CARRIE BOW CAY 337
-- 1986b Cylindroleberididae of the western North Atlantic and the northern Gulf of Mexicoand zoogeography of the Myodocopina (Ostracoda) Smithsonian Contr Zool 425 1-139
--- 1987 Ostracoda from the Skagerrak North Sea (Myodocopina) Proc BioI Soc Washington100 876-891
--- and A C Cohen 1978 Dantyinae a new subfamily of Ostracoda (Myodocopina Sarsiellidae)Proc BioI Soc Washington 91 490-508
--- and C E King 1965 A new species of luminescent Ostracoda from Jamaica West IndiesMicropaleontology II 105-110
Lie U 1968 A quantitative study of benthic infauna in Puget Sound Washington USA in 1963-1964 Fiskedirektoratets Skrifter serie HavUnders0kelser Bergen 14 229-556
Macintyre 1 G R R Graus P N Reinthal M M Littler and D S Littler 1987 The barrierreef sediment apron Tobacco Reef Belize Coral Reefs 6 1-12
Miiller G W 1893 Ueber lebensweise und entwick]ungsgeschichte der ostracoden SitzungsberichteKoeniglich Preussischen Akad Wiss Berlin 23 355-381
Okada Y and K Kato 1949 Studies on luminous animals in Japan III Preliminary report on thelife history of Cypridina hilgendorfi Bull Biogeog Soc Jap ]4 21-25
Poulsen E M 1962 Ostracoda-Myodocopa I Cypridiniforrnes-Cypridinidae Dana Report 57 1-414
-- 1965 Ostracoda-Myodocopa 2 Cypridiniformes-Rutidermatidae Sarsiellidae and Aste-ropidae Dana Report 65 1-484
Remington R D and M A Schork 1970 Statistics with applications to the bio]ogical and healthsciences New Prentice-Hall Inc Englewood Cliffs New Jersey 418 pp
Riitz]er K and I G Macintyre 1982 The habitat distribution and community structure of thebarrier reef complex at Carrie Bow Cay Belize Pages 9-45 in K Riitzler and I Macintyre edsThe Atlantic Barrier Reef Ecosystem at Carrie Bow Cay Belize I Structure and communitiesSmithsonian Contr Mar Sci 12
Sousa W P 1984 The role of disturbance in natura] communities Ann Rev Ecol Syst 15 353-391
Spracklin B 1982 Hydroidea (Cnidaria Hydrozoa) from Carrie Bow Cay Belize Pages 239-252in K Riitzler and 1 Macintyre eds The Atlantic Barrier Reef Ecosystem at Carrie Bow CayBelize 1 Structure and communities Smithsonian Contr Mar Sci 12 239-252
Teeter J W 1975 Distribution of Holocene marine Ostracoda from Belize Pages 400-498 in KF Wantland and W Pusey III eds Studies in geology No2 Belize Shelf-carbonate sedimentsclastic sediments and ecology Am Assoc Petrol Geol Tulsa Oklahoma
Wilson C B 1913 Crustacean parasites of West Indian fishes and land crabs with descriptions ofnew genera and species Proc US Nat Mus 44 189-277
DATE ACCEPTED February 6 1989
ADDRESS Division of Life Science Los Angeles County Museum of Natural History 900 ExpositionBoulevard Los Angeles California 90007 (affiliation Biology Dept University of California LosAngeles)
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