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Page& 131-139 in R. H. Stround, ed. Stemming the of coastal fish habitaTloss. National Coalition for Marine Conservation, Savannah, Georgia. 1992.

New England’s Rivers and

Terry A. Haines

The Atlantic salmon (Salmo salar) is an Shortly after Europeans settled in North anadromous fish native to the north Atlantic America, salmon stocks began to decline from . Adults feed and grow in the Davis a combination of overharvest, blockage of Straits between Labrador and Greenland access to spawning areas by dams and other (Mills 1989), and migrate to coastal rivers obstructions, and degradation of quality between about 40“ and 70’ north latitude for resulting from logging and agricultural prac­ reproduction (Netboy 1974). In the United tices and municipal and industrial wastes States of America, Atlantic salmon originally (Dunfield 1985, Netboy 1974, Stolte 1981). By occurred in at least 28 rivers (Fig. l), from the the late 1940s to the early 195Os,the Atlantic St. Croix on the - salmon population of the United States con­ border to at least the Housitonic in Connecti­ sisted of only a few hundred fish returning to cut, and probably as far as the Hudson River five rivers in Maine (Beland 1984). The in New York (Dunfield 1985). Estimates of Anadromous Fish Conservation Act was populations range from 300.000 (USFWS) passed in 1965 to preserve existing stocks and 1984) to 1.1 million fish (Dunfield 1985). to reestablish salmon populations in rivers

Figure I. Atlantic salmon distribution in New England over time. Bold lines indicate rivers having populations of returning salmon. (sources: USFWS (1984). and A. Mcistcr (personal communication).

131 132 STEMMING THE TIDE OF COASTAL FISH HABITAT LOSS where populations had been extirpated Embryos develop in the gravel during (USFWS 1984). After more than 25 years of winter and hatch in April to early May after recovery about 3,000 to 7,000 Atlantic salmon 175 to 195 days of incubation. Alevins remain now return to 16 rivers in New England (Lewis in the gravel for about six weeks after hatching 1991). Fewer than 1,000 of these are of wild and obtain nourishment from the yolk. Fry origin and the remainder are from introduc­ emerge from the gravel from mid-May to mid- tions of hatchery-reared parr or smelts (J. June and disperse to nursery areas. After two Marancik, U.S. Fish and Wildlife Service, to six years of stream residence, salmon parr personal communication). The continued undergo physiological transformation to the recovery and existence of New England’s smelt stage, which is capable of survival in Atlantic salmon population necessitates the water. Smelts migrate down stream during presence of suitable environmental conditions spring and swim to the feeding grounds in the for all life stages from returning adult to north Atlantic to continue the life cycle. emigrating smolt. In this paper, 1 review the riverine habitat conditions Atlantic salmon Mature Salmon require for survival and reproduction and the effects of human activities on these conditions. Once they have approached the freshwater environment, and possibly before, salmon are Atlantic Salmon Life Cycle dependent on olfaction to find the proper migration route (Mills 1989). There is disagree­ The life history of Atlantic salmon was ment on the nature of the involved odors, but recently reviewed by Danie et al. (1983) and Stabell(1984) concluded that odors produced Mills (1989). After feeding and growing in the by the fish, themselves, pheromones, are most north Atlantic for one to three years or important. Pollution from copper and zinc more, most salmon (refer to Table 1 for mining halts upstream migration of salmon terminology) return to their natal streams to (Saunders and ‘Sprague 1967), possibly by reproduce. Salmon congregate in , interference with odor reception. Experiments bays, and river mouths in the early spring and with Pacific salmon demonstrated that fish then migrate up the rivers to the spawning would home to the source of an organic areas, apparently following olfactory cues to chemical to which they had been exposed as locate these areas. Spawning takes place in smohs (Cooper et al. 1976). Whether this is mid-October to mid-November in gravel areas true homing, the ability of artificial chemicals having moderate current. Individuals that to modify homing behavior was demonstrated survive spawning (kelts) either drop down and raised the possibility that industrial and stream immediately after spawning, or over- municipal wastes discharged into Atlantic winter in the stream and migrate down stream salmon rivers could reduce homing and thus the following spring. reproductive success. Salmon are required to negotiate various obstacles along the length of the migration Table 1. Terminology of Atlantic salmon Ilye stages. route and are capable of surmounting a Source: Alian and Rber (1975). vertical barrier of 2.5 to 3.5 m (8 to 12 ft) Term Dejnilion under favorable conditions. Fish ladders or lift facilities can allow fish passage around most Embryo From fertilization of egg to hatching Alcvin From hatching to absorbance of yolk dams, but fishway efficiency in passing Fry From absorbance of yolk to dispersal from Atlantic salmon varies widely. Efficiency has rcdd been estimated at 85 to 95% (mean 92%) for Parr From dispersal from redd IO outmigration the , Maine, which is believed Smelt Fully-silvered juvenile migrating to the sea Post- From departure from river to onset of wide to be very significant for population status if smoh annulus formation after first winter in the sea the barrier is near the mouth of the river (J. Salmon All fish from onset of wide annulus formation Marancik, U.S. Fish and Wildlife Service, per­ after first winter in the sea (grilse= one sea- sonal communication). Dams can also inter­ winter salmon) fere with migration through reduction in water Keh Salmon from spawning umil it re-enters the sea current. Adult salmon are rheotactic and STEMMINGTHETIDEOFCOASTAL FISHHABITATLOSS 133 require a minimum stream velocity of 0.3 to progress up most rivers, which may have 0.6 m/set (I to 2 B/se@ to continue movement current speeds greater than 4.5 km/hr (3 up stream (Weaver 1963). mph), salmon must have access to resting Another problem from dams and turbine areas where current velocity is C 61 cm/s (2 operation is the supersaturation of water with ft/s) along the way. In addition, holding pools atmospheric gases, primarily nitrogen, a of similar current velocity, but larger and phenomenon reviewed by Wolke et al. (1975). deeper than resting pools, are required in the Briefly, air may be entrained in falling water or vicinity of spawning areas. A holding pool in water passing through a turbine and dis­ surface area of 2.8 m2 (3.4 yd2) per adult is solved at concentrations in excess of atmo­ required (McLaughlin and Knight 1987). The spheric saturation. Gas emboli then form in ideal holding pool has a depth of 1.8-3.6 m blood and tissues of fish exposed to the gas- (6-12 ft) at low flow (McLaughlin and Knight supersaturated water, resulting in death. Such 1987). Channelization of river beds for an event killed 200 Atlantic salmon (10% of drainage and flood control may destroy such the run) in the St. , New habitat. Brunswick, in 1968 (MacDonald and Hyatt The mean temperature of the has 1973). increased 0.5”C (IOF) since the industrial Stream impoundment may also alter water revolution, perhaps as a result of the green- temperature patterns, which are key factors in house effect, although natural variation the delineation of the geographic range of cannot be absolutely ruled out (Kerr 1990). If Atlantic salmon (Danie et al. 1983). Salmon emissions of greenhouse gases continue at the spawning streams are not found south of present rate, global temperatures may increase about 40’ north latitude (in the northern I OC (2°F) above current temperatures by the hemisphere) because summer temperatures are year 2025, and 3°C (2.5’F) by 2100 too warm. Salmon generally enter rivers in (Houghton et al. 1990). All may not late spring or early summer when tempera­ experience uniform temperature increases. tures exceed 5°C (40OF); migration ceases Temperature increases are expected to be when temperatures exceed 25’C (76’F) (Elson greater at higher than at lower latitudes (Smith 1975. McLaughlin and Knight 1987). If water 1990). Meisner (1990) predicted that a 4.1 ‘C discharged through a dam is drawn from the (7OF) increase in July and August air surface of the impoundment, stream tempera­ temperature will reduce brook trout (Sal­ ture may increase rapidly in the spring and velinus fontinalis) habit in two southern reach higher summer maxima. If water is Ontario, Canada, streams by 30 to 42%. Thus, drawn from deep in the impoundment, stream the amount of suitable stream habitat for temperatures may be increased above normal Atlantic salmon may be reduced by global during winter and decreased in summer. High warming, and rivers in the southern part of the summer stream temperatures may also result range may become too warm for parr survival from cooling water discharge from steam (Power 1990). electric generating stations and industrial In addition to changes in temperature, facilities, reduced ground water inflow because global climate change may alter annual and of withdrawals for domestic use, and removal seasonal precipitation patterns and thus runoff of bankside vegetation during timber harvest­ and stream flow (Hengeveld 1990). Even if ing. Restricted upstream passage, coupled precipitation amount were unchanged, in- with the absence of water currents, increased creased mean annual air temperature would water temperature, and the possibility of gas result in decreased runoff and thus decreased bubble disease, make dams a major impedi­ flow (Regier and Meisner 1990). Frenette et al. ment to upstream migration of Atlantic (1984) showed that variability in the number salmon (USFWS 1984). of Atlantic salmon parr in a Quebec river was Atlantic salmon are capable of swimming at directly related to stream flow, and Havey and sustained speeds of 2.2 km/ hr (1.5 mph) and Davis (1970) found that the number of can reach a maximum burst speed of 24 landlocked Atlantic salmon parr in a Maine km/hr (IS mph) (Elson 1975). To make stream was directly related to rainfall. 134 STEMMING THE TIDE OF COASTAL FISH HABITAT LOSS

Embryo oxygen content. Stability of stream beds is important for Spawning occurs in the fall, generally from survival of embryos to hatching. Hydraulic mid-October to early November, when water forces typically mold stream beds into temperature has declined to 7-lO“C(43-50’F). alternating riffles and pools. Streambed Eggs are deposited in depressions (redds) materials may shift somewhat during flood- excavated and covered with gravel by females. water flows, and may modify pool configura­ Redds are usually constructed in areas where tion occasionally, but the natural channel is water velocity is increasing, especially at the relatively stable (Elson 1975, Jordan and tail of a pool or head of a riffle. Preferred Beland 1981). Channelization of streams and areas are in average water depth of 38 cm ( 15 increased runoff from watershed alterations in) and with average current velocity of 53 may increase flow, destabilize stream chan­ cm/s (21 in/s) (Beland et al. 1982). Substrate nels, disrupt redds, and kill salmon embryos. particle size is usually 1.2 to 10 cm (0.5 to 4 in) in diameter. Parr Embryos develop in the gravel for 175 to 195 days (Danie et al. 1983). Permeability of Atlantic salmon embryos hatch typically in the spawning substrate is critical for embryo late April or early May, and fry emerge from survival. Water exchange must be sufficient to the gravel a month later (Jordan and Beland maintain adequate dissolved oxygen concen­ 1981). The fish, now called Parr, then disperse tration. Standard water velocity, measured at and occupy territories in riffles and runs 20 cm (8 in) depth in salmon redds, ranges (Gustafson-Greenwood and Moring 1990). between 74 and about 5,000 cm/h (28 to 1,900 Current velocity at the fish’s nose (focal in/h) (Jordan and Beland 1981). but embryos velocity) is considered the most important may fail to hatch in redds with velocity < 600 factor defining habitat use of salmon parr cm/h (235 in/h) (Peterson 1978). If the stream (deGraaf and Bain 1986). Parr select focal bed contains more than 20% sand (particle size velocities of 5 to 35 cm/s (2 to 14 in/s), and 0.06 to 2.2 mm (0.002 to 0.09 in)), redd larger fish select higher velocities (Morantz et permeability is insufficient (Peterson 1978). al. 1987). McLaughlin and Knight (1987) and Human activity in the watershed may result Elson (1975) reported that substrate particle in siltation of stream beds and reduced sub­ size was important; small parr (4 to 7 cm (1.6 strate permeability. Siltation often results to 2.8 in) total length) preferred areas with from watershed activities such as agriculture, particle size < IO cm (3.9 in) diameter and logging practices, construction of roads, pipe- large parr (total length > 7 cm (2.8 in) total lines, and buildings, and from streambank length) preferred particle sizes from 7 cm up to erosion (Holland 1975). Watt (1988) attributed 25 cm (2.8 to 10 in) or larger, depending on the the decline in Atlantic salmon catch at the length of the fish. Watershed activities that head of the of Fundy to intensive deposit silt in stream beds, or channelization agricultural development that resulted in that reduces riffle/pool distribution and re- siltation of rivers in the area. Grant et al. moves coarser materials, reduce parr habitat. (1986) found that stream crossings by timber Territoriality among fish allocates space harvesting vehicles reduced biomass of sal­ and food resources to ensure adequate growth monid fishes in Canadian streams, most likely and to reduce predation (Symons 1974). because of siltation of downstream reaches. Territory size is a function of the size of the Dissolved oxygen levels for embryo incuba­ fish, bottom roughness (visual isolation), and tion and hatching are optimal at or near satura­ food availability (Kalleberg 1958, Symons tion. The limiting level for early embryos is 1971). Larger salmon parr will defend their 1.14 mg/ L (1.14 ppm), for mature embryos 5.9 territories against smaller Parr, which may mg/L, and for hatching eggs and alevins 8 reduce salmon production in a stream mg/L (McLaughlin and Knight 1987). Pu­ (Symons and Heland 1978). Salmon parr feed trescible organic matter from domestic or on invertebrate drift (Danie et al. 1983) and industrial waste discharge or from farm thus may compete for food with other animal wastes may reduce stream dissolved salmonid and non-salmonid fishes. Mac- STEMMING THE TIDE OF COASTAL FISH HABITAT LOSS 135

Crimmon et al. (1983) found that salmon parr The major causes of pollution in streams in growth was suppressed by the presence of the six New England states are silt, nutrient brook trout, and Gibson and Dickson (1984) enrichment, and organic enrichment (Table 2). found that salmon parr growth increased Industrial and municipal wastes have been when brook trout were removed from a greatly reduced through stringent enforcement stream. Symons (1976) reported that salmon of clean water legislation, so that many rivers parr would defend their territories against in New England are again capable of other species, but none of the species seemed supporting salmonid fishes (EPA 1990). Toxic to have a competitive advantage in obtaining pollutants may reach Atlantic salmon nursery food. Sayers (1990) found that the introduc­ streams from industrial and municipal waste tion of salmon parr into a stream with resident discharge, aerial application and runoff of brook trout caused the trout to move to forest and cropland pesticides, runoff from deeper, slower-flowing stream reaches, and croplands, mining activity, and atmospheric that intra-specific competition for habitat deposition of acids. among salmon was at least as important as Organochlorine pesticides previously caused inter-specific competition. Nevertheless, the serious losses of Atlantic salmon (Elson 1967, introduction of other fish species that may Elson et al. 1973, Locke and Havey 1972); compete with salmon parr for food may well however, these pesticides have been banned reduce production of salmon in rivers. from use and the replacement pesticides are Salmon parr are vulnerable to a variety of much less toxic to fish (Haines 1981, Elson et fish and avian predators. Fish predators al, 1973, Zitko et al. 1970). Metal mining has include northern pike (Erox lucius), chain been a serious problem in New Brunswick and pickerel (&ox niger), burbot (Lora iota), and Newfoundland, Canada (Elson et al. 1973), brown trout (Salmo trurta) (Bley 1987, Mills but is not now a problem in New England. 1989). Important avian predators are the However, proposals are pending to open at belted kingfisher (Megaceryle alcyon) and least three metal mines in Maine, which could common merganser (Mergus merganser) (El- have adverse effects on Atlantic salmon. son 1962). Human intervention in the introduc­ Atmospheric deposition of strong acid (acid tion or protection and enhancement of rain) has caused serious losses of Atlantic predators may reduce survival of salmon Parr. salmon in Norway, Sweden, and the United

Table 2. River miles not meeting wafer quality srandards by srate and cause. Source: EPA (1990).

Cause

Total Impaired Percenr Organic Slate Miles Miles Impaired Sill Nutrients Bacteria Enrichmenr

CT 8,400 298 3.5 12 163 176 156 ME 3 I.672 394 I .2 I50 I64 198 I91 MA 10,704 933 8.7 - 386 879 394 NH 14.544 381 2.6 - - 368 I31 RI 724 92 12.7 - 20 53 35 VT 5,162 628 12.2 465 327 238 388 Total 71.206 2.726 3.8 627 I.060 I.912 I .295

Cause

Suspended FIOH Habitat Stale Merals Pesticides Solids Salinirj, Alteration Modtjicarion PH Thermal

CT 98 - II - - III - - ME 3 - I4 - - - - - MA 169 20 41 I8 - - 5 - NH ------RI 74 ------VT 27 - - - 257 409 I9 474 Total 371 20 66 I8 257 520 24 474 136 STEMMING THE TIDE OF COASTAL FISH HABITAT LOSS

20 I 0

z 15 0 -

l­ a I­ 10 - a_ - 0 w 5 CK cl_

> A - 0 a APRSO JUNSO OCT90 DECSO n

7.0 B F 20 I 1 0 6.5 z 15 0 6.0 - t- a c­ 10 - a - 0 W 5 @L n

> 1 - 0 a APRSO JUNSO OCT90 DECSO n DATE

Figure 2. Temporal change in stream pH (solid line) and daily precipitation (vertical bars) in Sinclair Brook (A) and the upper (B). STEMMING THE TIDE OF COASTAL FISH HABITAT LOSS 137

Kingdom (ICES 1989, Rosseland et a!. 1986). 2-3s mortality (Ruggles 1980). Smelt passage In , acidification has caused the through electric generating turbines, however, elimination of salmon runs in I3 rivers in may cause mortality rates up to 40 or 50% , Canada, and reductions in the (Ruggles 1980). Semple and McLeod (1975) runs in an additional 18 rivers (Watt 1986). cite a mortality of 16.5% for Atlantic salmon Effects in the United States have not been so smelts at a hydroelectric facility in Nova drastic. The New England region does receive Scotia. acidic deposition (NADP !990), but most accessible Atlantic salmon habitat is not Summary chronically acidic (Haines 1980, 1987). Some streams tributary to mainstem salmon rivers As anadromous fish, Atlantic salmon do acidify episodically, depending on precipita­ require suitable freshwater habitat for a!! life tion and climatic conditions, axrd salmon parr stages from returning adult to emigrating populations may occasionally be reduced in smelt. The most significant habitat problems these streams (Haines et a!. 1990). For for salmon survival are dams, siltation of example, in 1990 the Upper Narraguagus stream substrate, and water pollution. As River reached pH levels as low as 5.3, whereas progress is made in restoring access of salmon Sinclair Brook, a tributary stream, reached to historic habitat, other problems in survival pH levels as low as 4.6 (Fig. 2). The passage of of salmon or reproduction may become more the new Clean Air Act in 1990 should greatly important. Stream acidity from acidic deposi­ reduce the risk of serious future losses of tion is not presently a serious problem in the Atlantic salmon in New England rivers. United States, but it could reduce salmon survival in some tributary streams. Global SmoIt climate change has the potential to affect Atlantic salmon survival seriously in fresh In New England, Atlantic salmon parr water by increasing stream temperatures and remain in freshwater for two to three years reducing flow. until they reach a size of 125 to 150 mm (5 to 6 in) total length (Bley 1987, Mills 1989). The Literature Cited transformation from the parr to smelt life Allan, l., and J. Ritter. 1975. Salmonid terminology. J. stage involves a complex mix of biochemical, Cons. Perm. lnt. Explor. Mer. 31293-299. physiological, morphological, and behavioral Beland, K., R. Jordan, and A. Meister. 1982. Water depth and velocity preferences of spawning Atlantic changes that prepare the fish for life in the sea salmon in Maine rivers. N. Am. J. Fish. Mgt. (Hoar 1979. Seaward migration occurs 2:1 l-13. primarily at night and is triggered by rising Blcy, P. 1987. Age, growth, and mortality of juvenile water temperatures, reaching a peak at 9 to Atlantic salmon in streams: a review. U.S. Fish 10°C (46 to 50’F) (Bley 1987, Ruggles 1980). Wild]. Serv. Biol. Rep. 87(4). 25 pp. Cooper, J., A. Scholz. R. Horral. A. Haslcr, and D. Predators of smelt, in addition to those listed Madison. 1976. Experimental conlirmation of the for Parr, include the eel (Anguillu rosrraru), olfactory hypothesis with homing, artificially im double crested cormorant (Pholucrocorax printed coho salmon (0. kimrch). J. Fish. Rzs. Bd. auritas), otter (Lutra lutra), and mink Can. 33:703-7 10. Dank. D., J. Trial, and J. Stanley. 1983. Species profiles: (Lurreolu vison) (Ruggles 1980). life histories and environmental requirements of The influence of dams on smelt emigration coastal fish and invertebrates (north Atlantic) is similar to that already described for Atlantic salmon. FWS/OBS-82/ I I. U.S. Fish Wildl. returning salmon. Because impoundments Serv., Washington, D.C. delay migration, they increase exposure of dcGraaf, D.. and L. Bain. 1986. Habitat use by and pref­ erencesof juvenile Atlantic salmon in two Newfound- smelts to predation and possibly effeci land rivers. Trans. Am. Fish. Sot. Il5:671-681. desmoltification (Saunders 1960). Passage of Dunfield, R. 1985. The Atlantic salmon in the history of smelts over dam spillways has been studied North America. Can. Sp. Pub. Fish. Aquat. Sci. 80, extensively for Pacific salmon in the north- I81 pp. Elson, P. 1962. Predator-prey relationships between fish- . Fish less than 20 cm (8 eating birds and Atlantic salmon. Bull. Fish. Ra. in) total length are seemingly able to withstand Bd. Can. 133.87 pp. vertical drops up to 90 m (300 ft) with only Elson, P. 1%7. Effect on wild young salmon of spraying STEMMING THE TlDE OF COASTAL FISH HABITAT LOSS

DDT over New Brunswick forests. J. Fish. Rcs. Bd. Intergovernmental Panel on Climate Change. Can. 24173l-767. Cambridge University Press, Cambridge, United Elson, P. 1975. Atlantic salmon rivers, smolt production Kingdom. and optimal spawning: an overview of natural pro­ ICES. 1989. Report of the study group on toxicological duction. In: J. Bohnc and L. Sochasky (cds.), New mechanisms involved in the impact of acid rain and England Atlantic Salmon Restoration Conference. its effects on salmon. C.M. 1989/M:4 Ref. E. Inter- Sp. Pub. 6:96-l 19. Int. Atl. Salm. Fed. St. Andrew% national Council for the Exploration of the Sea, N.B., Canada. Copenhagen, Denmark. Elson, P.. A. Mcistcr. J. Saunders, R. Saunders, J. Jordan, R. and K. Beland. 1981. Atlantic salmon spawn­ Sprague, and V. Zitko. 1973. Impact of chemical ing survey and evaluation of natural spawning pollution on Atlantic salmon in North America. In: success. Project AFS-20-R. Final Performance M. Smith and W. Carter (cds.), Proceedings of the Report. Maine Sea-Run Atlantic Salmon Commis­ international Symposium on the Atlantic salmon: sion, Bangor, Maine. Management, Biology and Survival of the Species. Kallcberg, H. 1958. Observations in a stream tank of ter­ Sp. Pub. 4(1):83-110. Int. Atl. Salm. Fed., St. ritoriality and competition in juvenile salmon and Andrews. N.B., Canada. trout (Salmo solar L. and S. rrurra L). Rep. 39:55- EPA. 1990. National Water Quality Inventory 1988. 98. Inst. Freshwat. Rcs., Drottningholm. Sweden. Report to Congress. EPA 4404-90-003. Environ­ Kerr, R. 1990. Global warming continues in 1989. Science mental Protection Agency, Washington, D.C. 247~52I. Frenettc, M., M. Caron, P. Julicn, and R. Gibson. 1984. Lewis, R. 1991. Can salmon make a comeback7 Interaction cntrc Ic debit et Its population de tacons Bioscience416-10. (Suitno sabr) de la riviere Matamec, Quebec. Can. J. Locke, D. and K. Havty. 1972. Effects of DDT upon Fish. Aquat. 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