Anadromous Brook Charr, Salvelinus fontinalis: Opportunities and Constraints for Population Enhancement

ROBERT J. NAIMAN, STEPHEN D. MCCORMICK, W. LINN MONTGOMERY, and RODERICK MORIN

Introduction Throughout most of their natural range, hancement and utilization in marine sys­ they enter salt water, where accessible, tems remains largely unexplored (Power, Success of enhancement programs de­ and grow rapidly while at sea. They have 1980). pends largely on the efficient use of natu­ well developed homing abilities, migrat­ The objectives of this article are to ral factors rendering a species suitable for ing seasonally at precise periods. At sea summarize studies initiated by Gibson a particular form of culture and manage­ they stay near their natal river, remaining and Whoriskey (1980) on anadromy in ment (Bardach et al., 1972). Most efforts in salt water for periods usually no brook chaIT, and continued by us, on the to enhance anadromous species have greater than 4-5 months. Within their na­ North Shore of the Gulf of St. Lawrence been directed toward commercial species tive freshwater range, they do not appear near Sept-lies, Quebec; to compare our with proven marketability. These spe­ to compete significantly with native results with those reported for other pop­ cies, however, are not necessarily the salmon or trout for space and resources. ulations; and to make recommendations most efficient in dealing with vagaries of Lastly, chaIT are popular recreational for enhancement of the fishery. For 5 the natural environment. Inability to con­ fishes and could have some commercial years we have examined the ecology of trol natural environmental factors intro­ potential. At present, brook chaIT are of anadromous and freshwater populations duces significant unpredictability into importance in recreational freshwater of brook chaIT, and have experimentally fisheries enhancement and aquaculture. fisheries and aquaculture, but their en- explored their growth and survival in sea­ Anadromous brook chaIT, Salvelinus water. Aspects of this study have been fontinalis, have some characteristics fa­ presented elsewhere on induction of vorable for enhancement and aquaculture Robert J. Naiman is Director, Center for Water anadromy and experimental sea ranching which will be reviewed in this article. and the Environment, Natural Resources Re­ (Gibson and Whoriskey, 1980; Who­ search Institute, University of Minnesota, Duluth, MN 55811. Stephen D. McCormick is riskey et aI., 1981a, b), parasites (Black, with the Department of Zoology, University of 1981; Black et aI., 1983), migration and California, Berkely, CA 94720. W. Linn Mont­ gomery is with the Department of Biological Sci­ ecology (Montgomery et aI., 1983), and ences, Northern Arizona University, Flagstaff, osmoregulatory physiology (McCormick AZ 86011. And, Roderick Morin is with the and Naiman 1984a, b, 1985a, b; Mc­ ABSTRACT-We present an overview of Department of Fisheries and Oceans, Artic Bio­ Cormick et al. 1985). These studies, studies conducted during 1978-84 on logical Station, Ste. Anne de Bellvue, Quebec H9X 3R4, Canada. combined with previous ecological in­ anadromous brook charI', Salvelinus fonli­ natis, near Sept-lies, Quebec, Can. Studies on experimental sea ranching are inte­ grated with those on the biology ofanadro­ mous stocks, results are compared with those reported for other anadromous and nonanadromous stocks, and recommenda­ tions are made for population enhance­ ment. Anadromous brook charI' exhibit short but synchronized migratory move­ ments, can enter brackish warer at a small size, remain in coastal waters, grow rapidly while at sea, home effectively, and provide a potentially high yield to recre­ ational and commercial fisheries. We sug­ gest that anadromous brook charr, and possibly other members of the genus Salvetinus, are reasonable candidates for local enhancement of coastal or estuarine salmonid fisheries and for sea ranching in northern latitudes.

49(4), 1987 1 feeding on a wide range of organisms,

50°20' depending upon their size and the size of D available prey. Although most populations of brook charr are restricted to fresh water, many coastal rivers in their native territory of northeastern North America contain RIVIERE a LA TRUITE anadromous brook charr. In northern lati­ tudes, brook charr migrations are charac­ terized by spring emigrations of 2 to 4­ year-old fish whose coastal sea residence lasts 2-4 months (White, 1940; Wilder, 1952; Dutil and Power, 1980; Cas­ tonguay et aI., 1982). In the southern portion of its range, the timing of emigra­ tion is more variable, often occurring in

o 2 3 4 the fall (Mullan, 1958). km Assessment of Potential for Enhancement Initial Experiments Figure I.-Location of principal study sites. An experiment to induce anadromy in a population of nonanadromous wild brook charr was conducted in 1978 and 1979 near Sept-Iles. This area is a sub­ arctic boreal forest biome (mean annual Table 1.-Physiochemical characteristics of the study sites. temperature = 1°C) composed of black Parameter Rivieire a la Truite Matamek River Moisie River spruce, Picea mariana; white spruce, P. Stream order 4 6 9 glauca; and balsam fir, Abies balsamea. Watershed area (km2) 36 673 19,871 Mean width (m) 15.6 51.7 208.7 The rivers are ice covered from about Mean depth (m) <1.0 1.8 25 November to April. There is a strong Mean annual discharge (m3/sec.) 1.5 13.7 466.1 Substrate Gravel Gravel/cobble Sand/cobble freshet in April and May when -50 per­ Temperature range (0G) 0.1-16.0 0.1-220 0.1-21.8 Annual degree days (oG/year) 2.000 2.225 2.219 cent of the annual discharge occurs. The pH range ? 4.8-6.0 6.3-7.1 aquatic growing season is short (-105 Alkalinity (as GaGa in mg/L) ? 0-0.5 0-2.1 Forest canopy development Mostly open open open days), and the rivers accumulate only about 2,100 degree-days annually (Table I). Brook charr from two river systems were chosen for study: An anadromous vestigations of freshwater brook charr in cessfully throughout the world. Brook population from the lower Moisie River the same watersheds and with informa­ charr inhabit well-oxygenated streams and its tributary Riviere it la Truite, and a tion from the literature, provide the basis and lakes, preferring temperatures below freshwater population from the Matamek for evaluating the opportunities for, and 20°C. They spawn in late summer or au­ River (Fig. I). The Matamek River popu­ constraints upon, the population en­ tumn, the date varying with latitude and lation does not migrate to sea due to a hancement of anadromous brook charr. temperature, over gravel beds in head­ large waterfall, 0.7 km from the ocean, water streams and rivers. Mature fish blocking upstream movement to suitable General Life History may travel a considerable distance up­ spawning areas, thus precluding anadro­ Brook charr are endemic to North stream to reach the spawning grounds. mous habits_ The three rivers differ sig­ America, distributed under natural condi­ Unlike lake charr, Salvelinus namay­ nificantly in physical dimensions but not tions from the Atlantic provinces south to cush, and artic charr, Salvelinus alpinus, in most chemical characteristics (Table I; Long Island, N. Y., in the Appalachian brook charr are short-lived with wild in­ Naiman et aI., 1987) or seasonal timing Mountains south to Georgia, west in the dividuals seldom attaining 5 years of age_ of environmental events (Naiman, 1982, upper Mississippi and Great Lakes Sexual maturity is usually reached at age 1983). drainages to Minnesota, and north to 3, but some individuals may mature at Initially, nearly 1,800 brook charr, age Hudson Bay (Scott and Crossman, age 2. Maximum size is about 5 kg. In I+ to 5+, were captured from the 1973). They have been introduced suc­ general, brook charr are carnivorous, Matamek River (identified with serially

2 Marine Fisheries Review numbered Carlin tags if > 10 cm fork variously with temperature, river dis­ 1978 1979 SEA-RUN RIVER SEA-RUN RIVER length (FL) or fin clipped if ~ 10 cm FL), charge (particularly spring floods), lunar CHARR CHARR CHARR CHAAR transported to the Matamek River estuary cycles, tides, and migrations by other from late May to early July, and released species. In the Moisie River the duration (Gibson and Whoriskey, 1980; Who­ of emigration appears to vary with the riskey et ai., 1981 b). The mean number pattern of discharge. In 1980, when dis­ of days at liberty for different age classes charge rose to an early maximum and ranged from 63 to 98. Fish were recap­ then declined steadily, the migration was tured in autum (late August and Septem­ largely completed within 2 weeks; in ber) as they returned to fresh water. Over 1982, when rains caused three secondary 2 years, 34.0 percent of the released fish discharge peaks following the early June 3.5 were recaptured. Best returns were in the maximum, the migration was extended 2+ and 3+ age classes with 38.0 and over 5 weeks (Montgomery et ai., 1983). 62.1 percent recaptured, respectively. Movements of brook charr into cool, Movement of transplanted fish to other spring-fed creeks of Cape Cod, Mass., nearby rivers was < I percent; predation seemed to correlate with high summer by birds and other fish was not evaluated. temperatures of estuarine waters (Mul­ All age classes included sea-run brook lan, 1958). Migratory activity of brook charr, but the largest percentages of sea­ charr on the Gaspe peninsula, Quebec, run charr occurred in older and larger correlates with lunar cycles (Castonguay AGE CLASS fish. Growth was substantially better in et ai., 1982). Migrations of Moisie River Figure 2.-Growth, expressed as sea-run charr than in fish remaining in charr to coastal waters coincide with percentage increase of body freshwater (Fig. 2), presumably a func­ simultaneous movements of several other weight, and mean daily growth rate are shown by age group for tion of increased food supply at sea. Tag­ species (Montgomery et aI., 1983), sug­ tagged brook charr for 1978 and ging severely stunted growth, especially gesting that environmental cues may be 1979. The 1978 sea-run charr in younger and smaller fish, and probably similar for all species. Water tempera­ data are calculated from Gibson suppressed anadromy. Nevertheless, tures in the river and the sea are similar at and Whoriskey (1980). Vertical lines above bars represent one tagged sea-run charr grew between 0.8­ this time (-8-1O°C), reducing the ther­ I S.E.; numbers are the sample 3.5 percent d- , which was clearly supe­ mal barrier to migration. size. Negative values result from rior to tagged fish in the freshwater popu­ Although it is apparent that move­ handling and tagging stress. lation (0.4 percent day), and they had ments are often rapid and directional, de­ Adapted from Whoriskey et al. greater condition factors than their fresh­ tails of migratory behavior are sparse. (l981b). water counterparts. Gonadal maturation Our own studies (Montgomery et al. 1) was suppressed in sea-run charr com­ are typical, involving capture, marking pared with fish remaining in fresh water, and recapture at a single trap site in the and there was a significantly larger per­ river and at several locations in the estu­ centage of female sea-run charr. ary. In the Moisie River system, one fish days if movements were strongly direc­ These results suggested that an inte­ travelled from Riviere a la Truite to the tional. grated research program could demon­ estuary (-15 km) in <2 days, while sev­ Data on total distances traversed by strate the viability of this technique eral others were recaptured in the estuary migrants are similarly tentative. Records where moderate enhancement of native after < 10 days. Anadromous arctic charr indicate that anadromous brook charr salmonid production was desired. In and whitefish, Coregonous sp., in the move 30-50 km upriver in the Moisie 1980 we begun a program investigating coastal Beaufort Sea have a net move­ River (MacGregor, 1973), and Cas­ movements, feeding, growth, survival, ment of 3-6 km d- I along the shore tonguay et ai., (1982) sighted schools of production, reproduction, and physiolog­ (Craig, 1984). Sockeye salmon, Oncor­ anadromous brook charr 48 km upstream ical adaptation to seawater for wild hynchus nerka, smolts travel 10-15 km from the mouth of the St. Jean River, anadromous brook charr. d- I in Babine Lake, B.C., and at much Quebec. Moore (1975) found migratory Movements greater rates in rivers (Brett, 1983). It Arctic charr a maximum of 50 km upriver seems unlikely that the trip to the Moisie on Baffin Island, N.W.T. Anadromous northern populations of River estuary would require more than 2 Studies indicate that brook charr at sea brook charr tend to leave streams and remain close to their natal rivers. Cas­ move to estuaries and coastal marine tonguay et al. (1982) recorded only two waters immediately after the peak in the IMontgomery, W. L., R. J. Naiman, S. D. Mc­ Cormick, F. G. Whoriskey, and G. Black. Biol­ strays to adjacent rivers 19 and 45 km spring freshet (White, 1940; Wilder, ogy of anadromous brook charr (Salvelinus fOn/i­ from the tagging location in the St. Jean 1952; Dutil and Power, 1980; Cas­ nalis) in the Moisie River, Quebec. Unpub!. River. During 2 years of experimental manuscr. W. L. Montgomery, Department of tonguay et ai., 1982). Nevertheless, Biological Sciences, Northern Arizona Univer­ tagging only three recaptures « I per­ movements to and from rivers correlate sity, Flagstaff, AZ 860 II. cent of total recaptures) were made away

49(4), 1987 3 from the tagging site in the Matamek Table 2.-Comparisons of brook charr feeding by the percent volume of taxa in stomach contents for regions of the Matamek River system. Indicated taxa are major components of diet. Streams are presented River estuary (Gibson and Whoriskey, in Increasing order of size. Fish were sampled in June and July.

1980; Whoriskey et aI., 1981 b). All were Stream Lake Estuary recaptured from an intense recreational Taxa/group S' G T M5 R B M E fishery in the Moisie River and had No. fish 28 51 32 111 9 24 29 18 moved 12-39 km. White (1941) and Fork length (em) 10-15 10-15 10·15 8·18 10-15 15-20 10·20 15 Smith and Saunders (1958) took mosr of Zooplankton 19 their recaptures within 5-8 km of the river Insects mouths. Fish generally moved <10 km Simuliidae. 12. p 4 2 14 16 Chironomidae, I. p 1 6 11 from tagging sites in Richmond Gulf Chaoborus. I 25 (Dutil and Power, 1980). The brook Total Diptera. I, p 11 26 19 25 6 trout's square tail with a low aspect ratio Trichoptera 40 27 12 36 9 4 14 is not designed for efficient high speed Ephemeroptera 4 11 5 5 16 45 36 Pleeoptera 6 4 14 3 2 swimming; the more slender somewhat forked tails of other charr are better Total insect 55 53 57 63 50 55 59 3 adapted for long migrations and pelagic Terrestrial insects 20 35 52 7 16 2 cruising (Power, 1980). This informa­ Amphipoda 45 tion, in concert with the observations by Fish 30 5 46 White (1942) that brook charr wander in 'Abbreviations and data sources: S = Sherry Creek. G = Gallienne Creek. T = Tehinieamen (O·Connor. 1974); schools in shallow waters along shore, M5 = Matamek River below 5th Falls (Gibson. 1973); R = Lake MacRae (Carseadden. 1970); B = Lake Bill suggests that anadromous brook charr re­ (O'Connor. 1971); M = Lake Matamek (Saunders. 1969). and E = Matamek River estuary (Whoriskey et ai.. 1981b). main inshore and close to their river of 21 = larvae. p = pupae origin. As in arctic charr (Craig, 1984) the tendency of brook charr to remain close to home rivers may be influenced by verse prey as they groW. Cladocera and by small aquatic insect larvae and terres­ pockets of brackish (5-28%0) and rela­ copedoda constituted 9-31 percent of the trial or aquatic insect adults, both of tively warm (8°-13°C) water along the food volume in June of emerging brook which have considerable refractory coastline during the open-water season. trout fry from the Matamek River exoskeletal materials. In sea water, This estuarine band varies in width and (Williams, 1981). Dipteran larvae, brook trout feed mainly on fleshy foods depth with freshwater input to coastal mainly simuliidae and chironomidae, such as fishes (particularly sand lance, waters, prevailing winds, and topo­ constituted 42-67 percent of the food vol­ Ammodytes sp.), mysids, and amphipods graphic features. In shallow waters (2-3 ume. In July and August the diet shifts to (Table 2; Whoriskey et al. 1981 b; Mont­ m) the entire water column is usually organisms captured at the water surface, gomery et al. I). Other dietary studies of brackish; in deeper waters a two-layered including both winged aquatic insects brook charr in salt water generally report system develops, with a lens of surface and terrestrial insects (Morin et al., 1982, piscivorous feeding (White, 1942; brackish-water overlying a wedge of and unpublished data). Greendale and Hunter, 1978; Dutil and colder marine water. A variety of factors The diets of juvenile and adult brook Power, 1980). White (1940, 1942) re­ (e.g., wind, topography) influence how charr differ between stream, lake, and es­ ported that brook charr in the Moser far the warm, brackish-waters and the tuarine environments (Ricker, 1932a, b; River estuary relied heavily on young brook charr extend seaward, but in most Carlander, 1969). These differences in eels and saltwater isopods, while am­ cases we have observed the seaward limit feeding are reflected in Table 2 based on phipods, nereid worms, and freshwater is < 10 km (Naiman et al., personal ob­ studies throughout the Matamek River insects constituted the remainder of the servation) . system. Diptera larvae and trichoptera diet while those in salt water consumed at are major components of diets in streams. least seven species of marine fishes. Feeding In 3rd to 5th order streams (Strahler, Differences between fresh and salt­ Brook charr are visual, opportUniStic 1957), brook charr of all sizes consume water feeding also relate to the abun­ predators, whose food habits change in mainly terrestrial invertebrates at the dance and seasonal availability of prey. relation to seasonal prey abundance, for­ water surface, particularly in late sum­ Studies in Quebec indicate that resident aging locality, and effects of agonistic mer. In large streams such as the Moisie fish in rivers feed regularly or heavily interactions with other fishes (Power, River, brook charr are more piscivorous only during the 6-week mid-summer pe­ 1980). When brook charr fry emerge and depend less upon organisms at the riod of insect growth and emergence from the spawning gravel, they initially water surface (Montgomery et aLI). (Gibson and Galbraith, 1975). In con­ feed on minute drifting prey (Williams, Prey consumed by brook charr in sa­ trast, anadromous brook charr have ex­ 1981; Walsh et aI., In press). Subse­ line waters differ in size and probably tensive food reserves available through­ quently, they feed during daytime along nutritional value from those in fresh­ out their residence in coastal waters. stream edges, consuming increasingly di- water. Freshwater foods are dominated Whoriskey et al. (1981 b) found substan-

4 Marine Fisheries Review Table 3.-Comparlson of age-weight and growth increments for freshwater and adJa­ cent estuarine populations of brook charr. Weight is shown In grams with instanta­ A . 6. N.S. .. ~ neous weight increments In parenthesis. • Mean weight and .. 6. 6. instantaneous weight increment 6. 6.6. 6. • • Location Lat. 2+ 3+ 4+ 5+ · •• • 0 Richmond Gulf. Quebec' 56'N 6. River 19 (0.8) 42 (0.5) 71 [5TIMATED· ACE5. Estuary 51 (1.3) 181 (0.8) 399 (0.5) 647 ,. 2· 3· , 4· Sept-lies. Quebec2 50'N . B Matamek Lake 22 (1.0) 59 (0.6) 110 (0.5) 188 v·1.50-0.01)( 221 Matamek River 42 (0.7) 85 (1.0) e.- • 6. 6. Matamek River estuary 66 (0.7) 126 (0.6) 227 • 6. Maisie River estuary 160 (0.8) 346 (0.7) 718 . 6.6. 6. 0 3 6. 6. • Moser River, Nova Scotia 45'N · 6. •• River 62 (0.4) 90 (0.7) 174 (0.6) 309 • .. Estuary 129 (0.7) 249 (0.3) 348 (0.6) 657 · 3.6 'Data from Dutil and Power (1980); length converted to weight using. for river fish. a . • " condition factor of 1.1 '~ 3.2 C 2Data from Saunders and Power (1970). Whoriskey et al. (1981b). and Montgomery et al. ~ 2.8 (text footnote 1). • ~ 2.4 6. 3Data from White (1940). 6.6. • i:: 2.0 6. 6. 6. it 1.6 • 6. .. <::> • 6. ~ 1.2 .. . y • 0.02x - 1.49 ).... 0.8 • • :::: 0.4 '< 0·. 6. t:;:) 0.0 tial numbers of amphipods and sand water chaIT populations. In extensive re­ 120 140 160 180 200 220 240 lance in the Matamek River estuary, and views of the growth of brook chaIT, FORK LENGTH (mm) both foods were available in great abun­ Bigelow (1963) and Carlander (1969) re­ Figure 3.-Growth rate, by weight dance in the Moisie River estuary from ported that sea-run chaIT attain a maxi­ (A) and length (8), of anadromous brook chaIT in the Moisie River are April to October (Montgomery et al. \ mum weight of about 4.5 kg, no greater presented as a function of initial size Flesh color of anadromous chaIT re­ than their maximum size reported from and age. Average daily growth (C) is flects food habits, and may serve as an freshwater. Unfortunately, no data were presented as a function of fork length indicator of perceived quality for human available on the age of such fish. In gen­ and age. The relationship between consumption. Increasing age of sea­ eral, growth data are sparse for larger and growth in weight and fork length (A) is not significant (P > O. I; ranched chaIT from the Matamek River older brook chaIT (> I kg, >6 years of r = -0.28), but the relationships was accompanied by increasing fre­ age) and for southern populations, partic­ between growth in length (8), daily quency of yellow or pink flesh and COITe­ ularly the anadromous populations that growth (C), and fork length are sig­ lated with increased dependence on once inhabited Long Island, N. Y. and the nificant (P <0.05; r = -0.58 and marine foods (Gibson and Whoriskey, large New England rivers (Power, 1980). 0.71, respectively). Circles = 1+, dots = 2+ , triangles = 3+ , and dia­ 1980). All I + aged fish, 78.3 percent of However, instantaneous growth incre­ monds = 4+. the 2+ , 57.4 percent of the 3+ , and 11.8 ments between age classes (Table 3) are percent of the 4+ fish had white flesh. In highest for anadromous brook chaIT of the 2+, 3+, and 4+ age classes the per­ Hudson Bay and the northern Gulf of St. centages with yellow flesh were, consec­ Lawrence. These regions have shorter and weight declined with increasing ini­ utively, 3.1. 7.0, and 5.9; with pink growing seasons and colder marine con­ tial size and age (Fig. 3B), although ab­ flesh, 14.9, 28.7, and 58.8; and with ditions than in the more southerly range solute rates (g/day) increased signifi­ deep pink flesh, 3.7, 7.0, and 23.5. of brook chaIT. The high growth rates in cantly (P >0.01) with length (Fig. 3C). Larger specimens of anadromous chaIT this area, however, may be an artifact of These data provide a conservative evalu­ from the Moisie River also had colored size and age since northern freshwater ation of growth since sampling con­ flesh, although no quantitative data were populations are smaller at any given age straints prevented study under fully collected. Arctic chaIT with red flesh than southern populations. An increase in marine conditions. The daily growth have the greatest market demand in growth rate (due to anadromy) will result rates from the Moisie River study com­ Labrador, followed by chaIT with pink in greater growth rates of smaller fish and pare favorably with results from the flesh; chaIT with white flesh are only used greater weight increments at any age. Matamek River, and are significantly locally (Andrews and Lear, 1969). Growth rates were calculated in 1980 greater than growth rates in fresh water for tagged Moisie River brook chaIT at Growth and Survival (Saunders and Power, 1970; Gibson, large for more than 30 days between tag­ 1973). Tagged brook chaIT introduced to Brook chaIT at sea exhibit enhanced ging and recapture, as growth was sup­ the Matamek River estuary grew at 0.8 ­ growth typical of anadromous salmonids. pressed during shorter intervals by the 3.5 percenUday by weight while tagged Table 3 illustrates this pattern by compar­ tagging procedure (Montgomery et al. l ). brook chaIT remaining in freshwater grew ing growth in adjacent sea-run and fresh- Rates of growth relative to both length at 0.4 percenUday (Fig. 2).

49(4), 1987 5 Several studies (Wilder, 1952; Dutil tagged and released; 557 were 101-150 ulation production. and Power, 1980; Whoriskey et al. mm FL, 251 were 151-200 mm FL, and Despite the apparent abundance of 1981 b; Castonguay et aI., 1982) indicate 156 were 201-350 mm FL. The effi­ flowing waters in northern watersheds, that, at least for some age or size classes, ciency of the fishery (IJ..) was estimated to only a small fraction of those waters actu­ brook charr experience enhanced growth be 22 percent for 101-150 mm fish, 45 ally contribute to fish production, mak­ through anadromy by feeding on marine percent for 151-200 mm fish, and 31 per­ ing the marine phase of brook charr life foods. Exceptions to this trend are usu­ cent for all sizes. Survival rate(s) be­ history even more significant. Fish are all y found in younger (I +, 2+) or tween years was estimated to be 31, 20, absent from first and second order smaller fish (Fig. 2, 3A, B, C). Moisie and 25 percent for the three size groups, streams and sparse in third order streams River brook charr consistently enter sa­ respectively. on the North Shore of the Gulf of St. line water after they attain a length of Estimates of survival for the sea-run Lawrence (Naiman et aI., 1987; Morin l 2: 15 cm FL (Montgomery et al. ). Virtu­ population of the Moisie River compare and Naiman, unpubl. data). These ally all growth rates for the smaller 1+ favorably with data from unexploited streams are subject to nearly complete and 2+ fish are below means predicted freshwater charr populations in the freezing in winter and experience sub­ from rates determined for fish 2: 15 cm Matamek watershed and with seasonal stantial variations in flow and tempera­ FL. This indicates that size classes which return rates for sea-ranched char in the ture during summer. First to third order remain in essentially fresh water, despite Matamek estuary. Gibson (1973) esti­ streams constitute 23 percent of the total being in the estuary, fail to attain the ex­ mated minimum survival to be 46 and 16 surface area of streams in the Moisie pected rapid rate of growth. In addition, percent based on recovery of brook charr River system (Naiman, 1983), but have fish ::s9 cm FL are rarely captured in the marked in a previous year at two sites in negligible fish production. Significant estuary, although they are common in the lower Matamek River. The lower es­ fish production in the Matamek system the river. These apparent limitations in timate (16 percent) was attributed to a occurs only in streams 2:4th order size at migration correspond to the greater proportion of tagged yearlings (O'Connor and Power, 1976). These known size limitations of brook charr which were thought to incur higher mor­ constitute 70 percent of the stream sur­ salinity tolerance (discussed below). tality during winter. O'Connor and face area in that system, but even this is Considered together, these data indicate Power (1976) estimated survival from an overestimate of productive habitat. that 0+ fish remain in the river, while year-class structures of four stream popu­ Given the depth distribution and habitat larger I+ and older animals move to the lations of brook charr in the Matamek requirements of fry and yearling, signifi­ estuary and constitute the sea-run compo­ watershed. Survival rates between suc­ cant charr production is possible in <40 nent of brook charr in anadromous popu­ cessive year classes of charr aged I year percent of the total lotic surface area. In lations (Montgomery et al.; I Whoriskey and older ranged between 17 and 37 per­ short, these northern river systems can­ et aI., 198Ib). cent. In our induced anadromy experi­ not normally support highly productive The survival of anadromous brook ments, minimum recovery rates over salmonid populations; anadromous popu­ charr has not been extensively studied. 2 years were 29 and 40 percent for charr lations circumvent to some degree the However, a number of sources suggest released in the Matamek River estuary limitations of food, space, and tempera­ that it may be favorable in comparison (Gibson and Whoriskey, 1980; Who­ ture (Gibson and Galbraith, 1975; Gib­ with freshwater populations. Saunders riskey et aI., 1981 b). son et aI., 1976; 0'Connor and Power, and Smith (1964) reported that yearling 1976; Walsh et aI., In press) during criti­ brook charr stocked in estuaries yielded Production cal mid-summer periods by moving to an average return to anglers of 28 percent The growth attained in our induced coastal areas. compared with an average of 13 percent anadromy project is particularly interest­ To evaluate the viability of an en­ for yearlings and 4 percent for younger ing since growth and production in the hancement program, it is essential to charr stocked in the stream. Dutil and Matamek River drainage network are know, first, the biological productivity of Power (1980) recaptured 56 percent of among the lowest reported throughout anadromous brook charr populations and tagged fish during studies in Richmond the geographic range of brook charr second, how much of that production can Gulf, and White (1941) reported returns (Saunders and Power, 1970; O'Connor be successfully harvested. Neither deter­ of 9-40 percent for different size classes and Power, 1976; Power, 1980). Charr mination has been made for an anadro­ of sea-run brook charr in the Moser production in lakes of the Matamek River mous brook charr population. Neverthe­ River, Nova Scotia. watershed ranges from only 0.5 to 2.2 less, given the observed growth rates of Survival of sea-run charr of the Moisie kg/ha/year (Carscadden, 1970; Saunders brook charr aged 2+ to 7+ in the Moisie River was estimated from tagged fish at and Power, 1970; O'Connor and Power, River estuary, production of those year large at the end of 1980, and recaptured 1973) and in streams leading into classes in freshwater populations (e.g., over the following 3 years by anglers Matamek Lake it is 14.5-66.4 kg/ha/year O'Connor and Power, 1976) could ap­ (Montgomery et al. I). Calculations were (O'Connor and Power, 1976). The proximately double if they moved to sea made according to Ricker (1975) assum­ growth of charr in the Matamek River is and incurred similar mortality. Such an ing constant angling pressure between most rapid during the first 2 years of life, estimate is based only on the difference years. Nine hundred sixty-four fish were accounting for 70-90 percent of total pop­ between sea and freshwater growth and

6 Marine Fisheries Review does not include possible increases in re­ Table 4.-Age-specific maturity for Matamek River brook charr from the Second and Fifth Falls, and from the Moisie River. Values in percent; M = male, F = female. Numbers In parentheses are sample sizes. cruitment by anadromous chaIT due to in­ Age creased growth, fecundity, and spawning frequency. Measures of yield from north­ 1+ 2+ 3+ 4+ ern commercial and native fisheries on Site M F M F M F M F Matamek River 1 anadromous populations of Atlantic Second Falls 0 (5) 29 (17) 29 (17) 73 (26) 91 (32) 50 (12) 100 (6) salmon, Arctic chaIT, brook chaIT, and Fihh Falls 41 (49) 0 (29) 37 (43) 81 (37) 69 (13) 100 (24) 88 (8) whitefish (Power, 1966, 1980; Power Maisie River2 and Lejeune, 1976; MacCrimmon and Anadromous only 20 (10) 0 (4) 50 (20) 52 (25) 80 (5) 72 (18) 100 (3) 100 (5) Freshwater and Gots, 1980) also suggest that this esti­ anadromous 10 (197) 3 (163) 33 (93) 27 (94) 67 (6) 67 (21) 100 (3) 100 (5) mate of potential for enhanced produc­ 'From Gibson et 01. (1976). tion is conservative. 2From Montgomery et al. (text footnote 1). Reproduction Despite extensive observations, we were unable to detect any significant dif­ ferences in choice of spawning site, re­ factors such as food, habitat, population of brook chaIT. Only the anadromous productive behavior, fertility, early on­ density, temperature, and salinity, may stock from the Koksoak River, Ungava togeny, or early life history between act indirectly (through size or growth Bay, Quebec, seems to increase fecun­ anadromous and nonanadromous stocks rate), or directly, on the onset of matura­ dity more rapidly than other populations, of brook chaIT (unpublished data). Power tion. In highly productive streams, brook and that only occurs at lengths >40 cm (1980) has described these reproductive chaIT mature as early as I + or 2+ years FL. However, when slopes of fecundity­ aspects in detail. Our discussion here of age; in northern or unproductive rivers weight regressions are compared for centers around maturation schedules, fe­ maturation begins at ages >3+ (Power, freshwater and anadromous brook chaIT cundity, and life history strategies. 1980). Relati vely late maturation also in the same region, the slopes for Moisie While age is often reported as a COITe­ characterizes anadromous brook chaIT River fish (2.8 in 1972, in MacGregor, late of maturation, other factors also in­ (White, 1940; Castonguay et aI., 1982), 1973; 3.0 in 1980, Montgomery et al. I) fluence its onset. Under laboratory or and anadromous and experimentally exceed those reported for fish from lakes hatchery conditions, size is a more im­ transplanted brook chaIT show signs of and streams in the Matamek River portant determinant of maturation in delayed gonadal development upon drainage (1.7-2.4; Fig. 4). Thus, anadro­ brook chaIT than either age or growth reentering freshwater (White, 1940; mous chaIT may gain an advantage in rate, explaining 97-99 percent of the vari­ Whoriskey et al. 198Ib). Dutil and terms of increased fecundity beyond that ation in maturation (McCormick and Power (1980) observed that final ripen­ predicted from simple increases in body Naiman, 1985b). Under accelerated ing of gonads of anadromous brook chaIT size (Fig. 4). growing conditions, brook chaIT mature occurs in freshwater. Freshwater brook charr in northern re­ and spawn in their first autumn, at an age Reproduction by anadromous brook gions have different life history strategies of 10 months and a weight of 60 g (Carl­ chaIT in the Moisie River system is than anadromous stocks (Power, 1980). son and Hale, 1973; McCormick and largely delayed until the fish attain age In northern streams the fish grow slowly, Naiman, unpubl. data). Other physiolog­ 2+ and return to spawning tributaries at not all fish mature at the same age, and ical features (e.g., nutritional state and lengths > 15 cm FL (Montgomery et there is the possibility of multiple spawn­ health) also influence time of maturation. aLI). Females lagged behind males in at­ ing so that the reproductive potential of a Schedules of maturation normally differ taining maturity (Table 4). Similar pat­ year class is spread over more than one between sexes, with the proportion of terns were observed in Moisie River fish year (Fig. 5A). Sex ratios shift in favor of mature males greater at small sizes than that had not been tagged. It is surprising females in the older age groups. A few that of mature females (Carlson and that maturity schedules for tagged fish males are long lived, and the oldest fish Hale, 1973; Power, 1980; McCormick from the Moisie River are comparable to in a population may often be a large male and Naiman, 1985b). This difference is those for tagged Matamek River chaIT (Power, 1980). characteristic of many fishes and other (Gibson et aI., 1976), as maturation was With the addition of a migratory phase poikilothermic vertebrates, and may re­ much lower in tagged Matamek River to the life cycle and a separation of late to the nature of mating systems and fish of the same age (Whoriskey et al., spawning and nursery areas from feed­ to differential energetic investment in re­ 1981 b). The only clear difference be­ ings grounds, a life cycle of the type in productive products (Bell, 1980). tween the two populations was that Fig. 5B is found in anadromous popula­ Brook chaIT mature over a wide range Matamek River females tended to lead tions. Anadromous migration provides of sizes and ages. This probably results males in percentage maturation at ages more living space and results in better from genetic differences between stocks greater than I + (Table 4). growth, possible lower mortality, larger combined with phenotypic responses to Power (1980) found little difference in fish, and better fecundity. Annual environmental variation. Environmental interpopulation weight-specific fecundity spawning usually occurs once maturity is

49(4), 1987 7 120

100 "' ~ ti eo <:: '­ ~ ~ 60 ~ ij~,,_~s ~ ~ 40 ~~~ex Matamek Lake & Lake MacRae 20 Malamek River 2 nd Foils & 5 th Falls

Fork Lenqfh (em)

Figure 4.-Size-related increases in NORTHERN UNEXPLOITED STREAMS SOUTHERN AND INTERMEDIATE fecundity for Quebec brook charr. AREAS. ANADROMOUS All intercepts of linear regression equations for fecundity on FL have been adjusted to zero. Projection of Figure 5.-The life history strategy of nonanadromous brook charr in northern lines on abcissa gives the size range streams is contrasted with that of an anadromous stock in the same area. Adapted of fish used for regression. Data from Power (1980). sources are: Moisie River, 1972 (MacGregor, 1973); Moisie River, 1980 (Montgomery et aI., text foot­ note I); Lake X and Lake MacRae in Matamek River watershed McCormick and Naiman (I 984b) (O'Connor, 1971); and Matamek demonstrated that survival of brook charr River (Gibson et aI., 1976). '­ '­ between 6.0 and 32.0 cm FL in 32%0 sea­ '­ ).. '-,­ .... 2 ""­ 400 ~ water was size dependent (r = 0.77), as ',t~ 0.64 y·465.7-3.27~ i':i was the ability to regulate plasma ions ...... ~ 2 ""­ C> (Na+, Cl-, K+, Mg+ ) and plasma os­ ""-""­ '­ ~ molarity (Fig. 6). Age, by itself, had no attained. Multiple spawning is normal so significant influence on seawater survival ;--'--"':;;------;,~S ---:2'::-0--:2~S--:''::-0---::,,~ 350 it'" FORK LENGTH (em) that the reproductive potential of a year of brook charr. Size dependent hypoos­ class is spread over 2-3 years. In combi­ moregulation may be the result of more Figure 6.-Size dependent survival nation, these factors make anadromy an favorable surface area-to-volume ratios after 20 days (-) and plasma osmo­ important alternate life history strategy in larger fish. Although seawater survival larity after 4 days (-) of exposure to for brook charr with oceanic access. increases throughout all sizes tested (6.0­ 32%0 seawater. Seawater hazard rate is a measure of the probability of Physiological Adaptation 32.0 cm FL), fish> 15 cm FL achieved a survival (number of deaths/days at to Seawater near-maximum rate of survival. risk), with survival rate increasing as In addition to ontogenetic changes as­ hazard rate approaches zero. Data Movement of brook charr from fresh sociated with size, hypoosmoregulation originally presented in McCormick water to salt water requires not only a is also affected by gonadal maturation of and Naiman (l984b). behavioral (migratory) impetus but also a males (McCormick and Naiman, 1985a). physiological tolerance to salinity. The Seawater survival of mature males is latter depends primarily on the ability to lower than that of mature females or im­ 2 regulate plasma ions in a hyperosmotic mature fish of similar size in spring, sum­ Mg+ ) and osmolarity in seawater during medium. For a variety ofteleosts, includ­ mer, and autumn (mature brook charr autumn is significantly reduced relative ing salmonids, salinity tolerance and hy­ were not tested in winter). Furthermore, to that of mature females. Mature fe­ poosmoregulatory ability have been mean survival time of mature males in males and immature brook charr of both shown to change as a function of size and seawater is reduced by >50 percent in sexes exhibit no such decline in autumnal life history stage (Parry, 1958; Conte and autumn during the normal spawning pe­ seawater survival. Wagner, 1965; Conte et aI., 1966; riod. The ability of mature male brook The decreased salinity tolerance of Nordlie et aI., 1982). charr to regulate plasma ions (Cl- and males during gonadal maturation signi-

8 Marine Fisheries Review Table 5.-Phylogenlc comparison of slze-dependent salinity tolerance and seaward Method of seawater acclimation also migration In salmonlds. affects survival. Brook charr acclimated Salinity tolerance Size al Size at 75% seaward for 1-2 weeks at intermediate salinities Salinity Method of Duration survival migration Refer­ (10-25%0) before transfer to 32%0 sea Species (ppt) acclimation (days) (em) (em) ances water had only 18 percent mortality after Oncorhynchus Pink salmon 32 Direct >14 <6.0 2.5-4.0 (1.2) 20 days vs. 85 percent mortality for those Chum salmon 32 Direcl >14 <6.0 3.2-7.0 (1.2) transferred directly (McCormick, un­ Chinook salmon 30 Direcl 30 6.5 6.0-10.0 (3.4) 30 Gradual 30 4.2 pubI. data). This is in agreement with our Coho salmon 30 Direct 30 7.0 10.0-11.0 (5.4) observations that brook charr may spend Salmo up to a month in an estuary before migrat­ Atlanlic salmon 29 Direct 44 13.8 12.7-15.2 (6,2) Steelhead lroul 30 Direct 30 16.0 15.0-20.0 (7,8) ing to sea (Montgomery et al. I), and that at estuarine sites characterized by high Salve/inus Brook charr 32 Gradual 20 19.0 15.0-18.0 (9,10) salinity, gill Na+, K+-ATPase activity

11 ~ Weisbart, 1968; 2 = Scott and Crossman, 1973; 3 = Wagner el aI., 1969; 4 = Healey, and hypoosmoregulatory ability are ele­ 1980; 5 = Conte et aI., 1966; 6 ~ Johnslon and Saunders, 1981; 7 = Conte and Wagner, vated (McCormick et aI., 1985). In addi­ 1965; 8 = Wagner et aI., 1963; 9 = McCormick and Naiman, 1984b; 10 = Montgomery et aI., lexl foolnole 1. tion, relatively small increases in salinity (e.g., 28%0 vs. 32%0) can have dramatic effects on salmonid survival (Jackson, 1981). Because the decision to use brook fies a potential negative effect on males and migratory behavior characteristic of charr in enhancement programs for in enhancement programs. In fact, poor salmon smolts (White, 1940; Wilder, anadromous salmonids will undoubtedly seawater survival (or preference) may ex­ 1952), but silvering of brook charr does be made relative to the use of other plain the existence of greater female: not indicate imminent seawater entry salmonids, it is instructive to compare the male sex ratios in most populations of (Black, 1981). Unlike smolting relative salinity tolerances of salmonids anadromous and sea-ranched brook charr salmonids, brook charr do not display (Table 5). Size dependent hypoosmoreg­ (White, 1940; Wilder, 1952; Whoriskey photoperiod-induced increases in gill ulatory abilty is a common feature of all et aI., 1981 b; Castonguay et aI., 1982; Na+, K+-ATPase activity or hypoos­ salmonids (Parry, 1958; Conte and Wag­ Montgomery et al. I). moregulatory ability (McCormick and ner, 1965; Conte et aI., 1966; Weisbart, The influence of seasonal daylengths Naiman, 1984a, b). Furthermore, there 1968; Farmer et aI., 1978). As might be on developmental events has a substan­ are no significant differences (P > 0.05) expected, the size at which salinity toler­ tial impact on the osmoregulatory physi­ in gill Na+, K+-ATPase activities or ance is achieved is similar to that at ology of salmonids. Photoperiod entrains plasma thyroxine levels between anadro­ which normal seaward migration occurs the timing of maturation in salmonids mous and nonanadromous brook charr (Table 5). Species of the genus Oncor­ (Billard et aI., 1978; McCormick and from adjacent rivers (McCormick et aI., hynchus have relatively greater salinity Naiman, 1985b) and will therefore influ­ 1985). From these and other results we tolerance at any size than Salmo sp. ence seawater survival of brook charr have concluded that physiological (e.g., Atlantic salmon and rainbow through male maturation (McCormick changes preparatory for seawater entry trout), followed by Salvelinus sp. (see and Naiman, 1985a). In Atlantic salmon, that are observed in smolting salmonids Rounsefell, 1958). We have hypothe­ Salmo salar, and Pacific salmon, are undeveloped in brook charr. sized that greater exploitation of the sea Oncorhynchus spp., the transition from While photoperiod appears to affect has been accompanied by adaptations to resident parr to migratory smolt is cued salinity tolerance of brook charr only decrease the impact of size dependent ion by photoperiod, presumably acting through maturation, other environmental transport in a hyperosmotic environment through the hypothalmus-pituitary sys­ factors will have direct impact. Though (McCormick and Naiman, I984b). Other tem. Known as smoltification, this proc­ largely unexplored in brook charr, the in­ things being equal, enhancement of ess includes seasonal (usually spring­ teraction of temperature and salinity af­ anadromous brook charr will require time) changes in appearance, behavior, fects ion transport in other salmonids longer rearing in fresh water so that a morphology, and body composition, and (Rao, 1969). Ionic equilibrium of larger body size is achieved before intro­ increases in gill Na +, K+-ATPase activ­ presmolt Atlantic salmon after seawater duction to seawater. ity and hypoosmoregulatory ability (see exposure was achieved more quickly at Exploitation of reviews by Hoar, 1976; Folmar and an acclimation temperature of IO°C than Dickhoff, 1980; Wedemeyer et aI., 1980; at ISC (Virtanen and Oikari, 1984). Anadromous Brook Charr and McCormick and Saunders, In press). Poor osmoregulatory capacity at low For at least 5,000 years northern peo­ Some of these changes may be mediated temperature could explain high mortality ples have fished for brook charr (Mac­ by surges in plasma thyroxine (Folmar of brook charr and Arctic charr overwin­ Crimmon and Gots, 1980). Nevertheless, and Dickhoff, 1980). Anadromous brook tered in seawater net pens (Saunders et throughout their native range, there are charr often possess the silver coloration aI., 1975; Wandsvik and Jobling, 1982). only four historical records of anadro­

49(4), 1987 9 mous brook charr being harvested com­ Table 5.-Perceived positive and negative aspects associated with the enhancement 01 anadromy in brook charr. mercially. In the 1800's they were netted Parameter Positive aspects Negative aspects in the ocean near the Magdalen Islands, Life history 1 Short, synchronized migrations 1 Must be close to seawater for anadromy traits 2. Remain in coastal areas 2. Requires overwinter habitat in northern rivers Quebec, during summer and pickled for 3. Rapid growth at sea 3. 2-3 yrs before migration in northern latitudes export to the West Indies where, if in 4. Multiple spawnings after age 2+ 5. Minimum straying choice condition, they brought a higher price than Atlantic salmon (Perley, 1852; Physiology 1 Tolerance to low pH and low tempera­ 1 Difficulty adapting to full-strength seawater tures until 15-18 em FL Lanman, 1873). Anadromous brook 2. Ability to'enter brackish water at a small 2. Poor salinity tolerance of mature males charr were also commercially exploited size in Ungava Bay, Quebec, in Laborador by Genetics 1 Genetically plastic throughout range; pos­ 1. None known the Hudson's Bay Company in the 19th sibility for selective breeding Century (G. Power2) and in Richmond Economics 1 Low costs for fishery development and 1. Typical resource management problems as­ maintenance sociated with anadromous fisheries Gulf, Quebec, between 1962 and 1964 2. Potentially high yield for recreational fish­ (Power and Lejeune, 1976). On the Kok­ ery 3. High quality fish soak River, Quebec, the native harvest 4. Improved social situation for rural com­ munities has been estimated at 15,500 kg/yeail. 5. Available to anglers with average in­ Anadromous charr currently are netted comes for local consumption wherever they are present in sufficient numbers, but they are not generally numerous enough to be of widescale commercial importance, since most natural populations consist of only a few thousand fish. ational fishing (MacCrimmon and Gots, factors are essential for potential use of The main economic value of anadro­ 1980). In coastal areas the angling effort this species for enhancement and for mous brook charr probably lies in provid­ for anadromous brook chaIT can be sub­ commercial sea ranching efforts. ing recreation. In contrast to occasional stantial. For example, in the Moisie There are, however, some negative as­ commercial and native exploitation, River, Quebec, 18 percent of the fish pects to consider (Table 6). First, brook anadromous charr have consistently been tagged as part of our migration study in charr must be close to seawater to induce a favorite with anglers, notably in New this remote area were returned by anglers anadromy. Apparently they do not make England and the Maritime region of (Montgomery et al. I). long (>75 km) migrations to sea. They Canada (Perley, 1852; Bigelow, 1963). Summary require overwintering habitat in fresh In the past, much of the recreational an­ water and rearing habitat for 2-3 years gling in eastern Canada and the north­ We suggest that anadromous brook before migration from northern rivers. eastern United States was provided by chaIT and other members of the genus Second, brook charr have difficulty in Atlantic salmon. However, the world­ Salvelinus, have inherent traits making adapting to full-strength seawater before wide catch had declined 42 percent be­ them reasonable candidates for enhance­ reaching a size of 15 cm FL. Three fac­ tween 1967 and 1982 despite a several­ ment programs (Table 6). In northern lat­ tors appear to exert the strongest limita­ fold increase in fishing effort (RASA, itudes they make short, synchronized mi­ tions on the salinity tolerance of brook 1983), and the decline continues. Some grations to coastal areas during the charr: I) Size is of primary importance; 41 percent of coastal streams in New ice-free season, they remain close to brook charr :515 cm FL will probably not Brunswick have sea-run charr in suffi­ shore where they are not subject to inter­ survive well at salinities greater than cient quantities to provide a sport fishery. ception by foreign vessels, they undergo 25%0. 2) Gradual acclimation at interme­ In Newfoundland, anadromous brook rapid growth while at sea, they begin re­ diate salinities eases osmotic stress and charr are second only to Atlantic salmon production at about 2 years and repro­ increases survival. 3) The negative effect as a quarry for anglers (Scott and Cross­ duce every 1-2 years thereafter, and they of male maturation on salinity tolerance man, 1964). Moreover, in recent years, exhibit the strong homing ability charac­ is likely to preclude introduction of ma­ about 21.7 million brook chaIT have been teristic of other salmonids (Fig. 7). ture fish into seawater, particularity in distributed annually from government Brook chaIT are able to tolerate a wide autumn_ Spring releases ofjuveniles may hatcheries in the United States and variety of environmental conditions in­ circumvent some or all problems associ­ Canada to meet public demand for recre­ cluding low food levels and low water ated with poor seawater survival of ma­ temperatures, and are more resistant to ture males. Eventually, development of low pH than other salmonids (Muniz and late-maturing stocks, sex control, or ster­ 2Geoff Power, Department of Biology, Univer­ sity of Waterloo, Waterloo, Ontario N2L 3G1, Grarde, 1974). Moreover, they appear to ilization may prove advantageous. The Canada. Personal commun. have population dependent differences in greater degree of size dependent salinity 3N.Q.I.A. 1976. Research to establish present life histories suggesting a genetic plastic­ tolerance in brook charr requires a longer levels of harvesting by native peoples of northern Quebec. Part II. A report on the harvests by the ity which offers substantial opportunities freshwater rearing phase than for other Inuit of northern Quebec. 230 p. for selective breeding. These positive anadromous salmonids. Since freshwater

10 Marine Fisheries Review TR IBUTARY RIVER ESTUARY OCEAN

JUNE EMERGENCE 3 - 6 cm (AGE 0+) ~ '\es Stream Edge, J,s,e"" Rapid Grawth an Terrestri aI WINTER Insect Faods RESIDENCE t 7- 10 cm MAY-JUN Summer Residence ~ (AGE 1+) Slow Grawth

'" ~t w'" - I AUC-5EP Drifting Invertebrate Foods Summer Residence, Brackish 1O-1~ cm MAY-JUN Rapid, Synchronized ~ Surface Water (AGE 2+) Migration (2-10 days) Moderate Growth on Insect Foods, Enhanced Fecundity I AUC - SEP / l1.N1NG 9 Extended Summer Residence ( OCT - NOV) MAY-JUN Rapid, Synchronized - Migration \ >"t,~ E~a~e~~o~~ ~c~d2!Y (2: AGE 3+) Migration (2-10 days) Limited Residence __ __ Limited Residence d ------d,S RETURN AUC-SEP; 9'S SEP-OCT Near Shore Rapid Growth Sand Lance, Mysids

Figure 7,-The life history of anadromous brook charr studied by our group near Sept-lies, Quebec, is summarized, See text for explanation,

rearing is often limiting or costly, this is ment technique (Gibson and Whoriskey streams where anadromous brook charr an important consideration in choosing a 1980; Whoriskey et aL 1981b). How­ once flourished and use of local native species for enhancement. It will be nec­ ever, certain aspects can be modified to populations for hatchery stock to supple­ essary to judge the limitations of salinity improve the production and return of ment existing populations. tolerance in brook chaIT against other anadromous chaIT. These modifications However, several characteristics of merits for enhancing salmonid produc­ are applicable to both population en­ brook charr will, in our judgment, make tion to properly evaluate their potentiaL hancement for the purpose of expanding them a relatively poor choice for com­ Finally, there are the typical resource fisheries potential and to commercial sea mercial net pen culture in high salinity. management problems associated with ranching, Relative to other species, brook charr re­ anadromous fisheries where fish often Survival could be improved by gradual quire a longer rearing period in fresh use habitats under the jurisdiction of sev­ acclimation to seawater, releasing fish water prior to entrance into seawater. eral agencies with different management only during the period of natural migra­ Gradual acclimation to seawater in com­ priorities, tion to sea, using large estuaries with the mercial net pen culture would in many Considering both the positive and neg­ proper geomorphology (mixed vs. strati­ cases be costly or infeasible, The poor ative factors, we recommend that anadro­ fied water, etc.), and using fish in good performance of mature males in sea mous brook chaIT be used in local en­ condition. Harvesting fish could be im­ water and the relatively small size of mat­ hancement programs where the objective proved by recapture at a counting fence, uration in both sexes will limit the period is to provide a high quality recreational or some other structure that concentrates of seawater culture, These factors will fishery, This may include habitat im­ individuals soon after reentry into fresh work against the economic viability of provement, supplemental stocking to aid water in autumn. Growth rates at sea net pen culture. previously overexploited populations, could possibly be enhanced by choosing One final consideration is that in the improved management to protect the sea­ marine sites having physical attributes to southern part of the brook charr's range sonal movements and reproduction of the insure abundant food resources and by (i.e., near Cape Cod, Massachusetts), population, or the development of a selective breeding of an anadromous faster growth could reduce the length of hatchery (or genetic) program to exploit stock. Although we are not aware of any freshwater residence. However, in those life history characteristics that appear negative genetic aspects, genetics could areas they would face increased numbers useful for population enhancement. For have major consequences and should be of competitors, warmer marine waters instance, the initial experiments with in­ considered, At this point we recommend which they do not tolerate well, and habi­ ducing anadromy (sea ranching) demon­ that enhancement be limited to the use of tat degradation, North of Cape Cod, con­ strated the potential for using brook chaIT local stocks, Two basic enhancement ditions are more favorable for enhance­ in a moderate effort, high yield enhance­ strategies are habitat restoration of ment of anadromous brook charr, We

49(4), 1987 JJ suspect that enhancement will be limited 37-48. Elsevier/North Holland Biomed. waters. Fish. Mar. Servo Tech. Rep. 777, Press, Amsterdam. II p. to that region. Black, G. A. 198\. Metazoan parasites as Healey, M. C. 1980. The ecology of juvenile In summary, from our research and ex­ indicators of movements of anadromous brook salmon in Georgia Strait. British Columbia. In perience we believe that anadromous charr (Salvelinus fontinalis). Can. J. Zool. W. J. McNeil and D. C. Hinsworth (editors), 59: 1892-1896. Salmonid ecosystems of the North Pacific, p. brook chaIT, and possibly other members W. L. Montgomery, and F. G. 203-230. Oreg. State Univ. Press, Corvallis. of the genus Salvelinus, are candidates Whoriskey. Jr. 1983. Abundance and Hoar, W. S. 1976. Smolt transformation: for enhancement programs in suitable re­ distribution of Salmincola edwardsii evolution, behavior and physiology. J. Fish. (Copepoda) on anadromous brook trout, Res. Board Can. 33:1234-1252. gions. At present, their potential has not Salvelinus fontinalis (Mitchill), in the Moisie Jackson, A. J. 1981. Osmotic regulation in been fully explored. Based on informa­ River system. Quebec. J. Fish BioI. rainbow trout (Salmo gairdneri) following tion presented here, we suggest that 22:567-575. transfer to seawater. Aquaculture 24: 143-15 \. Brett, J. R. 1983. Life energetics of sockeye Johnston, C. E., and R. L. Saunders. 1981. brook chaIT be considered for relatively salmon. In P. A. Wayne and S. I. Lustick Parr-smolt transformation of yearling Atlantic low-effort, high-yield enhancement pro­ (editors). Behavioral energetics, p. 29-67. salmon (Salmo salar) at several rearing Ohio State Univ. Press. Columbus. temperatures. Can. J. Fish. Aqua!. Sci. grams in suitable areas, with economic Carlander, K. D. 1969. Handbook of freshwater 38:1189-1198. gains accruing to the local populace. fishery biology. Vol. \. Iowa State Univ. Lanman, C. 1873. The salmonidae of eastern Press, 752 p. Maine, New Brunswick, and Nova Scotia. Carlson, A. R., and J. G. Hale. 1973. Early U.S. Commisso Fish. Rep. 2:219-225. Acknowledgments maturation of brook trout in the laboratory. MacCrimmon, H. R., and B. L. Gots. 1980. Prog. Fish. Cult. 35:150-153. Fisheries for charrs. 839. In E. K. Balon This research was assisted by the ef­ Carscadden, J. E. 1970. Ecology of brook trout, (editor), Charrs: Salmonid fishes of the genus Salvelinus fontinalis in two lakes in the upper Salvelinus, p 797-839. Dr. W. Junk Publ., forts of many technicians and colleagues. Matamek River system, Quebec. M.Sc. The Hague, 928 p. We thank F. G. Whoriskey, Jr., G. A. thesis, Univ. Waterloo, Ont., 67 p. MacGregor, R. B. 1973. Age, growth and Black, G. Walsh, P. Heinermann, Castonguay, M., G. J. FitzGerald, and Y. Cote. fecundity relationships of anadromous brook 1982. Life history and movements of trout, Salvelinus fontinalis (Mitchill), in the J. Critchley, A. McNeil and E. T. Mont­ anadromous brook charr, Salvelinus Moisie River, Quebec. Undergrad. Hon. gomery for field assistance, S. Peterson fontinalis, in the St. Jean River, Gaspe, Thesis, Univ. Waterloo, Ont. for discussions of economic and social Quebec. Can. J. Zool. 60:3084-309\. McCormick S. D., and R. J. Naiman. 1984a. Conte, F. P., and H. H. Wagner. 1965. Osmoregulation in the brook trout, Salvelinus aspects of fisheries, and M. C. Healey, Development of osmotic and ionic regulation fontinalis. I. Diel, photoperiod and growth M. E. McDonald, G. Power, and R. L. in juvenile steelhead trout (Salmo gairdneri). related physiological changes in freshwater. Compo Biochem. Physiol. 14:603-620. Compo Biochem. Physiol. 79A:7-16. Saunders for discussions of anadromous J. Fesler and C. Gnose. 1966. _-:-,--_ and . 1984b. Osmoreg­ salmonid biology and for substantially Development of osmotic and ionic regulation ulation in the brook trout, Salvelinus improving the manuscript. The concept in juvenile salmon (Oncorhynchus kisutch). fontinalis. II. Effects of size, age and Compo Biochem. Physiol. 18: 1- I5. photoperiod on seawater survival and ionic of inducing anadromy in brook chaIT was Craig, P. C. 1984. Fish use of coastal waters of regulation. Compo Biochem. Physiol. initially formulated by R. J. Gibson. This the Alaskan Beaufort Sea: a review. Trans. 79A:17-28. research was supported by the Matamek Am. Fish. Soc. 113:265-282. ____ and 1985a. Hypo- Dutil, J. D., and G. Power. 1980. Coastal osmoregulation in an anadromous teleost: in­ Research Program of the Woods Hole populations of brook trout, Salvelinus fluence of sex and maturation. J. Exper. Zool. Oceanographic Institution, NOAA Of­ fontinalis, in Lac Guillaume-Delisle 234:193-198. (Richmond Gulf) Quebec. Can. J. Zool. ____ and . 1985b. Some deter­ fice of Sea Grant, the Tai-Ping Founda­ 58:1828-1835. minants of maturation in brook trout, Salveli­ tion, the WHOI Education Office, and Ie Farmer, G. J., J. A. Ritter, and D. Ashfield. nus fontinalis. Aquaculture 43:269-278. Ministere du Loisir, de la Chasse et de la 1978. Seawater adaptation and parr-smolt ____ , , and E. T. Mont­ transformation of juvenile Atlantic salmon, gomery. 1985. Physiological smolt character­ Peche du Quebec. This is Contribution Salmo salar. J. Fish. Res. Board Can. istics of anadromous and non-anadromous No. 5957 of the Woods Hole Oceano­ 35:93-100. brook trout (Salvelinus fontinalis) and At­ graphic Institution and No. 107 of the Folmar, L. c., and W. W. Dickhoff. 1980. The lantic salmon (Salmo salar). Can. J. Fish. parr-smolt transformation (smoltification) and Aquat. Sci. 42:529-538. Institution's Matamek Research Station. seawater adaptation in salmonids: A review of and R. L. Saunders. In press. selected literature. Aquaculture 21: 1-37. Preparatory physiological adaptations for Gibson, R. J. 1973. The interrelationships of marine life of salmonids: Osmoregulation, Literature Cited brook trout, Salvelinus fontinalis (Mitchill), growth and metabolism. Am. Fish. Soc. and juvenile Atlantic salmon, Salmo salar L. Symp. Andrews, C. W., and E. Lear. 1956. The Ph.D. Dissert., Univ. Waterloo, Ont., Can., Montgomery, W. L., S. D. McCormick, R. J. biology of arctic char (Salvelinus alpinus L.) 163 p. Naiman, F. G. Whoriskey, and G. Black. in northeastern Labrador. J. Fish. Res. Board ____ and D. Galbraith. 1975. The rela­ 1983. Spring migratory synchrony of Can. 13:843-860. tionships between invertebrate drift and salmonid, catastomid, and cyprinid fishes in Bardach, J. E., J. H. Ryther, and W. O. salmonid populations in the Matamek River, Riviere a la Truite, Quebec. Can. J. Zool. McLarney. 1972. Aquaculture. Wiley- Quebec, below a lake. Trans. Am. Fish. Soc. 61 :2495-2502. Intersci., N.Y., 868 p. 104:529-535. Moore, J. W. 1975. Distribution, movements, Bell, G. 1980. The costs of reproduction and , P. C. Kerkhoven, and R. L. and mortality of anadromous arctic char, their consequences. Am. Natur. 116:45-76. Haedrich. 1976. The fecundity of unexploited Salvelinus alpinus L., in the Cumberland Bigelow, H. B. 1963. Salvelinus fontinalis. In brook trout poulations in the Matamek River, South area of Baffin Island. J. Fish BioI. Fishes of the western North Atlantic, p. Quebec. Naturaliste Can. 103:417-423. 7:339-348. 525-542. Mem. Sears Found. Mar. Res. I. ____ and F. G. Whoriskey. 1980. An Morin, R., R. J. Naiman, and M. Legault. 1982. Billard, R., B. Breton, A. Fostier, B. Jalabert, experiment to induce anadromy in wild brook Influence of riparian vegetation and woody and C. Weil. 1978. Endocrine control of the trout in a Quebec river on the North Shore of debris on the ecology of brook trout fry, teleost reproductive cycle and its relation to the Gulf of St. Lawrence. Naturaliste Can. Salvelinusfontinalis.ln R. J. Naiman (editor), external factors: Salmonids and cyprinids 107:101-110. The Matamek research program: Annual re­ models. In P. J. Gaillard and H. H. Boer Greendale, R., and J. G. Hunter. 1978. Feeding port for 1981, p. 177-190. Woods Hole (editors), Comparative endocrinology, p. of freshwater fishes in estuarine and coastal Oceanogr. Inst. Tech. Rep. WHOI-82-29.

12 Marine Fisheries Review Mullan. J. W. 1958. The sea-run or "salter" ____ and R. Lejeune. 1976. Le potentiel Wagner, H. H., F. P. Conte, and J. L. Fesler. brook trout (Salvelinus fon/inalis) fishery of de peche du Nouveau Quebec. Cahiers de Ge­ 1969. Development of osmotic and ionic regu­ the coastal streams of Cape Cod, Massachu­ ographie de Quebec 20:(50):409-428. lation in two races of chinook salmon, setts. Mass. Div. Fish. Game, Bull. 17: 1-25. Rao, G. M. M. 1969. Effect of activity, salinity, Oncorhynchus tshawy/scha. Compo Biochem. Muniz, I. P., and M. Grarde. 1974. Overleving and temperature on plasma concentrations of Physiol. 29:325-341. ar olike arter lakesefisk: vonn fra et surt vass­ rainbow trout. Can. J. Zool. 47:131-134. ____, R. L. Wallace, and H. K. Camp­ drag. [Survival of various species of salmonid RASA. 1983. The Atlantic salmon crisis: A po­ bell. 1963. The seaward migration and return fish in water from an acid water-course.] In F. sition statement. Restoration Atl. Salmon of hatchery-reared steelhead trout in the Alsea H. Brgekke (editor), Hydrokjemiske og hy­ Am. (RASA), Inc., Hancock, N.H., 5 p. River, Oregon. Trans. Am. Fish. Soc. drologiske rapporter frq NIVA, p. 29-39. Ricker, W. E. 1932a. Studies of speckled trout 92:202-210. SNSF-proj. IR 3/74. (Salvelinus fOn/inalis) in Ontario. Ont. Fish. Walsh, G., R. Morin, and R. J. Naiman. In Naiman, R. J. 1982. Characteristics of sediment Res. Lab. Publ. 44:69-110. press. Daily rations, diel feeding activity and and organic carbon export from pristine boreal 1932b. Studies of trout producing distribution of age-O brook charr (Salvelinus forest watersheds. Can. J. Fish. Aquat. Sci. lakes and ponds. Ont. Fish. Res. Lab. Publ. fOn/inalis) in two subarctic streams. Environ. 39: 1699-1718. 45:113-167. BioI. Fish. 1983. The annual pattern and spa­ 1975. Computation and interpreta­ Wandsvik, A. and M. Jobling. 1982. Overwin­ tial distribution of aquatic oxygen metabolism tion of biological statistics of fish populations. tering of migratory Arctic charr, Salvelinus in boreal forest watersheds. Ecol. Monogr. Fish. Res. Board. Can. Bull. 191:382 pp. alpinus, (L.) reared in salt water. J. Fish BioI. 53:73-94. Rounsefell, G. A. 1958. Anadromy in North 20:701-706. ____ , J. M. Melillo. M. A. Lock, T. E. American salmonidae. U.S. Fish Wildl. Servo Wedemeyer, G. A., R. L. Saunders, and W. C. Ford. and S. R. Reice. 1987. Longitudinal Fish. Bull. 58:171-185. Clarke. 1980. Environmental factors affecting patterns of ecosystem processes and commu­ Saunders, J. W., and M. W. Smith. 1964. Plant­ smoltification and early marine survival of nity structure in a subarctic river continuum. ing brook trout (Salvelinus fOn/inalis) anadromous salmonids. Mar. Fish. Rev. Ecology 68: 1139-1156. [Mitchill], in estuarial waters. Can. Fish Cult. 42(6):1-14. Nordlie. F. G., W. A. Szelistowski. and W. C. 32:25-30. Weisbart, M. 1968. Osmotic and ionic regula­ Nordlie. 1982. Ontogenesis of osmotic regu­ Saunders, L. H. 1969. Ecology of brook trout, tion in embryos, alevins, and fry of the five lation in striped mullet. Mugil cephalus L. 1. American smelt and arctic charr of Matamek species of Pacific salmon. Can. J. Zool. Fish BioI. 20:79-86. Lake, Quebec. Ph.D. Dissert. Univ. Water­ 46:385-397. O·Connor.1. F. 1971. Ecology of brook trout, loo, 130 p. White, H. C. 1940. Life-history of sea running American smelt and American eel in two lakes ____, B. C. Muise, and E. B. Henderson. brook trout, Salvelinus fon/inalis, of the in the Matamek River system, Quebec. M.Sc. 1975. Mortality of salmonids cultured at low Moser River, Nova Scotia. J. Fish. Res. thesis. Univ. Waterloo, 84 p. temperature in seawater. Aquaculture 5:243­ Board Can. 5:176-186. 1974. On the brook trout (Salveli­ 252. ____. 1941. Migrating behaviour of sea IIUS fon/inalis) production and lcophic dynam­ ____ and G. Power. 1970. Population running Salvelinus fon/inalis. J. Fish. Res. ics of four streams in the Matamek River sys­ ecology of the brook trout, Salvelinus fOn/i­ Board Can. 5:258-264. tem. Quebec. Ph.D. Dissert.. Univ. nalis, in Matamek Lake, Quebec. J. Fish. 1942. Sea life of the brook trout Waterloo. 201 p. Res. Board Can. 27:413-424. (Salvelinus fontinalis). J. Fish. Res. Board and G. Power. 1973. Homing of Scott, W. B., and E. J. Crossman. 1964. Fishes Can. 5:471-473. brook trout (Salvelinusfon/inalis) in Matamek occurring in the fresh waters of insular New­ Whoriskey, F. G., R. J. Naiman, and P. H. Lake. Quebec. J. Fish. Res. Board Can. foundland. Royal Ont. Mus., Life Sci. Cont. Heinerrnann. 1981a. Steelhead trout (Salrno 30:1012-1014. 58:1-124. gairdneri) on the North Shore of the Gulf of ____ and . 1976. Production and 1973. Freshwater St. Lawrence near Sept-Isles, Quebec. Can. J. by trout in four streams in the Matamek water­ fishes of Canada. Fish. Res. Board Can., Fish. Aquat. 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