THE UNIVERSITY OF NORTH CAROLINA W A T E R R E S O U R C E S R E S E A R C H I N S T I T U T E

N O R T H CAROLINA STATE UNIVERSITY T H E UNIVERSITY O F N O R T H CAROLINA at RALEIGH at CHAPEL HILL

124 Riddick Building North Carolina State University Raleigh, North Carolina, 27607

June 7 , 1971

TO : Whom It May Concern

F RON : David M. Howells

SUBJECT: I n s t i t u t e Report E?o, 49 - "Migration and Metabolism i n a Stream Ecosystem,'"y Charles A. S. Hall

lJhi le t h i s r e p o r t s and i n t e r p r e t s f indings on f i s h migra t ion and stream metabolism i n New Hope Creek, i t has a much broader a p p l i c a t i o n t o Piedmont streams i n general ,

I4r. Hal l ' s conclusions and recornmendatfons, pages xv t o xix, r e l a t e s t o d iu rna l v a r i a t i o n s i n d issolved oxygen and importance of pre-da-m sampling, b a r r i e r s t o f i s h migrat ion, and stream c l a s s i f i c a t i o n f o r research and o the r s c i e n t i f i c purposes.

Enclosure

MIGRATION AND JXIETABOLIISM I N A STREAM ECOSYSTEM

by

Charles A, S. Hal l

A t h e s i s submitted t o t h e f a c u l t y of the Universi ty of North Carolina a t Chapel H i l l i n p a r t i a l f u l f i l l m e n t of requirements f o r the degree of Doctor of Philosophy i n t h e Department of Zoology, September 1970.

TQe work upon which t h i s pub l i ca t ion i s based was supported i n p a r t by funds provided by t h e Office of Water Resources Research, Department of t h e I n t e r i o r , through the Water Resources Research I n s t i t u t e of the Universi ty of North Carolina a s authorized under t h e Water Resources Research Act of 1966.

Pro j e c t No. B-007-NC Matching Grant Agreement mo. 14-01-0001-1933

Professor H. T. Odum - Thesis Advisor Professor Charles M. Weiss - Projec t Direc tor

Department of Environmental Sciences and Engineering School of Public Health

Universi ty of North Carolina a t Chapel

February 1971 c 'I

TABLE OF CONTENTS

. . . . . . . . . . . . . . . . . . . . . . . . . . LIST OF TABLES, ix

. . . . . . . . . . . . . . . . . . . . . . . . . LIST OF FICTURFS. xi

INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . I.

Theories f o r Migrat ion. . . . . . . . . . . . . . . . . . . . 2 P4igratiion t o a r ~ i d unfavorable condi t ion , Migration and reproduct ion , Migration and optimal use of f l u c t u a t i n g environments.

. . . . . . . . Role of Migrat ing Animals i n Mineral Cycl ing. 7

Previous S tud ie s on t h e Movements of F ishes . . . . . . . . . 8 Movements t o t a l l y wi th in one s t ream, Movements of f i s h e s i n s t reams wi th ad jo in ing l akes , Movements of f i s h e s be- tween f r e s h and salt water , Movements of f i s h e s i n t h e open sea .

. . . . . . . . . . . . . . . . . . . . Statement of Purpose. 14

. . . . . . . . . . . . . . . . . Descr ip t ion of Study Area. 14 Q u a l i t a t i v e energy flow diagram f o r migra t ion i n New Hope Creek.

MATERIALSANDMETHQDS. . . . . . . . . . . . . . . . . . . . . . . 2 2

. . . . . . . . . . . . . . . . . Physical and Chemical Data. 2 2 C h a r a c t e r i s t i c s of t h e sampling s t a t i o n s , Discharge, Stream morphology - depth and width, I n s o l a t i o n , Stream temperature, To ta l phosphorus i n water , Phosphorus i n organisms, To ta l n i t rogen i n water , Stream conduc t iv i ty , Discharge of leaves .

M e t a b o l i c S t u d i e s . . . . . . . . . . . . . . . . . . . . . . . 31 Dissolved oxygen, Winkler method; Dissolved oxygen, Gal- van ic probe method; Di f fus ion r a t e s ; Gross community metabolism: Two s t a t i o n a n a l y s i s , S ingle curve method; Estimate of metabol.ism from pH changes; Computer program f o r e s t ima t ing community metabolism from d i u r n a l oxygen curves .

Fish Sampling Procedures and Apparatus. . . . . . . . . . . . 53 Design of weirs and traps, Check on possible sampling bias in up a.nd down traps, Check on ra:e of fish escape from traps, Sampling modifications for low water, Sampling modifications during high waters, Methodology of handling species and species groups for analysis, baily fish sampling procedure.

Physical Data. . . . . . . . . . . . . . . . . . . . . . . . 7 4 Stream morphology, Stream level and discharge rate, Stream temperatures, Light intensity at the surface of the stream, Leaf discharge, Total phosphorus, Nitrogen, Stream conductivity.

Metabolicstudies. . . . . . . . . . . . . . . . . . . . . . . 90 Daily variations in oxygen, Annual variations in metabo- lism, Spatial variations in metabolism, Annual and spatial variations in P/R ratio.

FishMovements. . . . . . . . . . . . . . . . . . . . . . . . 110 Analysis of all species considered together: Principal sampling station, April 1968 - June, 1970, Seasonal variations in movements, Cumulative occurrence of species vs. cumulative occurrence of individuals, Diversity of moving animals, Movements at other stations on New Hope Creek, Movements at Morgan Creek; Analysis by each spe- cies: Numerical and weight contribution of each species to migration, Seasonal patterns of movements for each taxonomic class, Evidence of spawning condition of fish at different times of the year, Recaptures of marked fish, Tagged fish returns analysis, Daily concentration of moving animals.

DISCUSSION. 174 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seasonal Patterns of Metabolism. . . . . . . . . . . . . . . . 174

P/R ratio and heterotrophic regime.

Spatial Distribution of Metabolism. . . . . . . . . . . . . . 175 Dilution of resources with depth.

Comparison With Some Other Studies. . . . . . . . . . . . . . 179 Patterns of Fish Movement. . . . . . . . . . . . . . . . . . . 182

Movements of different species, Movements and floods, Movements of juvenile fishes, Differential movements of different-sized fish.

Com~arisons of Ewrgy Rwlgets . . . . . . . . . . . . . . . . 182 Energy of ruaning water, Energy of bi~logical metabolism, Energy of insolation, Energy of fish metabolism, Energy of migration,

Net Contributions of Migration to Headwaters and Turnover Rate. . . . . . . . . . . . . . . . . . . . . . . . . . . . .I92

Comparisons of migration in New Hope Creek with salmon migration.

. Possible Adaptive Values of Migrations in New Hope Creek. .I93 Migration as a coupling function, Interaction of yield and organization.

Some Other Animal Migrations and Environmental Energy Patterns . . . . . . . . . . . . . . . . . . . . . . . . . . . 196

. . . . . New Hope Creek Watershed Annual Phosphorus Budget. 197

. . . . . . . . . . Analog Simulation of a Migration Model. .202 Analog results and discussion.

LITERATURE CITED. . . . . . . . . . . . . . . . . . . . . . . . . 212 APPENDIX A: DIFFUSION PROCEDURES USED IN NEW HOPE CREEK METABOLISMSTUDIES . . . . . . . . . . . . . . . . . . . . . . . . 227

AESTRACT

Fish migrat ion and t o t a l stream metabolism were s tud ied i n New

Hope Creek, North Caro l ina , from Apr i l , 1968 t o June, 1970. Up-

s t ream and downstream movement of f i s h e s was monitored us ing weirs

with t r a p s . Most of t h e 27 spec i e s had a c o n s i s t e n t p a t t e r n of

l a r g e r f i s h moving upstream and sma l l e r f i s h moving downstream. Both

upstream and downstream movements were g r e a t e s t i n t he sp r ing . For

example, i n t h e s p r i n g of 1969, a d a i l y average of 7 f i s h weighing a

t o t a l o f 1081 grams were caught moving upstream, and 17 f i s h , weighing

a t o t a l o f 472 grams, were caught moving downstream. Although more

moved downstream than up, t h e l a r g e r average s i z e of t h e f i s h moving

upstream r e s u l t e d i n a l a r g e t r a n s f e r of f i s h mass upstream.

Diurnal oxygen s e r i e s were run t o measure t h e metabolism of t h e

aqua t i c community. Gross photosynthes is ranged from 0.21 t o almost

9 g m-2 day-1 ~ ~ / m ~ / d a ~ ) , and community r e s p i r a t i o n from 0 . 4 t o

13 g m-2 day-1 a t t h e p r i n c i p a l sampling s t a t i o n and both were

h ighes t i n t h e s p r i n g . Area va lues of metabolism were s i m i l a r f o r

d i f f e r e n t p a r t s of t h e s t ream, bu t both product ion pe r volume and

r e s p i r a t i o n pe r volume were much l a r g e r nea r t h e headwaters than

f a r t h e r downstream. This was appareqt ly due t o t h e d i l u t i n g e f f e c t

of t h e deeper water downstream. Migration may allow populat ions t o

t ake advantage of such d i f f e rences i n p r o d u c t i v i t y by maintaining

young f i s h i n a r eas of high p r o d u c t i v i t y .

An energy diagram was drawn comparing energies of insolation,

currents, photosynthesis, respiration, fish populations, and migra-

tions. Parts of this model were simulated on an analog computer.

Input energies from insolation and stream flow were similar. About

0.14 percent of the total respiration of the stream was from fish

populations, and over one year about 0.01percent of the total energy

used by the ecosystem was used for the process of migration. If it

is assumed that upstream migration is necessary to maintain upstream

stocks, which may be periodically decimated by droughts, the migration

energy has an amplifying value of 14.

ACKNOWLEDGEMENTS

The d i s s e r t a t i o n was done under t h e superv is ion of Howard T.

Odum and Charles M. Weiss, with f a c i l i t i e s a t New Hope Creek pro-

vided through t h e cour tesy of t h e Duke Fores t Administrat ion and

with cooperat ion of t he North Caro l ina Wi ld l i f e Resources Commission

by cour tesy of Harry Cornel l . E l izabe th Mcblahan, Edward J. Kuenzler,

and Joseph Bailey served on t h e superv isory committee.

F inancia l support was provided by t h e Water Resources Research

I n s t i t u t e , Univers i ty of North Carol ina, Grant B-007-NC (Office of

Water Resources Research, United S t a t e s Department of t h e I n t e r i o r )

t o Charles M. Weiss, 2nd AEC Contract AT-(40-1)-36666, H. T. Odum,

p r i n c i p a l i n v e s t i g a t o ~ , and an a l l o c a t i o n from t h e North Carol ina

Computer Center ,

Thomas P. Stevenson, Wayne Frankl in , John Floyd and o t h e r s

a s s i s t e d i n t h e o f t e n arduous f i e l d work and d a t a processing. John

Gum and o t h e r s a t tl7e Univers i ty of North Caro l ina Computation

Center a ided with d i g i t a l computer programs,and Larry Burns and Fred

W a f aided wi th an analog program. Tony Owens of t h e Department of

Environmental Sciences and Engineering, Univers i ty of North Caro l ina

d id phosphorus and n i t rogen ana lyses . Dennis Whigham provided S nso-

l a t i o n c h a r t s . Joseph Bai ley of Duke Univers i ty aided i n t h e

i d e n t i f i c a t i o n of f i s h e s .

Peter Larkin, of the Institute of Ecology, University of

British Columbia, and David Narver, of the Fisheries Research

Board of Canada,provided suggestions and funds for salmon studies

at Vancouver, Nanairno, and Babine Lake, British Columbia in the

summer of 1969.

My advisor, Dr. Howard T. Odum, was the impetus and nucleus

for the excitement in ecology that I have experienced at the

University of North Carolina for the past three years. I an

grateful for having had this opportunity.

LIST OF TABLES

Tables

Some C h a r a c t e r i s t i c s of t h e Various Sampling S t a t i o n s

D r i f t of Oxygen Recorder Over One o r Seve ra l Days

Modif icat ions i n Basic Trapping Procedure

Catch of F i s h i n 'Sideways' Traps

Floods i n New Hope Creek That Affected Sampling

Fishes Captured i n New Hope Creek and Groupings Used t o S impl i fy Analysis

Organisms Other Than F i s h Captured i n New Hope Creek

Depth and Width P r o f i l e f o r 300 m Below Concrete Bridge, May 23, 1970

Depth and Width P r o f i l e f o r 1.8 km Above Concrete Bridge S t a t i o n , A p r i l , 1969

Depth and Width P r o f i l e f o r 900 m Above Wood Bridge S t a t i o n , May 13, and 23, 1970

Depth and Width P r o f i l e For t h e Zone l O O O m Above Blackwood Sampling S t a t i o n , May 18, 1970

Light I n t e n s i t y a t Surface of New Hope Creek

T o t a l Phosphorus (Dis solved and suspended) i n New Hope Creek a t Concrete Bridge S t a t i o n

Nitrogen Compounds i n New Hope Creek

T o t a l Community Metabolism f o r New Hope Creek, Concrete Bridge S t a t i o n , Apr i l , 1 9 6 8 - ~ a ~ , 1970

T o t a l Community Metabolism f o r New Hope Creek, Wood Bridge S t a t i o n , June, 1968-August, 1969

Page --, -

17. T o t a l Community Metabolism f o r New Hope Creek, Blackwood S t a t i o n , Febrxary, 1969-February, 1970

18a. Average Dai ly F i s h Movements by Month

18. Summary of Trap Catches a t Wood Bridge and Jungle S t a t i o n s , New Hope Creek

19. Summary of Trap Catches a t Morgan Creek

20. Minimum, Maximum and To ta l Mass and T o t a l Numbers of Each Species o r Group Sampled a t P r i n c i p a l S t a t i o n , Mew Hope Creek

21. Average Mass of Animals Moving a t P r i n c i p a l S t a t i o n

22 . Evidence of Spawning Conditlcn

23. Recapture of Marked F i s h

24. Recapture of Tagged F i s h

25. Concentrat i o n ( ~ a i l y ) o f Moving Organisms

26. Concentrat ion of Phosphorus a t D i f f e r e n t S t a t i o n s on Same Dates

27. Metabolism i n Some Other Unpolluted Streams

28. Metabolism of Some Se lec t ed Lakes and Maine Waters

29. Annual Movement of Phosphorus i n New Hope Creek: June 14 , 1968- ~ u n e 13 , 1969

A-1 . D i f fus ion Constants Derived from Diurnal Oxygen Data

A-2. Predic ted Values f o r Di f fus ion Constant f o r Ne7~ Hope Creek Above Concrete Bridge S t a t i o n Using Formula Based on Average Depth and Veloc i ty

A-3. Basis f o r Calcu la t ions of Di f fus ion Coef f i c i en t from Dome Measurements

A-4. Es t imates of Di f fus ion Constant ( K ) Obtained Using t h e Dome Method f o r Representa t ive Pools and R i f f l e s Above Concrete Bridge S t a t i o n

LIST OF FIGURES

Page --.--

1. Locat ion of sampling s t a t i o n s on New Hope and Morgan Creeks, North Carol-ina.

2 . A t y p i c a l r i f f l e s t r e t c h of New Hope Creek, l o c a t e d j u s t above t h e Concrete Bridge S t a t i o n .

3. Energy c i r c u i t diagram f o r migra t ion i n New Hope Creek.

4. Symbols used i n energy network diagrams, from H. T , Odum (1967a) .

5. Water s t a g e vs . discharge .

6 . T o t a l mass of l eaves (dry weight ) discharged pe r day i n s t ream flow a t Concrete Bridge S t a t i o n vs . s t a g e l e v e l ( o r d i n a t e ) i n cent imeters above zero flow.

7. Cork and tub ing device t o f i l l oxygen b o t t l e with- ou t a i r mixing.

8. Comparison of probe and Winkler oxygen values over a 24 hour pe r iod , J u l y 25, 1969, a t Concrete Bridge S t a t i o n .

9. V a r i a t i o n i n oxygen meter readings wi th cons tan t d i sso lved oxygen and varying temperature.

LO. Comparison of d i f f e r e n t d i f f u s i o n cons t an t s ob ta ined i n t h i s s tudy .

11. S i m i l a r i t y of oxygen curves one hour ' s flow d i s t ance a p a r t a t Blackwood S t a t i o n , February 14 , 1-970.

12 . S i m i l a r i t y of oxygen curves one h o u r ' s flow d i s t ance a p a r t a t Concrete Bridge S t a t i o n , February 14, 1970.

13. A represen ta t ive sample of s i n g l e s t a t i o n analys is f o r community metabolism i n New Hope Creek, February 14, 1970, a s conducted and p l o t t e d by the UNC CALCOMF p l o t t e r .

14. Various l i n e s drawn t o represent daytime r e s p i r a t i o n .

15. Carbon dioxide t i t r a t i o n of New Hope Creek water f o r metabolic s t u d i e s .

16. Est imation of metabolism i n New Hope Creek, Diurnal pH method.

17. Ear ly design of f i s h weir (~ecember , 1968).

18. Design of f i s h weirs used i n New Hope Creek.

19. Big Pool sampling s t a t i o n , looking downztream during normal sp r ing flow.

20. "Sid.eways" f i s h sampling arrangement.

2 Design of f i s h t r a p used i n New Hope Creek.

22. a . "sideways" and b . "double reverse" weirs used. t o t e s t poss ib le sampling b i a s i n normal t r a p arrangement.

23. New Hope Creek d.uring drought (~ep tember , 1968) . 24. New Hope Creek a t Concrete Bridge S t a t i o n during

f lood.

25. Overrun of weir during severe f lood a t Big Pool S t a t i o n on Apr i l 14 , 1970.

26. Daily water s tage l e v e l ; i n cm above zero flow, of New Hope Creek a t Concrete Bridge S ta t ion .

27. Mean d a i l y temperatures f o r New Hope Creek during t h i s study.

28. Typical d iu rna l oxygen curve f o r spring, Concrete Bridge S ta t ion .

29. Typical d iu rna l oxygen curve f o r spring, Wood Bridge S ta t ion .

x i i i

30. Typical d iu rna l oxygen curve f o r spr ing , Blac1cwood S t a t i o n .

31. Typical d iu rna l curve f o r l a t e f a l l , Concrete Bridge S ta t ion .

32. Typical d iu rna l curve f o r winter , Wood Bridge S t a t i o n .

33. Typical d iu rna l oxygen curve f o r l a t e f a l l , Blackwood S ta t ion .

34. Annual- v a r i a t i o n i n metabolism, Concrete Bridge S ta t ion , New Hope Creek, Apr i l , 1968 - May, 1970.

35. Annual v a r i a t i o n i n metabolism, Wood Bridge S ta t ion , New Hope Creek, June, 1968 - August, 1969,

36. Annual v a r i a t i o n i n metabolism, Blackwood S ta t ion , New Hope Creek, February, 1969 - February, 1970.

37. Seasonal v a r i a t i o n of photosynthesis r e s p i r a t i o n r a t i o a t Concrete Bridge S ta t ion .

38. Average d a i l y migrat ion by month.

39. Cumulative species versus cumulative individuals t rapped a t p r i n c i p a l sampling s t a t i o n ; only f i s h e s a r e included.

40. Upstream and downstreax movement of each species o r species group i n New Hope Creek by s i ~ e i n t e r v a l .

41. Average d a i l y movement f o r each number, by species .

42. Seasonal p a t t e r n s of i n s o l a t i o n under a hardwood canopy, Duke Fores t , near New Hope Creek.

43. Growth of tagged f i s h , New Hope Creek.

44. Annual movement and metabolism of f i s h popula- t i o n s i n the headwaters of New Hope Creek above t h e Concrete Bridge.

45. Energy flow diagram f o r upstream (middle s e t of modules) and downstream (lowermost s e t of modules) of New Hope Creek.

46. Diagram of phosphorus flows i n New Hope Creek watershed.

47. Energy flow diagram f o r analog computer model.

xiv +r

A

48. Analog symbols represent ing the energy pathways i n Figure 47.

49. Analog output of energy pulse generator.

50. Analog simulat ion of annual energy acc rua l t o populat ions of f i s h e s i n New Hope Creek.

A-1. Use of c l e a r p l a s t i c dome t o measure diff 'usion constant .

- 2 Use of p l a s t i c dome t o measure d i f fus ion .

M i e i o n and Metabolism in a Stream Ecosystem - ---.. -"- - "- -" - ----

Preserving and Enhancing the Qualities of the Waters of North Carolina

In the period April, 1968 to June, 1970 an intensive investi-

gation was made in New Hope Creek, in the stretch where it flows

through Duke Forest, to establish the relationship between fish

migration and the total stream metabolism. New Hope Creek at this

particular point may be the only stream in the Research Triangle

area of North Carolina where studies of relatively natural conditions

can be carried out. Nearly all other streams in the region are

either polluted or are too small for any extended studies. The

location within Duke Forest, with controlled access made the region

particularly desirable for studies in what is essentially a natural

outdoor laboratory.

The basic investigation consisted of monitoring up and down-

stream movement of fishes, using weirs with traps. Of the 27 species

collected, most had a consistent pattern of the larger fish moving

upstream and smaller fish moving downstream. Movement in both

directions was greatest in the spring. For example, in the spring

of 1969, a daily average of 7 fish weighing a total of 1087 gms

were trapped moving upstream and 17 fish weighing a total of 472 gms

were caught moving downstream. Although more fish moved downstrean

than up, the larger average size of the fish moving upstream resulted

in a larger transfer of fish mass upstrsam. Associated with the fish

movement studies, the metabolism of the aquatic communiJiy was determined

using the technique of diurnal oxygen measurements. Gross photo-

synthesis ranged from 0.21 to almost 9 g/m2/day and community respira-

tion from 0.4 to 13 g/m2/day. All measurements of this nature were

highest in the spring. Both production and respiration per volume

were much larger near the headwaters than farther downstream. This

was apparently a result of the diluting effect of the deeper dater

downstream. Migration appeared to allov .the fish population to ta,ke

advantage of such differences in productivity by rnzlntaSning :joung

fish in areas of high productivi5y.

A total energy diagram was devi-ed cornparin& energies of insola-

tion, currents, photosynthesis, respiration, fish gopula tions and

migrations. When this mode? was simulated on an analog computer, it

was determined that input energies frorr, insolation and streax flow

were similar with about 0.14 percent of the total respiration of the

stream derived from fish ponulations. Over a period of a year a>out

0.01 percent of the total energy used by the ecosysiem N a s consumed

in the process of migration. It can be a s a m c d ihai t h 2 upstream

migration is necessary to maintain upstream fish stocks xhich may be

periodically decimated by drought conditions. Migration el?ergy appears

to have an amplifying value of 14.

There a r e c e r t a i n l e s sons t h a g may be l ea rned from t h e preceding

i n v e s t i g a t i o n , which a r e of value f o r p re se rv ing and enhancing t h e

q u a l i t i e s of t h e waters of North Caro l ina . As found i n New Hope Creek

and probably f o r most o t h e r streams of piedmont North Carol ina, t h e

d i u r n a l d i sso lved oxygen v a r i a t i o n may be q u i t e l a r g e up t o t h r e e o r

f o u r m g / l . Thus, c r i t e r i a f o r oxygen i n any s tream must be e s t a b l i s h e d

a s a minimum pre-dawn value s i n c e t h i s could very w e l l be t h e l i m i t a -

t i o n f o r any aqua t i c organism r e q u i r i n g oxygen s i n c e they must l i v e i n

t h e s t ream 24 hours a day. The d a i l y f l u c t u a t i o n s i n oxygen were

found t o be g r e a t e s t i n shallow water . This c h a r a c t e r i s t i c may be

of cons iderable s i g n i f i c a n c e i n water q u a l i t y dec is ions s ince :

( 1 ) a s s t reams become more shallow dur ing summer low waters , t h e

d i f f e r e n c e between day and n igh t oxygen values become l a r g e r ;

( 2 ) upstream, t h e r e a r e gene ra l ly more shallow reg ions of t h e s t ream

wi th g r e a t e r day-night d i f f e r ences i n oxygen content .

Since aqua t i c organisms us ing d isso lved oxygen r e q u i r e more a t

h igher temperatures , suxmer condi t ions , t h e r e f o r e , may c r e a t e a

c r i t i c a l circumstance due t o ( a ) lowering t h e s o l u b i l i t y of oxygen

i n water and (b ) i nc reas ing t h e oxygen requirements of organisms and

( c ) i nc reas ing t h e d a i l y f l u c t u a t i o n of oxygen a s b i o t i c components

of t h e s t ream ecosystem become more crowded i n shal lower water . It i s

t h e r e f o r e i n d i c a t e d t h a t t h e oxygen requirements f o r streams be s e t

f o r minimum condi t ions a t one hour before s u n r i s e during per iods of

h ighes t temperature and/or lowest waters , g e n e r a l l y i n August. It

would thus be indicated that if any pollution is suspected in a

stream, it becomes even more critical to establish the pre-dawn

oxygen level with reference to the quality of the particular body

of water.

' Regional planning of aquatic wastes disposal should take into

account the potentially greater stress that is imposed on the shallower

regions of streams and rivers. This implies that the establishment

of regional plans for economic growth, a basic principle should be

one of not introducing industries and waste disposal facilities on

the headwaters of rivers.

It was also determined from the investigation on New Hope Creek

that many fishes in Piedmont streams have distinct patterns of move-

ment. These may be necessary for optimizing the reproductive potential

fish populations that are available for restocking of an area that may

naturally or otherwise loose fish population. It may be a wise manage-

ment policy to aid this movement by removing unnecessary stream obstruc-

tions. A localized area of pollution in a stream may be detrimental

to more fish than just those in the immediate vicinity. The entire

reproductive potential for a large area of a stream may be lost as

migrating fish attempt to move through a polluted region. This con-

sideration should be taken into account in stream pollution studies

and may be particularly critical during the March to May period of

fish migration. Utilizing information gathered in the study, predic-

tions for a repopulation of an area that has been totally depleted

xix

of a fish population due to pollution indicates that it would take

about 2$ years to re-establish the pre-pollution population. This

estimate could be used in the economic assessment of pollution damage.

The nutrient balance established for New Hope Creek as it flows

through Duke Forest, with particular emphasis on the cycling of

phosphorus, indicated the value of a natural ecosystem for retaining

vital nutrients. Protection of water sheds have thus two values,

one for maintaining stocks of nutrients in valuable locations such

as forests and keeping the same nutrients out of oligotrophic streams

where they might cause undesirable eutrophication if they should be

released.

The value of ilTew Hope Creek to the studies in the basic metab-

olism of a stream cannot be overemphasized since so few unpolluted

streams are available for such studies. Maintenance of this stream

in its natural state as an outdoor laboratory for the Triangle Uni-

versities requires that it receive a stream classification under the

North Carolina system of stream classification which will give it

ad.equate protection.

INTRODUCTION

Animal migra t ions a r e a conspicuous and important phenomenon i n

many ecosystems of t h e world. Myriads of popular a r t i c l e s have been

w r i t t e n about t h e migra t ions of f i s h e s and b i r d s , and t h e s c i e n t i f i c

l i t e r a t u r e i s f u l l of d a t a on t h e s e and o t h e r migrants .

This s tudy cons iders migra t ion a s a func t iona l component of a

stream ecosystem by r e l a t i n g f i s h movements t o stream metabolism.

Seasonal p a t t e r n s of metabolism and f i s h migra t ion were measured i n

f i e l d s t u d i e s i n New Hope and Morgan Creeks, Orange and Durham Counties,

North Caro l ina , from Apr i l , 1968 t o June, 1970. The r e s u l t s were

compared with t h e movement p a t t e r n s of some o the r spec i e s i n the b io -

sphere a s r epo r t ed i n t h e l i t e r a t u r e .

What i s t h e r o l e of migra t ion i n t h e many and v a r i e d ecosystems

i n which it i s found? Under what condi t ions do groups of animals

t h a t migra te have s e l e c t i v e va lue over o t h e r groups t h a t do not migrate?

How much energy i s requi red t o migrate , and can enerqy be gained by

migrat ion? What e f f e c t does migra t ion have on t h e ecosystem of which

it i s a component and v i c e v e r s a ? What percentage of an ecosystem's

energy budget i s t i e d up i n maintaining a migratory component? This

s tudy cons iders t h e above ques t ions f o r a small warm-water stream i n

t h e piedmont reg ion of North Carol ina.

Theories For hl igrat ion

There may be s e l e c t i v e advantages f o r migrat ion p a t t e r n s which

lead t o success of t h e migrants and s u r v i v a l of t h e systems which

support migrants . Consider previous s t u d i e s which d i scuss

migra t ion a s a mechanism f o r improving t h e chances of s u r v i v a l of

t h e populat ion.

Non-reproductory migratory movements may be undertaken f o r t h e

sake of s e l f o r spec i e s p re se rva t ion (Ijeape, 1931). Three p r i n c i p a l

types a r e : a l imen ta l , o r having t o do with food; c l i m a t i c , o r having

t o do wi th extremes i n c l imate ( p a r t i c u l a r l y tempera ture) , and

gametic, o r having t o do wi th reproduct ion. Heape considered t h a t

t h e s e d i f f e r e n t migra t ion types a r e o f t e n r e l a t e d : "In a l l animals

which experience a gametic migra t ion , a r e t u r n journey i s involved

which i s d i r e c t l y concerned with e i t h e r c l i m a t i c o r a l imenta l

condi t ions ." According t o him, t h e r e t u r n journeys o f t e n can be

considered nomadic; and non-gametic migra t ions a r e considered, as a

r u l e , spasmodic o r due t o exc'eptional condi t ions--al though he d i scusses

on t h e next page r e g u l a r seasonal movements of a r c t i c animals which

move i n response t o "not cold so much a s want of food."

bligration t o Avoid Unfavorable Condit ions

Al lee e t a l . (1949, p. 539) s t a t e t h a t an organism has but t h r e e -- choices when exposed t o adve r s i ty : it may d i e , a d j u s t , o r migrate .

3

Thus,in t h e i r d i scuss ion of f l u c t u a t i o n s i n environmental cond i t i ons ,

migra t ion i s considered a ~ e c h a n i s m f o r removing t h e organism from

unfavorable circumstances. The reason f o r r e t u r n dur ing more

f avo rab le circumstances i s not a s c l e a r l y s p e l l e d out .

Migrat ion and Reproduct ion ---.. "-,- - -- Migrat ion may b r ing f i s h e s back t o a r eas i n which t h e i r ances t r a l

eggs developed. "In most i n s t a m e s t h e r e i s a l s o a seasonal o r

p e r i o d i c a l (non-spawning o r l a r v a l ) migra t ion a f f e c t i n g t h e immature

and mature" (Meek, 1916). He cons iders migra t ions t h a t occur from

deep t o shallow reg ions i n a l ake , movements up and down r i v e r s , and,

p a r t i c u l a r l y , movements i n va r ious loca t ions i n t he sea . In t h e

ocean, he says , t h e r e i s a genera l movement i n toward shore f o r

spawning, followed by d i s p e r s a l seaward. This p a t t e r n r ecu r s each

year with inc reas ing amplitude a s t h e young mature. The r e s u l t i s

t h a t t h e o l d e s t f i s h e s d i s p e r s e f a r t h e r from shore during non-spawning

t imes. Heape (1931) g ives many examples of f i s h e s , b i r d s , and

mammals with ex tens ive migratory movements f o r reproduct ion without ,

however, saying why an animal should migrate t o reproduce.

Migrat ion and Optimal Use o f F luc tua t ing Environments

Mayr and Meise (1930), a s quoted i n Cox (1968), suggested t h a t

competi t ion f o r food, p r i n c i p a l l y a s a r e s u l t of reproduct ive excess ,

i s t h e f a c t o r favor ing t h e development of mechanisms al lowing seasonal

occupat ion of a r e a s with a l t e r a t i o n s of favorable and unfavorable con-

d i t i o n s . A r i go rous approach to t h i s problem has been undertaken by

s tuden t s of S. C . Kendeigh (S iebe r t , 1949; West, 1960; Cox, 1961;

4

Zimmerman, 1965). These s t u d i e s i nves t iga t ed the energy balance o f

migra t ing b i r d s i n terms of energy r equ i r ed t o migrate and energy

gained by being i n d i f f e r e n t p l aces a t d i f f e r e n t t imes. S i ebe r t

concluded t h a t southward migrat ion f o r t he s l a t e - co lo red junco and t h e

whi te - throa ted sparrow was a metabol ic neces s i ty . West came t o t h e

same conclusion f o r t h e t r e e sparrow, bu t d id not f i n d t h a t t h e

nor thern migra t ion gained an improved energy balance. Cox (1961)

found t h a t r e s i d e n t t r o p i c a l f i nches would ga in l i t t l e by northward

migrat ion. Zimmerman, however, concluded t h a t t h e d i c k c i s s e l gained

an improved energy balance i n both i t s nor thern and southern movements.

Cox (1968) suggested d ivergent adapta t ion by both morphological

and e tho log ica l means. Given i n t e r s p e c i f i c o r i n t e r g e n e r i c

competi t ion, animals may broaden t h e i r n iche by exp lo i t i ng , f o r

example, d i f f e r e n t food sources; o r , they may broaden t h e i r niche by

e x p l o i t i n g s p a t i a l l y d i f f e r e n t environments. Cox showed t h a t wi th in

taxonomic groupings (usua l ly o rde r s o r f a m i l i e s ) , culmen (a p a r t of

t h e beak) length v a r i a b i l i t y among spec i e s was much g r e a t e r f o r b i r d

groups t h a t d id not have a high frequency of migratory members.

Thus some b i r d groups diverged by e x p l o i t i n g d i f f e r e n t food sources w i th in

a s i n g l e environment, and oth.ers moved t o d i f f e r e n t a r eas . There

may be a l i m i t t o food n iche divergence a t which animals must begin

t o e x p l o i t new phys ica l environments. llechanisms f o r t h i s a r e discussed

by Cox.

Migratory p a t t e r n s of animals a s soc i a t ed with Texas e s t u a r i e s

a r e considered i n r e l a t i o n t o t h e primary product ion and environmental

food supply by Odum and Nosltins (1958), Simmons and Hoese ( l959) ,

H e l l i e r (1960), Odum and Yilson (1962), Copeland ( l965) , and Odum

(1969). These s t u d i e s emph2size how t h e very l a r g e s p r i n g product ion

of t h e s e a r eas a r e u t i l i zec l by migrat ing animals, e s p e c i a l l y during

t h e i r j uven i l e s t a g e s , and how t h e migratory p a t t e r n s a r e such

t h a t maximvm use i s made o f t h e pu l se i n energy i n those ecosystems

during t h e l a t e sp r ing , The migrations themselves a r e seen as a

mechanism t o even out t h e flow of energy i n t h e system and d i s t r i b u t e

energy and n u t r i e n t s . Odum (1959) s t a t e s t h a t "Seasonal a.nd

d iu rna l migra t ions not only make p o s s i b l e occupation of reg ions

which would be unfavorable i n t h e absence of migrat ion but a l s o

enable animals t o maintain a h igher average d e n s i t y and a c t i v i t y

r a t e . "

Another at tempt t o exp la in t h e reproduct ive migrat ions of

animals i n r e l a t i o n t o s e l e c t i v e advantages f o r t h e migrat ing

popula t ion and e n e r g e t i c c h a r a c t e r i s t i c s of environments is by

Margalef (1963, 1968). He d i scusses d i f f e r e n t degrees of matur i ty

i n ecosystems. Margalef de f ines ma tu r i t y i n terms of t h e degree

of o rgan iza t ion of t h e ecosystem, which i s not n e c e s s a r i l y r e l a t e d

t o chronologica l age. According t o him, l e s s mature ecosystems a r e

l e s s e f f i c i e n t i n t h e i r u s e of energy and support l e s s biomass on

t h e same energy flow. Thus t h e r e i s an excess of a v a i l a b l e energy

t h a t may be exported. More mature systems, with a g r e a t com-

p l e x i t y of b i o l o g i c a l i n t e r a c t i o n s and r e s u l t a n t g r e a t e r e f f i c i e n c y i n

energy use , produce no, o r a t l e a s t l e s s , excess energy.

Margalef cont inues with t h e argument t h a t those ind iv idua l o r -

ganisms t h a t have developed behavior p a t t e r n s lead ing t o reproduct ion

6 8'

i n l e s s mature ecosystems have l e f t behind more o f f s p r i n g and, t h e r e -

f o r e , a r e s e l e c t e d f o r . He g ives examples of animals t h a t tend t o &B

spend t h e i r a d u l t l i f e i n more mature a r e a s and reproduce i n l e s s

mature a r eas o r send l a r v a e o r reproduct ive elements i n t o them.

Some examples a r e : migra t ing b i r d s f l y i n g t o p o l a r reg ions t o r e -

produce, ben th i c animals sending l a r v a e i n t o l e s s mature p lanktonic

environments, and c lupe id f i s h e s t h a t spawn i n l e s s mature p a r t s of

t h e coas t of Spain and spend t h e i r adu l t l i v e s i n more mature

reg ions . Even t h e seemingly enigmatic s i t u a t i o n of e e l s and salmon

can be explained i n t h i s way, he says , s i n c e t h e s p e c i f i c reg ions t h a t

both a d u l t animals i n h a b i t a r e more mature than t h e s p e c i f i c h a b i t a t

of t h e l a r v a e of t h e r e s p e c t i v e f i s h e s .

McLaren (1963) found i n models of migrat ing zooplankton t h a t

t h e energy saved by l i v i n g one-half o f t h e day i n co lder water ,

where metabolism was l e s s , was g r e a t e r than t h e energy used i n t he

process of migrat ion. The energy gained by t h i s process could

then be used f o r growth and reproduct ion .

Ricard (1968) cons iders animal migra t ions a s an i n t e g r a l p a r t

of b i o l o g i c a l rhythm. He sugges ts t h a t migra t ion serves a s a

mechanism f o r r e g u l a t i n g popula t ion numbers of spec i e s such as

swallows, s i n c e many members of a popula t ion a r e l o s t during m i -

g r a t i o n . This may a l s o be t r u e f o r lemmings, although t h e i r movement

cannot be considered a t r u e migra t ion s i n c e t h e lemmings do not

r e t u r n . Ricard a l s o sugges ts t h a t animals move t o d i f f e r e n t a r eas

where t h e i r food i s s easona l ly more abundant. He gene ra l i ze s :

"One must conclude, t h e r e f o r e , t h a t migrat ion i s not t h e only s o l u t i o n

7

t o t h e problem of t h e balance between animals and food resources , bu t

t h a t it i s t h e one t h a t e x i s t s 8t t h e p re sen t time."

According t o F. R , Harden Jones (1969) migra t ions a r e "an

adap ta t ion f o r abundance by making t h e most of a va r i ed environment."

In g iv ing a d e t a i l e d a n a l y s i s of migratory p a t t e r n s of f i v e groups

of f i s h e s , he cons iders how t h e s e p a t t e r n s have evolved t o a i d i n

t h e u t i l i z a t i o n 05 var ious food sources.

Fos t e r (1969), i n reviewing p o s s i b l e causes f o r t h e development

of migra t ion i n f i s h e s , cons iders t h e p o s s i b i l i t i e s of changes i n

food a v a i l a b i l i t y , c l ima te , s a l i n i t y , and topography over geologic

time. The i n t e r a c t i o n of e x p l o i t a t i o n of new resources with the need

f o r t h e a d u l t s o r eggs t o s t a y wi th in c e r t a i n phys io logica l l i m i t s

may have s e t t h e s t a g e f o r t h e f i r s t f i s h migra t ions .

One common f a c t o r i n a l l t h e s e previous s t u d i e s i s t h e r o l e of

migra t ion i n i nc reas ing t h e flow of energy, o r decreas ing t h e energy

l o s s , t o popula t ions involved. Movements away from energy-consuming,

food-poor, cold reg ions i n t he win te r , a s wel l as t o energy-r ich

a reas of high p roduc t iv i ty , can be considered i n t h e s e terms. The

energy c o s t of migrat ion has been considered by I d l e r and Clemens

(1959) , McLaren ( l963) , and Bre t t (19 70) .

Role of Migrat ing Animals i n hlineral Cycling

Among t h e f i r s t au thors t o cons ider t h e p o t e n t i a l of migrat ing

animals f o r r ecyc l ing o r important l i m i t i n g minera ls was Juday e t - a l . (19521, who specula ted upon t h e r o l e of dead salmon i n br inging - phosphorus and o t h e r minerals t o t h e s t ream-lake ecosystems of t h e

F *

salmon's e a r l y l i f e h i s t o r y . Quan t i t a t i ve work on t h i s was under+ I,e:l

by Donaldson (1967) and Krokhin (1967) who demonstrated t h e very l a r g e b..

r o l e dead salmon had i n supplying s u f f i c i e n t l y high l e v e l s of

phosphorus t o maintain p r o d u c t i v i t y of sockeye lakes a t a s u f f i c i e n t

l e v e l t o support l a r g e runs o f salmon.

Many f u r t h e r examples may be present i n o t h e r f i s h e s , v e r t i c a l l y

migra t ing plankton, and migrat ing b i r d s . With t h e tremendous i m -

por tance of small amounts of some t r a c e elements now being recognized

(Hutchinson, 1957; Goldman, 1969), p o s s i b i l i t i e s do e x i s t f o r

migra t ions t o con t ro l c r i t i c a l n u t r i e n t s .

Previous S tudies on the Movements of Fishes

Nearly a l l s t u d i e s of f i s h migra t ion wi th in f r e s h water have

occurred with spec i e s t h a t a r e a s soc i a t ed e i t h e r with lakes o r t h e

ocean. Only a small amount of t h e t o t a l information a v a i l a b l e con-

cerns f i s h e s t h a t spend a l l t h e i r t ime wi th in one f resh-water stream.

Movements To ta l ly Within One Stream

Many s t u d i e s have been conducted over t h e years t o s tudy f i s h

movements i n s t reams. Bangham and Bennington (1938) r e p o r t a r e -

cap tu re of only about 11 percent of f i s h e s seined and marked i n a

warm-water Ohio stream. Three cen t r a rch ids (smallmouth bass , green

sun f i sh and rock bass) had much h ighe r (19-20) percentages of t a g

r e t u r n s than d i d o t h e r spec i e s . No marked f i s h were recovcr~c; In

ad jacent one-mile s e c t i o n s of s t reams loca ted above and below the

marking a rea . They concluded from t h e s e s t u d i e s t h a t f i s h i n t h e i r

s t reams moved about very l i t t l e . Fur ther evidence f o r t h i s view,

9

most of i t based on r e t u r n s of tagged f i s h by s p o r t fishermen, i s

suppl ied i n Sco t t (??d.9$ f o r rock bass i n Indiana and by Tate (19fl9)

f o r smallmouth bass i n some small s t reams i n Iowa, Allen (1951) found

l i t t l e seasonal movement of t r o u t i n New Zealand, Gerking (1959) con-

cluded t h a t most f r e s h water f j s h e s had l imi t ed home ranges, and

Gunning and Shoop (1961) found l i t t l e s h o r t range movement i n stream

dwell ing American e e l s .

Other i n v e s t i g a t o r s have come up wi th o t h e r conclusions. S t e fan ich

(1952) found some f i s h e s t h a t had moved and some t h a t were s t a t i o n a r y

i n a Montana cold-water stream. Brown (1961) found s i m i l a r r e s u l t s

f o r warm-water f i s h i n Ohio. Bjornn and Mallet (1964) found very

d i s t i n c t p a t t e r n s of sp r ing upstream movements and f a l l downsteam

movements f o r n a t i v e popula t ions of c u t t h r o a t t r o u t and Dolly Varden.

Some of t h e s e f i s h had t r a v e l e d a t l e a s t 50 t o 60 miles . Considerably

g r e a t e r numbers of f i s h were recaptured i n a r eas o u t s i d e of t he

o r i g i n a l cap ture a r e a than wi th in . Behmer (1964) found d i f f e r e n t

p a t t e r n s of movement f o r d i f f e r e n t warm-water f i s h e s i n Iowa, i n -

c luding some movements of 40 mi les . Hunt (196A) r epor t ed t h a t wild

brook t r o u t i n Laurence Creel:, Wisconsin, used upstream reaches of

t h e creek f o r spawning much more than they used downstream areas ,wi th

t h e in fe rence t h a t t h e t r o u t moved upstream t o spawn. He a l s o

found cons iderable d i s p e r s i v e movements of young t r o u t , gene ra l ly

i n a downstream d i r e c t i o n . S h e t t e r (1368) found complicated p a t t e r n s

of t r o u t movement i n t h e Au Sable River i n Michigan. Many d id not

move; some move up and some moved down, with no p a r t i c u l a r seasonal

p a t t e r n ev ident . The p a t t e r n s d i f f e r e d from one watershed t o another .

i C

S h e t t e r l s s tudy, a s a l l those l i s t e d so f a r , i s based on r e c a p t u r t

of marked f i s h e i t h e r by se in ing , e l e c t r i c shocking, o r ang le r r e -

t u rns .

One answer t o t h e s e complicated p a t t e r n s of movement ( including

no movement) i s suppl ied by Funk (1955), who suggested t h a t many

stream f i s h e s have both a mobile and a sedentary popula t ion of each

spec ies . As i n o t h e r s t u d i e s , h i s work ind ica t ed va r i ed p a t t e r n s of

f i s h movements. Some f i s h moved up, some moved down,and some d id not

move a t a l l . This was t r u e both f o r spec i e s groups and f o r d i f f e r e n t

i nd iv idua l s w i th in a spec i e s . A l l important f i s h spec i e s showed a

g r e a t e r tendency t o move i n t h e sp r ing than during t h e summer. Re-

s u l t s of f i s h movements i n t h e f a l l were va r i ed .

Unfortunately, almost a l l of t h e s e d a t a a r e heavi ly b iased by

t h e sampling procedures. hfuch more f i e l d work was done i n summers

than a t o the r t imes of t h e year . More sampling was done i n a reas

r e a d i l y a c c e s s i b l e t o v e h i c l e s , hence angl ing p re s su re a l s o tended

t o be concent ra ted a t t h e s e a r eas causing b i a s of r e s u l t s toward

r ecap tu re s i n t h e a r ea of o r i g i n a l cap ture . Some s t u d i e s considered

r ecap tu re s wi th in t h e same pool a s r e p r e s e n t a t i v e of no movement,

o t h e r s included a l l f i s h captured wi th in one mile of t h e sampling s i t e .

The o v e r a l l p i c t u r e f o r streams t o d a t e i s confusing. A s i m i l a r

conclusion i s reached i n a l i t e r a t u r e survey by Carpenter (1967).

Movements of F ishes i n Streams with Adjoining Lakes

On t h e o t h e r hand t h e r e i s a f a i r l y c o n s i s t e n t p a t t e r n of f i s h

spawning runs from lakes and ponds t o inf lowing o r outflowing s treams.

Stream dwell ing brook t r o u t moved upstream i n t h e f a l l ; brown and

11 4

rainbow t r o u t were p r i n c i p a l l y captured moving upstream i n the spr ing

and summer (She t t e r , 19%) . Suckers, which were t h e most important C

f i s h captured dur ing t h i s s tudy i n terms of numbers and mass, were

captured moving downstream i n t h e sp r ing and upstream i n t h e f a l l .

S h e t t e r sugges ts t h a t t h i s i s probably a spawning run from lakes t h a t

a r e l oca t ed above t h e counting weir . Northern p ike gene ra l ly moved

downstream, and o t h e r f i s h e s had l e s s c o n s i s t e n t p a t t e r n s . During

t h e one year of S h e t t e r v s s tudy, approximately equal numbers of f i s h

were captured moving upstream a s down. No d a t a were given a s t o t h e

s i z e of t h e f i s h e s moving upstream and down.

Raney and Webster (1942) and Ra.yner (1942) found runs of

spawning common white suckers and rainbow t r o u t i n an i n l e t t o

Skanea te les Lake, New York. The suckers moved upstream i n Apr i l and

May and back downstream sometime l a t e r i n ?lay. The rainbow t r o u t

migrated i n t o t h e s t ream dur ing t h e second and t h i r d week i n Apri l

and appeared t o s t a y i n t h e stream f o r f i v e days t o two months.

Other s t u d i e s done i n Michigan us ing two-way f i s h wei rs (Carbine

and S h e t t e r , 1943) showed t h a t t r i b u t a r y s t reams cont r ibu ted many

small brook t r o u t t o t h e main stream of Hunt Creek and t h a t l a r g e

sp r ing runs of suckers and redhorses moved downstream from Houghton

Lake i n t o Muskegon River. Some of t h e suckers and redhorses r e tu rned

upstream; but t h e ma jo r i t y , apparent ly d.id no t , and many dead spent

f i s h were observed j u s t a f t e r spawning. S imi la r r e s u l t s were obtained

a t Lake Gogebic. Large upstream runs of suckers and rainbow t r o u t

were captured i n a two-way weir i n s t a l l e d a t t h e mouth of t h e P l a t t e

River where it e n t e r s Lake Michigan. About 20 t imes more f i s h were

captured moving up t h e P l a t t e t han down. Most of t h e f i s h movement

occurred dur ing t h e month of Apr i l and was apparent ly a s soc i a t ed

with spawning. In t h e Brule River of Wisconsin (Niemuth, 1967),

heavy runs of excep t iona l ly l a r g e brown t r o u t moved out of Lake

Superior during t h e summer and f a l l f o r spawning. Large numbers of

t h e s e f i s h d ied a f t e r spawning, although some r e tu rned t o t h e l ake

t h e fol lowing sp r ing . Young t r o u t s tayed i n t h e r i v e r f o r about

two yea r s , then moved down t o the lake. Warner (1959) found t h a t

landlocked salmon moved downstream from l a r g e l akes i n Maine t o

spawn and t h a t t h e major i ty r e tu rned t o t h e lakes a f t e r spawning.

Perhaps t h e most i n t e n s i v e s tudy of t h e r e l a t i o n of lake-dwelling

f i s h and spawning s treams has been conducted by Martman e t a l . (1962) -- i n Loon Lake, B r i t i s h Columbia, wi th n a t u r a l l y occurr ing rainbow t r o u t .

Both i n l e t and o u t l e t streams were used f o r spawning, a l though t h e

i n l e t s t ream was used much more heavi ly . Both spawning runs apparent ly

had a l a r g e m o r t a l i t y of spawning f i s h .

A common c h a r a c t e r i s t i c i n most of t h e s e s t u d i e s i s t h a t more f i s h

a r e captured going from t h e l akes i n t o t h e streams than v i c e versa . -- Since a l l weirs used f o r t h e s e s t u d i e s had mesh s i z e s t h a t allowed

juven i l e f i s h t o pass , t h e movements f o r t h e t o t a l popula t ions a r e

unknown. There may b e a s u b s t a n t i a l r e t u r n of small f i s h . In add i t i on ,

most of t h e s e s t u d i e s i n d i c a t e t h a t t h e movements of f i s h i n t o the

streams were a s s o c i a t e d with spawning a c t i v i t i e s and t h a t a l a r g e

percentage of t h e spawning f i s h f a i l e d t o r e t u r n t o t h e l ake from which

they o r i g i n a l l y came.

Movements of F ishes Between Fresh and S a l t Water

Some spec i e s of f i s h t h a t move between f r e s h and s a l t water have

been s tud ied in t ens ive ly . The movements of salmon a.nd e e l s have

been reviewed by Harden Jones (190R), and t h e g e ~ e s a l l i f e h i s t o r y

p a t t e r n s of t h e s e f i s h e s i s wel l known. Banks (1969) has reviewed

t h e l i t e r a t u r e on t h e movement of salmon from the sea t o t h e i r

spawning grounds. Fishes such a s salmon, t h a t spend t h e i r adu l t

l i f e i n s a l t water but spawn i n f r e s h waters , a r e known as anadro-

mous; whi le t hose t h a t do t h e r eve r se , such as e e l s , a r e known as

catadromous.

The l i f e h i s t o r y of s e v e r a l o t h e r A t l a n t i c anadromous f i s h ,

such a s alewives, shad, and s t r i p e d bass , a r e reviewed by Bigelow

and Schroeder (1953), Talbot and Sykes (1958), and Mi l l e r (1969).

S tud ie s of the'movements of brook t r o u t between f r e s h and s a l t water

have been done by Smith and Saunders (1958, 1967, 1968) on Prince

Edward Is land . Sumner (1962) s tud ied t h e movements of c u t t h r o a t

t r o u t between f r e s h and s a l t water i n Oregon. Many o the r s ea

f i s h e s , such a s ta rpon and snook, t r a v e l f r e e l y between f r e s h and

s a l t water i n movements apparent ly not d i r e c t l y connected with

spawning (Breder, 1948).

Movements of F ishes i n t h e Open Sea

The movements of P a c i f i c salmon on t h e open sea have been

summarized by Manzer (1960), Neave (1964), and Royce e t a l . (1968). -- These papers p re sen t evidence f o r extremely fa r - ranging movements of

some ind iv idua l f i s h t h a t may encompass almost t he e n t i r e P a c i f i c

Ocean. The f i s h e s appear t o fo l low f a i r l y wel l def ined rou te s ,

o f t e n i n a broad c i r c u l a r p a t t e r n , and r e t u r n t o t h e i r parent

s t reams from two t o seven yea r s a f t e r t h e i r en t rance i n t o t h e sea .

The movements of c t h e s f i s h e s a r e i n many cases not wel l

known. S t rasburg (1969) and Royce (1967) r e p o r t t h a t a v a i l a b l e

evidence i n d i c a t e s a movement t o t h e no r th of b i l l f i s h e s i n

summer and a r e t u r n southward i n win ter . Neave and Hanavan (1960)

found a northward movement of many spec i e s from May t o August and

September. Mather (1969) r e p o r t s east-west A t l a n t i c migrat ions of

b l u e f i n t una and seasonal north-south movements of white marl in .

Seasonal north-south movements f o r s eve ra l spec i e s of no r theas t

P a c i f i c Ocean s o l e have heen repor ted by Alverson e t a l . (1964). -- F. R. Harden Jones (1968) summarized much of t h e a v a i l a b l e evidence

concerning movement of many North Sea f i s h e s t o and from breeding

and winter ing grounds.

Statement of Purpose

Many of t h e s e previous s t u d i e s show migra t ion of f i s h e s t o be

prominent. Presumably, t h e s e movements involve cons iderable amounts

of energy, poss ib ly enough t o be i n f l u e n t i a l i n c o n t r o l l i n g , d i r e c t l y

o r i n d i r e c t l y , t h e main flows of energy wi th in t h e i r ecosystems. To

s tudy t h i s p o s s i b i l i t y more f u l l y r equ i r e s measurements of migra t ion

and energy budgets i n t h e same ecosystem, i n o rde r t o determine t h e i r

r o l e s and r e l a t i v e magnitudes. This was done f o r a New Hope Creek,

a small s t ream loca t ed i n Duke Fores t , North Carol ina.

Descr ip t ion of Study Area

New Hope Creek i s a r e l a t i v e l y small piedmont s t ream loca t ed i n

Orange, Durham, and Chatham count ies , North Caro l ina (Figure 1 ) . I t s

Figure 1. Location of sampling s t a t i o n s on New Hope and

Morgan Creeks, North Carol ina. Each s t a t i o n was given a mnemonic

name. S t a t i o n 1 i s "Way up"; 2 i s "Horsefield"; 3 i s lfBlackwoodw;

4 i s "Weight l i m i t 10"; 5 i s "Wood Bridge"; 6 i s "Jungle"; 7 i s

'Toncre te Bridge", a l s o "Big Pool" s t a t i o n i s loca t ed about 100

meters upstream from "Concrete BridgeM; 8 i s nP-66ff; and 9 i s

"Pipeline." "Blackwood", "Wood Bridge", and "Concrete Bridge" -

"Big Pool" s t a t i o n s , numbers 3, 5, and 7 , were most heavi ly

sampled. M i s t h e l o c a t i o n s a p l e d on Morgan Creek.

waters flow i n t o New Nope River and then i n t o Haw River and Cape

Fear River . The p r i n c i p a l s t ~ t d y a r e a i s l oca t ed i n t he Korst ian

Div is ion of Duke ForesC hetween Chapel H i l l and Durham. The stream

i n t h i s reg ion i s chamc?er ized by a moderate g rad ien t (3.96 m km-')

and v i r t u a l lack of p o l l u t i o n . The average width i s about 5 m and

the average depth i s about 0.4 m. Rocky r ap ids a l t e r n a t e with deep

l a r g e pools (Figure 2 ) . The water i s normally c l e a r , although

t h e stream becomes t u r b i d during f loods .

New Hope Creek i s r e l a t i v e l y unaf fec ted by man's a c t i v i t i e s

and has t h e b i o l o g i c a l c h a r a c t e r i s t i c s of a d i v e r s e and hea l thy

stream. Larvae of mayfl ies , s t o n e f l i e s , caddis f l i e s and many o t h e r

i n s e c t s a r e abundant i n t h e r i f f l e s and t h e f i s h l i f e i s d ive r se .

The North Caro l ina Divis ion of Inland F i s h e r i e s has c l a s s i f i e d t h e

creek a s a "Robin-Warmou.th" stream (Carnes, Davis and Tatum, 1964)

and cons iders t h e s t ream t o be one of t h e b e s t f i s h i n g streams i n

t h e Deep-Haw watershed. However, f i s h i n g p re s su re i s l i g h t i n t h e

po r t ion of t h e creek s tud ied . Much of t h e watershed l i e s wi th in t h e

Duke Fores t and t h e r e s t runs through f o r e s t e d a reas with an

occas iona l farm. Very s l i g h t add i t i ons of domestic sewage e n t e r

from s e v e r a l sources near t h e headwaters. About 3.8 km below t h e

s tudy a r e a , however, t r e a t e d sewage from t h e town of Durham e n t e r s t h e

creek. Very low oxygen ( < 1.0 ppm) was occas iona l ly found below t h e

po in t of sewage a d d i t i o n dur ing t h i s s tudy.

New Hope Creek, l i k e many o t h e r piedmont streams of North Caro l ina ,

i s s u b j e c t t o extreme f l u c t u a t i o n s i n water l e v e l s . During the two

years s tud ied summer water flows dropped t o almost zero, although

Figure 2. a . A t y p i c a l r i f f l e s t r e t c h of New Nope Creek,

l oca t ed j u s t above t h e Concrete Bridge s t a t i o n . b. A t y p i c a l pool

of New Hope Creek, l oca t ed j u s t above t h e Big Pool sampling

s t a t i o n . This p a r t i c u l a r pool i s over 150 m long and 16 m wide.

numerous l a r g e pools remained. F a l l , win ter and sp r ing f loods were

f a i r l y f requent and r a i s e d t h e water flow t o a s much a s 14.2 nS

second-' (400 cu. f t . second-'). During t h e s e per iods t h e stream

expanded wel l beyond t h e banks and t h e water became q u i t e muddy.

Morgan Creek, l oca t ed j u s t west of Chapel Hill, i s a smal le r

stream which flows i n t o Univers i ty Lake,a 70 ha a r t i f i c i a l i m -

poundment, about 3 km below t h e s tudy s i t e . .Morgan Creek above

Univers i ty Lake i s a l s o v i r t u a l l y unpolluted and sha.res many phys ica l

c h a r a c t e r i s t i c s and spec i e s of f i s h with New Hope Creek.

Q u a l i t a t i v e Energy Flow Diagram f o r Migration i n New Hope Creek -.

Figure 3 shows a q u a l i t a t i v e energy flow diagram f o r migrat ion

i n New Hope Creek. The symbols used a r e those developed by H. T.

Odum (1967a, 1967b, 1969; Figure 4 ) . Q u a n t i t a t i v e d a t a on some of

t h e s e flows a r e made a v a i l a b l e l a t e r i n t h i s t h e s i s .

The u l t i m a t e source of t h e energy t h a t runs the hydrology and

t h e biology of New Hope Creek i s , of course, t h e sun. Energy flows

from t h e sun t o green p l a n t s i n t h e water , such as ben th i c a lgae ,

aqua t i c macrophytes, and pseudophytoplankton. Energy i s then

t r a n s f e r r e d through food chains t o t h e f i s h populat ions. Sun energy

a l s o e n t e r s New Hope Creek in ' d i r ec t ly through t h e Duke Fores t t r e e s ,

which drop t h e i r l eaves i n t o t h e water , and through t h e organisms t h a t

feed on t h e s e leaves . About one-half of a l l t h e energy requi red t o run

the b io logy of t h e stream e n t e r s i n t h i s fash ion . The s to rage tanks

r ep re sen t t h e accumulation of organic ma te r i a l produced by t h e primary

producers t h a t i s no t immediately used by higher t r o p h i c l e v e l s . An

obvious example i s t h e accumulation of dead leaves on t h e bottom of t h e

Figure 3 . Energy c i r c u i t diagram f o r migrat ion i n New Hope

Creek. See t e x t f o r explanat ion.

Figure 4 . Symbols usccl in e r x r g y ne twork d i a ~ r a ~ : ; , F,-o;:I 1:. T.

Oduni, (1967a).

ENERGY SOURCE PASSIVE ENERGY HEAT SINK S'TQRAGE

POTENTIAL PURE ENERGY &'%'OEM G A T E GENERATING WORK RECEPTOR

SELF-MAINTAlfJ I NG . PLANT ECONOiiI f C CONSUMER POPULATIONS TEAf4SACTOR

POPULATION

stream, many of which a r e not eaten u n t i l t he fol lowing spr ing .

Migrat ion of f i s h e s and o t h e r organisms i s represented by t h e

dashed l i n e s connecting t h e popula t ions of f i s h e s . The usage of energy

a t any one p l ace i n t h e stream i s t o a c e r t a i n ex t en t dependent upon

t h e r e l a t i o n of t h a t p a r t o f t h e s t ream with c u r r e n t s and o t h e r p a r t s

of t h e stream. An i n d e f i n i t e number of such product ion- f i sh popula t ion

s e t s could be drawn rep resen t ing d i f f e r e n t p a r t s of t h e stream.

MATERIALS AND METHODS

The genera l p l an f o r t h e s tudy over a 27-month per iod

included s t u d i e s of upstream and downstream migra t ion a t s eve ra l

double-weir s t a t i o n s and measurements of photosynthesis and

r e s p i r a t i o n i n va r ious s e c t i o n s of t h e stream us ing changes i n

oxygen concent ra t ion .

Physical and Chemical Data

C h a r a c t e r i s t i c s of t h e Sampling S t a t i o n s

Nine sampling s t a t i o n s f o r oxygen and/or fish-movement ana lys i s

were e s t a b l i s h e d on New Hope Creek and one on Morgan Creek. The

l o c a t i o n s o f t h e sampling s t a t i o n s a r e given i n Figure 1, and some

c h a r a c t e r i s t i c s o f each s t a t i o n a r e given i n Table 1.

Discharge

Current v e l o c i t i e s were measured with an A. O t t (#I36241 'pigmy1

cu r ren t meter, To ta l stream flow was determined by measuring the

r a t e of flow e i t h e r i n about. 1 2 p o i n t s i n a g r i d p a t t e r n i n t h e s t ream

(during f lood s t a g e s ) o r i n t h e middle of fou r 38 cm p ipes through

which a l l water flows under t h e concre te br idge . The t o t a l d i scharge

was c a l c u l a t e d a s t h e summation of each flow r a t e t imes the c ross

s e c t i o n a l a r e a represented by t h a t flow r a t e . Daily s t a g e measurements

were maintained, and a graph of s t a g e versus d ischarge was cons t ruc ted

Table 1. Some C h a r a c t e r i s t i c s o f t h e Var ious Sampling S t a t i o n s

S t a t i o n Ki lometers above Stream Bottom

Erwin Road width Type

1. Way up

2. H o r s e f i e l d

3 . Blackwood

4. WL 1 0

5. Wood Bridge

6. J u n g l e

7. Big Pool

18.0 3.0 (a ) sand and g r a v e l

14.3 4 .0 (a ) sand and g r a v e l

12.5 8.3 s i l t and sand and b o u l d e r s

10.6 lO.O(a) s i l t

6 .0 b o u l d e r s and g r a v e l

10.0 b o u l d e r s and g r a v e l

3.3 14.2 bou lders and g r a v e l

8. Concrete Bridge 3.2 5.1 ha rd rock

9. P i p e l i n e (b) 5.0 (below) 15.0 s i l t and s l u d g e

Morgan Creek ---- 4.0 sand and b o u l d e r s

a. e s t i m a t e d

b. below o u t l e t from Durham Sewage Treatment P l a n t

24

(Figure 5) from which d a i l y flow r a t e s were read. Discharge a s

m3 s c was computed a s 0.02832 times d ischarge a s cubic f e e t

per second ( c f s ) . Stream Morphology: Depth --- and Width "..

Stream width and depth were measured a t 50 o r 100 m i n t e r v a l s

f o r one o r two km above each major oxygen sampling s i t e . Each

i n t e r v a l between t h e Concrete Bridge and t h e Wood S t a t i o n s was

marked o f f with a 50 m s t r i n g . I n t e r v a l s a t o t h e r l oca t ions were

determined by pacing o f f 100 m. A marked t a p e was stretched. across

a t each loca t ion ; t h e width was measured and depths were taken a t

1 m i n t e r v a l s . The average depth f o r each stream i n t e r v a l was

computed a s t h e a r i t h m e t i c mean o f a l l t h e depth measurements i n

t h a t i n t e r v a l .

The average width and depth f o r t h e s e c t i o n of stream over

which water flowed during one hour was c a l c u l a t e d from:

where D i s t h e d ischarge of t h e stream a t t h a t time i n m3 h r - l , L

i s t h e length i n m of each stream bed segment. Wn i s t he width

i n m of t h e stream a t each success ive sampling l o c a t i o n (50 o r 100 m),

Idn i s t h e average depth i n m a t t h a t l oca t ion , and n i s t h e t o t a l

number of sample segments necessary f o r [(w,) (On) (L,)] t o equal

one hour ' s water d i scharge . The t o t a l l ength o f n stream segments

was t h e length of stream through which t h e water flowed i n one hour.

Once t h i s t o t a l l ength was found t h e average of a l l width and depth

Figure 5. Water stage vs. discharge. Abscissa = stage level

in inches (cm) above zero flow. Ordinate = discharge in 103 m3

day-l. The break before the last two values occurs as the stream

overflows its banks.

measurements i n t h a t i n t e r v a l was a l s o found. These va lues weri

used f o r c a l c u l a t i o n s of stream metabolism.

Time i n t e r v a l s f o r water masses t o flow between two p o i n t s

was computed from stream morphology a s fo l lows:

where t i s t h e time, i n hours , f o r t h e water mass t o flow t h a t

d i s t a n c e , D is t h e d ischarge i n m3 h r - l , i s t he mean width o f

t h e s t ream i n t h e i n t e r v a l between t h e two p o i n t s , and i s t h e

mean depth i n t h a t same i n t e r v a l . Time i n t e r v a l s were a l s o , on one

occasion, checked with dye. The t u r b u l e n t and va r i ed na tu re o f t h e

s t ream made t h i s method d i f f i c u l t , s i n c e t h e mass of dye i n t he

c u r r e n t t r a v e l e d much f a s t e r than s i d e eddies . The r e s u l t s were about

40 percent lower t han t h e morphology method, bu t were not d i f f e r e n t

enough t o e f f e c t metabol ic c a l c u l a t i o n s .

I n s o l a t i on

Est imates of r e l a t i v e amounts of i n s o l a t i o n pene t r a t ing

t r e e canopies a t two reg ions of New Hope Creek were made with a Weston

M ~ d e l 756 i l l umina t ion meter'. This measured t o t a l i nc iden t sun l igh t

i n foot-candles.

Measurements were made on a completely c loudless day. Est imates

of r e l a t i v e sun energy reaching New Hope Creek a t d i f f e r e n t l o c a t i o n s

were made by sampling every 100 meters f o r a d i s t a n c e of 1 km above

oxygen sampling s i t e s . Each s p e c i f i c l o c a t i o n was determined by

pacing o f f approximately 100 m , t hen t ak ing a reading a t t h e c e n t e r

of t h e s t ream j u s t above t h e s u r f a c e of t h e water. Since t h e

i n s o l a t i o n was patchy, t h e l igh t r e c e p t o r was moved i n an a r c a t

a rmqs l eng th and an average reading was taken.

In add i t i on , t o t a l i n s o l a t i o n , both i n a c l e a r f i e l d and under

a hardwood canopy, was obtained wi th an Epply pyrohel iometer from

t h e I n t e r n a t i o n a l Bio logica l Program s i t e loca ted i n another s e c t i o n

of Duke Fores t (Blackwood d i v i s i o n ) which i s about 200 m from t h e

watershed of t h e headwaters of New Hope Creek (Figure 1, n o r t h of

S t a t i o n 1 ) .

Stream Temperature

Temperature on each sampling d a t e was msasured, gene ra l ly i n

t h e l a t e a f te rnoon, us ing a s tandard l abo ra to ry thermometer.

Diurnal temperatures taken with t h e oxygen-temperature recorder

were co r r ec t ed a s explained i n t h e s e c t i o n on "Metabolic S tudies . "

A l l d i u r n a l temperatures above 5' C va r i ed dur ing t h e day. Since t h e

l a t e r a f te rnoon temperatures were, almost without except ion, about

2" C warmer than t h e average temperature f o r t h e day, average tempera-

t u r e s f o r days on which d i u r n a l temperatures were no t run were com-

puted a s t h e l a t e a f te rnoon temperature minus 2.

To ta l Phosphorus i n Water

A l l phosphorus and n i t rogen a n a l y s i s were based on FWPCA (1969).

To ta l phosphorus i n s t ream water was determined us ing a Technicon

Auto-analyzer with a 660 mu f i l t e r and a 5 cm flow c e l l . Samples

were c o l l e c t e d i n 100 m l polyethylene b o t t l e s t o which 40 mg of Hg

2 F

l i t e r - ' had been added a s a p re se rva t ive . The samples were f r o z e n

u n t i l analyzed.

Samples were d iges t ed i n an au toc lave with p e r s u l f a t e and

s u l f u r i c ac id . Phosphorus a n a l y s i s was by co lor imet ry fol lowing

stannous c h l o r i d e r educ t ion and t h e formation of a phosphomolybdate

complex.

Phosphorus i n Organisms

Est imates o f t o t a l phosphorus contained i n s eve ra l spec i e s of

hardwood leaves fol lowing abscission (Woodwell, 1970) and i n a

mixed f o r e s t (Gosz e t a l . , 1970) were averaged t o g ive approximate -- values (0.041 percent P dry weight) f o r leaves f l o a t i n g down New

Hope Creek.

Est imates of t o t a l phosphorus i n f i s h were taken from va lues

suppl ied by Vinogradov (1953) and Donaldson (1963). These were

approximately 0.3 percent P by weight f o r many spec i e s of f i s h and

0.4 percent P f o r whole sockeye salmon, r e spec t ive ly . An approximate

va lue of 0.35 percent wet weight was used f o r c a l c u l a t i o n s i n t h i s

t h e s i s .

Tota l Nitroeen i n Water

To ta l n i t rogen was a l s o analyzed on t h e Technicon Autoanalyzer.

Samples were taken from t h e same b o t t l e s a s f o r P a n a l y s i s , and

analyzed c o l o r i m e t r i c a l l y fol lowing d i g e s t i o n with a s u l f u r i c a c i d

s o l u t i o n conta in ing potassium s u l f a t e and mercuric s u l f a t e . The

b lue c o l o r measured r e s u l t s from t h e a d d i t i o n of a l k a l i n e phenol,

sodium hypoch lo r i t e and sodium n i t r o p r u s s i d e .

Stream Conductivity

The conductivity of water samples from New Hope Creek was

determined with a Yellow Sprin.gs Instruments Company Model 31

conductivity bridge.

Discharge of Leaves

Estimates of (dry weight) leaves flushed downstream were made

from June 13, 1968 to June 12, 1969. During normal water levels

the leaves that accumulated on the upstream side of the 1/4

inch (0.6 cm) hardware-cloth weir were removed every day or two.

During flood levels, when it was impossible to maintain the weirs,

leaves were sampled by holding a 50-foot (16.4 cm) fish seine with

a mesh size of 0.4 cm across the stream for 15 minutes (sometimes

less during exceptionally heavy flow). The weight of leaves moving

downstream in 24 hours was calculated assuming constant flow. On

days during which leaf discharge was not measured, estimates were

obtained by reading values from the graph of water stage versus

leaf discharge (Figure 6). This was possible because of the nearly

linear relation of total leaves discharged to the water stage

when plotted on semi-log paper. Although there was a tendency for

greater leaf discharge for a given water level to occur during the

fall, this was not sufficiently consistent to use seasonal correc-

tions in reading the graph. The calculations made using these

data did not require precise measurements.

Figure 6. To ta l mass of leaves (dry weight) discharged per

day i n s t ream flow a t Concrete Bridge S t a t i o n vs . s t a g e l e v e l

(o rd ina t e ) i n cent imeters above zero flow.

5 0 7 5 STAG E

Metabolic S tudies

E n t i r e ecosystems, l i k e ind iv idua l organisms, produce and use

energy t o main ta in l i f e . T h i s process can be measured by determining

t h e t o t a l amount of oxygen, o r carbon d ioxide , produced and consumed.

The fol lowing s e c t i o n desc r ibes how t h e s e gases rer re measured i n

New Hope Creek and a r e used t o e s t ima te metabolism,

Dissolved Oxygen, Winkler Method

Est imates of community metabolism f o r New Hope Creek were made

from d iu rna l v a r i a t i o n s i n d isso lved oxygen and pH. Oxygen was

measured both by t h e az ide modi f ica t ion of t h e Winkler method and

by an automatic f i e l d temperature and oxygen r eco rde r (Rustrak

Model 192) used with e i t h e r a Yellow Spring Instrument #5419 probe

o r Rustrak #I921 probe.

The Winkler de te rmina t ions were made fo l lowing Standard Methods

(American Publ ic Heal th Assoc ia t ion , 1965). For t he d i u r n a l

s t u d i e s , samples of water were taken every two o r t h r e e hours f o r

24 hours a t s tandard s t a t i o n s . A simple tube device minimized

oxygen d i f f u s i o n from t h e a i r dur ing f i l l i n g of t h e 300 m l sampling

b o t t l e (Figure 7 ) . The sampling b o t t l e clamped t o t h e end of a

s t i c k was he ld wi th incu r ren t tube about 15 cm below t h e water

sur face . Dupl ica te samples were taken wi th in about two minutes of

each o the r . A l l r e agen t s were added i n t h e f i e l d and t i t r a t e d wi th in

12 hours i n t h e labora tory . Welch (1968) found no d i f f e r e n c e i n

d u p l i c a t e oxygen samples when one was t i t r a t e d immediately and the

o the r 24 hours l a t e r . Nater temperatures were taken with a s tandard

F i g u r e 7. Cork and t u b i n g d e v i c e t o f i l l oxygen b o t t l e wi thou t

a i r mixing.

3 3

l abo ra to ry thermometer. Percent s a t u r a t i o n was ca l cu la t ed from

t h e oxygen s o l u b i l i t y va lues of Churchi l l e t a l . , (1962); they -- a r e i n t e rmed ia t e t o o t h e r va lues i n t h e l i t e r a t u r e ,

Dissolved Oxygen, Galvanic Probe Method

Oxygen concent ra t ion were measured a t one s t a t i o n with an

automatic r eco rde r i n s t a l l e d i n a s t reamside shed. The ch ie f

advantage of t h i s method was t h e tremendous savings i n e f f o r t t o

o b t a i n a d i u r n a l curve. Only one hour o r l e s s per day was requi red

t o s e t up and s t anda rd ize t h e instrument aga ins t Winkler determina-

t i o n s , read t h e c h a r t , and e n t e r t h e d a t a on punch cards. A

t y p i c a l d i u r n a l sequence us ing Winklers r equ i r ed about 28 hours. I n

add i t i on , a continuous record was obtained so t h a t non-typical water

masses could be i d e n t i f i e d . One disadvantage of t h e probe was t h a t

only one s t a t i o n could be sampled on a given day with t h e equipment

a v a i l a b l e . The membrane e l e c t r o d e may have been l e s s accu ra t e than

Winkler de te rmina t ion because of d r i f t , however, s i n c e t h e probe

averages oxygen va lues i n va r ious water masses flowing over i t , it

may be a t r u e r r e p r e s e n t a t i o n of s t ream oxygen. Figure 8 shows

t y p i c a l r e s u l t s of O2 es t ima te s us ing both methods. The maximum

d e v i a t i o n i n t h i s case was only about 0.3 gm3, which i s wi th in

extreme ranges of d u p l i c a t e Winklers.

P a r t i c u l a r c a r e was necessary t o avoid seve ra l sources of

e r r o r i nhe ren t i n t h e f i e l d record ing u n i t . The probe was water-

v e l o c i t y dependent, and it was necessary t o p l ace t h e probe i n

water t h a t had a v e l o c i t y of a t l e a s t 0.5 meter pe r second o r t h e probe

Figure 8. Comparison of probe and Winkler oxygen va lues over

a 24 hour pe r iod , J u l y 25, 1969,at Concrete Bridge S t a t i o n . Open

c i r c l e s a r e average of d u p l i c a t e Winkler samples, t h e range of

which i s represented by a v e r t i c a l l i n e . Data from ga lvanic probe

and r eco rde r a r e t r i a n g l e s connected by s o l i d l i n e . The maximum

d i f f e r e n c e between t h e two de termina t ions i s 0 . 4 5 mg 1-I (g m-3),

which i s wi th in t h e range of d u p l i c a t e Winkler samples.

would u s e oxygen f a s t e r than t h e water could resupply it. During

pe r iods of low flows rock j e t t i e s were cons t ruc ted t o concent ra te

t h e flow of t h e major p a r t o f t h e s t ream on t h e probe. The c u r r e n t

v e l o c i t y was assumed s u f f i c i e n t i f manual movement of t h e probe

i n t h e water d i d not i nc rease t h e reading on t h e meter. A

mechanical a g i t a t o r t h a t increased flow over t h e probe was used

during pe r iods of extremely low stream flow.

Other p o t e n t i a l sources of e r r o r were t h e co r r ec t ions f o r

e f f e c t s of temperature on t h e probe. The manual t h a t comes with

t h e Yellow Springs Model 51 oxygen meter s t a t e s t h a t t h e temperature

response of t h e oxygen probe i s about 5 percent per degree

cent igrade . In o t h e r words, f o r each degree higher t han t h e

c a l i b r a t i o n temperature, t h e probe would read about 5 percent t oo

high.

A check on t h i s temperature e f f e c t was made by p u t t i n g t h e probe

and a thermometer i n t o a conta iner completely f i l l e d with water a t

s eve ra l d i f f e r e n t oxygen l e v e l s . The conta iner was then cooled with

an i c e ba th while a magnetic s t i r r e r kept a cons tan t flow over t h e

probe. Temperature and oxygen readings were recorded. As t h e

temperature of t h e water dropped, t h e reading of t h e oxygen meter

a l s o dropped even though t h e oxygen content of t he water remained

cons tan t , s i n c e t h e con ta ine r was a i r t i g h t and no d i f f u s i o n of oxygen

could occur . Af te r t h e temperature of t he water approached zero,

warm water was put i n t h e water ba th and t h e temperature r a i s e d t o

t h e o r i g i n a l value. Di f fus ion was checked by determining i f t h e new

oxygen reading a t t he o r i g i n a l temperature was t h e same a s t h e o r i g i n a l

oxygen reading, and small co r r ec t ions were made. Sample r e s u l t s

obtained by t h i s method are presented i n Figure 9. Resul t s fo-

changes i n a i r readings with temperature were s i m i l a r , The

temperature c o r r e c t i o n f o r t h e oxygen probe p e r degree change in

temperature i s t h e s lope o f t h e l i n e of t h e graph, which i s ex-

pressed a s :

where Oc is t h e cor rec ted oxygen reading i n mg O2 rl, 0, i s t h e

uncorrec ted oxygen va lue , t i s t h e d i f f e r e n c e i n temperature from

t h a t a t s t anda rd iza t ion , and S i s t he d a i l y average s a t u r a t i o n va lue .

This c o r r e c t i o n was en tered i n t o t h e computer a lgori thm.

Temperature e f f e c t s on t h e record ing u n i t were checked by

p u t t i n g t h e whole u n i t i n a r e f r i g e r a t o r whi le leav ing t h e probe a t

room temperature. No changes i n reading occurred. The c h a r t was

read a s percentage of f u l l s c a l e , and the fol lowing equat ion was

used t o c a l c u l a t e oxygen concent ra t ions (mg 1 - I ) where 0, i s

oxygen concent ra t ion ,

i n mg 1-I a t c a l i b r a t

Ct i s c h a r t reading ,

- 1 i n mg 1 a t time t , Oc i s oxygen concent ra t ion ,

ion time c , a s determined by Winkler t i t r a t i o n ,

i n percentage of f u l l s c a l e , a t time t and Cc

i s cha r t reading , i n p r c e n t a g e of f u l l s c a l e , a t 1-hr: t ime of c'yygen

c a l i b r a t i o n . Since t h e reading of t h e oxygen r eco rde r i s l i n e a r l y

p ropor t iona l t o t h e concent ra t ion of oxygen i n water (Gulton

Figure 9. Var i a t ion i n oxygen meter readings with cons tan t

d i sso lved oxygen and varying temperature. Laboratory de termina t ion

was done May 11, 1969, The s lope of t h i s l i n e r ep re sen t s t h e

temperature c o r r e c t i o n necessary t o g e t t r u e readings a t temperatures

o t h e r than t h a t a t which t h e probe was c a l i b r a t e d . Ca l ib ra t ion was

a t 24' C and a t oxygen s a t u r a t i o n f o r room temperature (8.33 mg 02

1 ) T r i ang le s r ep re sen t descending temperatures , c i r c l e s r ep re sen t

r i s i n g tempera tures , and p o i n t s r ep re sen t a l a t e r decrease t o room

temperature. The s l o p e a t oxygen va lues l e s s than s a t u r a t i o n was i n -

v e r s e l y p ropor t iona l t o t h e percent s a t u r a t i o n .

Industries, Bulletin no. M26802), the above simple proportion will

give the oxygen concentration mg 1-I (g ~ n - ~ ) . Comparison of

temperature-corrected scale readings and IYinkler oxygen values

over a period of several days indicates that drift was relatively

small (Table 2).

Table 2 . D r i f t sf Oxygen Recorder Over One o r Several Days

Numb er Observed reading , Reading i f of temperature t h e r e were D i f fesence

days s i n c e cor rec ted no d r i f t i n i t i a l

Date c a l i b r a t i o n mg 1 -1 mi3 1

-1 mg 1 -1

June IS 1 7.47

J u l y 2 1 2 6.73

J u l y 25 4 5.67

Apr i l 23

Apri l 25

May 29

Diffusion Rates

Oxygen movos from a i r t o water and water t o a i r according t o

Dal ton ' s law of p a r t i a l p ressures . Correc t ions must be made i n

aqua t i c metabol ic s t u d i e s f o r t h i s . I t i s necessary t o know both

t h e percent oxygen s a t u r a t i o n of t h e water and t h e d i f f u s i o n

cons tan t t o make t h e s e co r r ec t ions .

Three months were t r i e d f o r determining t h e d i f f u s i o n

cons tan t on New Hope Creek: t h e d i u r n a l curve method (Odum, 1956;

Odum and Hoskins, 1958), t h e stream morphology method (Churchi l l

e t a l . , 1962) and t h e dome method (Hall and Day, 1970), The d iu rna l -- curve method gave r e s u l t s o f t e n much h ighe r than the o t h e r two

methods and was not used f o r t h i s s tudy. The reason f o r t h e high

va lues was t h a t n ight t ime r e s p i r a t i o n was not cons tan t , a pre-

r e q u i s i t e f o r t h e accu ra t e u s e of t h i s method (Odum and Wilson,

1962; Owens, 1969).

For t h i s s tudy t h e stream morphology method, which averages pool

and r i f f l e va lues , was used t o determine t h e d i f f u s i o n cons tan t ; t h i s

cons tan t v a r i e d from day t o day a s t he water l e v e l changed (Figure 10) .

The dome method gave s i m i l a r r e s u l t s , s i n c e New Hope Creek i s about

equal ly d iv ided between pools and r i f f l e s . A more complete t rea tment

of t h e d i f f u s i o n s t u d i e s i s given i n Appendix A.

Figure 10. Comparison of d i f f e r e n t d i f f u s i o n cons tan ts

ob ta ined i n t h i s s tudy. Crosses represent es t imates based on

t h e s t ream morphology method; t r i a n g l e s r ep re sen t es t imates made with

dome method; and c i r c l e s represent e s t ima te s made from d i u r n a l curves

s e l e c t e d f o r s u b s t a n t i a l d iu rna l range. The stream morphology

method was used f o r c a l c u l a t i o n s made f o r t h i s s tudy. See t e x t f o r

f u r t h e r explanat ion.

Gross Community Metabolism

Primary product ion and t o t a l r e s p i r a t i o n of t h e l i v i n g organisms

i n New Hope Creek were measured us ing d i u r n a l v a r i a t i o n s i n

metabol ic gases (Odum, 1956; Odum and Hoskins, 1958; Odum and Wilson,

1962; Beyers e t a l . , 1963). The b a s i s f o r t h e s e measurements i s -- t he fundamental equat ion f o r photosynthes is (or r e s p i r a t i o n ) :

Thus, t h e t o t a l c r e a t i o n and u t i l i z a t i o n of organic compounds i s pro-

p o r t i o n a l t o t h e amount of C02 and 02 being produced and consumed.

Some v a r i a t i o n s i n t h e r e l a t i o n of oxygen t o energy occur when

p r o t e i n s o r f a t s a r e being u t i l i z e d i n s t e a d o f sugars , o r when t h e r e

a r e l a g s i n one process r e l a t i v e t o another . For t h e s e reasons it

i s most accu ra t e t o cons ider t h e metabolism i n terms of oxygen

without conver t ing t o carbon o r c a l o r i c va lues .

Two S t a t i o n Analysis

The most accu ra t e e s t ima te s of photosynthesis and community

r e s p i r a t i o n can be obtained f o r a stream with t h e "two s t a t i o n ' ?

method of oxygen a n a l y s i s (Odum, 1956; Owens, 1969). The two s t a t i o n

a n a l y s i s i s based on the a c t u a l change i n oxygen occurr ing a s a water

mass flows from one reg ion of t h e stream t o another . Thus, changes

over a c l e a r l y def ined a r e a can be measured and r a t e s of change determined

from d i f f e r e n c e s between t h e upstream and downstream oxygen measure-

ments of t h e same water mass.

This method was gene ra l ly imprac t ica l on New Hope Creek because

of t h e n e c e s s i t y of sampling a t t h r e e t o f i v e loca t ions t h a t were t o o

f a r a p a r t . However, on one occasion two s t a t i o n a n a l y s i s was run

run on New Hope Creek a t both t h e Blackwood S t a t i o n and t h e Concrete

Bridge S t a t i o n . Tho r e s u l t s (Figure 11 and 12) i n d i c a t e t h a t i n t h e s e

s h o r t s t r e t c h e s of Mew Hope Creek, equivalent t o t h e d i s t a n c e water

flow i n about one hour, t h e metabolism of one p a r t i s s i m i l a r t o

another .

S ingle Curve Method -.

Where upstream and downstream d iu rna l curves a r e s i m i l a r , one

may use a s i n g l e s t a t i o n curve a s a f i r s t approximation (Odum,

1956). This procedure was used f o r New Hope Creek. The b a s i c

procedure i n e s t ima t ing stream metabolism by t h i s method i s t o

measure oxygen and temperature i n t h e f i e l d every two o r t h r e e

hours us ing e i t h e r Winkler oxygen methods o r a ga lvanic probe,

e i t h e r with o r without recorder . The d a t a a r e then p l o t t e d

(Figure 13) . The f i r s t d e r i v a t i v e i s computed f o r t h e s e oxygen

changes and aga in p l o t t e d us ing t h e same time sca l e . Correct ions

f o r d i f f u s i o n a r e made by adding t h e product of t h e d i f f u s i o n

cons tan t and t h e s a t u r a t i o n d e f i c i t t o t h e rate-of-change curve.

I f t h e r e were no b i o l o g i c a l o r chemical a c t i v i t y i n t he water

being s tud ied , t h e r e would b'e only t h e change i n t h e oxygen con-

c e n t r a t i o n s over t h e day due t o temperature changes a f f e c t i n g

s a t u r a t i o n va lues . The rate-of-change curve would be near zero

f o r t h e e n t i r e day. However, b i o l o g i c a l r e s p i r a t i o n tends t o i t , w i l r

t he oxygen i n t h e water throughout t h e day and n i g h t , and t h e photosyn-

t h e s i s o f green p l a n t s r a i s e s t h e oxygen dur ing t h e day. Thus, a

Figure 11. S i m i l a r i t y of oxygen curves one hour ' s flow d i s t a n c e

a p a r t a t Blackwood S t a t i o n , February 14, 1970. Winkler de te rmina t ions

were done i n dup l i ca t e . The t r i a n g l e s a r e oxygen concent ra t ions a t

t he upstream s t a t i o n ; t h e s o l i d l i n e connects t h e i r averages. The

c i r c l e s a r e oxygen concent ra t ions a t t h e downstream s t a t i o n , and t h e

broken l i n e r ep re sen t s t h e average of t hese . MN i s midnight.

F igure 12. S i m i l a r i t y o f oxygen curves one hour ' s flow d i s t a n c e

a p a r t a t Concrete Bridge S t a t i o n , February 14, 1970. Symbols and

l i n e s used a r e same as f o r Figure 11.

Figure 13. A r e p r e s e n t a t i v e sample of s i n g l e s t a t i o n a n a l y s i s

f o r community metabolism i n New Hope Creek, February 14, 1970, a s

conducted and p l o t t e d by t h e UNC CALCOMP p l o t t e r . The upper graph

i s t h e average of d u p l i c a t e Winkler de te rmina t ions taken every t h r e e

hours. Each t r i a n g l e r ep re sen t s a s i n g l e sample. The second graph

i s of temperature taken every t h r e e hours. The t h i r d graph i s t h e

percent s a t u r a t i o n of t h e average of t h e two Winklers a t t h e

temperature of t h e sample. The lower graph shows t h e r a t e of change

( f i r s t d e r i v a t i v e ) of t h e oxygen samples. The l i n e with t h e tri-

angles shows t h e r a t e of change co r rec t ed f o r d i f f u s i o n o f oxygen

across t h e s u r f a c e of t h e water . The gross photosynthesis of t h e

water mass represented by these water samples i s ind ica t ed by the

a rea s t i p p l e d . Gross community r e s p i r a t i o n i s est imated a s t he a r e a

cross-hatched. The d i f f u s i o n cons tan t i s i n g O 2 rn-3 h r - I atmosphere-',

and t h e depth i s average depth i n meters f o r one hour ' s flow d i s t a n c e

above sampling s t a t i o n .

c h a r a c t e r i s t i c rate-of-change curve i s produced, r i s i n g during day-

l i g h t and f a l l i n g a t n i g h t , o f t e n l e v e l i n g a s t he amount of oxygen

t h a t d i f f u s e s i n equals t h e amount of oxygen being used by r e sp i r ing

organisms (Figure 13) .

Although daytime r e s p i r a t i o n tends t o lower t h e amount of oxygen

i n t h e wa?er, t h i s i s masked by t h e inc rease i n oxygen caused by

photosynthesis . Thus, a r e a l mcasure of r e s p i r a t i o n during the day-

t ime is impossible by t h i s method. In order t o overcome t h i s

d i f f i c u l t y , it was o r i g i n a l l y suggested (Odum, 1956; Odum and

Hoskins, 1958) t h a t da.ytime r e s p i r a t i o n shoul d be approximately

equal t o n ight t ime r e s p i r a t i o n , and t h a t a l i n e drawn on t h e r a t e -

of-change curve a t t h e average n ight t ime r e s p i r a t i o n r a t e would

approximate daytime r e s p i r a t i o n (Figure 14a) . Fur ther refinement

of t h i s method (Odum and Wilson, 1962) t akes i n t o account t h e varying

na tu re of daytime r e s p i r a t i o n which i s g r e a t e r toward t h e end of t h e

day when temperatures and oxygen l e v e l s a r e h igher . Thus, a s lop ing

l i n e drawn from t h e pre-dawn low po in t on t h e rate-of-change curve

t o t h e pos t - sunse t minimum (Figure 14b) i s probably a more accu ra t e

r e p r e s e n t a t i o n of what i s occurr ing i n na ture . In almost a l l curves

analyzed f o r t h i s s tudy , t he .pos t - sunse t ra te-of-change po in t i s lower

than t h e pre-dawn p o i n t , i n d i c a t i n g g r e a t e r r e s p i r a t i o n during t h e l a t t e r

p a r t of t h e day.

Fu r the r s t u d i e s (So l l i n s , 1969; Odum, Nixon and DiSalvo, 1970)

have ind ica t ed t h a t daytime r e s p i r a t i o n may be considerably higher

due t o h igher oxygen l e v e l s and pho to re sp i r a t ion . Thus, t h e ac tua l

daytime r e s p i r a t i o n curve may d i p down cons iderably as suggested i n

Figure 14. Various l i n e s drawn t o r ep re sen t daytime

r e s p i r a t i o n : a . cons tan t daytime r e s p i r a t i o n a t t h e l e v e l of

average n ight t ime r a t e s (from Odum and Hoskins, 1958); b.

varying daytime r a t e s s i m i l a r t o vary ing n ight t ime r a t e s (Odum

and Wilson, 1962); c . hypothe t ica l curve assuming r e s p i r a t i o n

p ropor t iona l t o oxygen concent ra t ion (So l l i n s , 1965); and, d.

hypo the t i ca l curve co r r ec t ing f o r pho to re sp i r a t ion (Odum, Nixon,

and DiSalvo, 1970). Corrected rate-of-change curve from Wood

Bridge S t a t i o n , New Hope Creek, October 4 , 1968. Daytime r e s p i r a -

t i o n as represented by l i n e b was used i n t h e present s tudy.

G 12

T I M E

Figure 14 c and d. This oxygen consumption i s obviously compen-

s a t e d f o r by a g r e a t e r amount of oxygen being concurren t ly pro-

duced by photosynthesis , a s t h e oxygen l e v e l i n t h e water r i s e s

during t h e day. Thus, community metabolism during t h e day may be

cons iderably g r e a t e r than during t h e n igh t . However, u n t i l some

adequate means f o r measuring pho to re sp i r a t ion becomes a v a i l a b l e ,

t h e method of connecting t h e pre-dawn p o i n t by a s t r a i g h t l i n e t o

t h e pos t - sunse t p o i n t is , a t l e a s t , o b j e c t i v e and may c l o s e l y

r ep re sen t a l l community r e s p i r a t i o n except pho to re sp i r a t ion i n

green p l a n t s . This procedure has been used f o r a l l a n a l y s i s of

New Hope Creek da ta .

The g ros s community r e sp i r a t ion was est imated by i n t e g r a t i n g

t h e a r e a between t h e zero rate-of-change l i n e and t h e d i f f u s i o n -

co r r ec t ed r e s p i r a t i o n curve over 24 hours (Figure 13). Gross photo-

syn thes i s was est imated by i n t e g r a t i n g t h e a r e a between t h e daytime

r e s p i r a t i o n l i n e and t h e daytime rate-of-change curve (Figure 13) .

The i n t e g r a t i o n s can be done by counting squares on graph paper o r

by us ing a planimeter .

Est imate of Pdetabolism from pH Changes

An e s t ima te of community metabolism was made us ing t h e d iu rna l

pH method (Beyers e t a l . , 1963). The product ion of carbon d ioxide -- by t h e r e s p i r a t i o n of l i v i n g organisms produces carbonic a c i d by

t h e fo l lowing formula:

I C02 + H20 $ H2C03 , (o the r carbon compounds)

Thus, r e s p i r a t i o n lowers t h e pH of t h e water , and photosynthes is ,

by t ak ing C 0 2 out o-C fhe water , r a i s e s t h e pH. Since t h e i n t e r -

a c t i o n of var ious ca7:bon compounds i n n a t u r a l waters i s extremely

complicated and s u b j e c t t o unknown buf fe r ing , a p r i o r i coord ina t ion - -- of pH and amounts of C02 produced o r u t i l i z e d i s v i r t u a l - l y

impossible. However? t h i s r e l a t i o n can be determined empir ica l ly by

t i t r a t i n g t h e water of i n t e r e s t with d i s t i l l e d water of known C02

concent ra t ion (Beyers e t a l . , 1963). A sample t i t r a t i o n of New -- Hope Creek water wi th carbon d ioxide-sa tura ted d i s t i l l e d water i s

suppl ied (Figure 15) .

The change i n r e l a t i v e amounts of carbon d ioxide i n t h e water

can then be determined by reading t h e pH-CQ2 graph. To determine

abso lu t e va lues of C02 i n t h e water r e q u i r e s s epa ra t e de te rmina t ions

of t o t a l C02 a t t h e s t a r t of t i t r a t i o n . However, t h i s i s no t

necessary s i n c e t h e metabol ic de te rmina t ions a r e based on changes i n

C02, no t on abso lu t e va lues .

Est imates of t o t a l product ion and r e s p i r a t i o n from changes i n

C02 were made by a procedure s i m i l a r t o t h a t used f o r oxygen. P l o t s

were made of r e l a t i v e amounts of CQ2 i n t h e water over 24 hours

(Figure 16) . The f i r s t d e r i v a t i v e of t h i s was p l o t t e d a s a nega t ive

func t ion t o make t h e r e s u l t s compatible with oxygen da t a which, of

course, behave i n an oppos i te fash ion . Daytime r e s p i r a t i o n was

es t imated according t o t h e methodology d iscussed i n the previLjli

s e c t i o n , and t o t a l photosynthesis and r e s p i r a t i o n were determined by

i n t e g r a t i n g t h e same a r e a s a s d iscussed f o r oxygen. A sample

de te rmina t ion i s included (Figure 16) . No c o r r e c t i o n f o r d i f f u s i o n

Figure 15. Carbon d ioxide t i t r a t i o n of New Hope Creek water

f o r metabol ic s t u d i e s . The a b s c i s s a r e p r e s e n t s t h e carbon d ioxide

i n t h e water sample added t o t h a t p re sen t a t t h e s t a r t of t h e

t i t r a t i o n .

Figure 16. Est imation of metabolism i n New Hope Creek, Diurnal

pH method. February 21, 1969. The upper graph r ep resen t s pH over

24 hours i n New Hope Creek, a l s o changes i n t h e carbon dioxide content

of t h e water . The lower graph i s t h e r a t e of change based on t h e above

carbon d iox ide va lues . Gross photosynthes is and community metabolism

a r e es t imated a s i n Figure 13. No co r rec t ions were made f o r d i f f u s i o n

of C02, b u t t h e r e s u l t s of t h i s method (Gross product ion = 1.33

gm m-3 day-', Resp i r a t ion = 2.0 gm m-3 day-l) agree f a i r l y wel l wi th

uncorrec ted- for -d i f fus ion oxygen e s t ima te s of 1.5 and 2.2 gm mw3 day-l

r e spec t ive ly .

G O % / L I T E R

C SUNRISE

was made, but outward d i f fus ion of C02 a t n igh t may inc rease t h e

es t imated community r e s p i r a t i o n .

- Computer Program f o r Est imating Community Metabolism from Diurnal Oxygen

4

Curves

Since t h e c a l c u l a t i o n s involved i n t h e s e determinat ions a r e long,

t ed ious , and s u b j e c t t o human e r r o r , u se was made of t h e I n t e r n a t i o n a l

Business Machine M ~ d e l 70 d i g i t a l computer and Calcomp p l o t t e r ,

l oca t ed a t t h e Tr i -Univers i ty Computation Center and t h e Univers i ty

of North Carol ina, r e spec t ive ly . These a r e l inked by cab le and a r e

completely coordinated. Appendix B i nc ludes a flow cha r t of t h e

program, t h e computer program, i n PL/1, and i n s t r u c t i o n s f o r e n t e r i n g

da t a .

F i sh Sampling Procedures and Apparatus

Design of Weirs and Traps - Weirs t o measure movements of s t ream f i s h e s were cons t ruc ted

fol lowing t h e b a s i c p lan of S h e t t e r (1938). Since many d i f f e r e n t

l o c a t i o n s were sampled over a cons iderable t ime s c a l e , v a r i a t i o n s i n

t h e b a s i c s e t u p occurred with d i f f e r e n t bottom types and evolu t ion of

design. A summary of t h e d i f f e r e n t modi f ica t ions used is given i n

Table 3 .

The b a s i c p l an f o r t h e 'hardware-cloth" (wire screening) weir

(Figures 17 and 18) was t o s t r e t c h a 0.6 cm (1/4") mesh b a r r i e r ac ros s

t h e s t ream a t an angle such t h a t migrat ing f i s h e s would be funneled

i n t o t r a p s placed a t e i t h e r s i d e of t h e stream. The lower 30 cm o r

so of t h e screening was bent a t a 90 degree angle t o t he v e r t i c a l and

placed on t h e rock-cleared stream bottom. Rocks were then placed on

Table 3 . Modificat ians i n Basic Trapping Procedure

Date Locat ion New Modification

Apr i l 10, 1969

Apr i l 15, 1968

June 5, 1968

June 14, 1968

October 22, 1968

February 1 2 , 1969

May 6, 1969

September 21, 1969

February 21 , 1970

Concrete

Concrete

Wood S t a t i o n

Jungle S t a t i o n

Jungle S t a t i o n

Big Pool S t a t i o n

Wood S t a t i o n

Big Pool S t a t i o n

Big Pool S t a t i o n

F i r s t day sampled. Downstream only.

Upstream and downstream t r a p s i n s t a l l e d .

Upstream t r a p only i n s t a l l e d

Upstream t r a p i n s t a l l e d

Upstream and downstream t r a p s i n s t a l l e d

~ a r g e - s i z e d ( 6 6 X 132 X 132 cm) t r a p i n s t a1 l e d (down) . Upstream and downstream t r a p i n s t a l l e d

Pipe-weir i n s t a l l ed

Large-sized t r a p s i n s t a l l e d up and down

Figure 18. Design of f i s h weirs used i n New Hope Creek.

a . Arrangement of weir and t r a p s . b. Hardware c l o t h weir .

c. S t e e l p ipe weir wi th wooden suppor ts , p l a s t i c spacing c o l l a r s

and hardware c l o t h s e a l with bottom of stream.

t h e top of t h i s screening, and t h e e n t i r e boundary of t h e screen

and s tream bottom Iws checked f o r ho les through which f i s h miphf:

pass.

In t h e summer of 1969, a more e l abo ra t e weir was c m s t r u c t c d

(Figures 18 and 19) . T??is weir was designed t o overcome t h e pro-

blem of leaves accumula.tjng i n t h e upstream s i d e of t h e hardld~are

c l o t h weir during high water. S t e e l e l e c t r i c a l condirit p ipes

spaced a t 1.5 cm i n t e r v a l s by p l a s t i c c o l l a r s were used fol lowing

the recommendations of Feimers (1966).

A f i s h - t i g h t border wi th the bottom of t h e stream was c rea t ed

by bending a 1 / 2 m wide s e c t i o n of hardware c l o t h over t h e lowermost

p ipe and by p i l i n g rocks on e i t h e r s i d e of t h i s . The p ipes were

he ld i n p o s i t i o n by cement-anchored wooden frames t h a t allowed

t h e p ipes t o be i n s e r t e d and removed. Hardware c l o t h t r ap -en t r ance

cones provided en t rance t o t h e a c t u a l t r a p s .

Traps were placed a t both ends of t h e weir ; one designed t o

ca tch f i s h moving upstream and t h e o t h e r downstream (Figure 18) .

Frames f o r t h e t r a p s were c o n s t r u c t e d o f 2 b 2 c m by 66 o r 132 cm

p ieces of aluminum (Figure 2 1 ) . This was covered with 0.6 cm wire

c l o t h t o form a r e c t a n g u l a r t r a p . A cone was cons t ruc ted a t one end

t h a t f i t t e d i n a corresponding cone i n t h e weir t o form a t i g h t f i t

t h a t could be e a s i l y separa ted f o r f i s h removal. The o t h e r end of

t h e t r a p contained a 10 cm by 10 cm spout with door t o f a c i l i t a t e

f i s h removal. A l a r g e r door was a l s o included.

Figure 19. Big Pool sampling s t a t i o n , looking downstream

during normal s p r i n g flow. The t r a p t o t h e l e f t c e n t e r of t h e

p i c t u r e ca tches f i s h moving upstream and t h e t r a p t o t h e r i g h t

c e n t e r ca tches f i s h moving downstream. The f a r r i g h t bank of

p ipes has been l i f t e d t o show underwater arrangement.

Figure 20. Y3idewaysf1 f i s h sampling arrangement. The en t rance

cone t o t h e t r a p i s v i s i b l e i n t he c e n t e r o f t h e t r a p . Also v i s i b l e

a r e t h e p l a s t i c pipespacing c o l l a r s and t h e method of anchoring t h e

p ipe suppor ts with rocks and concre te .

Figure 21. Design of f i s h t r a p used i n New Hope Creek. The

frame i s made from 2 by 2 cm by 66 o r 132 cm s e c t i o n s o f aluminum.

The s c r e e n i n g i s 1/4 inch (0 .6 cm) hardware s c r e e n , heirl t o t h e frame

w i t h aluminum "pop" r i v e t s . A cone o f w i r e s c r e e n i n g on t h e we i r

f i t s i n s i d e t h e s i m i l a r cone i n t h e t r a p . The smal l box i s a

spou t f o r removing smal l f i s h .

Check on Possib1.e Sampling Bias i n Up and Down Traps - - ..--- - The c o n s i s t e n t l y g r e a t e r ca tch i n f i s h mass moving upstream com-

pared t o f i s h mass moving downstream r a i s e d t h e p o s s i b i l i t y of

sampling b i a s - - t h a t f i s h e n t e r t h e upstream t r a p more r e a d i l y than

the downstream t r a p . This was considered u n l i k e l y , a s a c e r t a i n

a d d i t i o n a l amount of e f f o r t would have been needed by t h e f i s h t o

swim up i n t o t h e t r a p ; whereas, a f i s h e n t e r i n g t h e downstream t r a p

could d r i f t pa s s ive ly wi th t h e c u r r e n t . However, t h i s p o s s i b i l i t y

was checked by two means, t h e "sideways" t r a p and t h e "double

reverse1 ' t r a p (Figures 20 and 2 2 ) . The sideways t r a p was designed

so t h a t f i s h e n t e r i n g from e i t h e r upstream o r downstream would have

t o e n t e r t h e a c t u a l t r a p sideways t o t h e c u r r e n t . This was con-

s idered t h e p r i n c i p a l check on t r a p b i a s . In t heo ry , i f t h e r e were

s u b s t a n t i a l e r r o r introduced by t h e f ac ing of t h e weirs t h i s would

become apparent by extreme v a r i a t i o n s from t h e expected r a t i o of

f i s h moving upstream t o f i s h moving down. However, dur ing the use

of t h e s e t r a p s about t h e same r a t i o of upstream t o downstream move-

ment occurred a s dur ing normal sampling a t s i m i l a r t imes of t h e

year (Table 4 ) .

A f u r t h e r check on t r a p b i a s was t h e use of t h e "double r eve r se"

t r a p s i n which f i s h moving upstream would have t o t u r n around and

e n t e r t h e t r a p moving downstream, and v i c e v e r s a . No l a r g e f i s h a t -- a l l were caught moving ~ ~ s t r e a m dur ing t h e use of t hese t r a p s , and

only a few were caught moving downstream even dur ing per iods of ex-

pected f i s h movement. Thus, perhaps t h e u l t i m a t e t e s t of t r a p b i a s

Figure 22. a. "sideways" and b. f fdouble reverse" wei rs used

t o t e s t p o s s i b l e sampling b i a s i n normal t r a p arrangement.

Table 4. Catch of F ish i n 'Sideways1 Traps

Date Number up Number down bQass up Mass down

1970

Apr i l 23

Apr i l 24

Apr i l 25

Apr i l 26

May 18

May 19

May 20

May 26

To ta l s 8 9 61 3052 1674

f a i l e d and t h e p o s s i b i l i t y of t r a p b i a s remains, although the

r e s u l t s of t h e l~sideways" t r a p s i n d i c a t e s t h a t t h i s i s un l ike ly .

Check on Rate of F ish Escape from Traps - The t rapping procedure used i n t h i s s tudy i s based on the

assumption t h a t it i s much e a s i e r f o r a f i s h t o swim i n t o t h e t r a p s

than t o s w i m ou t of them, The cone en t rances make t h i s seem

i n t u i t i v e l y t r u e . However, t h i s was checked by a s e r i e s of 18 s e t s

of experiments i n which f i s h t h a t were a l r eady caught i n t h e t r a p s

were l e f t i n them f o r an add i t i ona l 24 hours. Thus t h e r a t e of t r a p

escape was measured.

A t o t a l of 132 f i s h weighing 10,364 g was placed i n t h e t r a p s .

Of t h e s e , 96 f i s h weighing 7747 g were recovered. Thus, over t h e

24-hour i n t e r v a l , 36 f i s h weighing 2517 g escaped. Since each f i s h

o r i g i n a l l y t rapped would have been i n t h e t r a p s an average of 1 2

hours (one-half t h e t r app ing t ime i n t e r v a l ) it was assumed t h a t t h e

escape r a t e f o r a normal day would be h a l f of t he 24-hour escape r a t e .

Correc t ing t h e above d a t a f o r t h i s g ives an average d a i l y escape

r a t e of 13.6 percent f o r numbers of f i s h and 12.3 percent f o r mass of

f i s h . No important d i f f e r e n c e i n escape r a t e s f o r d i f f e r e n t t r a p s was

noted except t h a t t h e smal le r (66 cm by 66 cm by 132 cm) t r a p , when

used t o ca tch f i s h moving downstream, had a g r e a t e r escape r a t e than

o the r t r a p s . This t r a p was rep laced i n February, 1969, by a l a r g e r

one from which v i r t u a l l y no f i s h escaped. Since the escape ra tes kqeie

not l a r g e , no co r r ec t ions were made i n t h e d a t a used f o r ana lys i s .

S a m ~ l i n ~ Modif icat ions f o r Low Water

During t h e summsr o f 1968 and t h e summer and f a l l of 1969 very

low r a i n f a l l condi t ions ex i s t ed i n c e n t r a l North Carol ina, and

extremely low flows i n New Hope Creek r e s u l t e d (Figure 23). However,

l a r g e pools remained throughout t h e stream; and i f t h e r u n of f i s h e s

captured i n t h e s p r i n g o f 1969 i s acy i n d i c a t i o n , t h e f i s h populat ions

were not adverse ly a f f ec t ed . During t h e s e per iods the Big Pool weir

was no longer f u n c t i o n a l because t h e water dropped below t h e l e v e l

of t h e t r a p en t rances . Sampling was conducted only a t t h e Concrete

Bridge s i t e , where adequate water depth was p re sen t .

Sampling Modif icat ions During High Waters --.--

New Hope Creek was sub jec t t o extreme f looding condi t ions during

t h e f a l l , win ter , and sp r ing months (Figures 24 and 25) . The magnj-

tude of some of t h e flows made any sampling of migra t ion impossible;

however, over t h e course of t h i s p r o j e c t va r ious sampling modi f ica t ions

were made t o i nc rease t h e l e v e l a t which f i s h counts could be made.

Table 5 g ives t h e maximum flow a t which sampling could be maintained

during t h e s tudy and t h e da t e s a t which sampling was impossible be-

cause of h igher water .

I n i t i a l l y t h e he ight of t h e weir was increased with add i t i ons of

wire c l o t h . This reached a p r a c t i c a l l i m i t a t a s t a g e l e v e l of about

46 cm above zero flow, and even l e s s during per iods of heavy l ea f

flow.

In February, 1969, t h e sampling s t a t i o n was moved t o a wider

po r t ion of t h e stream so t h a t a given inc rease i n water flow would

cause l e s s of an inc rease i n water he ight . Thus sampling was i n i t i a t e d

a t t h e Big Pool s i t e , loca ted about 100 m above t h e Concrete Bridge

Figure 23 . New Hope Creek dur ing drought (September, 1968) .

a . R i f f l e a r e a a t Wood Bridge sampling s t a t i o n . b . Pool a r ea

below Wood Bridge S t a t i o n .

Figure 24. New Hope Creek a t Concrete Bridge S t a t i o n during

f lood. a. shows r i f f l e a r e a j u s t above f i s h sampling s t a t i o n

(Apri l 14, 1970). b. shows l a r g e pool between the concre te

b r idge and t h e Concrete Bridge sampling s t a t i o n (March 21, 1970).

F i g u r e 25. a . Overrun of we i r d u r i n g s e v e r e f l o o d a t

Big Pool S t a t i o n on A p r i l 14, 1970. b . D e s t r u c t i o n o f t r a p p i n g

a p p a r a t u s , A p r i l 14, 1970.

Table 5. Floods i n New Hope Creek That Affected Sampling

Date of s t a r t Maximum s t age l e v e l Maximum stage t h a t Day weir of flood i n cent imeters above could be sampled inoperable

zero flow a t t h a t t ime i n cm above zero flow

May 27 Oct. 19 Nov. 12 Nov. 19 Dec. 4

Jan. 21 Feb. 9 Feb. 28 March 3 March 18 March 24 June 16 Aug. 6 Oct. 2

Jan. 30 Feb. 2 Feb. 17 Feb. 25 March 5 March 21 March 31 Apr i l 14

s i t e . This provided a means of sampling t o a s t a g e l e v e l of about

76 cm, i f l e a f flow was small . During high waters a s e r i e s of rock

and wire suppor ts were u t i l i z e d t o maintain t h e weir i n an upr ight

pos i t i on . Removal of f i s h and genera l maintenance were performed two,

t h r e e , o r fou r t imes each day during flows t h a t approached t h e l i m i t

of sampling. During t h e f a l l of 1969 t h e en t rance cone poin t ing

downstream was l i n e d with a s t i f f c l e a r p l a s t i c shee t so t h a t

clogging of t h i s cone with leaves would not occur.

The f i n a l major modi f ica t ion t o a i d i n sampling during high

water was t h e cons t ruc t ion of t h e p ipe weir d i scussed previous ly .

This allowed sampling t o a s t a g e l e v e l of about 76 cm, even during

per iods of moderately heavy l e a f d i scharge . Sampling was impossible

during higher s t a g e l e v e l s , and t h e t r a p s and p ipes were removed

t o avoid t o t a l d e s t r u c t i o n of t h e apparatus . Floods of t h i s magni-

tude r a r e l y happened during per iods of expected heavy migrat ion.

Methodology of Handling Species and Species Groups f o r Analysis

Due t o some u n c e r t a i n t i e s i n taxonomy dur ing t h e beginning of

t h i s s tudy , and t o t h e n e c e s s i t y of s imp l i fy ing t h e l a r g e amounts

of d a t a f o r a n a l y s i s , a l l organisms were placed i n t o one of 2 1

taxonomic groups a s l i s t e d i n Table 6. A l l taxonomic groups of more

than one spec i e s a r e l i s t e d below.

pickerels sf^ inc luded both chain p i c k e r e l (Esox n i g e r ) and r e d f i n -- p i c k e r e l (Esox americanus), but t h e r e d f i n p i cke re l was captured only - as a r a r i t y . F l a t bul lhead ( I c t a l u r u s p la tycephalus) has r ecen t ly

been subdivided i n t o two spec i e s , I. p la tycephalus and I . brunneus. - -

They a r e considered a s one spec i e s f o r t h i s s tudy , although both

Table 6. Fishes Captured in New Hope Creek and Groupings Used

to Simplify Analysis.

Analyzed as: Common name a Scientific name a

B. crappie

Bluegill

B. H. Chub

Bullhead

Chubsucker

Creek chub

Darter

H.F. shiner

Madtom

Pickerel

Redhorses

Pirate perch

Black crappie

Bluegill

Bluehead chub

Flat bullhead

Snail bullhead

Creek dhubsucker

Creek chub

Johnny darter

Piedmont darter

Highfin shiner

Margined madtom

Chain pickerel

Redfin pickerel

Smallfin redhorse

V-lip redhorse

Pirate Perch

Pomoxis nigromaculatus (~eseur) -

Lepomis macrochirus Rafinesque

Hybopsis leptocephalus (Girard) -

Ictalurus platycephalus (Girard)

Ictalurus brunneus (Jordan)

Erimyzon oblongus (blitchi 11)

Semotilus atromaculatus (Mitchill)

Etheostoma nigrum Rafinesque

Percina crass a (Jordan and Brayton)

Notropis altipinnis (Cope)

Noturus insignis (Richardson)

Esox niger (Le~eur) -- Esox americanus Gmelin

Moxostoma robustum (Cope)

Moxostoma collapsum (Cope)

Aphrododerus sayanus (Gilliams)

a Nomenclature used follows A List of Common and Scientific Names --- - o f Fishes from the United States and Canada, 1960, American Fisheries - -- - Society publication No. 2., Waverly Press, Inc., Baltimore,l02 p.

Table 6 . Continued

Analyzed a s : Common name a S c i e n t i f i c name a

Pumpkinseed Pumpkinseed

sun f i sh Redbreast sun f i sh

s h i n e r White s h i n e r

Sandbar sh ine r

M . s h i n e r Whitemouth sh ine r

Others American e e l

Bow f i n

Gizzard shad

Green sun f i sh

Largemouth bass

Lepomis gibbosus (Linnaeus)

Lepomis a u r i t u s (Linnaeus)

Notopis a lbeo lus Jordan -- Notropis s cep t i cus (Jordan and

G i l b e r t )

Notropis a lborus Hubbs and Raney

Anguilla r o s t r a t a (LeSueur) -- Pmia ca lva Linnaeus -- Dorosoma cepidianum (LeSueur)

Lepomis cyanel lus Rafinesque

Micropterus salmoides (Lacepede)

Speckled k i l l i f i s h Fundulus ra thbuni Jordan and Meek

a r e common i n New Hope Creek. 'Redhorses' inc lude both v - l i p

redhorse (Moxostoma collapsum) and t h e sma l l f in redhorse (Moxo-

stoma robustum). About two-thirds of t h e redhorses captured were

v - l i p . The p a t t e r n s of movement of both spec i e s were not obvious-

l y d i f f e r e n t .

'Larger N o t r o p i s ~ n c l u d e d seve ra l spec i e s t h a t were morph-

o l o g i c a l l y q u i t e s i m i l a r t o an unt ra ined eye. The most abundant of

t h e s e was t h e whi te sh ine r , Notropis a lbeolus although t h e

sandbar s h i n e r , Notropis s cep t i cus , and probably seve ra l o t h e r

spec i e s , were a l s o taken. Dar te rs were gene ra l ly Johnny d a r t e r ,

Etheostoma nigrum, although some Piedmont d a r t e r s , Percina c r a s s a ,

were a l s o captured.

'Crayf i sh1 included members of up t o four spec ies l i s t e d a s

being p re sen t i n New Hope Creek (Hobbs, mimeographed). No attempt

was made t o s e p a r a t e t hese i n t o spec ies . 'Frogs ' included seve ra l

spec i e s , and ' T u r t l e s ' included f o u r spec ies . 'Others ' included

largemouth bass , Micropterus salmoides; green sun f i sh , Lepomis

cyanel lus ; bowfin, -- Amia ca lva ; g izzard shad, Dorosoma cepedianum;

and snakes, wooddrnks, muskrats, l a r g e bugs and var ious o t h e r organ-

isms. Table 7 g ives an a n a l y s i s of 'o ther lorganisms encountered.

Dai ly F i sh Sampling Procedure

The f i s h were removed from t h e t r a p s and ind iv idua l ly weighed.

Smaller f i s h were marked by a f i n c l i p d i s t i n c t i v e f o r each s t a t i o n .

Fish l a r g e r than about 80 g were tagged with ind iv idua l ly numbered

d a r t t a g s (Floy Tag and Manufacturing Company). The l i v e f i s h were

re turned t o t h e water about 50 m upstream o r downstream from t h e wei rs ,

i n t h e d i r e c t i o n they were moving.

Table 7 . Organisms Other Than Fish Captured i n New Hope Creek

Common name S c i e n t i f i c name Number captured

Dragonfly nymph Order Odonata 1

Walking s t i c k Anisomorpha. s p . - -- 1

Water bugs 2

Crayf i shes Cambarus spp . - 446 Procambarus s p . -

Newt Notopthalmus v i r idescens 2

Toads Bufo spp . -- Snapping t u r t l e Chelydra se rpen t ina

Pa in ted t u r t l e Chrysemys p i c t a a

Mud t u r t l e

Musk t u r t l e

Kinosternon sp . - Sternothaerus s p . -

Black (Racer) snake Coluber c o n s t r i c t o r

Common water snake Nat r ix sipedon

Wood duck Aix sponsa P

a . 62 t u r t l e s of t h e fou r types

RESULTS

Physical Data

Stream Morphology

New Hope Creek, i n t h e one km s t r e t c h above Concrete Bridge

S t a t i o n , averages 11.6 m i n width and 0.45 m i n depth i n a normal

spr ing . Measurements taken a t t h i s and o t h e r s t a t i o n s a r e given

i n Tables 8-11. From t h e s e measurements i t becomes apparent t h a t

t h e g r e a t e s t depth i n New Hope Creek, a t l e a s t i n t h e a r eas sampled,

i s above t h e Wood Bridge, and t h e l e a s t depth i s above Blackwood

S t a t i o n . A l l depth va lues taken on d i f f e r e n t d a t e s were cor rec ted

t o ' s tandard water l e v e l t (50 cm above zero f low) , which was the nor-

ma1 sp r ing flow.

Stream Level and Discharge Rate

Stage l e v e l a t t h e Concrete Bridge S t a t i o n va r i ed from a minimum

of 0 cm above t h e l e v e l of no flow t o a maximum of more than 100 cm

above t h e l e v e l of no flow. In spr ingt ime normal s t a g e l e v e l s were

about 50 cm. Figure 26 g ives d a i l y water l e v e l s f o r t h e 27 months of

t h i s s tudy.

Dai ly d ischarge r a t e s va r i ed from summer and drought l e v e l - ~ ) f

0 m3 daym1 t o sp r ing f lood l e v e l s of a t l e a s t 7 l o 5 m3 day-' o r

251 cub ic f e e t second-l.

Table 8. Depth and Width P r o f i l e f o r 300 m Below Concrete Bridge,

May 23, 1970

Meters below s t a t i o n Width

m

Mean dep th

m

5 0 14.9 0.55

100 10.9 0.37

150 8.4 0.26

200 4.3 0.23

250 5.7 0.37

300 11.8 0.45

Average 9.33 0.45

Cor r ec t i on t o s t anda rd flow (See t e x t f o r exp lana t ion)

- Average dep th a t s t anda rd f low 0.54

Table 9. Depth and Width P r o f i l e f o r 1.8 km Above Concrete Bridge,

S t a t i o n , Apr i l , 1969

Meters above s t a t i o n Width

m

Mean depth

m

Average 0.445 a t s tandard flow

Table 10. Depth and Width P r o f i l e f o r 900 m Above Wood Bridge

S t a t i o n , May 13, And 23, 1970

Meters above s t a t i o n Width

Mean depth

Average

Correc t ion t o s tandard flow . 0.07

- Average depth a t s tandard flow 0.68

Table 11. Depth and Width P r o f i l e For t h e Zone 1000 m Above Blackwood

Sampling S t a t i o n , May 18, 1970

Distance above s t a t i o n Width

m Average depth

m

Average 7.1 0.23

Correc t ion t o s tandard flow 0.05

Average depth a t s tandard flow 0.28

Figure 26. Dai ly water s t a g e l e v e l , i n cm above zero flow, of

New Hope Creek a t Concrete Bridge S t a t i o n .

Stream Temperatures

The minimum temperature recorded i n New Hope Creek was 0" C

during January, 1969 and 1970. A maximum temperature of 28" C was

recorded on J u l y 21, 1969. Average d a i l y temperatures f o r t h e

s tudy per iod a r e presented i n Figure 27.

Light I n t e n s i t y A t Surface of Stream

The i n t e n s i t y of l i g h t s t r i k i n g t h e su r f ace of New Hope Creek

was measured on a completely c loud le s s day a t 100 m i n t e r v a l s f o r

1 km above both t h e Blackwood S t a t i o n and t h e Concrete S t a t i o n

(Table 12) . The r e s u l t s i n d i c a t e t h a t when l e a f canopy i s f u l l

about 11 percent of t h e s o l a r l i g h t energy e n t e r s t h e aqua t i c

ecosystems a t each s t a t i o n with ranges from 9 t o 66 percent of t h a t

above t h e f o r e s t . The r e s u l t s a t t h e Blackwood S t a t i o n may be

b iased by t h e one very l a r g e va lue (6080 foot -candles) which was

not r e p r e s e n t a t i v e f o r t h a t s t r e t c h .

Leaf Discharge

Measured d a i l y l e a f d i scharge a t t h e Concrete Bridge S t a t i o n

va r i ed from zero t o about 825,000 g. The amount discharged pe r day

was l i n e a r l y p ropor t iona l t o , w a t e r s t a g e when p l o t t e d on semilog

paper (Figure 6) . Estimated t o t a l monthly l e a f d i scharge i s given

i n Table 29, found i n t h e "Discussion."

Total Phosphorus

Water samples analyzed f o r t o t a l phosphorus i n d i c a t e t h a t New

Hope Creek has about t h e same amount of phosphorus (Table 13) a s many

o the r f reshwater environments summarized i n Hutchinson (1957). Values

ranged from not d e t e c t a b l e ( l e s s than 0.005 ppm) t o 0.26 ppm (or mg 1 - l ) .

Figure 27. Mean d a i l y temperatures f o r New Hope Creek dur ing

t h i s s tudy.

q- P- *,L a

*-4

LA-

ti:.

Table 12. L igh t I n t e n s i t y a t S u r f a c e o f New Hope Creek

Meters L i g h t Date above I n t e n s i t y and oxygen sampling f o o t - Time Loca t ion s t a t i o n c a n d l e s

Sep t . 26, 1969 Open F i e l d : S t a r t 1 1 : 4 0 - 1 2 ~ 1 8 B l ackwood S t a t i o n

100

200

300

400

500

600

700

800

900

1000

Average

Open F i e l d : F i n i s h

S e p t . 26, 1969 Open F i e l d : S t a r t 1*3:!?0-14:OO C x L c r e t e BY, l g e

S t a t i o n

Table 13. To ta l Phaphorus (Dissolved and Suspended) i n New Hope

Creek a t Concrete Bridge S t a t i o n

Date Water s t age Tota l phosphorus (ppm) Manual Autoanalyzer

1968

Aug. 29 5

Sept . 18 1

Sept . 23 0

Oct. 25 18

Nov. 9

Nov. 24

Dec. 16

Dec. 22

Jan . 8 3 3

Jan. 20

Jan . 24

Jan . 28

Feb. 3

Feb. 4

Feb. 9

Table 13. Continued

Date Water s t a g e Tot a1 phosphorus (ppm) Manual Autoanalyzer

1969 (Continued)

Mar. 3

Mar. 5

Mar. 7

Mar. 8

Mar. 10

Mar. 16

Mar. 19

Mar. 21

Mar. 26

Apr. 2

Apr. 6

Apr. 10

Apr. 11

Apr. 17

Apr. 20

Apr. 25

May 8

May 14

May 18

June 4

J u n e 11

J u l y 2

J u l y 1 3

0.270

0.050

0.030

0.020

0.020

0.060

0.100

0.030

0.020

Not d e t e c t a b l e

0.050

0.020

0.060

0.020

0.250

0.130

0.060

0.060

0.060

0.160

0.040

0.080

Normal va lues were i n t h e range of 0.02 t o 0.1 ppm and t h e average

f o r a l l samples taken was 0.06 ppm. No seasonal t r end was ev i -

dent and samples o f t e n v a r i e d widely from one sampling d a t e t o

t h e next . Nor was t h e r e any cons i s t en t r e l a t i o n between r i v e r

d i scharge and phosphorus concent ra t ions .

Nitrogen

Severa l t o t a l n i t rogen analyses were done and t h e d a t a a r e

presented he re f o r p o s s i b l e u s e a s base l i n e d a t a (Table 14) .

The r a t i o of N t o P v a r i e d from 6.5:l t o 210:l.

Stream Conduct ivi ty

The r e s u l t s of t h r e e s t ream conduc t iv i ty de te rmina t ions a t

d i f f e r e n t t imes of t h e year were from 63 t o 200 um ohms.

Metabolic S tud ie s

The r e s u l t s of a l l metabol ic s t u d i e s a r e given i n Tables 15

t o 17 and i n Figures 28 t o 36 and i n t h e Appendix.

Daily Var i a t ions i n Oxygen

On a l l days on which oxygen was measured t h e oxygen showed some

man i fe s t a t ion of t h e expected d iu rna l curve, t h a t i s , it r o s e dur ing

t h e day l igh t hours and dropped a t n igh t . Within t h i s genera l p a t t e r n

many v a r i a t i o n s were observed (Figures 28-33; Appendix C ) .

Annual Var i a t ions i n Metabolism

Photosynthesis i n New Hope Creek a t t h e Concrete Bridge S t a t i o n ,

which was t h e s t a t i o n most heav i ly sampled, va r i ed from about 0.21

m N 0 O P N y g g g g g g . z .

t o 8.85 g 02 mm2 day-l (0.58 t o 10.88 g O2 mm3 d a y m l ) . Gross

community r e s p i r a t i o n v a r i e d from 0.39 t o 13.40 g O 2 mm2 day-l

(0.94 t o 16.30 g O2 rn-3 dayw1). Typical d i u r n a l curves f o r t h e

Concrete Bridge, Wood Bridge, and Blackwood s t a t i o n s f o r t imes of

g r e a t e s t and l e a s t metabolism (spr ing and l a t e f a l l o r win ter ) a r e

given i n Figures 28-33. Annual r e s u l t s i n which t h e same months

f o r d i f f e r e n t yea r s a r e lumped, a r e p l o t t e d i n Figures 34-36.

Thus New Hope Creek has an annual cyc le of metabolism t h a t

r e p e a t s f a i r l y c o n s i s t e n t l y from one yea r t o t h e next . Both

photosynthesis and r e s p i r a t i o n a r e l e a s t i n t h e winter . Primary

product ion inc reases a s t h e season progresses , reaching a peak

i n March and Apr i l when l i g h t s t r i k i n g t h e s u r f a c e i s a l s o maxi-

mal. Resp i r a t ion fo l lows a s i m i l a r p a t t e r n but remains high

throughout t h e summer. Very high r e s p i r a t i o n i s a s soc i a t ed with

high p r o d u c t i v i t y and/or high temperatures and low water. March

1970 had a g r e a t e r metabolism than a t any o t h e r t ime s tud ied , and

the h ighes t metabolism recorded (March 13) was a s soc i a t ed with a

small f lood. Tables 15-17 g ive va lues f o r each day sampled.

S p a t i a l Var i a t ions i n Metabolism

On a l l days s tud ied , volume metabolism was g r e a t e s t a t t h e

Blackwood S t a t i o n . Except during t h e l a t e r summer, t he /'bod Bridge

S t a t i o n has t h e l e a s t volume metabolism (Tables 15 - 1 7 ) . Areal

va lues were gene ra l ly s i m i l a r on any one d a t e a t a l l s t a t i o n s . The

r e l a t i v e l y high a r e a l product ion and r e s p i r a t i o n a t Wood Bridge

S t a t i o n during t h e summer may be a r e s u l t of an erroneous water

Figure 28. Typical d i u r n a l oxygen curve f o r sp r ing , Concrete

Bridge S t a t i o n .

Figure 30. Typical d i u r n a l oxygen curve f o r s p r i n g ,

Blackwood S t a t i o n .

Figure 31. Typical d i r u n a l curve f o r l a t e f a l l , Concrete Bridge

S t a t i o n .

- SU?iRlSE. .--,---

* SUNSET I k'-"- Y - - f - - - - - - - - - F - -

.00 5.00 9.00 33.0 17.0 21.0

Figure 32. Typical d iu rna l curve f o r win ter , Wood Bridge

S t a t i o n .

Figure 33. Typical d i u r n a l oxygen curve f o r l a t e f a l l , Blackwood

S t a t i o n .

' CCR.

. O

N CO

0

G-l d

0

a Ln

N

N Ln

A

N M

0

a

0, d

Table 15. Continued

Wink1 e r Gross Community Gross Community o r Depth product i o n r e s p i r a t i o n product i o n r e s p i r a t i o n

Date probe m g 0, m-3 day-' g O2 m-3 day-l g 07 m-2 day-' g 07 m-2 day-'

J u l y 27 P 0.45 0.99

Aug. 25 W 0.37 0.77

Oct. 3 P 0.65 1.23 4.28

4 P 0.44 1.36 4.15

5 P 0.41 0.74 2.54

Nov. 16 P 0.35 1.22 3.34

2 1 W 0.36 0.60 1.58

1970

Feb. 14 W 0.40

Mar. 13. P 0.50 6.30 7.15 3.15 3.58

Figure 34. Annual variation in metabolism, Concrete Bridge

Station, New Hope Creek, April, 1968 - May, 1970. The solid line '

connects means of gross photosynthesis for each month and the

broken line connects means of community respiration for each

month. The vertical bars are 1 standard deviation from the mean.

The horizontal axis is months.

Figure 35. Annual variation in metabolism, Wood Bridge Station,

New Hope Creek, June, 1968 - August, 1969. The solid line connects

measurements of gross photosynthesis and the other line connects

measurements of community respiration.

F i g u r e 36. Annual v a r i a t i o n i n metabolism, Blackwood S t a t i o n ,

New Hope Creek, February, 1969 - February 1970. The s o l i d l i n e

connec t s measurements o f g r o s s p h o t o s y n t h e s i s and t h e o t h e r l i n e

connec t s measurements o f community r e s p i r a t i o n .

depth, s i n c e dur ing low waters t he water does no t flow r a p i d l y

enough t o be inf luenced by t h e deeper 'upstream water which was

included i n e s t ima te s of average water depth. When no flow

a t a l l was p re sen t , mean depth measurements were taken i n t h e

pool sampled i t s e l f , e l imina t ing t h i s source of e r r o r .

Annual and S p a t i a l Var i a t ions i n P/R Rat io

New Hope Creek a t t h e Concrete Bridge S t a t i o n e x h i b i t s an

annual v a r i a t i o n i n t h e r a t i o of photosynthes is t o r e s p i r a t i o n

(Figure 37). The stream a s a whole i s more au to t roph ic i n t h e

spr ing and becomes inc reas ing ly he t e ro t roph ic during t h e summer.

Only r a r e l y was t h e stream running e n t i r e l y upon energy produced

t o t a l l y wi th in i t s boundaries.

F ish Movements

The d a t a f o r f i s h movements a r e presented i n Table 18 t o 25

and Figures 38 t o 41, and i n Appendix D.

Analysis of A l l Species Considered Together: P r inc ipa l Sampling S t a t i o n

A ~ r i l 1968 - June 1970

During t h e 27 months (785 days) of t h i s s tudy , f i s h movement was

sampled on 455 days. During t h i s per iod 6,043 f i s h and o t h e r organisms

were captured i n t h e t r a p s a t t h e p r i n c i p a l sampling s t a t i o n , 2,655

moving upstream and 3,379 moving downstream, f o r a d a i l y average of

5.8 organisms moving upstream and 7,4 moving downstream. The l i v e

mass ( l i v e weight) of t h e organisms moving upstream was 187,927 g and

moving downstream was 93,092 g f o r a d a i l y average of 421 g and 209 g ,

r e spec t ive ly . F i sh alone accounted f o r 170,229 g moving upstream

llla

Figure 37. Seasonal v a r i a t i o n of photosynthesis r e s p i r a t i o n

r a t i o a t Concrete Bridge S ta t ion .

and 47,964 g moving downstream.

Thus f o r t h i s s tudy 1.27 t imes more organisms were captured

moving downstream than upstream, and 2.02 times more mass of

organisms was captured moving upstream. Although more animals

moved down than up, t h e l a r g e r s i z e of t hose moving up con t r ibu ted

t o a n e t movement of mass upstream. For f i s h alone, 3.58 times

more mass was sampled moving upstream. A number of very l a r g e

snapping t u r t l e s moving downstream con t r ibu ted heav i ly t o t h e

d i f f e r e n c e between t h e t o t a l mass of f i s h e s moving downstream

and t h e t o t a l mass of a l l organisms.

More organisms moved up than down on 183 days; more moved

down than up on 172 days; and t h e movement was equal o r zero on

94 days. A g r e a t e r mass of organisms was captured moving upstream

on 239 days; a g r e a t e r mass was captured moving downstream on

145 days; and t h e movement was equal o r zero on 65 days.

Seasonal Var i a t ions i n Movements

Table 18a and Figure 38 summarize by month t h e movements of a l l

organisms captured a t t h e p r i n c i p a l sampling s t a t i o n . The maximum

number and weight of organisms sampled was i n t h e sp r ing months of

t h e t h r e e yea r s sampled. The g r e a t e s t mass moving was, c o n s i s t e n t l y ,

i n March and Apr i l , and t h e g r e a t e s t number of animals moving was i n

Apr i l , May, and June. Movement was much l e s s during low water i n t h e

l a t e summer and during t h e win ter months. The p a t t e r n of movements

was q u i t e s i m i l a r from one year t o t h e next , although movements i n

1969 were g r e a t e r than movements during 1968 o r 1970.

Table 18a. Average Daily Fish Movements by Month

Number Average Average Average Av e r ag e of number number mass mass

Date 1968 - 1970

days moving moving moving sampled UP down UP g

moving down g

April 2 1 2 4 347 3

June 2 5 14 5 3 27 130

J u l y 17 11 2 11 1 5 8

August 15 5 1 7 4 3 9

September

October 1 5 8 4 184 6 4

November 2 0 3 3 170 42

December 17 0 1 2 0 13

January 1 9 0 1 6 1 11

February 18 1 3 301 2 9

March 16 4 9 1204 275

Apri l 30 8 3 2 2280 927

Table 18a. Continued

Number Average Average Average Average of number numb e r mass mass

Date 1968 - 1970

days moving moving moving sampled UP down g

moving down g

May 3 1 8 13 358 310

June 2 4 9 8 319 222

Ju ly 15 3 2 177 122

August

September

October

November

December

January

February

March

Apr i 1

May

June

Figure 38. Average d a i l y migra t ion by month. Mass i s i n grams

moving p e r day. Numbers a r e represen ted by l i n e s and mass by ba r s .

Cumulative Occurrence of Species vs . Cumulative Occurrence of Ind iv idua l s

New spec ie s were con t inua l ly added t o t h e t o t a l a s sampling pro-

gressed , inc luding a warmouth which was captured on t h e l a s t day

t h a t samples were run and which had not been encountered previous ly .

For f i s h t h e p l o t of cumulative number of spec i e s versus t h e log

of t h e cumulative number of i nd iv idua l s (Figure 39) was remarkably

s t r a i g h t and c o n s i s t e n t with t h e o r e t i c a l spec i e s organiza t ion

suggested by Odum, Cantlon, and Kornicker (1960). The l i n e may

come up s l i g h t l y a s found i n some s t u d i e s (Preston, 1963).

D ive r s i t y of Moving Animals

A t o t a l o f 44 spec i e s were encountered i n t h e 6,034 animals

t rapped a t t h e p r i n c i p a l s t a t i o n during t h i s study. Of t h e s e 4,416

were f i s h of 27 spec i e s . The d i v e r s i t y of t h e s e organisms a s measured

by D 1 = ( s - l ) - l logeN (Margalef, 1968), where S i s the number of

spec i e s and N i s t h e number of i nd iv idua l s , was 43/8.6052, o r 5.0,

f o r a l l animals and 26/8.3929 o r 3.1 f o r f i s h alone.

Movements a t Other S t a t i o n s on New Hope Creek

Table 18 summarizes t h e f i s h sampling d a t a f o r t h e Wood Bridge

and Jung le S t a t i o n s . The movement p a t t e r n s a r e , i n gene ra l , s i m i l a r

t o t h o s e observed a t t h e p r i n c i p a l sampling s t a t i o n . However, t h e lack

of downstream sampling a t t h e s e s t a t i o n s be fo re October,1968 makes

a n a l y s i s of some of t h e s e d a t a l e s s u s e f u l . When these statioAi5 a v L L i .

sampled s imultaneously with t h e p r i n c i p a l s t a t i o n , a movement of t o t a l

mass a t l e a s t as g r e a t a s a t t h e Concrete Bridge S t a t i o n is apparent .

Figure 39. Cumulative spec i e s versus cumulative ind iv idua l s

t rapped a t p r i n c i p a l sampling s t a t i o n ; only f i s h e s are included.

The movements a t t h e Jungle S t a t i o n may have been inf luenced by

t h e p o s i t i o n of t h e t r a p i n t h e middle of a very l a r g e pool r a t h e r

than i n an i n t e r p o o l r i f f l e , a s a t t h e Concrete Bridge and Wood

Bridge s t a t i o n s .

Movements a t Morgan Creek

Data f o r Morgan Creek a r e given i n Table 19. Local s p o r t s

fishermen i n t e f e r e d with sampling a t Morgan Creek during t h e

sp r ing and almost no sampling was accomplished during per iods of

expected l a r g e migrat ion. Nevertheless , Morgan Creek a l s o shows

a g r e a t e r movement of animals upstream than down, although t h e

r e s u l t s a r e l e s s pronounced than those f o r New Hope Creek. A s i n

New Kope Creek, t h e movements were g r e a t e s t i n t h e s p r i n g .

Analysis by Each Species

Twenty-seven spec i e s of f i s h and 16 spec i e s of o the r organisms

g r e a t e r t han 1 g were captured dur ing t h i s s tudy . No spec i e s was

captured a t o t h e r l oca t ions t h a t was not a l s o captured a t t h e p r i n c i -

p a l sampling s t a t i o n , with t h e except ion of one small spec i e s of

s h i n e r caught a t Morgan Creek , thay may o r may not have been Notropis

a l t i p i n n i s . Most o f t h e f i s h captured i n New Hope Creek were a l s o

captured i n Morgan Creek. The only except ions were bowfin, chain

p i c k e r e l , green s u n f i s h , largemouth bas s , speckled k i l l i f i s h ,

t h r e a d f i n shad and piedmont d a r t e r , A l l bu t t h e p i c k e r e l were en-

countered i n New Hope Creek only a s a r a r i t y .

Numerical and Weight Cont r ibut ion of Each Species t o Migration - The maximum, minimum, and average weight, a s wel l a s t h e

t o t a l number and mass, o f t h e more important spec i e s encountered

M d M N M In O O l n N CO N r l U 3 V )

*. 6, .I

rl N N

at the principal sampling station are given in Table 20. V-lip

and smallfin redhorses together were, by far, the most important

in terms of mass. Turtles, redbreast sunfish, flat bullheads,

and chain pickerel also contributed heavily to the total mass.

Frogs (including tadpoles), whitemouth shiners, white shiners,

crayfish, bluehead chub, and redbreast sunfish were most frequently

encountered. The average size moving upstream and downstream of

each species is given in Table 21. A more detailed analysis of

each species by size interval and by upstream or downstream move-

ment at the principal sampling station for the entire sampling

period is presented in Figure 40.

It is apparent from this information that for almost all fish

species, there is a tendency for larger individuals to move upstream

and for smaller individuals to move downstream. This is particularly

evident for black crappies, bluegill sunfish, flat bullheads, creek

chubsuckers, pumpkinseed sunfish, redhorses, white shiners, and

redbreast sunfish. Creekchubs and darters show no particular pat-

tern, highfin shiners and whitemouth shiners had greater upstream

movement for all sizes, madtoms and pirate perches of all sizes

moved downstream more than up. All larger species showed the large

fish upstream--small fish downstream pattern.

Smaller clayfishes were captured moving downstream more f~..

quently than up. Larger crayfishes moved in both directions about

equally. Turtles of all sizes were caught moving downstream more

often than up.

Table 20. Minimum, Maximum and T o t a l Mass and T o t a l Numbers o f

Each S p e c i e s o r Group Sampled a t P r i n c i p a l S t a t i o n ,

New Hope Creek

Minimum Maximum T o t a l T o t a l Average S p e c i e s weight weight number weight weight

Black c r a p p i e 1 .0 210 64 4457.5 69.6

Bluegi 11 0.5 194 266 1869.5 7.0

Bluehead chub 1 .0 80 404 4204.6 10.4

Bul lhead 1 .0 689 2 03 14416.2 71.0

Chubsucker 1 .0 345 202 4544.0 22.4

Creekchub 1 .0 8 4 7 2 798.5 11.0

D a r t e r 1 .0 4 109 161.0 1.4

Highf in s h i n e r 0.5 7 139 270.5 1 .9

Margined madtom 1 .0 3 2 120 1107.5 9.2

P i c k e r e l 1 .0 738 217 13394.5 61.7

P i r a t e p e r c h 1.0 12 8 1 309.0 3 .8

Pumpkinseed 1.0 287 120 1982.0 16.5 s u n f i s h

Redhorses 1.0 1363 328 150799.7 459.7

Redbreas t s u n f i s h 0.5 167 394 15299.5 38.8

White s h i n e r 1.0 3 5 543 361 2.9 6.6

Whitemouth s h i n e r 1.0 6 868 960.5 1.1

C r a y f i s h e s 1.0 44 446 5242.0 11.7

Table 20. Continued

Minimum Maximum Tota l Total Average Species weight weight number weight weight

Frogs 1 .0 500 881 5359.7 "6.0

Turt l e s 6.0 4000 62 35679.7 575.4

Table 21. Average Mass of Animals Moving a t P r i n c i p a l S t a t i o n

Average Average Species mass up mass down

Black c rapp ie

B lueg i l l

B%uehead chub

Bullhead

Chubsucker

Creekchub

Darter

Highfin s h i n e r

Madt om

P icke re l

P i r a t e perch

Pumpkinseed sun f i sh

Redhorses

Redbreast s u n f i s h

White s h i n e r

Whitemouth s h i n e r

Crayf i sh

Frogs

Turt 1 es

Others

&/UMBER DOWN MUT':BEB UP

fdUP?B ER DOWN

I N U M B E R DOWN NUMBER lJP

NUMBER BONK NUMBER UP

%-a -,,--.-,.-. L o I 187 _ _ __ __l_-..l .__-_-_--I--.. -

NUMBER DOWN Ll

NUMBER L'F

l o o

NUMBER DOWN NUMBER lj?

Seasonal P a t t e r n s of Movements f o r Each Taxonomic Class

Each taxonomic group was analyzed f o r seasonal t r ends i n move-

ments (Figure 41 asd Appendix D ) . From t h e s e it is obvious t h a t t h e

overwhelming bulk of t h e movement f o r a l l taxonomic groups, with

t h e p o s s i b l e except ion of c r a y f i s h , occurs i n t h e spr ing . There is

i n some f i s h e s continued, although smal le r , movements throughout

t h e summer; and f o r chain p i cke re l a secondary s e r i e s of movements

f o r t h e f a l l . The p a t t e r n f o r most f i s h e s i s repea ted from year

t o year . The cen t r a rch ids a r e almost never encountered during t h e

co lder months. Some important movements f o r each group a r e noted

below:

Black crappie: Crappies had one of t h e l a t e s t movements of any

spec i e s , gene ra l ly not moving u n t i l l a t e May o r June; however, a few

small i nd iv idua l s were caught moving downstream i n t h e sp r ing of

1969 and 1970. The movements i n 1968 were l a r g e r than i n e i t h e r

of t he o t h e r two years .

B lueg i l l : B lueg i l l s moved p r i n c i p a l l y i n Apr i l and May. Very

l a r g e numbers of small f i s h were caught moving downstream i n 1969

and 1970.

Bluehead chub: In t h e sp r ing of 1968 and 1970 these f i s h were

one of t h e most c o n s i s t e n t upstream movers; bu t i n t h e sp r ing of 1969,

t h e movements were much sma l l e r and were not a s d i s t i n c t l y

upstream.

Creek chubsuckers: These f i s h were not d i s t i ngu i shed from red-

horses u n t i l March of 1969. Heavy movements of t h e s e f i s h upstream

occurred i n March and Apr i l of 1969 and 1970, and a l a r g e movement

Figure 41. Average d a i l y movement f o r each number, by s p e c i e s .

F u l l names and s c i e n t i f i c name f o r each f i s h a r e given i n Table 6 .

Lines a r e numbers of f i s h and ba r s a r e mass, i n g , months a r e A p r i l ,

1968, through June, 1970. Each spec i e s i s on a s epa ra t e page.

Black c rapp ie Figure 41 a

B lueg i l l Figure 41 b

Bluehead chub Figure 41 c

Creek chub Figure 41 d

Creek chubsucker Figure 41 e

Dar te rs Figure 41 f

F l a t bul lhead Figure 41 g

Highfin s h i n e r Figure 41 h

Mad t om Figure 41 i

P icke re l Figure 41 j

P i r a t e perch Figure 41 k

Pumpkinseed Figure 41 1

Redbreast s u n f i s h Figure 41 m

Redhorses Figure 41 n

White s h i n e r Figure 41 o

Whitemouth s h i n e r Figure 41 p

Crayf i sh Figure 41 q

Frogs Figure 41 r

T u r t l e s Figure 41 s

Others Figure 41 t

'm o r - .

P I R A T E PERCH

T- r y -

WHITE

(V z. t-i

is, w -,

. iITEMr!iJTH SHINER

'II) or- 7

t OTMERS

downstream of small f i s h e s occurred i n 1970.

Creek chub: Creek chubs were most f r e q u e n t l y encountered i n

Apr i l and May o f 1968, and were r a r e l y sampled l a t e r . Nearly a l l

movements were i n Apr i l and May, with a s l i g h t upstream b i a s .

Dar te rs : Dar te rs moved upstream dur ing t h e sp r ing and r a r e l y

a t any o t h e r t ime.

F l a t bu l lhead: These f i s h were caught a t nea r ly a l l t imes

of t he year . Peaks i n movements occurred i n t h e warmer months.

Highfin sh ine r : These l i t t l e f i s h were t h e most numerous

f i s h i n t h i s s tudy. Movements i n both d i r e c t i o n s were g r e a t e s t

i n Apr i l , May, and June.

Madtom: Madtoms moved g r e a t e s t i n May and June. In 1968 and

1970, movements were more up than down.

Redhorses: The two spec i e s of redhorses completely dominated

t h e mass of f i s h e s i n New Hope Creek migra t ions . Movements were

l a r g e a t almost a l l seasons of t h e year , wi th some diminuation

during t h e summer and very l a r g e peaks i n March and Apr i l . A

small f l ood i n December of 1969 caused heavy movements even i n

t he winter . Small redhorses were no t caught very o f t en .

Redbreast sun f i sh : The,se c o l o r f u l f i s h e s were caught during

a l l warmer months of t h e year . Heaviest movements occurred i n

March, Apr i l , and May, Movements from year t o year were s i m i l a r

i n magnitude . Pumpkinseed sun f i sh : These f i s h were r a r e l y caught except i n

Apr i l , May, and June, although some smal le r f i s h e s were captured

moving downstream i n March.

P i r a t e perch: P i r a t e perch s t a r t e d t h e i r annual movements

be fo re most o t h e r f i s h e s , a s e a r l y a s January i n 1969. They

were r a r e l y caught a t t imes o t h e r than t h e sp r ing , a l though

f loods i n t h e f a l l of 1969 may have s t imula ted t h e secondary

movements noted then.

Chain p i c k e r e l : P ickere l moved throughout t he year with

peaks i n both t h e f a l l and spr ing . Smaller p i c k e r e l were f r equen t ly

captured i n t h e summer. Thei r preda tory h a b i t s may in f luence t h e

year-round movements noted.

White s h i n e r s : Movements were g r e a t e s t i n Apr i l and May, and

1968 and 1970 were more important than 1969. Smaller movements

continued throughout t h e year . The l a r g e s t recorded movements

were a s soc i a t ed with the f loods which occurred i n October, 1968,

a f t e r a long drought.

Crayf i sh : Crayf i sh were a c t i v e throughout t h i s s tudy with a

peak movement dur ing t h e spr ing of 1969. Movements were gene ra l ly

more upstream than down except during t h a t time.

Frogs: Frogs were caught from t ime t o t ime, most f r equen t ly

i n t h e sp r ing a s tadpoles moving (or being swept downstream) down-

stream. Some l a r g e b u l l f r o g s were a l s o captured moving i n both

d i r e c t i o n s .

T u r t l e s : T u r t l e s were caught i n t h e sp r ing and summer, and

not a t a l l dur ing t h e r e s t of t h e year . Some very l a r g e snapp ing

t u r t l e s con t r ibu ted t o a l a r g e t r a n s f e r of mass downstream dur ing

t h e spr ing .

155

Others: Miscellaneous organisms a l s o moved most heav i ly

during t h e spr ing . Sometimes t h e cap tu re of a l a r g e muskrat o r

a number of snakes con t r ibu ted t o heavy movement during o t h e r t imes

of t h e year . In genera l t h e r e was g r e a t e r movement downstream than

up f o r t h e s e a s so r t ed c rea tu re s . Large-mouth bass , of s p o r t

f i s h i n g i n t e r e s t , exh ib i t ed movements similar t o o the r cen t r ach ids

but on a sma l l e r s c a l e ; a few l a r g e bas s moved upstream i n t h e

sp r ing and small ones moved downstream a t v a r i o u s t imes of t h e year ,

o f t e n i n t h e f a l l .

Evidence of Spawning Condit ion of F ish a t D i f f e ren t Times o f t h e Year

Records were kept of s i g n s of reproduct ive a c t i v i t y f o r t h e

f i s h sampled. These s i g n s inc lude : breeding t u b e r c l e s , s easona l ly

b r i g h t co lo r s , and t h e a c t u a l d i scharge of eggs o r m i l t , Table

22 g ives t h e s e r e s u l t s f o r a l l spec i e s where t h e information is

a v a i l a b l e . Signs of breeding cond i t i on were only noted i n t h e

spr ing , and were i n v a r i a b l y a s soc i a t ed wi th heavy movements of t h a t

spec ies . F ish i n obviously r i p e condi t ion were taken almost i n v a r i -

ab ly moving upstream, and spent f i s h were always taken moving down-

stream.

Recaptures of Marked Fish

Of t h e 6 ,043 f i s h and o t h e r organisms captured a t t h e p r i n c i p a l

sampling s t a t i o n , 417, o r 6.9 pe rcen t , were marked from previous

Table 22. Continued

Species (number, i f more than 1) Date D i rec t ion moving Condition noted

Creek chubsucker (Cont)

(many 1

Creekchub

1969

Mar. 30

Mar. 31

Apr. 6

Apr. 9

1970

Apr. 11

Apr. 11

Apr. 12

Apr. 19

Apr. 20

Apr. 25

Tubercles

Tubercles

Tubercles

Tubercles

Discharged eggs

Tubercles , rosy-colored, discharged m i l t

Tubercles

Tubercles and rosy- co lored

Concave be1 l y

Tubercles

Table 2 3 . Recapture of Marked Fish

Number of Number of Tota l To ta l Percent Percent marked f i s h marked f i s h number number recaptured recaptured recaptured recaptured moving moving moving moving

Species moving up moving down up down UP down

Black c rapp ie

B lueg i l l

Bluehead chub

Bullhead

Chubsucker

Creekchub

Dar t e r

Highf i n s h i n e r

Madt om

Pickere l

P i r a t e perch

Pumpkinseed sun f i sh

Table 23. Continued

I Number o f Number of Total Total Percent Percent marked f i s h marked f i s h number number recaptured recaptured recaptured recaptured moving moving moving moving moving up moving down up down UP down

I Redhorses 16 13 248 82 6.5 15.9

Redbreast sunf ish 15 2 1 212 181

White sh ine r 2 1 54 329 213

Whitemouth shiner 13 26 578 286

Crayfish 11 9 190 256

Frogs 0 0 18 863

Others 3 1 5 2 532 5.8 .2

This very low recap tu re r a t e i n d i c a t e s t h a t , i n genera l , 'home

range1 movements (Gerking, 1959) were no t being in t e rcep ted - -o r

poss ib ly t h a t t h e f i s h became very t rap-shy a f t e r one encounter.

Tagged Fish Returns Analysis

During t h i s s tudy l a r g e r f i s h were marked with numbered

p l a s t i c d a r t t a g s (Floy Tag and Manufacturing Company), and 75

of t h e s e were recaptured (Table 2 4 ) . Recapture p a t t e r n s were

va r i ed , bu t many ind iv idua l f i s h were captured moving upstream and

recaptured moving downstream s h o r t l y t h e r e a f t e r . Fish were some-

times recaptured a t t h e same l o c a t i o n and moving i n t he same

d i r e c t i o n without having been captured moving i n the o t h e r d i r e c t i o n .

These gene ra l ly occurred only during i n t e r v a l s i n which t h e weir

was disassembled i n t h e in te r im. F ishes marked a t t h e p r i n c i p a l

s t a t i o n were r a r e l y recaptured a t another , although t h e o the r

s t a t i o n s were sampled much l e s s f r equen t ly .

Daily Concentrat ion of Moving Animals

Each spec i e s was analyzed f o r number of i nd iv idua l s moving up-

stream o r downstream during a given day. This may be some ind ica t ion

of t h e tendency f o r t h e f i s h . t o school , and may have t h e o r e t i c a l i m -

p l i c a t i o n s a s t o t h e b e s t way f o r a given mass of f i s h t o be moved

from one p l ace t o another .

A l l t a x a t r a v e l e d more f r e q u e n t l y a s i nd iv idua l s than i n any

o t h e r numerical a s s o c i a t i o n , and i n groups of two more than i n any

l a r g e r groups (Table 25). Thus , a l l t a x a gene ra l ly appear not t o

t r a v e l i n schools , During per iods of heavy migra t ion , however, some

spec i e s were captured i n numbers of 10 o r more p e r day. This was t r u e

of bluehead chub, r edb reas t sun f i sh , b l u e g i l l sun f i sh , redhorses ,

a ' a , a, M C, M cd cd n C,

m o m 4 4 N

m m o o \ D * * b M 4 4 N

N N N

k .

% 9 $ & 2 2 2 2 2 2 k k k k & E X . 3 2 E Z

Table 25. Concentrat ion (Daily) of Moving Organisms

Times Organisms moving i n groups of

0 1 2 3 4 5 6 7 - 9 10 - 19 20 o r more

Black c r app ie

868

B l u e g i l l

841

Bluehead chub

714

Bullhead

755

Chubsucker

810

Creekchub

848

Dar te r

825

a, k

8 k 0

0 N

0, d

I

0 l-4

cn 1

I-.

\O

In

d

M

c.l

d

0

v, a, v, k 0 C 5 0, a, v) d I-.

DISCUSSION

This s tudy of migra t ion and metabolism of t h e New Hope Creek

stream system allows t h e two t o be r e l a t e d s o we may i n f e r some of

t h e r o l e s t h a t migra t ion may p lay i n s t ream metabolism and t h e ways

i n which t h e migrat ions may t a k e advantage of programs of l i f e

support . These comparisons may be made by examining t h e seasonal

t iming of events , th'e s p a t i a l d i s t r i b u t i o n s , t h e n u t r i e n t s processed,

and t h e energy involvements of each p a r t .

Seasonal P a t t e r n s of bletabolism

New Hope Creek has a sharp peak i n primary product ion i n t h e e a r l y

spr ing (Figures 34-36). This peak i s a s soc i a t ed with high l e v e l s of

r e s p i r a t i o n t h a t cont inue throughout t h e summer and e a r l y f a l l . There-

f o r e , a s t h e season progresses , t h e stream becomes inc reas ing ly dependent

upon o u t s i d e sources of energy and/or energy s to rages . This g r e a t e r

usage than product ion of energy c o n s t i t u t e s a condi t ion of he te ro t rophy.

P/R Rat io and Hetero t rophic Reg'ime

The g r e a t e r r e s p i r a t i o n than photosynthes is observed i n New Hope

Creek (Figures 34-36) i s o f t e n c h a r a c t e r i s t i c of woodland streams t h a t

a r e dependent on al lochthonous d e t r i t u s f o r some o r a g r e a t d e a l o; t h e i .

energy supply (Smith, 1966). Hoskin (1959) found s i m i l a r p a t t e r n s i n

o the r s t reams of North Caro l ina . The p r i n c i p a l supply of t h i s d e t r i t u s

t o New Hope Creek is probably l e a f f a l l and organic runoff from

t h e surrounding f o r e s t . A s shown by some experiments I made a t

t h e Pennsylvania S t a t e Univers i ty , mayfl ies and s t o n e f l i e s , both

abundant i n New Hope Creek, qu ick ly reduced dead leaves of many

spec i e s t o ske l e tons of vascu la r t i s s u e . Such ske l e ton ized

leaves were o f t e n observed i n New Hope Creek during t h e l a t e

spr ing and summer months. Other sources of e x t e r n a l l y suppl ied

energy may be f o r e s t i n s e c t s dropping i n t o t h e stream, organic

substances i n runoff , and seve ra l minor sources of domestic

p o l l u t i o n t h a t e x i s t near t h e headwaters (Research Tr i ang le

Regional Planning Commission, 1968). These may be l e s s i n

summer when d ischarge i s small

S p a t i a l D i s t r i b u t i o n o f Metabolism

A s shown i n Tables 15-17 and Figures 34-36 t h e p roduc t iv i ty and

r e s p i r a t i o n p e r u n i t a r ea a r e f a i r l y uniform i n t h e zones of New A

Hope Creek s t u d i e s although t h e r e was a t l e a s t a t h r e e - f ~ l d range of

depth and volume metabolism. Table 11 sugges ts t h a t t h e percentage +

of sun energy reaching t h e creek s u r f a c e i s s i m i l a r i n t h e two

s t r e t c h e s s tud ied (Blackwood S t a t i o n and Concrete Bridge S t a t i o n ) .

Comparison of phosphorus b y . s t a t i o n i s made i n Table 26, but no d i f f e r -

ences were found t h a t c o n s i s t e n t l y c o r r e l a t e d with d i f f e r e n c e s i n

volume metabolism.

D i lu t ion of Resources with Depth a

Many previous au thors have found an inve r se e f f e c t of depth

and t h e p r o d u c t i v i t y of waters . Rawson (1952; 1960) with d a t a from

l a r g e borea l l akes i n Canada ind ica t ed t h a t t h e a r e a l product ion

of n e t plankton, benthos, and f i s h was inve r se ly propor t iona l t o

t h e depth of t h e l akes . Shallower lakes were more product ive i n

t h e s e h igher t r o p h i c l e v e l s . Steeman Nielsen (1957) found an in -

v e r s e r e l a t i o n s h i p of p roduc t iv i ty and depth of t h e euphotic zone

i n t h e sea . Bailey (1967) found an inve r se r e l a t i o n between depth

and primary product ion i n t h e Sacramento-San Joaquin es tuary .

Demersal oceanic f i s h e r i e s tend t o be concentrated on r e l a t i v e l y

shallow banks and nea r shore a r eas (Bigelow and Welsh, 1924;

Alverson, 1964).

In New Hope Creek with s i m i l a r metabolism pe r u n i t a r e a , i n -

c rease i n water depth between s t a t i o n s diminished t h e concent ra t ion

of metabolism pe r u n i t volume. I t may be reasoned t h a t food resources

f o r f i s h were a l s o d i l u t e d . I f so t h e shallow zones may have more

concent ra ted food f o r young f i s h .

In New Hope Creek t h e r e were two mani fes ta t ions of t h i s change

i n water depth. These a r e changes from deeper, downstream a reas t o

more shallow upstream regions , and changes a t any one p l ace a s

t h e water drops during t h e summer. Much o f t h e energy t h a t e n t e r s

a system remains t h e same no ma t t e r what t h e depth, f o r both l i g h t

and leaves e n t e r a s t ream on a square meter b a s i s . In add i t i on , t h e

amount of energy a v a i l a b l e t o ben th i c p l a n t s would be l e s s i n

deeper reg ions because of e x t i n c t i o n with depth. Thus t h e t o t a l energy

t o support organisms does not change much with depth. I t does be-

come more concent ra ted , however, and perhaps more a v a i l a b l e t o

food chains. This e f f e c t during summer low waters may be p a r t i a l l y

o f f s e t by a l e s sen ing of t h e t o t a l l i g h t energy input t o the stream

as t h e t o t a l water a r e a becomes smal le r .

Both photosynthesis and community respiration in New Hope

Creek varied seasonally with a spring maxima and a secondary peak in

the fall (Figures 38-40). The possible causes may be seasonal

variations in minerals, solar energy, temperature and water level

changes.

Daily records of insolation under a deciduous forest canopy

kept at the International Biological Program Site (Figure 42) showed

seasonal patterns of insolation with a peak in early spring that

corresponds with the peak of observed values of gross primary pro-

duction (Figures 34-36). The peak of photosynthesis was in March

rather than in June, due to the shading effect of overhead trees,

which leaf during the middle of April. A second, smaller peak in

primary production in the fall also corresponded with an increase

in light following leaf fall.

Neither dissolved phosphorus (Table 13) nor any of the important

forms of nitrogen (Table 14) showed any consistent seasonal variations

that were correlated with seasonal variations in metabolism.

The seasonal variation of community respiration at the Concrete

Bridge Station (Figure 34) showed two peaks, one during the high

solar energy input in the spring and one during fall low waters.

Therefore, apparently neither the primary production nor respira-

tion was controlled by temperature which had maximum values in

late summer.

Comparison With Some Other Studies

The areal metabolism was generally lower than values obtained

in other studies (Table 27). The metabolism was within ranges of

Figure 42 . Seasonal p a t t e r n s of i n s o l a t i o n under a hardwood

canopy, Duke F o r e s t , near New Hope Creek. Pyroheliometer d a t a i s

from t h e I n t e r n a t i o n a l Bio logica l Program s i t e loca ted a few hun-

dred meters t o t h e n o r t h of t he upstream watershed of New Hope Creek.

Table 27. Metabolism i n Some Other Unpolluted Streams

Gross To ta l community Production r e s p i r a t i o n

Location Time g m-2 day- l g m-2 day''

Birs, Switzer land a 1946, Apr i l 11-12

Kljasma, Russia a 1929, J u l y 21

I tchen , England b Apr i l - October

IOrdinary1 s tream i n North Caro l ina (var ious t imes of year ) 1956 - 1957

I v e l , England c ' t y p i c a l 1 s i n g l e curve a t two l o c a t i ons Elay 1964

Spring Creek, Pennsylvania d E n t i r e year 1-17 1.5-13

This s tudy Concrete Bridge S t a t ion 0.21-8.85 0.39-13.40

a. Quoted i n Odum, 1956

b. Hoskins, 1958

c. Owens, 1969

d. Cole, 1969

Pa.tterns of F ish Movements

The annual peak i n f i s h movemelts was close1 y cnrrcl a t e d w; t l ~

t h e annual peaks i n g m s s phntosvnthcs is , community s c s p j r a t i o n a ~ d

t h e end of win ter f loods (Figure 34-36, and 26) . These may be a

s e l e c t i v e p a t t e r n i n t h e f i s h e s t o schedule t h e i r own time of high

energy usage with t h e time of maximum t o t a l energy a v a i l a b i l i t y

i n t h e environment. Among t h e l a r g e f i s h e s caught i n t h i s s tudy ,

t h e r e was no c l e a r - c u t p a t t e r n f o r maximum f i s h growth apparent a t

any one time of t h e year (Figure 4 3 ) . Storage and l a g processes i n

t h e s t ream may smooth out t h e pu l se i n food a v a i l a b i l i t y a t second

and t h i r d t r o p h i c l e v e l s over t he season when t h e young f i s h e s

a r e ready t o t a p t h e food cha ins . The cont inuing high l e v e l s of

r e s p i r a t i o n a f t e r t h e pu l se i n primary product ion i n d i c a t e sus-

t a i n e d b i o l o g i c a l a c t i v i t y i n summer and f a l l .

Movements of D i f f e ren t Species

Some smal le r spec i e s , such a s d a r t e r s and p i r a t e perch, with

a high s u r f a c e t o volume r a t i o and r e s u l t a n t high f r i c t i o n , do not

show a l a r g e upstream movement. The energy l o s t i n rnigrati.cn may he

g r e a t e r than t h a t gained.

Movements and Floods. - The cu r ren t a s an a u x i l i a r y energy source t o moving animals both

a i d s i n t h e planned movements downstream and inc reases energy demands

on animals holding t h e i r p o s i t i o n . Observations dur ing high water

sugges ts a c t u a l washing of f i s h downstream i s not important .

Table 28. Metabolism o f Some S e l e c t e d Lakes and Maine Waters

--a-

Gross Coinmn i t y p roduc t i o n r e s p i r a t i o n

Loca t ion Time g m-2 day- l - 2 - 1 - J3 _m_md2~ -* _ . - _ - --.

Eniwetok A t o l l a

Texas Bays b

S t u a r t Farm Pond c Durham, N. C .

Lake P*lichigan d 3.2 km from s h o r e 6.4 km from s h o r e

Sacramento-San Joaqu in d e l t a e

Midsummer 1954

Various t i m e s 1957

J u l y 13-14 1968

1 1

i3rackish Ponds blorehead C i t y , I.?.C, f ' Average f o r y e a r

a. Cdum and Odum, 1355

b . Oduix and Wilson, 1958

I

c. Odum and Wilsor., 1958 I

I

d. Planny and Hal 1, 1969 g . Sum of n e t p r o d u c t i n a and n i g h t t ime r e s p i ~ z - t i ? n

e . Ba i ley , 1970

f . Odum e t a l . , 1970

Figure 4 3 . Growth of tagged f i s h , New Hope Creek. The l e f t

of each p a i r of p o i n t s r ep re sen t weight a t f i r s t cap ture , and t h e r i g h t

po in t r e p r e s e n t s weight a t second (or t h i r d ) capture . Decrease i n

weight f o r some f i s h probably r ep re sen t s spawning lo s ses .

bc = black crappie ; ecp = chain p i c k e r e l ; f b = f l a t bu l lhead;

r b = r edb reas t sun f i sh ; r h = redhorses .

The g r e a t e s t d i scharge occurred during t h e months of February and

March, b u t few animals were sampled i n t h e downstream t r a p dur ing

t h i s per iod . For example, on February 2, 1970, t h e r e was a medium-

s i z e d f lood t h a t r a i s e d t h e water from 50 t o 65 cm above zero flow.

During t h e s p r i n g o r summer months t h i s would have been accompanied

by an increased movement both upstream and downstream. However,

on t h i s d a t e , when t h e water temperature was only 7" C , t h e r e was

no recorded movement e i t h e r upstream o r downstream. This p a t t e r n

was repea ted on many occasions dur ing cold weather. Apparently

t h e f i s h move downstream only by i n t e r n a l program.

Movements of J u v e n i l e Fishes

Many more very small f i s h e s may move downstream than were

measured i n t h i s s tudy, s i n c e t h e mesh s i z e on t h e weir and t r a p s

was l a r g e enough t o l e t any f i s h sma l l e r than 1 g pass through.

Plankton n e t s hung i n t h e cu r r en t on s i x s epa ra t e days during pe r iods

of heavy migra t ion of l a r g e r f i s h caught only one small d a r t e r . More

ex tens ive sampling could poss ib ly g ive very d i f f e r e n t r e s u l t s . On

May 15, 1970, leaves p l a s t e r e d on t h e s i d e of t h e downstream t r a p formed

a b a r r i e r i n which was observed a school of about 250 t i n y (1.3 cm)

f i s h . Complete keying was impossible but they appeared t o be some

spec i e s of Notropis. How o f t e n t h i s occurs when t h e movement is

not observed i s a mat te r f o r another s tudy.

D i f f e r e n t i a l Movements of Di f fe ren t -S ized Fish

The gene ra l ly upstream movement of l a r g e r f i s h e s and gene ra l ly

downstream movement of sma l l e r f i s h e s observed i n t h i s s tudy r a i s e

some i n t e r e s t i n g q u e s t i o n s about usage o f a v a i l a b l e energy by a

p o p u l a t i o n of animals . The upst ream movements a r e obv ious ly t i e d

i n w i t h r e p r o d u c t i o n , which i m p l i e s t h a t t h e r e may be r e a s o n s t o

b r i n g t h e p o t e n t i a l progeny upst ream. One r e a s o n would be t o

d i s t r i b u t e t h e g e n e t i c s t o c k o v e r t h e s t ream. The v e r y smal l f i s h e s

w i t h l a r g e surface- to-volume r a t i o s a f f e c t i n g f r i c t i o n cannot s w i m

upst ream a g a i n s t t h e c u r r e n t , b u t t h e l a r g e ones can and l o , and

t h e s m a l l ones can and do move downstream w i t h t h e c u r r e n t . Thus,

t h e upst ream m i g r a t i o n s of t h e a d u l t s may b e n e c e s s a r y a s a g e n t s

f o r s t o c k maintenance and gene d i s p e r s a l . S i n c e t h e s p r i n g p u l s e

i n energy a v a i l a b l e p e r volume a t t h e ups t ream s t a t i o n i s con-

s i d e r a b l y g r e a t e r t h a n t h e p u l s e a t t h e downstream s t a t i o n , it would

be more advantageous t o have t h e most r a p i d l y growing smal l s t a g e s

l o c a t e d upst ream.

T h i s r a i s e s t h e q u e s t i o n o f why t h e f i s h move back d o m s t r e a m .

The l a r g e number o f sub-one-year c l a s s f i s h moving downstream i n

s p r i n g i n d i c a t e s a d i s p e r s a l of many f i s h e s a f t e r spending one y e a r

u p s t r e a n . Th is may b e an ad.apta t ion t o p r e v e n t p o p u l a t i o n p r e s s u r e s

between t h e new y e a r c l a s s of j u v e n i l e s and o t h e r f i s h e s upst ream

which i n c r e a s e t h e i r a c t i v i t i e s a s t h e wa te r warms. Because o f

geomet r ic a d a p t a t i o n t o r o c k s , c u r r e n t s , and microenvironments l a r g e

f i s h may e x p e r i e n c e l e s s s t r e s s i n t h e deeper w a t e r s , N e l l i e r (1962)

found s m a l l f i s h e s moving t o deeper wa te r s a s t h e y grow. Another

p o s s i b i l i t y was c o n s i d e r e d by Margalef (1968), who commented on t h e

movement o f animals a s t h e y grow o l d e r from t h e h i g h l y p r o d u c t i v e

r e g i o n s o f j u v e n i l e growth t o more s t a b l e environments . Downstream

157 reg ions i n New Hope Creek may be, t o a f i s h a t l e a s t , more s t a b l e ,

s i n c e they a r e not s u b j e c t t o t h e extreme d iu rna l v a r i a t i o n s in

oxygen t h a t occur i n t h e upstream, more shal low reg ions d u r j n g t h e

low water s t a g e s of summer drought. In add i t i on , t h e deep pools

i n t h e downstream regions provide insurance aga ins t complete

a n n i h i l a t i o n during extreme droughts. There may be a t r a d e o f f

of high p r o d u c t i v i t y versus a more s t a b l e environment t h a t i s b e s t

u t i l i z e d by sending armies o f young t o t h e h ighly product ive reg ions

t o g e t a quick s t a r t i n l i f e , followed by d i s p e r s a l of t hose t h a t

su rv ive t o more s t a b l e regions. A s i n g l e small f i s h i s more ex-

pendable t han a l a r g e r one s i n c e t h e r e a r e many more of t h e former

and an ecosystem has inves ted l e s s of i t s energy resources i n it.

Comparisons of Energy Budgets

Consider next t h e energy budgets of t h e stream, i t s metabolism,

t h e f i s h e s and t h e i r migrat ions. To r e l a t e energy budgets of t h e

f i s h e s t o t h a t of o t h e r p a r t s of t h e system it i s convenient t o

express work processes i n t h e i r c a l o r i c form, s i n c e energy is a . common denominator f o r a l l p rocesses .

Energy of Running Water

The phys ica l p o t e n t i a l energy r e l eased i n turbulence ( E ) i n a

cubic meter of water flowing downhill between t h e Blackwood sampling

s t a t i o n and t h e Concrete Bridge S t a t i o n can be ca l cu la t ed a s equat ions

t h a t fo l low:

- 3 E i n kg-m m = (mass i n g) ( acce l e ra t ion due t o g r a v i t y i n n ~ e c - ~ )

(d i f f e r ence i n he ight i n m)

*

E i n Cal m-3 = ( lo3) (9.8) (45.7) (2.34 X l o m 3 Cal kg-m)-'

E i n Cal m m 3 = 1040 Cal m-3

For t h e y e a r a mean t ime of one day was r e q u i r e d f o r w a t e r

t o f low from Blackwood S t a t i o n t o t h e Concre te Bridge s t a t i o n , t h e r e -

f o r e t h e p h y s i c a l power d i s s i p a t i o n i s 1040 Cal m-3 day-1, o r about

350 Cal m m 2 day-'.

Energy o f B i o l o g i c a l Metabhlism

The mean b i o l o g i c a l metabolism i n t h e same c u b i c meter of

wa te r f lowing th rough t h e same zone o v e r t h e e n t i r e y e a r was found

t o be about 2.85 g oxygen p e r c u b i c mete r p e r day (Table 1 5 ) . S ince

about 3.5 C a l o r i e s a r e r e l e a s e d p e r gram o f oxygen metabo l ized

(Brody, 1945) , a t o t a l o f about 10.0 C a l o r i e s o f energy were used i n

r e s p i r a t i o n p e r day. Thus t h e system r e c e i v e s about 100 t imes more

energy from t h e work of c u r r e n t s a s from o r g a n i c f u e l s .

Energy o f I n s o l a t i o n

The energy of i n s o l a t i o n r e a d i n g through t h e canopy t o t h e s t ream

was e s t i m a t e d t o be 6 .7 3 l o 4 CaP m-3 d a y - l . Thus t h e s o l a r energy

budget i s abou t60 times thewater c u r r e n t c o n t r i b u t i o n .

Energy o f F i s h Metabolism .

A rough f i g u r e f o r t h e u s e o f oxygen by f i s h under normal

c o n d i t i o n s i s about 100 m l (0.143 g 02) h r - l kgq1 (Brown, 1957;

B r e t t , 1965) . Th i s i s q u i t e v a r i a b l e w i t h t empera tu re and a c t i v i t y

r a t e s b u t i s p robab ly c l o s e t o mean v a l u e s f o r New Hope Creek. There

i s about 18.3 g f i s h m-3 i n New Hope Creek (Carnes e t a l . , 1964) which --

would u s e (0.143 g 02) (365 d a y s ) ( 2 4 hours ) (18.3 a kg) o r 23

g O 2 m-3 year-1. For t h e e n t i r e watershed above t h e Concrete

Bridge, this would be, including additions and losses by migration,

about 4 g 02 m-2 year-1, or about 13.4 Cal m-2 year-l (Figure 44) .

Energy Used by Migrating Fishes -- .-- -- Consider three different ways to measure the energy used by

migrating animals such as fjslles: (1 ) one nay calculate the tatzl

work expended against f~lction;l.J. forces and/or that used in mnvjn~

the organisms against gravity. (2) One may convert the additima!

oxygen used during mig~?t;on to Caloric values. ConsJderah;e data

exists on oxygen use chiring diEferent levels o f a a c i ; v i t ~ ~ (Grown, 1957;

Brett, 1965, 19701, and estimates have bee? made for use during mi-

gration (Brett, 1970). These figures are about 100 ml O2 kg-I hr- 1

for fish at Isw levels of activity and about twice this during migra-

tions. (3) One may weigh fish and analyze them for different food

reserves at the beginning and end of their movements. This type of

work has been done for organisms that do not feed during their

spawning migration, such as salmon (Idler and Clemens, 1959).

The second method was used for these estimates. An additional

100 ml 02 kg hr-I was aBloted as the cost of migration. Multi-

plying this by the annual biomass moving upstream at the Concrete

Bridge (120 kg) by the period of major movements (three months or

2200hours) gives a total energy cost of 132,000 Cal, or 0.88 Cal

year-'. This is about 7 percent of the estimated fish m e t a b o ? ' ~ ~

and about 0.1 percent of the annual metabolism of the entire ecosystem.

If it is assumed that the upstream migration is necessary to maintain

the stocks of fish in the upstream position of the stream, this energy

used for migration has a multiplying effect of 14. All energy relations

are summarized in Figure 45.

Figure 44. Annual imvc!:ient arid r ~ c t ::holi s;; o f f i i l i p+~l ln t . i 011s

i n the hcadi rat r rs of Sew liopc Cree!< a!>ovc the Concrcip Cridge .

Numbers represcii: appr.oziii:~.te s:an,!i ng c r o ~ r , a r : n i i ~ i ni gr2T i c w , 7 - 1 r e s p i r a t i o n and food i n t a k e i n Cnl n- - y c o r . Food inti!:. i s

ca lcu la t ed to b a l ance o t h e r energy flop, s .

Figure 45. Energy f l o ~ diagram f o r upstream (nio'dle s e t of

modules) and downst reas ( l o i i emos t s e t o f ~ ~ ~ o d u l e s ) o f Scir iIo;~c

Creel:. Eietabolisrm i s i n Cal n-3 d a y - ? , 3s V O ~ U I O C d i f i e : - encc~ i n

metabolisin i s suggos t c d a s an j rnportant f a c t o r . Energy enters

t h e systcm a s sun energy, which p a s s e s t h r o i ~ g h food c h a i n s , 2nd

sireaiii fluw energy which a i d s i n the d i s t r i b u t i o n o f resources

and d i s p e r s a l of was t e s . Upstream ni igrat ion r e q u i r c s s d l i t i o n a l

energy t o overcome t h i s f low.

Net Contr ibut ion of Migration t o

Headwaters and Turnover Rate T

During t h i s s tudy an est imated 119 ,400gEsh and o t h e r animals

moved each year i n t o t h e reg ion above t h e Concrete Brid.ge, and 56,700

g moved ou t of t h i s a r ea . A n e t movement i n t o t h i s a r e a of 62,700 g

5 2 occurred. There i s approximately 1.6 10 m of s t ream above t h e

Concrete Bridge a s determined by f i e l d measurements and topographic

maps. Over t h i s one year per iod t h e r e was a n e t add i t i on o f 0.39

g animals m-2 o f water. This i s about 14 percent of t h e es t imated

f i s h s tanding crop of 2.78 g m q 2 (Carnes e t a1 . , 1964) . Summing -- t h e mass of animals leav ing t h i s a r e a and t h e mass of animals

en t e r ing t h e a r e a g ives 176,100 g , o r 1.1 g m-2 of animals (about

0 .8 g m-2 o f f i s h alone) involved i n migrat ing t o o r from t h e a r e a .

This is 40 percent of t h e es t imated s tanding crop of f i s h i n t h a t

reg ion (Figure 44 ) . The s tanding crop of a p a r t o f New Hope Creek would, according

t o t h e s e f i g u r e s , be rep laced i n 3.7 years by upstream migra t ion

a lone , 7.85 years by downstre& migra t ion a lone , o r i n 2.5 yea r s by

movement from above and below. This replacement r a t e has implica-

t i o n s f o r p r e d i c t i o n s i n r e l a t i o n t o p o l l u t i o n and f i s h k i l l s . These

r e l a t i o n s h i p s a r e summarized i n Figure 44 . Food i s c a l c u l a t e d t o *

balance r e s p i r a t i o n and migrat ion.

Comparison of Migration i n New Hope Creek With a Salmon Migration

Two lakes i n B r i t i s h Columbia, Owikeno and Long Lakes, were

chosen f o r a comparison with New Hope Creek. The t o t a l runs of

salmon a r e wel l known and have been v i r t u a l l y cons tan t wi th in

t h e per iod of record , i n d i c a t i n g t h a t p re sen t average run f i g u r e s

a r e about t h e same a s run f i g u r e s before heavy e x p l o i t a t i o n

The mean annual ca tch and escapement of a l l salmon (mostly

sockeye) has been about 500,000 f i s h . An e s t ima te of t h e average

weight f o r sockeye salmon i s about 2.5 kg f o r each f i s h ( I d l e r and

Clemens, 1959). Thus, about 1.25 - 109 g of f i s h en tered t h e lake

2 a r e a of 97.1 km o r would have en tered had t h e r e been no f i s h e r y .

This is about 12.5 g m-2, o r 21.4 t imes t h e con t r ibu t ion of migra-

t i o n t o t h e upstream reaches of New Hope Creek. However, t h i s mass

i s d i s t r i b u t e d throughout much deeper water , does not f eed , and

was produced from a much broader food base .

Poss ib l e Adaptive Values of iyiigrations i n New !-Iope Creek

Migration A s a Coupling Function . ~ '

Various p o s s i b i l i t i e s e x i s t f o r t h e s e l e c t i v e advantage of t y ing

toge the r var ious s e c t i o n s of t h e s t ream by animal migra t ion : (1)

Already d iscussed i s t h e r o l e of migrat ion i n t h e reproduct ion and

d i s p e r s a l o f j uven i l e s t ages of t h e spec i e s . (2) Various a r eas ~f

t h e s t ream may become devoid of f i s h e s due t o n a t u r a l d i s a s t e r s ,

such a s drought , summer low oxygen, severe p reda t ion , e t c . Migra-

t i o n provides a s t eady source of r eco lon ize r s t h a t can occupy empty

h a b i t a t s : (3) The migrat ion and reproduct ive system allows a

popula t ion t o be maintained i n a c u r r e n t . Any downstream d r i f t

may be compensated f o r by migra t ion . (4) Preda tors , moving through

t h e s t ream tend t o feed most heav i ly i n a r eas with l a r g e numbers

of prey s p e c i e s , and thus tend t o con t ro l p o s s i b l e excess ive

inc reases i n t h e s e s p e c i e s . (5) The con t r ibu t ion of minerals t o

upstream reg ions by migra t ing animals i s d iscussed elsewhere i n

t h i s s e c t i o n . (6) According t o Levins (1964) migrat ion has s e l e c t i v e

va lue i n t h a t it al lows s u f f i c i e n t in te rchange between popula t ions

s o t h a t l o c a l adapta t ion f o r shor t - range environmental f l u c t u a t i o n s

w i l l n o t become a very important f a c t o r which would reduce t h e

o v e r a l l f i t n e s s of t h e gene pool . However, t h i s does not reduce

t h e a d a p t a b i l i t y of t h e popula t ion a s a whole t o widespread changes

i n environment. This may be a f a c t o r i n t h e s e l e c t i o n of spec i e s

f i t n e s s i n s t reams such a s New Hope Creek i n t h a t t h e f i t n e s s of

t h e gene pool a s a whole i s maintained and no t wasted on non-

s e l e c t i v e adap ta t ion t o shor t - te rm l o c a l even t s , such a s s t r e s s dur ing

except iona l drought.

I n t e r a c t i o n of Yield and Organizat ion

New Hope Creek, l i k e many o t h e r complex systems, can be a r b i -

t r a r i l y d iv ided i n t o a subsystem expor t ing energy and another sub-

system rece iv ing t h i s energy, and i n r e t u r n supplying c e r t a i n

o rgan iza t iona l o r o t h e r s e r v i c e s t o t h e exp lo i t ed system i n a feedback

loop. Examples of t h i s r e l a t i o n would be: A prey Ifdonatingft a

c e r t a i n percentage of i t s energy resources t o p reda to r s i n exchange

f o r popula t ion r e g u l a t i o n ; f lowers provid ing bees with energy i n

return for pollination services; and farms su?plying cities with

food and receiving in exchange fertilizers, farm machinery,and

social services.

Such a system can be defined in New Hope Creek. The upper regions

of the stream export food energy to downstream regions and receive

in return genetic informatio? resources, populations of higher

trophic levels to utilize seasonal energy excesses, population

control, reseeding when necessary, and minerals. All of the above

are concentrated biological control agents and are effective in

relatively small amounts.

Margalef (1962, 1969; see also Deevey, 1969) has considered

the energy information exchange between two systems or subsystems

in some detail. A downstream system that has greater organization

(which Margalef calls 'mature') may be more efficient in its use

of energy. An upstream, less organized system (which Margalef calls

'imrnatureV)may not have the structural and organizational frame-

work for using energy as efficiently as the more mature system, and

as a result often loses much of its energy to export. If the more

organized system is able to utilize this energy lost by less organized

systemsit,inacertain sense, exploits the less organized system. In

return, the more organized system gives v~ information to the lcs

mature one, aiding it in becoming more efficient in its own use of

energy, and increasing its organization.

Although New Hope Creek is readily divisible into two segments

the upstream one supplying energy to the downstream one, and the

downstream one supplying information t o t h e upstream one, f u r t h e r

agreement wi th Margalef 's theory i s no t s u b s t a n t i a t e d . The i n d i c e s

of ma tu r i t y should, according t o Margalef, be h igher i n t h e downstream

reg ion and lower i n t h e upstream region . This was no t s u b s t a n t i a t e d

by i n v e s t i g a t i o n s . The upstream region had h igher volume product ion

(mean of 3 . 1 versus 1.4 g m-3 day'') and a lower r a t i o of product ion

t o r e s p i r a t i o n (0.4 versus 0 .6 ) , i n d i c a t i n g t h a t t he biomass sup-

por ted p e r u n i t o f photosynthes is was g r e a t e r . S tudies of pigment

r a t i o s (Motten, 1970) showed s l i g h t l y g r e a t e r D430/D665 f o r both

pools and r i f f l e s i n t h e upstream regions of New Hope Creek i n oppo-

s i t i o n t o Margalef 's theory . Thus, t h e energy-information i n t e r -

change theory may have v a l i d i t y a p a r t from any cons idera t ion of

r e l a t i v e ' m a t u r i t y . '

Some Other Animal Migrations and Environmental

Energy P a t t e r n s

Migration i n New Hope Creek may be an example of a widespread

phenomenon. A number of examples of migra t ion t o regions of high

p r o d u c t i v i t y f o r reproduct ion were considered i n t h e d i scuss ion - - fo r

example, the Texas Bays work of Odum and o t h e r s . Other examples

would be summer migrat ion of many b i r d s t o a r c t i c a r eas f o r reproduc-

t i o n dur ing t h e high energy input of very long day l igh t pe r iods , and

t h e heavy u t i l i z a t i o n of r i c h e s t u a r i n e and nearshore reg ions by

small salmon, p a r t i c u l a r l y chums and p inks . Fur ther ana lyses of p re -

s e n t migra t ion and energy a v a i l a b i l i t y p a t t e r n s , a s wel l as p o s t -

P l e i s tocene opening of niches and evolu t ion of lakes us ing sockeye

salmon, may be f r u i t f u l a r eas f o r a d d i t i o n a l r e sea rch .

New Hope Creek Watershed Annual Phosphorus Budget

F ish migrat ion upstream may p a r t i a l l y o f f s e t t h e downstream

t r a n s p o r t of minera ls . It i s important i n t h e mineral budgets

of salmon lakes i n Alaska (Donaldson, 1968) and i n Russia (Krokhin,

1967); and i n New Hope Creek, t h e r e l a t i v e l y l a r g e mass of f i s h

moving upstream q m a r e n t l y a l s o con t r jbu te s t o t h e mineral balance.

Some d a t a on t b e p h o s p 5 ~ r u s budget f o r New Hope Creek a r e ca l cu la t ed

i n Table 29 and summarized. f o r t h e watershed i r ? Figure 46.

Measurements were made from .Tune 14, 1968 t o June 13, 1969 of

phosphorus flows i n t h e water i n leaves and i n f i s h e s . The r e s u l t s

i nd ica t ed t h a t phosphorus discharged i n suspension o r s o l u t i ~ n i s ,

by f a r , t h e most important f a c t o r i n t h e movement of t h a t element.

Less than 0 . 2 percent of t h a t amount i s l o s t i n l e a f d i scharge

and even l e s s than t h a t a s f i s h and o t h e r animals moving downstream.

The amount of phosphorus brought upstream by f i s h e s was about one-half

of t h a t l o s t by l e a f d i scharge and l e s s than 0 .1 percent of t h a t

l o s t by s t ream discharge . Therefore, on an annual b a s i s , t h e con-

t r i b u t i o n of phosphorus t o t h e headwaters by f i s h was sma l l .

Some s t u d i e s have suggested t h a t upstream movements of inver -

t e b r a t e s may he lp o r e n t i r e l y compensate f o r t h e doh-nstream d r i f t

(Minckley, 1961; Bal l and Hooper, 1963; Hughes, 1970). Others have

emphasized t h e l o s s i n d r i f t . I n sec t d r i f t was not formally sampled

i n New Hope Creek; however, casua l observa t ions and seve ra l 24 hour

plankton n e t d r i f t sam2les i nd ica t ed t h a t i n s e c t biomass i n t h e

d r i f t recorded by Waters (1965) and Anderson and Lehmkhul (1968)

and a d j u s t i n g these t o t h e volume of d ischarge of New Hope Creek.

Table 29. Annual Movement of Phosphoms i n New Hope Creek: June 14 , 1968 - June 13, 1969.

To ta l To ta l Tota l T o t a l Wet weight P T o t a l P Month water d i s so lved and l e a f P o f animals i n animals animals in animals

d i s cha rge suspended d ischarge l o s t in-moving upC moving upd moving downC moving downd 104 m3 P dischargea g dry w t . l eavesb g g g wet w t . g

€! g

1968

June 14-30 14.04 7,570 478 0.2 5,560 19.46 2,210 7.84

J u l y 14.06 . 7,990 971 0 . 4 3,440 12.09 1,795 6.28

August 1 .93 104 764 0 .3 2,290 8.02 1,218 4.27

September 0 0 0 0 480 1.68 300 1.05

October 15.8 8,540 20,025 82.9 3,440 12.05 1,985 6.95

November 81.6 44,100 851,760 349.0 5,140 18.00 1,259 4.40

December 77.3 40,500 38,475 6 .6 620 2.16 403 1.41

8 a Monthly water d i s cha rge t imes mean t o t a l phosphorus concen t r a t i ons (5.4 . 10- g P g-l wa te r ) .

Monthly l e a f d i scharge t imes mean phosphorus conten t of leaves (4.1 . 10-4 g P g-l l eaves ) .

Monthly averages for each day times number of days i n month.

Monthly movements t imes mean phosphorus conten t o f f i s h (3.5 . g P g-l f i s h ) .

cd a, c , C L t J > M 0 m c d b o a ,

0 In I-. cn

0 N 00

.-, r. N

0

"i 01 M N

0 0 d e,

CO a

0

m d

d Ln Ln

-3

a d-

0 0 0 " d CO

N

L n m rl

ri . rl k

2

I-. N

a 0 N

M \L) rl

.-,

cn Ln

v, N

N M Ln

0 M M " N m d

N

I-. w cn

r. 10

M ri u3

m

N

d N N

m

rl a m

CO

CO N N " rl

ri rl cd cd t' 3 0 a b 2

Figure 46. Diagram of phosphorus flows i n New Hope Creek

watershed. Flows a r e i n grams of phosphorus p e r h e c t a r e of watershed

p e r year . The watershed has about 6800 h e c t a r e s above t h e Concrete

Bridge S t a t i o n . Phosphorus added i n r a i n f a l l was ca l cu la t ed a s about *.

1 p a r t s P i n r a i n f a l l (Donaldson, 1967; Cooper, 1969) t imes

l o 4 in2 h a - l t imes 0.95 m r a i n f a l l f o r t h e one yea r per iod (U .S.

Weather Bureau, Raleigh-Durham Ai rpo r t , N.C.), o r about 95 g phos-

phorus p e r h e c t a r e .

Annual cyc l ing of phosphorus through l e a f development and f a l l

was obta ined f o r deciduous f o r e s t i n Duke Fores t (Gar re t t , personal

communication). Using conversions from Gosz e t a l . (1970) and Rodin -- and Bazi lev ich (1965), t h i s r e p r e s e n t s about 1960 g P p e r ha pe r

yea r f o r l e a f l i t t e r and approximately 2880 g P p e r ha f o r t o t a l

l i t t e r .

Standing crop of phosphorus f o r f i s h e s was est imated from

stream sampling d a t a done by t h e North Caro l ina Wi ld l i f e Resources

Commission (Carnes e t a l . , 1964). Thei r va lue of 27.8 kg p e r ha -- (l758g f o r 0.154 a c r e s ) was m u l t i p l i e d by t h e approximate phospho-

r u s content of f i s h (0.35 p e r c e n t ) .

T o t a l phosphorus i n t h e t o t a l o rganic s tanding crop of Duke

Fores t was approximated by averaging va lues f o r 14 p ine and mixed

deciduous reg ions of t h e world given i n Rodin and Bazi levich (1965).

A mean va lue of 63 kg phosphorus p e r ha was used, as t h e watershed

of New Hope Creek i s about one-half deciduous and one-half p ine

(anonymous map suppl ied by t h e Duke Univers i ty School of F o r e s t r y ) .

Even assuming t e n c a t a s t r o p h i c f loods p e r year of t h e magnitude of

t h e l a r g e s t recorded i n New Hope Creek, only about 0.2 of one percent

of t h e mass of leaves l o s t would be l o s t a s i n s e c t d r i f t . Thus

phosphorus l o s s by i n s e c t d r i f t was considered smal l .

The r e s u l t s of t hese c a l c u l a t i o n s computed f o r t h e e n t i r e

watershed a r e given as a diagram us ing energy flow language (Figure

4 6 ) . Standing crops from l i t e r a t u r e va lues a r e included. The

l o s s of phosphorus by New Hope Creek (97 g ha - l ) compared t o t h e

s tanding crop i s smal l , and may be e n t i r e l y rep laced by amounts

added i n r a i n f a l l a lone.

This l o s s of phosphorus from Duke Fores t by New Hope Creek

was about t h e same a s va lues from o t h e r s t u d i e s repor ted i n Rodin

and Bas i lov i t ch (1965). This compares with a general l o s s of

from 0 . 9 t o 21 kg pe r ha pe r year f o r Ca, Mg, K , and Na f o r four

s t u d i e s summarized by Likens e t a l . (1967). Standing crops of -- phosphorus i n f i s h e s i s an important r e s e r v o i r during summer low

water flows. Migration may be important i n t h e mineral budget

by maintaining t h i s r e s e r v o i r .

. Analog Simulation of a Migration Model

An Electronic Associates, Inc. Model TR-20 analog computer

.was used to simulate the process of migration in New Hope Creek.

Figure 47 gives, in energy flow symbols( a model based on parts of

the energy diagram in Figure 45. Included are differentia-l equations

for the energy accrual to upstream and downstream populations of

fishes and approximate coefficients for energy transfer based on

New Hope Creek data.

The analog circuitry corresponding to the differential equations

is given in Figure 4 8 .

Analog Results and Discussion

Annual results of the model are given in Figures 49 and 50.

The former shows the input of energy into the upstream and downstream

compartments of the model after the function generator and associated

pots had been adjusted to give an energy input curve with a spring

peak similar to such curves observed in New Hope Creek. Figure 50

shows the rate of energy accrual to upstream, downstream, and upstream

and downstream combined populations of fishes with and without

migration.

Each population of fishes draws energy as a product of the

energy available in the environment and the number of fishes available

to exploit the energy source. As fishes move from one region of

the stream to another their ability to add to their energy supplies

changes, being greater in the upstream, more productive regions. Thus,

as the upstream or downstream population of fishes gains or loses

Figure 42 . Energy flow diagram for analog computer model. The

input of sun and organic energy is represented by the circle on the

left -hand side of the page. The curve drawn in the circle repre-

sents the annual energy input to upstream and downstream fish

populations with a peak in the spring corresponding to peaks in

photosynthesis and respiration that occur in New Hope Creek. The

storage symbols represent fish population biomass in upstream and

downstream regions which feed from the energy sources. The energy

input to the downstream population is, on a volume basis, only one-

third of the energy input to the upstream population. Arrows

drawn to heat sinks represent metabolism, or energy loss by the

second law of thermodynamics. The multiplier symbols represent

rates of food flow from the primary production and other input

sources to the fish populations. The lines connecting the fish

populations are migrations. The energy drain on fish migrating

against the current is represented by the heat sink attached to the

upstream migration line. Current-assisted downstream movement is

shown as a multiplier on the downstream migration route with an

input from the upper circle. Differential equations describing

the populations and transfer coefficients are included below the

figure.

Figure 4 8 . Analog symbols representing the energy pathways

in Figure 47. Symbols are standard analog notation. Triangles

represent summers (two inputs) or inverters (one input), triangles

with rectangles are integrators,six-sided figures are multipliers,

and the small circles are potentiometers, or lp0ts.l Lines drawn

between symbols are electrical lflowsl (actually differences in

potential), with an arrow pointing into a vertical line represent-

ing one-way flow and diagonal bars,off-on switches. The letters

VDFG stand for variable diode function generator. Numbers on each

symbol refer to actual numbers used on the computer. The upper set

of modules represents the energy-input pulse generator, the middle

set represents the upstream fish population, and the lower set

represents the downstream fish population.

Figure 49. Analog output of energy pu l se genera tor . The

annual i npu t of energy i n t o New Hope Creek was s imulated a t t h e

upstream s e c t i o n (a) and t h e downstream s e c t i o n (b) of t h e

stream, wi th a l a r g e sp r ing pu l se s i m i l a r t o one found i n New

Hope Creek.

Figure 50. Analog s imula t ion of annual energy acc rua l t o

populat ions of f i s h e s i n New Hope Creek. The annual energy flow

t o f i s h e s i n New Hope Creek a t t h e downstream s t a t i o n reg ion (a

and b) and a t t h e upstream region (c and d) i s represented without

migra t ion (b and c) and with migrat ion (a and d ) . Since a g r e a t e r

mass of f i s h moved upstream than downstream, t h e r e i s a n e t ga in

t o t h e upstream popula t ion and a n e t l o s s t o t h e downstream popu-

l a t i o n . The energy accrua l t o t h e t o t a l populat ion of f i s h e s i s

represented by t h e upper p a i r of l i n e s . Line o i s without migra-

t i o n and l i n e f i s with migra t ion . The t c t a l amount of energy

flowing t o t h e e n t i r e populati,on of s t ream f i s h e s i s g r e a t e r with

migra t ion than without .

i n d i v i d u a l s , t h e popula t ion ' s energy drawing power becomes g r e a t e r

o r l e s s . In Figure 5 0 , t h e lower s e t o f l i n e s r e p r e s e n t s t h e annual

energy acc rua l t o t h e downstream populat ion of f i s h e s . The upper

of t h e s e two l i n e s (a) r ep re sen t s energy acc rua l without migra t ion .

With migra t ion (b) , t h e energy accrua l i s l e s s s i n c e a g r e a t e r

mass of f i s h movcsupstream than downstream. On t h e o t h e r hand, t h e

upstream popula t ion gains energy with migrat ion a s a g r e a t e r mass

of f i s h move i n t o t h e upstream region than moves out ( l i n e c ,

without migrat ion; l i n e d , w i th ) . The t o t a l energy acc rua l t o t h e

e n t i r e popula t ion of f i s h e s i s g r e a t e r with migrat ion ( l i n e f )

than without ( l i n e e ) s i n c e t h e energy ga in t o t h e f i s h moving i n t o

t h e upstream region i s l a r g e r than t h e l o s s t o t h e downstream popu-

l a t i o n . This inc ludes t h e l o s s of energy due t o t h e cos t o f

upstream migra t ion .

Any model of na tu re s u f f e r s from s i m p l i c i t y , and t h i s one i s

no except ion. The g r e a t e s t de f i c i ency i n t h i s model i s t h e f a i l u r e

t o inc lude p rov i s ions f o r reproduct ion and growth of young. The

movement of a j uven i l e f i s h o r egg mass may have more p o t e n t i a l

f o r drawing energy than an equiva len t mass of l a r g e r f i s h . Another

shortcoming i s t h e Pack of d a t a on populat ion l e v e l s of f i s h e s a t

t h e two d i f f e r e n t s t a t i o n s . Only one downstream value was

a v a i l a b l e and had t o be used f o r both compartments. However, t h i s

model shows some cons is tency with t h e hypothesis f o r an adapt ive

r o l e of migrat ion i n complex s t ream ecosystems.

SUMMARY

1. Patterns of fish and other aquatic animal movements were in-

vestigated in two North Carolina streams from April, 1968

to June, 1970 using weirs with traps.

2 . Community metabolism was measured during the same period

using diurnal analysis of oxygen.

3. More animals were caught moving downstream than up, but more

than twice as much mass (three times for fish alone) of or-

ganisms was sampled moving upstream than down. This was true

for all locations sampled and for virtually all months of

the year.

4. A pattern of larger fishes moving upstream and smaller fishes

moving downstream was observed for virtually all species of

fish.

9. Migration was by far heaviest each year during the months of

blarch, April and May. This was true for all species considered

as a group and also for most species considered individually.

6. All physical evidences of spawning condition were observed during

periods of maximum movement for each respective species, and

ripe specimens (discharging eggs or milt) were almost invariably

sampled moving upstream. Spent individuals were sometimes

sampled moving downstream, but never upstream. Thus, it is

assumed that the movement patterns are connected with spawning

and t h a t f i s h move upstream t o spawn.

7 . The very low recapture (6.9 percent ) of marked i n d i v i d u a l s

i n d i c a t e s t h a t t h e movements a r e t r u e migrat ions and a r e n o t ,

i n gene ra l , merely i n t e r c e p t s of home-range movements. The

low r e t u r n s a l s o i n d i c a t e t h a t f o r many f i s h t h e upstream move-

ment i s not accompanied by a r e t u r n downstream movement.

The f a t e of t hese post-spawning ind iv idua l s i s n o t known.

8. Movement of f i s h e s i n t h e s t reams was increased by a r i s e i n

water temperatures during t h e s p r i n g months and by a r i s e i n

water l e v e l during a l l per iodsof t h e year except dur ing t h e

win te r .

9. Each spec i e s was more Likely t o t r a v e l one a t a t ime than by

two's o r more, bu t concent ra t ions of many ind iv idua l s t r a v e l i n g

toge the r , o r a t l e a s t on t h e same day, occurred more f r e - C-

quent ly than would be expected by chance a lone .

10. Metabolism f o r t h e e n t i r e aqua t i c community of New Hope Creek

was moderately low compared with o the r a r eas s tud ied . Gross

primary product ion a t t h e p r i n c i p a l sampling s t a t i o n ranged

from 0 . 2 t o almost 9 g O2 m - 2 day- l . Normal values were about

1 and 1.5 g O2 m-2 day- l , r e s p e c t i v e l y . Both gross primary

product ion and community r e s p i r a t i o n were g r e a t e s t i n Apr i l

and May, wtih another smal le r peak i n t h e autumn. Resp i r a t ion a

was n e a r l y always g r e a t e r than photosynthes is , i n d i c a t i n g t h a t

New Hope Creek i s dependent upon e x t e r n a l sources f o r much

o f i t s organic energy. *

11. Although volume metabolism was cons iderably g r e a t e r i n some

regions than in others, areal metabolism (i.e.,volume metabolism

corrected for depth) was remarkably constant throughout the

stream. The generally shallower upstream reaches of the stream

had the highest volume photosynthesis and respiration.

12. The hypotheses are presented that behavioral patterns of up-

stream migration for spawning have selective value in starting

juvenile fishes in a region where food resources are concentrated.

Later dispersal to other, often more stable and less stressful,

areas may also have selective advantage. Other potential

advantages accruing from migration include: recolonization

of defaunated regions, such as those following a drought;

overcoming displacement tencencies of a current; more efficient

population control of prey; genetic interchange; and distribu-

tion of minerals.

13. An energy diagram was drawn comparing calculations of metabolism

and fish migration. About 0.01 percent of the entire eco-

system's energy usage is contributed by the migrating animals.

14. About 40 percent of the estimated standing crop of fish above the

principal sampling station were involved in migration during

one year. Upstream migration alone could replace the population

of part of New Hope Creek in 3.7 years. Downstream migration

alone would take 7.9 years, and replacement from above and

below would take 2.5 years.

15. The l o s s of phosphorus from t h e watershed a b m c t h c Coacretc

Bridge by a l l processes connected with New Hope Creek was small

compared wi th t h e s tanding crop of phosphorus, and may ha.ve

been rep laced by r a i n f a l l . The mount of phosphorus rep laced

upstream by migrat ing f i s h was small considered on an annual

b a s i s , b u t dur ing t h e summer months upstream con t r ibu t ions

of phosphorus from t h e sp r ing and summer f i s h runs may be

important . I n a d d i t i o n , t h e s tanding crop of f i s h e s was

an important r e s e r v o i r of phosphorus.

16. New Hope Creek can be considered an energy-information exchange

system s i m i l a r t o many o t h e r such systems. Upstream regions l o s e

a c e r t a i n p a r t o f t h e i r energy r e se rves t o downstream popula-

t i o n s , and, i n r e t u r n , downstream populat ions supply gene t i c

information and con t ro l func t ions t o t h e upstream reaches .

17. Analog s imula t ion of f i s h popula t ions i n New Hope Creek with

and without migra t ion i n d i c a t e t h a t observed p a t t e r n s of

migra t ion can inc rease t h e energy acc rua l t o a populat ion of

f i s h e s .

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APPENDIX A

DIFFUSION PROCEDURES USED IN NEW HOPE CREEK

METABOLISM STUDIES

One of the most difficult tasks in determining community meta-

bolism from changes in oxygen is establishing the rate at which

oxygen enters or leaves the water if the water is not at 100 per-

cent saturation. Two papers in 1954 (Haney, 1954; Ammon, 1954)

summarized the knowledge about diffusion to date and concluded that

the amount of gas transfer could be calculated by equation (1):

where D is the diffusion rate per area, S is the saturation defi-

cit between water and air, z is the depth, and k is the gas transfer

coefficient defined on a volume basis. Since the rate of diffusion

is linearly proportional to the saturation deficit, no correction

of the constant is needed at different oxygen concentrations. See

Odum (1956) and Owens (1969) for a more detailed consideration of

this.

Diurnal Curve Method for Determining the Diffusion Constant

In the study in New Hope Creek, three methods were used to esti-

mate the diffusion coefficient. In the first, appropriate values were

substituted into the expression (2) supplied by Odum (1956):

where k is the diffusion constant in g oxygen rn-3 hr'l, qm is the

1 rate of change at a time before dawn in mg 1.' hr' , qe is the rate of

- 1 change at a time after sunset in mg 1-I hr , Sm is the percent ss turil-

tion deficit before d z ~ n at the time chosen for ca.lc112atj.ng q,, end Se

is the percent saturatkm deficit aftm sunset at the time chcse.7 fo-

calculating qe. (These terms must be chosen careftilly to avoLd non-

representative water masses .) If respiration at night is constant,

the change in dissolved oxygen will be proportional to the percent

saturation of the water. However, as pointed out by Odum and Vilson

(l962), hielch (l968), and Owens (l969), the respiration of an aquatic

ecosystem is not constant, especially in systems with many small

algal cells. In general, respiration is considerably higher in the

post-sunset hours than in the pre-dawn hours, due to daytime storages

in the cells (Odum et al., 1963), higher temperatures, and higher

oxygen values. Not correcting for different levels of respiration can

introduce serious error into the determination of k and will, in gene-

ral, give values that are too large. Also, as pointed out by Owens s,

(1969), the difference between the saturation values must be large to

give meaningful results, because of statistical error. This method

was used for New Hope Creek data on several days with smooth oxygen

curves showing no abrupt changes caused by different water masses or

sampling error. Table A-1 gives these results, which varied widely

from day to day.

A possible correction for the differences in respiration bras

given by Odum and Wilson (1962) :

where kl is the volume based diffusion constant, q is the rate of

change in g mm3 hr-I per gradient of 100 percent saturation deficit, .I

1 r is the independently determined respiration also in g m'3 hr- ,

and S is the percent saturation deficit. Neither Odum and FTilson

nor this investigator was able to apply this correction because of

insufficient independent data on respiration.

In open waters, Copeland and Duffey (1964) and Owens (1969)

used different diffusion constants at different times of the day

due to changes of circulation patterns with changes in wind and tem-

perature stratification. Since all of New Hope Creek flows through

more or less protected areas, either through a gorge or in areas

completely surrounded by trees, there were few wind effects apparent

over most of the stream. Hourly corrections of k due to differences

in wind velocity were considered unimportant.

Stream Morphology Method - of Determining - the Diffusion Constant ---.---

A second method of determing k uses the expression (4), originally

developed by Streeter and Phelps (1925) and improved upon by Churchiil

et al. (1962) --

The diffusion constant k2 in this expressi-on (4) is defined dif-

ferently from k of equations 1-3, k2 is the g m-3 hr-I diffusion when

the saturation deficit is one mg 1 where k2 is in days-', R equals

the hydaulic radius (approximately the depth), in feet, and V is the

cross sectional mean velocity. Corrections can be made for other

temperatures at 2.41 percent per degree C (Churchill p;t alL, 1962).

Two Diffusion Coefficients and Conversion - ---- - --.-

The diffusion equations for the two commonly used diffusion coeffi-

cients may be compared in equations (5) and (6).

D = kS = k (100 - percent oxygen saturation of stream) . (5)

Table A-1. D i f f u s i c n Cons tan t s Derived from D i u r n a l Oxygen Data

Date Locat ion Depth D i f f u s i o n c o n s t a n t m g rn-3 h r - l

1969 Mar. 29 Concrete Bridge 0 .55 2 .04

Apr. 1 1 1 t I 0 . 5 0 6 - 5 7

Apr. 25 Blac kwood 0.25 2 .50

May 1 6 Concrete Bridge 0.41

June 2

July 1

J u l y 25

1970 Feb. 14 I I I I 0 . 40

Feb. 1 4

23 1

where D is the total oxygen diffusion rate expressed as g 02 m-3 hr'l,

k is the diffusion coefficient as g m-3 hr-I 100 percent saturation

deficitmL (base e), and S is the percent saturation deficit. k is

the slope of a semilog plot (natural log) of diffusion with time.

The total diffusion process using k2 is:

where k2 is the re-aeration rate coefficient (days-', base lo), Cs is

the oxygen saturation concentration (mg 1-I or g m-3) at the prevailing

temperature and pressure, C is the actual stream oxygen concentration,

and D2 is expressed as g 02 m-3 daym1. k2 is the slope with time of

a semilog plot (base 10).

Solving (5) and (6) for k and k2, respectively, gives:

k = D s-' in g O2 m-3 hr-I per 100 percent saturation deficit ( 7 )

- 1 -1 -1 ,-3 k2 = D (Cs - C) in g O2 m-3 day g ( 8 )

The last formula reads "grams oxygen per cubic meter per day per grams

per cubic meter difference between oxygen saturation and stream oxygen

values . I 1

Dividing (8) by 24 gives diffusion per hour; thus, the time

intervals used in each formula are made the same. One hundred per-

cent saturation deficit expressed as grams per cubic meter is Cs.

Since the engineering formulation for re-aeration coefficients (k2)

is defined in terms of logl0, k2 must be multiplied by 2.3 to yield

the equation: 2.3 ' (k2) Cs

k = 24

in g m-3 hr-I per 100 percent saturation deficit (9)

Thus, with a knowledge of the oxygen saturation value for the time in

question, k2 can be expressed as k, and the formula of Churchill et al.

can be used in diurnal oxygen studies. A further correction must he

made for temperature, since Churchill's formula which is base.d only

on differences in stream depth and flow is temperature-dependegt

with different molecular activity rates. This correction: 2.41 percent Q

increase or decrease per degree above or below 20O C as a geometric

ratio, somewhat compensates for the increase in k as the saturation \i

value of water falls with increased tem~erature. For purposes of cal-

culation, the formula k2(T) = k2 (at 20' C) 1.0241 (T - 20) is used. Thus, k is not temperature-dependent due to changing saturation values,

while k2 is.

The area based diffusion coefficients obtained by this method

were divided by the average depth of the water reach being considered

to give values for volume diffusion corrections (equation 1). Table

A-2 gives results of diffusion constant estimates for different mean

depths, mean flow conditicns, and different stage levels for the

stretch of creek above the Concrete Bridge station through which water

flows in one hour.

Dome Measurements of Diffusion Coefficient - --- - The third method of determining diffusion constant is a direct

empirical approach using methods derived from the work of Copeland

and Duffey (1964), who determined k by comparing the changes in oxy-

gen concentrations in the water with changes under a clear plastic

dome. Similarly, k has been determined at the Institute of Marine

Sciences, University of North Carolina, by measuring the rate of

transfer of carbon dioxide across the water surface using an open

system with a plastic dome and an infrared C02 analyzer (Hall and

Day, 1970).

Table A-2. P r e d i c t e d Values f o r D i f f u s i o n Cons tan t f o r New Hope

Creek Above Concre te Bridge S t a t i o n Using Formula Based on Average

Depth and v e l o c i t y a

Average K2 day-' K k dep th ( p e r a r e a ) g m-2 h r - I g ~ n - ~ hr - l - l

m a t 20' C atmosphere-' atmosphere

a See e q u a t i o n s (4 and 9 ) .

In the present studies, the diffusion rate was measured directly

by determining the rate of oxygen re-entry into a clear plastic domz

after filling it with pure nitrogen. In theory, any gas can be used,

since, according to Dalton's law of partial pressure, the oxygen

would diffuse independently of the concentrations of other gases

present. However, experiments with CO and methane were abandoned 2

after these gases diffused into the water, allowing water to rise

too rapidly into the inside of the dome, changing gas values. Since

ordinary air is 78.09 percent nitrogen by volume, water in contact

with the atmosphere would be at about 78 percent saturation relative

to an atmosphere of 100 percent pure nitrogen. This relatively small

difference allows the diffusion of oxygen into the sphere to be stu-

died without interference by substantial loss of the atmosphere

within the sphere and the resultant changes of sphere volume.

The apparatus for these determinations were set up according

to Figures A-1 and A-2. The clear plastic dome was attached to a

plywood collar which floated the dome on the surface of the water.

An oxygen probe (Yellow Springs Instrument $15419) was inserted in

the top of the dome with an airtight seal. Thus, tk probe was

measuring the partial pressure of oxygen in the atmosphere within

the dome. The dome was then sunk within the stream, tilted at an

angle that allowed a substantial flow of stream water over the

probe. The flow was considered sufficient if additional manual

agitation of water over the probe did not give higher readings on the

meter. This was necessary because the probe requires a minimum flow

of water for accurate readings.

Figure A-1. Use of clear plastic dome to measure diffusion

constant. a. The dome is immersed into the water to equilibrate

the probe with oxygen in the stream. b. The water-filled dome

is turned upright, filled with N2, and floated on the water. The

hole in the dome, open to the atmosphere, insures a pressure of

1 atmosphere inside the dome after gas flow is stopped. c. The

diffusion constant, k, is computed from the rate of oxygen re-entry

into the dome in the water.

Once the probe had reached equilibrium with the water with a

constant reading for several minutes, the meter was adjusted to '10,'

which represented 100 percent of the partial pressure of the oxygen

in the water. Three Winkler oxygen determinations were taken at

this time. Thus, the '10' on the meter scale was equivalent to the

total concentration of oxygen in the water as determined by the

average of the three Winkler tests.

While still submerged, the dome was turned collar-side down.

Nitrogen was introduced into the dome through a hole in the side;

this procedure floated the dome on the water surface with an atmos-

phere of pure nitrogen, After a short period, the oxygen meter

would read near zero, as there was little or no oxygen within the

dome. An additional hole in the dome insured that the nitrogen

within the dome was at 1 atmosphere. The nitrogen was turned off,

the holes plugged, and time and meter reading records were kept as

oxygen diffused into the atmosphere of the dome from the water.

It is inaccurate to calculate rates from finite measurements

of change as a simple quotient. Instead, the integral form must be

used because the expression is non-linear. The derivation in Table

A-3, supplied by Dr. H.T. Odum, is required to provide the units

necessary for easy computation.

Diffusion Constants Used for These studies

A11 diffusion results were converted to a volume basis for

comparison. The results for diffusion constant determinations by

each method varied from one to another and within the three methods

used (Figure 10). The results of the stream morphology estimates

Table A-3. Basis for Calculations of Diffusion Coefficient from

Dome Measurements

It is desirable to find the area-based diffusion coefficient

for oxygen, K , in grams oxygen per square meter per hour per atmos-

pheres gradient.

K as defined above relates the flow of oxygen into the dome to

difference in partial pressure with equation (1).

3 = KA (pw - pd) (1)

where J is in grams of oxygen per hour, A is the area of dome in

square meters, pw is partial pressure of oxygen in water in atmos-

pheres, and pd is partial pressure of oxygen in dome as percent of

total pressure. Solving for K gives equation (2) and the desired

units. J

g m-2 atmosphere - 1 K = (2) A (P, - P ~ )

The weight of oxygen (Q) in air phase in the dome is given by

equation (3) which has geometrical density considerations. Q is in

grams of oxygen

where v is the volume of dome above waver in ml, /J is the density

of air at elevation and temperature, and p is partial pressure of

oxygen in dome in percent of total pressure.

Next write the differential equation stating that the rate of

change of the weight of oxygen in the dome's air phase is equal to

the flux of oxygen across the surface (J).

Table A-3 continued

Then substitute in equations (1) and (3).

Next integrate as in equation (7).

Integral tables show the left hand

$ a d; bx 1 = - In (a + bx) b

p is the variable x; substituting d

the integral expression 2

( 7

expression to be of the form

back in a, b, and p one finds d

+ I

To evaluate the integration constant, substitute p as initial 0

pressure when time is zero at start. Then

The final integral equation becomes

Table A - 3 . con t inued

Changing n a t u r a l l o g s t o base 10 l o g s and s o l v i n g f o r K one

o b t a i n s

For 100 p e r c e n t s a t u r a t i o n w i t h a tmospher ic oxygen a s t h e g r a d i e n t ,

m u l t i p l y by 0 .20s t h e oxygen f r a c t i o n of atmosphere. The c o n s t a n t

i n t h e above e q u a t i o n becomes 0.48 i n s t e a d of 2.3. Where v i s t h e

volume of dome i n m l , p i s t h e d e n s i t y of a i r i n mglml, A i s t h e

a r e a of dome i n s q u a r e m e t e r s , t i s t ime a f t e r s t a r t i n h o u r s , p,

i s p e r c e n t s a t u r a t i o n , and po i s p e r c e n t oxygen i n dome a t s t a r t .

R e s u l t s f o r f i v e e s t i m a t e s of d i f f u s i o n c o n s t a n t s u s i n g t h e

dome method a r e g i v e n i n Table A-4.

Table A-4. Estimates of Diffusion Constant (K) Obtained Using

the Dome Method for Representative Pools and Riffles Above Concrete

Bridge Station

Date Location Depth K k (average) g rnm2 hr-I g m-3 hr-I

at 0 percent at 0 percent saturation saturation"

1970 May 13 Riffle 0.55 0.95 1.73

May 22 Riffle 0.58 0.49 0.83

June 14 Riffle 0.35 0.178 0.712

May 21 Pool 0.55 0.10 0.18

June 16 Pool 0.35 0.036 0.079

Based on average depth for stream at that time and place.

were in between the pool and riffle estimates obtained with the dome.

The results of the diurnal curve method were scattered and often

higher than estimates made using the other two procedures. As men-

tioned in "Methods and Materials", the lack of correction for

differences in respiration would tend to give high values for k

obtained by the diurnal curve method where evening respiration is

higher than pre-dawn respiration, as is the case for New Hope Creek.

The stream morphology estimates of diffusion, which were

roughly substantiated by the dome measurements, were used for the

estimates of metabolism for this study.

Bailey (1970) found similar variations within and between

different diffusion determination methods, but concluded that for

the shallow reaches of the Sacramento-San Joaquin region, Churchill's

formula had reasonably good predictive value.

Diffusion Constants for Other Stations

A diffusion constant of 1 g m-3 hr-I was used for the Blackwood

station. This was based on estimates by the stream morphology method

of from 0.77 to 1.0 g mm3 hr-I and one estimate by the dome method

- 1 of 1.3 g m'3 hr . The area.based diffusion constants were similar

to the ones for the Concrete Bridge station, but the shallower nature

of the stream gave larger volume values.

The diffusion constant used for the Wood Bridge station was 0.4

g m-3 hr-I at all depths sampled for oxygen changes except during

very low water when a diffusion constant of 1 was used. These dif-

fusion constants are based on the areal values from the other stations.

A very deep pool located above the shallows at the station gives this

site the deepest average depth for all the oxygen stations, resulting

in the relatively small volume diffusion constant.