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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
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rl a m
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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.