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Il\)S-Fred'y\ -Ptow ~Ystem ·"' EFFECTS OF Y'.:..OW PA'rTERNS BELOW LARGE a stronger preference for overhead bank cover than do smaller DAMS ON STRE..bJi BENTHOS: ~ REVIEW though competition and territoriality may explain this James V. Ward ., difference. ,,.· Department of Zoology and Entomology 2) The been d~vei~ped is a habitat Colorado State University .. · Fort Collins, Colorado 80523 ioss of available trout I' ABSTRACT and the I sections investigated, The variously modified flow patterns below dams ~e considered in rela­ ' to exist between le cov;: and. standing tionship to the effects Qn ecological factors of importance to the benthic crop. this area is needed to allow the biologist communities of receiving streams. Species composition and diversity are coo­ the siderably modified by upstream impoundments. Benthic standing crop may be ~0 uantify the biological significance of dewatering enhanced or reduced, largely depending on the flow regime. Daily flow fluctu­ of various stream reaches. ations, if not too severe, may b~ associated with dense benthic populations as long as a relatively constant seasonal flow pattern is maintained. Little is known regarding subtle, sublethal effects of dams on life cycle phenomena and biotic interactions and more data are needed on current preferenda of import­ ant fish food species. Any flow regime which significantly reduces habitat diversity should be avoided. A diverse substrate with silt-free interstices will considerably reduce deleterious effects of periods of reduced flow, I 1: fluctuating flow and high current velocity. In establishing flow criteria for benthos, each dam must be considered individually. INTRODUCTION The productivity, diversity and composition of the stream benthic community is extremely important to the total functioning of the stream eco­ Needs • Soui€e.e.~ Il\)s-fred'Y\ -Ptow ~ystem. Besides providing a major source of food for stream fishes, macro­ CAMeY".i caJ.J ~lsh.c~cs Sec, efy; brnthos may be the best indicators of past and prevailing ecological condi­ JC/7(;,) tions. They are not as mobile as fishes and do not present nearly the taxonomic difficulties as microorganisms (in which congeneric species often h11ve widely divergent ecological requirements). In addition, a diverse h"r.thic community is able to process relatively large amounts of organic: lilllttcr • The term "benthos," as used in this paper, refers to the benthic ~Acroinvertebrates of streams. Until recently, most studies of biota below dams in North America con~ ~tlllHcd benthos only parenthetically and only in direct relationship to fish f>t>l•uhtions. In the past few years several invest-igators have concentrated t),.,h •tudics on benthic communities below dams, and their work bas provided ••ltllt1cmnl insight into the conditions prevailing; in these modified environ­ '"'111" lind the effects of these conditions on the structure and function of 1)1., lilT Nun ecosystem. 235 t~ ~ ,..... ~ . ........ :r.... _....,. .. "-...... -~ "[' ·~~- Iii!,·~·-- . l<i.!'."~ 'i~-p--·=~.Pf~z.'::' 7~.C;}-:tJtDli~-c~.-... ~~fit~~~~!/;:' ~ .. :~t.,l'-~ ~~'Itt- ~~~.,~~,_~,·~~~-,. 4 ~~-,.t:"'i~~-~."~:::;:~~~ .. .:; ... ~ ..- l, ,4;..~U,.·» ~~" ' ·~~ ...... _.}!,~~~""'::".~ ·'I '·' 236 237 ~ It is the purpose of this paper to review and synthesize ~rk relating course, in the interstices between substrate parti~les. Morphological adapta- to effects of variously modified £low patterns below dams on stream benthos tions of stream invertebrates, such as flattening and streamlining, enable and to elucidate interrelationships oetween discharge and other abiotic and them to avoid the current by being cryptic or by residing in the boundary biotic components important to the benthic community. layer. if The mosaic distribution of the benthic fauna of stony streams has long GENERAL EFFECTS OF CURRENT been recognized and is the primary reason it is so difficult to obtain certain types of field data with a high degree of statistical validity. The In discussing effects of current speed on stream benthos several factors mosaic distribution of the benthos results from the microdistribution patterns must oe kept in mind. Of primary :importance is the £act that many stream of current, substrate type, and food. The multitude of microhabitats allows invertebrates are highly adapted to conditions in running water, and a large a rich and varied benthic fauna to exist in stony streams. Conversely, any­ number are confined to lotic environments (rheostenic) because- of inherent thing which acts to reduce habitat heterogeneity will tend to reduce the num­ current requirements often associated with their respiratory physiology or ber of niches and consequently the benthos. feeding mechanisms (1). Certain caddisflies are able to tolerate much lower oxygen concentrations in flowing than in still water. (2). Some lotic species EFFECTS OF DAMS ON STREAM BENTHOS ,_..,..;-,,._.k'f-~ are restricted to running water because they have lost the ability to move their respiratory structures in the absence of current. Other organisms have Effects dams have on the benthic fauna of the receiving stream depend feeding mechanisms which depend upon current. Blackfly (Sfmuliidae) larvae, upon the type of dam (water storage, hydroelectric, etc.), which in turn which have cephalic .fans for filtering suspended organic particles fr.om the determines the flow pattern; the reservoir depth from which water is released; water, are unable to .feed below certain current velocities (3). Net-building the characteristic stream benthos of the region w1der consideration; and a caddisflies have preferred current velocities in relation to ~et construction, myriad of other £actors relating to the geochemistry, topography, and meteor­ and some species will not opin nets below a certain current velocity (4). ology. The trophic status, depth, retention time, temperature profile, ex­ Other benthic organisms are tied to certain current regimes because of their tent of drawdown and other limnological conditions obtaining in the reservo!r requirements for certain types of substrates (5). are of utmost importance. Characteristics relating to the physical nature ~f It was first suggested by Ruttner (6) that running water was physiologi­ the stream below the dam, such as channel morphology, may also be of critiC!al cally richer than still water. and both algae (7) and invertebrates (8) may. i~portance in determining effects on ~tream benthos. However, within temper­ ~thin limits, increase their metabolic rate as current increases. ate regions, temperature and flow regimes and their ramifications are often Stream organisms vary widely in their current preferences (9), and thie the factors of major importance to the benthos below dams. is generally, but not always, related to their ability to maintain their Effects of th~ temperature pattern below dams have been considered in "":..; l ,._ position against the current (10). Chutter (11) emphasizes the importance of dl'toH by the ~~riter elsewhere (12,13), and the only thermal ftActors to be separating effects of current and depth when studying current preferences of ronuidcred in the present paper are those influenced by the flow pattern. stream benthos. ~~~ to space limitations, discussion will be confined to research published It must be emphasized that many stream invertebrates, to a much greater · In •cicntific journals, and only selectively to processed technical reports degree tha4 most fishes, may not be exposed to appreciable current even tboUJb 811 d other unpublished works. Since Isom (14) recently reYiewed pertinent they reside in a rapid stream. Jaag and AmbUhl (8) have shown that even in. •ludJru ~ontaining information on effects of impoundments on benthic macro­ areas of rapid current, there is a thin zone, the boundary layer, above all Jnv~rt~bretcs in the Tennessee Valley, only limited reference will be made to solid surfaces in which there is negligible current. The thickness of this that hglon. Special influences, such as organic pollution or the release of IHJ.lt II b "dead water" zone, which varies according to velocity. viscosity and turbu­ u atances from the reservoir, ~1 not be considered here. lence, is a few mm. There are other dead water zones behind objects and, of .,.,~,,,('~~.. , ·~:' ' 239 238 ... TABLE 1. Effects'of flow patterns below dams o~ stream Diversity and Standing Crop benthos in North America north of Mexico Table 1 summarizes tfue effects of dams on the diversity and standing crop Probable b Reservoir of benthos in the receiving stream. Although only flow-related causes are Effect flow-related c d Reference cause Type Release Location listed, most authors indicated that other factors may also have had an influ­ st.Crop Diversity California 28 ence on the benthic community. However, all .~thors except Lehmkuhl (15) I NI constan:::y (CS) L considered the flow regime to exert an important, if not major, influence on L Wiscs.nsin 24 NI D constancy (R) the benthos. 25 constancy C L Ontario Whereas diversity, if indicated, was invariably reduced, benthic stand­ I(?) D L Colorado 12 ing crop may be either enhanced or reduced in streams below dams. The effect I D constancy S on standing crop is mainly dependent upon whether the dam results in a more L Colorado 23 D D low flow IH(S) constant or more fluctuating flow pattern. Increased flow constancy, in all 21 fluctuatione H L(?) Massachusetts cases indicated, resulted in an enhanced benthic community. D D L Arkansas 17 Apparently increased seasonal flow constancy may have a beneficial I D controlled HCSR release effect on standing crop even if associated vith short-term fluctuations, pro­ L Saskatchewan 15 viding these are not too severe. Thus Pfitzer. (16) reports an enhanced ben­ D D NI HIS thos below TVA dams despite great daily fluctuations in current velor.ity and L Utah 18 I D &eaaonal HI discharge. Hoffman and Kilambi (17) also report enhanced density of benthos constancy below hydroelectric dams in Arkansas compared wi~h natural streams, despite L Tenn.Valley 16 I NI NI UCR daily fluctuations ;~ current velocity and discharge. This they attribute to 19 I HI L Colorado D NI fluctuation a more stable substrate re~ulting from controlled water releases.
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