A LIMNOLOGICAL SURVEY OF PAXSON AND SUMMIT LAKES IN INTERIOR ALASKAY With Special Reference t o t h e Lake Trout and Lake Whitefish . George L Van Whye and James W . Peck Alaska Department of Fish and Game Division of Sport Fish Fairbanks, Alaska INTRODUCTION A survey of Summit and Paxson Lakes w a s initiated by the Alaska Department of Fish and Game, Sport Fish Division, in 1962. The objectives of the survey were: (1) evaluate t h e potential lake trout (Salvelinus namaycush) sport fishery, ( 2 ) determine the possibility of a commercial whitefish fishery, (3) gather information on growth r a t e s and a g e of maturity for t h e s e fish in Interior Alaska, and (4) provide limnologica 1 information for management of t h e s e populations. Lake trout and whitefish populations a r e found scattered throughout Interior Alaska l a k e s . The importance of t h e Lake trout a s a trophy sport fish has become very evident in the past five y e a r s while t h e economic value of t h e whitefish has not a s y e t been determined. Little prior information of t h e s e f i s h e s is available for t h i s region. Additional data on the life history of the round whitefish (Prosopium cylindraceum) w a s collected in conjunction with, but not presented in t h i s study and w a s reported in t h e form of a t h e s i s written a s partial fulfillment of t h e require- 1/ This work w a s carried out under Dingle-Johnson Federal Aid t o Fish Restoration, Project No. F-5-R-3 Job B between the d a t e s of June 1 , 1962 and May 1 , 19 64. ments for a Master of Science Degree from the University of ~ i c h i ~ a & / . LOCATION AND DESCRIPTION OF THE STUDY AREA Paxson and Summit Lakes lie a t the head of the Gulkana River in t h e Copper River Basin. This b a s i n i s located in southcentral Alaska a t approximately i a t i t u d e 63O N and longitude 145' W , between the 2,600 and 3,300 foot e l e v a t i o n s , and drains into the Gulf of Alaska, Figure 1 . Historically, both Paxson and Summit Lakes were formed a s a result of Pleistocene glaciation. Glaciers in t h e Chugach Mountains dammed t h e Copper River forming a large l a k e within the Copper River Basin. The present l a k e s and river valleys were carved out of deposited sediment after retreat of t h e i c e and drainage of t h e Copper River Basin. Between the river v a l l e y s , muskeg and marshes occupy depressions in t h e poorly drained, permanently frozen old l a k e floor. The s o i l type surrounding both lakes i s glacial moraine. The rock type is sedimentary in origin which, in many c a s e s , h a s become metamorphosed. Paxson and Summit Lakes drain a n area of approximately 3 ,000 square miles in the southern foothills of the Alaska Range. Tributaries a r e short, clearwater streams. Summit Lake i s drained by the Gulkana River which flows into Paxson Lake 15 miles downstream. LIMNOLOGY OF THE LAKES Physical Characteristics Morphometry of Paxson Lake. Paxson Lake has a surface area of 6 . 0 8 square miles, i s 1 0 . 3 miles long, and has a maximum width of 0 . 9 miles. The maximum depth i s 89 f e e t , and the mean depth is 35 f e e t . Paxson Lake exhibits a regular basin type bottom (Figure 2). The total a r e a between shore and the outer extreme of the 40-foot depth zone i s 3 . 6 8 square miles or 60 percent of t h e total surface a r e a . The greater part of the shoal area l i e s in the southern portion 01 t h e l a k e toward t h e outlet. 2/ Peck, James W. A comparative study of the growth of t h e round whitefish, Prosopium cylindraceum, in the Copper River Drainage, Alaska . 53 pages. F I G U R E 1. PORTION O F STATE OF ALASKA SHOWING STUDY AREA. Figure 2 . Paxson L a k e contour map. Roman numerals indicate sampling s t a t i o n s . The shore line of Paxson Lake i s quite regular and c o n s i s t s of sedimentary rocks which, in many c a s e s , have become metamorphosed. The substrate above 20 feet i s composed principally of sedimentary rocks t o 2 inches in diameter, interspersed with sand and some organic debris. Below 2 0 f e e t , the substrate i s chiefly c l a y with traces of organic debris. Morphometry of Summit Lake. Summit Lake h a s a s u r f a c e area of 6.38 square miles, i s 7 . 1 miles Long, and h a s a nlaximum width of 1.1 miles. The maximum depth i s 214 feet and the mean depth i s 70 f e e t . Summit Lake has both a n irregular shoreline and very irregular bottom a s shown in Figure 3. The total area between shore and t h e outer extreme of t h e 40-foot depth zone is 3.50 square miles or 55 percent of the total surface a r e a . The greater part of the shoal a r e a s a r e in the southern portion of the l a k e . Diversion of glacial streams in 1 9 2 9 , fed by the Gulkana glacier out of Gunn Creek, resulted in a clearing of the l a k e waters. The drainage basin has a typical a r c t i c continental weather regime with mild t o warm summers and s e v e r e winters. The mean maximum temperature is 3 7.3O F , and the mean minimum i s 1 6 . SO F The mean a r p u a 1 temperature i s 27.2' F. Ice i s usually present on both l a k e s from early November until early June and may obtain a thickness of 36 inches ( U . S Weather Bureau records). Mean annual precipitation i s 11 . 7 inches and the mean annual snowfall i s 48.6 i n c h e s . Seasonal variation in the amount of daylight varies from 20 hours in June to l e s s than 5 hours in December, giving a general frost-free period of about 67 d a y s . Paxson Lake, which i s below tfee l i n e , p o s s e s s e s a brushy t o wooded shoreline with alder and willow being the dominant brush forms and spruce and a s p e n being the dominant tree forms. . . Summit dake is above t h e tree l i n e , which for t h i s area i s about the 3 ,000-foot elevation. The shoreline vegetation is restricted t o willow and a lder brush with tundra-type ground cover. Part of the e a s t shore i s formed by t h e Richardson Highway road bed and l a c k s vegetation entirely. Most of the exposed shoreline i s composed of racks ranging in s i z e from 2 inches t o 2 feet in diameter. The substrate above 20 f e e t i s chiefly composed of large1 rubble. Some sand and rocks t o 2 inches in diameter a r e present. Organic material i s present in only small quantities in the very shallow waters of t h e north and s o u t h e a s t e n d s . Generally, the substrate below 20 feet i s rock or c l a y deposited by glacial outwash. A s shown in Table I,, Summit and Paxson Lakes a r e very similar, t h e Figure 3 . Summit Lake contour map. Roman numerals indicate sampling stations. TABLE 1. LAXE MORPHOMETRY DATA, PAXSON AND SUMMIT LAKES. P a x s o n Lake Summit Lake Elevation 2,600 f t , 3,300 f t . Area 6 . 0 8 sq. m i . 6.38 sq.mi. M a x i m u m width 0.9 mi. 1.1 m i . Maximum l e n g t k 10.3 m i . 7.1 m i . Maximum d e p t h 89 f t . 214 f t . Mean d e p t h 35 f t . 70 f t . P e r c e n t bottom area l e s s t h a n : 20 f t . d e p t h 3 2% 40 f t . d e p t h 5 5% 60 f t . depth 64% 80 f t . depth 8 5% 100 f t . d e p t h 9 1% 120 f t . d e p t h 9 4% 140 f t . d e p t h 9 7% 160 f t . d e p t h 98% 180 f t . d e p t h 99% 200 f t . d e p t h 99% main differences being in total volume of water and depths, Summit Lake being t h e greater of t h e two. Summit Lake h a s a mean depth of twice that 01 Paxson Lake, but the percent of shoal a r e a s i s practically the same. . Temperature - Paxson Lake Two temperature s t a t i o n s were maintained in 1963. Data were collected s i x times between June 30 and August 9 , 1 9 6 3 . Temperature profiles a r e shown in Figures 4 and 5 . The data indicated that this l a k e does not distinctly stratify during t h e summer months. However, on July 4, a distinguishable thermocline w a s present a t Station 11, but not a t Station I. This thermocline disappeared three d a y s later and may reflect a temporary abatement of prevailing winds. The a b s e n c e of stratification a t Station I on July 4 may be due t o the proximity of t h i s station t o the Gulkana River inlet. Warmer temperatures penetrated t o greater depths later in t h e s ummer . , Paxson Lake, due t o i t s location, s h a p e and orientation t o prevailing w i n d s , is subject t o continual turnover during i t s ice-free period. Based on o b s e d a t i o n s by local r e s i d e n t s , ice forms about the first of November and t h e break-up occurs during the first or second week of June. In 1962, freezeup occurred between November 4 and 9 . The d a t e of freeze-up in 1963 i s not known. Temperature - Summit Lake. Two temperature stations were maintained in 1963. Data were collected five times between July 2 and August 1 3 . Temperature profiles a r e shown in Figures 6 and 7. Data collected on July 2 immediately following spring break-up implied the presence of inverse stratification. The data a r e insufficient t o be conclusive and no winter temperature data were collected t o support t h i s implication. However, it is quite probable, based on studies of l a k e s a t t h i s and other latitudes with similar climatic conditions, that both Paxson and Summit Lakes experience some degree of inverse stratification during their periods of i c e cover. A weak thermocline w a s evident on July 9 a t Station 11. This thermocline disappeared prior t o July 2 1 . Prevailing winds blow along the long a x i s of Summit Lake. Although i t s fetch i s l e s s than that of Paxson Lake, Summit Lake is located above the tree line and i s more exposed; therefore, it should experience vertical mixing of a t l e a s t the magnitude found in Paxson Lake. Transparency. Light penetration w a s measured with the standard Secchi d i s c and w a s made in conjunction with plankton sampling. Readings in Paxson F I G U R E 4. WATER TEMPERATURES I N PAXSON LAKE, 1963 Station I T e m p e r a t u r e (OF) FIGURE 5. WATER 'IEMPERATURES I N PAXSON LAKE, 1963 Station I1 Temperature 3 50 400 450 (OF) 500 550 0 10 20 - 30 C, w V C C, 2 40 R 50 . ..- - - - -6/30/63 60 7/4/63 7/20/63 70 80 .......8/9/63 FIGURE 6. WATER TEMPERATURES I N SUMMIT LAKE, 1963 Station 1 30° Temperature (OF) 40° 45O 55O FIGURE 7. WATER TEMPERATURES IN SUMMIT LAKE, 1963 Station IT Temperature (OF) Lake ranged from 1 0 t o 1 4 f e e t . Summit Lake w a s more variable, with readings ranging from 5 t o 18 f e e t . Decreased transparency resulted from turbidity fluctuations, a s no correlation w a s observed between Secchi d i s c values and the quantity of plankton c o l i e c t e d . The highest turbidity in Summit Lake w a s observed following a period of high winds which could have produced sufficient mixing t o bring up the fine bottom sediments. In addition, Gunn Creek, the principal inlet, brings variable amounts of melt water from Gulkana Glacier into Summit Lake. Chemical Characteristics To obtain a quantitative chemical a n a l y s i s of Paxson and Summit Lake w a t e r s , samples were s e n t t o t h e University of Alaska for a n a l y s i s . Samples were collected a t t h e five-foot depth in the center of both lakes on October 2 3 , 1963 and a t that time, it was believed both lakes had experienced complete mixing. Therefore, t h e data a s shown in Table 2 should be representative of t h e entire lake . The chemical characteristics indicated that both l a k e s a r e soft water lakes; both experiencing slow buffering. The lack of sufficient carbonate in Paxson Lake is the probable c a u s e of the high C 0 2 content which may have been a temporary condition due t o the f a c t that the pH was alkaline. Both Lakes exhibited the high bicarbonate content which i s characteristic of soft water Lakes. Biological Characteris t i c s Plankton. Table 3 presents t h e results of plankton sampling conducted in both Paxson and Summit Lakes during the summer of 1 9 6 3 . Plankton w a s collected in 40-foot vertical h a u l s , using a Wisconsin type plankton net with Number 25 s i l k boiting cloth mesh. Plankton coliections were made in conjunction with temperature data collections in both l a k e s . The specific location of sampling stations a r e noted in Figures 2 and 3 . The mean settled volume of plankton per samples from Paxson Lake w a s twice that of t h o s e examined from Summit Lake. Two types of phytopiankton, Ceratium and Dinobryon, were found in Paxson Lake, but not in Summit Lake. The zooplankter Polyarthra was poorly represented in the Paxson Lake s a n ~ p l e sand w a s not found in samples from Summit Lake. It w a s observed that each organism present in both l a k e s w a s represented by a larger number of individuals in Paxson Lake. TABLE 2. WATER A N A L Y S I S OF PAXSON AND SUMMIT LAKES, O G M B E R , Summit --Pa%son 02 9.4 ppm 8.1 p p m C02 0.0 ppm 5.5 pH-th a l k a l i n i t y 9 . 0 ppm 0 . 0 pprn M.O. 50.0 pprn 6 4 . 0 pprn Total alkalinity 59.0 pprn 6 4 . 8 pprn pprn PH 7.8 7.4 ME4 N i t r o g e n 0.07 pprn 0 . 0 7 pprn NO2 -t- NOj N i t r o g e n 0.00 pprn 0.02 pprn PO4 0.12 ppm 0.10 pprn 0 - 0 0 pprn 0.46 pprn SO4 3.00 ppm 0 . 0 9 ppm Tannin 0.10 ppm 0 . 0 0 ppm Turbidity 0.00 ppm 2.00 ppm I 1963 TABU 3 . P A X S O N Dsite ---- Station 7/20 7/20 ITI I1 - TOTAL PLANKTON SAMPLE ANALYSIS, PAXSON AND SUMMIT LAKES, L A K E Mean S e t t l e d Volume ( c c ) 0.95 0.73 4.89 Most a b u n d a n t f o r m s . - S U M M I T Forms P r e s e n t Phytoplankton ~sterionelg* Ulothr ix* Coe l o s p h a e r i u m Microcystis Ceratium Dinobryon Aphanizomenon Anabaena Zooplanktoq Copepoda* Nauplius larvae* Polyart h r a Kellicottia Keratella 1 Each s a m p l e b a s e d o n two, 4 0 - f o o t v e r t i c a l h a u l s . * 1963~ Date 7/8 7 -- / 2 1 7/21. 7129 7/29 8/13 8/13 Mean S e t t l e d Volume ( c c ) -- Station L A K E I1 0.40 0.20 0.15 0 . 45 0.70 0.17 0.35 TOTAL 2.42 V I1 V V I1 V - Forms P r e s e n t 7 - Phytoplankton Aster ionella* U l o t h r ix* Microcys t i s * Coelosphaerium Anabaena A c t i n a s trum Zooplankton Keratella Kellicottia Copepoda* Nauplius l a r v a e Benthic Fauna. A total of 40 bottom samples w a s taken in the two lakes using a n Ekman dredge 22.8 cm by 22.8 cm in c r o s s section. Samples were obtained from various depths in the north, e a s t , and w e s t portions of Summit Lake and the north, south, and e a s t portions of Paxson Lake. Table 4 and 5 present results of the dredgings in t h e two l a k e s . Chironomidae larvae were the most abundant forms in both l a k e s . In Summit Lake, Trichoptera and a clam, Pisidium s p . , were a l s o abundant. In Paxson Lake, other abundant: forms were various oligochaetes, t h e gastropod Gyralus s p . , and Pisidium sp. Compared t o Summit Lake, t h e benthic fauna of Paxson Lake was richer in numbers and types of organisms. The only exception w a s the relative abundance of Trichoptera in Summit Lake. The greater abundance of oligochaeta and the presence of hirudinea and amphipods in Paxson Lake reflected t h e more organic nature of i t s substrate, a s opposed t o the fine c l a y in Summit Lake. Although not confirmed by measurement, individual organisms were observed t o be larger Fn Paxson Lake than in Summit Lake. This difference in s i z e w a s b e s t illustrated by the mollusks where s h e l l s of Paxson Lake mollusks were heavier than those in Summit, indicating a greater supply of carbonates . Except for Chironomidae larvae In Paxson Lake, the majority of organisms occurred a t depths of 20 feet or l e s s . Chironomidae iarvae in Paxson Lake were most abundant a t 60 f e e t , while maximum abundance in Summit Lake occurred a t 1 0 f e e t . In both temperature and substrate type, the sixty-foot depth in Paxson Lake w a s similar t o the ten-foot depth in Summit Lake. Rooted Aquatic Vegetation. Potamogeton i s the major plant type in both l a k e s , being represented by a number of different s p e c i e s . Rooted aquatic plants a r e very s c a r c e in Summit Lake and a r e restricted to the north and southe a s t shoal a r e a s . Rooted vegetation i s s o s p a r s e in Summit Lake that it should have almost no affect on the environment. Paxson Lake has abundant aquatic vegetation on every shoal except t h o s e which a r e subjected t o severe wave action. Potamogeton and Myriophyllum a r e the most abundant forms, providing a distinct habitat t y p e . Although no measurements of piant growth were made, eariy summer growth w a s extremely slow. Apparent1 y a temperature threshold was reached in l a t e summer after w h i c h plant growth proceeded rapidly. Accelerated growth TABLE 5 . BOTTOM FAUNA, SUMMIT LAKE, AUGUST 1963 G I V I N G ORGANISM T Y P E AND AVERAGED FREQUENCY O F OCCURRENCE A v e r a g e N u m b e r of O r g a n i s m s P e r S q u a r e M e t e r Dert h ( f t .) 2-5 TOTALS Chironomidae Diptera 41 3 adult 20 3 -- 5 pupae 125 45 -- 30 pupae 170 -- -- 2 0 pupae LOO 81 58 505 --- 3 648 PERCENT Oligochaeta --- L m n e . G y r a l u s 2. P i s i d i um -s*. 3 33 106 -- -- --- 140 93 138 323 2046 -- -- 41.4 Trichoptera 4.0 2.8 24.7 Total Per Depth 4.5 6.7 15.8 w a s observed during the l a s t week in July, 1963. At that time, water temperatures in the shaal a r e a s ranged f r o m 48.5O t o 52.5' F . Fish Sampling Methods. Fish populations were sampled by variable mesh gill n e t s and by bag s e i n e . The variable mesh gill n e t s were 1 2 5 feet long and 6 f e e t d e e p , consisting of 5 panels 25 feet in length, with mesh s i z e varying from 1 t o 3 . 5 inches stretch measure. In a n efIort t o reduce sampling b i a s , net s e t s were made along the shorelines a t predetermined locations with the small mesh toward shore. A s a result of differences in bottom contour, t h e Larger mesh sampled a t various depths. The bag s e i n e w a s 50 feet long and 6 feet d e e p with one-half inch bar mesh. Sampling w a s done throughout the s e a s o n t o c o l l e c t the young-ofthe-year f i s h . Because sampling w a s intensive, 7 8 7 hours in Paxson Lake and 6 7 7 hours in Summit Lake, it was doubtful that any s p e c i e s in the two lakes were not represented in the c a t c h . Relative Abundance. Red salmon (Oncorhynchus nerka) spawn in the e a s t fork of t h e Gulkana River and in tributaries above and below Paxson Lake. There i s a l s o limited red salmon spawning in Gunn Creek, the inlet: of Summit Lake. Adult salmon first appear in t h e two lakes soon after t h e i c e cover d i s a p p e a r s . In 19 62, spawning continued into October. Adult red salmon were taken incidentally in the variable mesh gill net c a t c h e s and were much more numerous in Paxson Lake than in Summit Lake. Both Paxson and Summit Lakes have a year-round population of juvenile red salmon. These fish were abundant and comprised t h e majority of the bag s e i n e c a t c h e s during August and September. Apparently most of the salmon fry were too small t o be captured in t h e s e i n e until August. The slimy sculpin (Cottus cognatus) w a s found t o be equally numerous in both l a k e s . Although it was larger and darker in Paxson Lake, none longer than 1 0 0 mm were observed in either lake. This s p e c i e s inhabits shallow waters and was readily captured with the bag s e i n e . A single long-nosed sucker (Catostomus catostomus) w a s captured in Paxson Lake. The resident population 01 this s p e c i e s w a s very s p a r s e . . Both l a k e s have small populations of burbot (Lota -lota) Only small individua 1s were t a k e n , a s the gear w a s unsuitable for taking t h e larger individuals. A few grayling (Thymallus arcticus) were taken from t h e shallower waters of both l a k e s . In Paxson Lake, the catch w a s restricted t o the-south end and near the outlet. In Summit Lake, the c a t c h w a s more widespread. The lake whitefish (Coregonus clupeaformis) was not found in Summit Lake, but w a s abundant in Paxson Lake. The rapid fall of the Gulkana River between t h e two takes may be a factor preventing t h e e a s y entry of lake whitef i s h into Summit Lake. The young-of-the- lake whitefish were taken in the bag s e i n e only during the early part of July. Later that month they completely d i s appeared from the c a t c h . This disappearance was apparently due t o a migration t o deeper water and may be correlated t o a r i s e in surface water temperature. Larger individuals were taken throughout the summer in gill n e t s . Thls s p e c i e s is taken occasionally on hook and line and supports a small, Local, s u b s i s t e n c e net fishery during the l a t e fall and early winter. The round whitefish (Prosopium cylindraceum) was found in both l a k e s . Young of the round whitefish were taken by bag s e i n e throughout the summer with the largest c a t c h e s being made a t night. Individuals of a g e 4 or older were taken in gill n e t s throughout the summer. This s p e c i e s i s a l s o taken occasionally by hook and l i n e , and supports a small, l o c a l , s u b s i s t e n c e net fishery when they move into the streams in the l a t e fall to spawn. The l a k e trout (Salvelinus namaycush) w a s present in both l a k e s . This s p e c i e s supports a relatively intense fishery during the summer months. Paxson Lake receives heavier fishing pressure due t o i t s good reputation and available boat f a c i l i t i e s . No lake trout were taken by bag s e i n e . Relative population data were collected for lake trout, two s p e c i e s of whitefish, grayling, and burbot on a c a t c h per unit effort b a s i s . Table 6 and 7 indicate a summary of gill net sampling in both l a k e s . To reduce fouling of the variable mesh gill n e t s by adult red salmon, no gill net sampling w a s conducted in Summit Lake before August 2 0 , or in Paxson Lake before September 6 . B y t h e s e d a t e s , adult red salmon were no longer present in the l a k e s in a n y appreciable numbers. The data indicated that lake trout populations in t h e s e l a k e s constituted a very simi!ar percentage of each l a k e ' s fish population. Catch per unit effort for l a k e trout in Summit Lake was slightly greater than in Paxson Lake and a greater catch per unit effort w a s achieved for grayling in Paxson Lake than in TABLE 6 . CATCH SUMMUXY U n i t of g e a r : - PAXSON LAKE, V a r i a b l e mesh g i l l n e t 1 2 5 ' x 6 ' Mesh: 1" - 3-1/2" s t r e t c h mesh Total hours f i s h e d : Total f i s h caught: 787 1,084 Total catch/unit e f f o r t : Species 1.377 Number Taken C a t c h / U n i t E f f o r t S/o Comp. Lake W h i t e f i s h (Coreqonus c l u p e a f o r m i s ) 5 27 Round FSnitef i s h (Prosopium c y l i n d r a c e u m ) 405 Lake T r o u t ( S a l v e l i n u s namaycush) 101 Grayling ( ~ h ~ m a l l uakr c t i c u s ) 1962 45 Burbot (-Lota l o t a ) One l o n g n o s e s u c k e r ( C a t o s tomus c a t o s t o m u s ) was t a k e n on S e p t e m b e r 20, 1 9 6 2 , i n P a x s o n L a k e , b u t n o t i n t h e above s e r i e s o f n e t s e t s . Red s a l m o n (Oncorhynchus n e r k a ) a n d s l i m y s c u l p i n ( C o t t u s c o q n a t u s ) have been o m i t t e d from b o t h t h e t a b l e and t h e c a t c h p e r u n i t e f f o r t c o m p u t a t i o n . NOTE: C a t c h p e r u n i t e f f o r t d e f i n e d a s number o f f i s h t a k e n p e r hour o f f i s h i n g w i t h above d e f i n e d u n i t of g e a r . TABLE 7 . CATCH SUMWU?Y U n i t of g e a r : - SUMMIT LAKE, 1962 V a r i a h l e mesh g i l l n e t 1 2 5 ' x 6 ' Mesh: 1" - 3 - 1 / 2 " s t r e t c h mesh T o t a l hours f i s h e d : Total f i s h caught: 677.5 995 L .469 Total catch/unit effort: Number Taken C a t c h / U n i t E f f o r t % Comp. Species Round W h i t e f i s h (Prosopium cylindraceum) 880 Lake T r o u t ( S a l v e l i n u s namaycush) 106 Grayling (Thymallus a r c t i c u s ) 5 Burbot (-Lota l o t a ) Red s a l m o n ( O n c o r h y n c h u s n e r k a ) and s l i m y s c u l p i n ( C o t t u s c o q n a t u s ) have been o m i t t e d from b o t h t h e t a b l e and t h e c a t c h per u n i t e f f o r t c o m p u t a t i o n . NOTE: C a t c h per u n i t e f f o r t d e f i n e d a s number o f f i s h t a k e n per h o u r o f f i s h i n g w i t h above d e f i n e d u n i t o f g e a r . Summit Lake. The catch per unit effort of burbot was approximately the s a m e for both l a k e s , hut sample s i z e s were too small t o afford meaningful camparisons. The round whitefish population of Summit Lake w a s more than t w i c e A M of Paxson Lake. This difference i s perhaps a result of competition in Paxson Lake between the two s p e c i e s of whitefish for food, s p a c e , or other resources. , The round whitefish population of Summit Lake w a s approximateIy equal to both s p e c i e s of whitefish in Paxson Lake. ECOLOGY OF LAKE TROUT AND WHITEFISH Age Determination Methods. Age a n d growth determinations were completed on l a k e trout and round whitefish from both Paxson and Summit Lakes, and lake whitefish from ~ ' a x s o nLake. All captured lake trout were used for a g e determinations. Sample s i z e s totaled 106 specimens from Summit Lake and 101 specimens from Paxson Lake. Due t o manpower and budgetary Limitations, it was decided t o limit the study sample of l a k e whitefish to t h e first 150 specimens captured. Age determinations were completed on 597 round whitefish from Summit Lake and 412 from Paxson Lake. Larger samples of round whitefish than l a k e whitefish.were used a s data were a l s o being prepared for a Masters t h e s i s on t h e a g e and growth of round whitefish. S c a l e s used in t h e growth a n a l y s i s were taken midway between t h e Lateral line and t h e origin of t h e dorsal fin on the left s i d e of t h e body. Plastic impressions of whitefish s c a l e s were made using a cold-press technique. Plastic impressions of l a k e trout s c a l e s were made using both hot-press and cold-press techniques. Optimum results were obtained using the cold-press technique. Four s c a l e s from e a c h f i s h were used a s a sample. Of t h e four, two s c a l e s of uniform s h a p e and characteristics were chosen. Measurements were made on both s c a l e s and t h e results were averaged. A g e determinations were made on a Bausch a n d Lomb model 42-63-58 micro-projector. A 2 . 7 ~l e n s w a s used for the whitefish s c a l e s and a 12x l e n s was used for t h e l a k e trout scales. Growth whs calculated by the formuis (Li/;,.) Sn, where Ln = fork length of fish a: ltk: yez: bt = fork length of fish a t time cf c q t u r e St = d i s t a n c e from s c a l e Ionus to posterior margin S, = d i s t a n c e from s c a l e foci.1~t o n'h annulus. 9: life The u s e of this l:rect proportion formula i s justified hy the s t r a ~ ~ h t - t i n e relationship which w a s found t o e x i s t between the avelage fork 1ang-i-hand the average s c a l e radius for each a g e group. Some deviation from t h i s relationship w a s observed in older a g e groups of l a k e trout, but this daviarlon included only about 1 0 percent of the f i s h in the s a ~ o l e s . Year C l a s s Composition. Figures 8 , 9 , ar,d ' .3 present sulnl-~ary:)I year c l a s s composition for l a k e trout and both speci?; of w h i t t . ~ i s h . The selectivity of the gear i s illustrated by the l ~ w £rec;~f =,r,cy of t > z y s u n s e r a g e groups of a l l s p e c i e s . It is interesting that the frequency of rcund waitefish G i Age-class VI w a s sharply reduced in both l a k e s (Figure C ) . Since the buik f:. t h e samples w a s composed of individuals from a g e s IV-IX, gear selectivity can be disregareed. Therefore, a g e cia ss VL may reflect a spawning year of unusually poor survival. Growth in Length The increment of yearly growth for lake trout in Sumniit snc: Pcxson Lakes i s tabulated in Tables 8 and 9 . In acJdition, a comparison of l a k e troat growth i s shown in Figure 11 As shown, growth was generally s L w s r in S u m ~ i t Lake than in Paxson Lake. To a g e V, growth r a t e s were similar, thaugh s1igh:ly slower in Summit Lake. During t h e s e y e a r s , the young l a k e trcut feed primariiy on insects and mollusks. The small difference in growth, t o a q e V , m a y be correlated with the more abundant invertebrate fauna in Pr7sson L a k e . . From a g e s V through IX, growth was much more rap i d In Ff3xson L a k e . This difference in growth, after the lake trout have s!?ifted to a f ~ s hd i e t , is p' ~ S a b l ycorrelaf2d t o the greater number of small f i s h in F , ~ x s a nLake. A s r~oi-l.!'i ~n t'?e food habits section presented l a t e r , red salmon fry were ut iil;7~3dcxferls:~e ! : ~ by Paxson Lake lake trout; while in Summit Lake, wb-.re red r;almon fly were m I c n FIGURE 8 . AGE C O M P O S I T I O N O F LAKE TROUT I N PFXSON AND SUlvlMIT LAKES, AS CALCULATED FROM G I L L N E T T f N G Aye group 1962, FIGURE 9. AGE COMPOSITIOI'J OF LIA-W WHITE&'ISII IPJ PAXSON, AS CALCULATdD FROM GILL NET SAMPLING A g e group 1962, FIGURE 10. AGE C O M P O S I T I O N O F RQUND WHITEFISH I N PAXSON AND SUMMIT LAKES, 1 9 6 2 , AS CALCULATED FROM GILL N E T SAMPW 4 / \ / r"/ - \,------------Su mmit Lake \ \ \ Paxson Lake \ 5 6 Age g r o u p 7 TABLE 8. CALCULATED GROWTH I N F O R K LENGTH (crn.) O F L A K E TROUT TAKEN FROM SUMMIT LAKE, 1962 AVERAGE LENGTH CALCULATED FROM EACH A G E GROUP A T END O F YEAR OF LIFE Number 0 f Fish 1 Weighted A v e r a g e 5.6 VII VIII XIV TABLE 9. CALCULATED GROWTH IN FORK LENGTH (cm.) O F LAKE TROUT TAKEN FROM PAXGON LAKE, 1962 AVEXAGE LENGTH CALCULATED FROM EACH AGE GROUP A T END O F YEAR O F LIFE Number of Fish I Weighted Average 6.Y II I11 IV V VI VIJ VI II IX 8 a XI1 XI I 12,2 18.0 23.6 29.0 36.3 42.9 49.3 51.1 59.4 63.5 73.7 78.0 F I G U R E 11. - GROWTH O F THE L4KE TROUT ( S a L v e l i n u s namaycush) OF S U M M I T AND PAXSON LAKES BASED ON AVERAGE CALCULATED FORK LENGTH F O R EACH AGE GROUP A,ge g r o u p - 30 - less numerous, l a k e trout were forced t o feed on the slimy sculpin. Therefore, after a g e V , production a n d growth of lake trout in t h e s e two l a k e s may well b e keyed t o the continued availability of red salmon fry. Beyond a g e X t h e number of f i s h from both l a k e s representing each a g e group w a s 'ioo small t o be of a n y u s e in depicting a n accurate increment of growth. Growth of the l a k e trout in Summit Lake may a l s o have been retarded somewhat by low water temperatures . Comparing the growth of f i s h in t h e s e two l a k e s with other l a k e s Generally, growth w a s slower involves c ~ n s i d e r a t i o nof many variables in Paxson and Summit Lakes than in l a k e s of lower latitudes. Growth comparison for Iarge Alaskan l a k e s a r e frequently made with Great Slave Lake and Great Bear Lake in C a n a d a , although t h e s e l a k e s a r e extremely large. . Great Slave Lake , located between latitude 61' and 63' north and longitudes 109O t o 117' e a s t , contains 20,500 square miles of surface and h a s a northern continental climate with a mean annual temperature of approxirna&ly 23O F. (Rawson, 1950). This temperature i s slightly below the 27.2O annual mean temperature in the Locality of Summit and Paxson Lak,0 a r e a s . Kennedy (1954) investigated population dynamics of l a k e trout in G e a t Slave Lake a n d reported a growth rate that w a s intermediate between that of Paxson a n d Snmmit Lakes. Miller and Kennedy (1943) a l s o investigated the l a k e trout in Great Bear Lake, a v a s t lake of 11,800 square mires located a t latitude 64O - 67O north and longitude l l g O - 123' w e s t . This l a k e is subjected t o severe climatic conditions with ice floes on the l a k e until l a t e July. In Great Bear Lake, the l a k e trout growth reporLed by Miller and Kennedy (1948) was exceedingly slow when compared t o Summit or Paxson Lakes. Age group XV, from Great Bear Lake, compared favorably with a g e group X in Summit Lake and a g e group VIII in Paxson Lake. Growth of the l a k e whitefish in Paxson Lake i s shown in TaSle 1 0 , and i s a l s o graphed for clarity in Figure 12. The annual increment of growth w a s nearly constant until a g e X. From a g e X through a g e XIII, a reduction in growth rate w a s evident. After a g e XIII, the sample w a s t o o small t o afford meaningful conclusions. In comparison, the growth rate for l a k e whitefish h " ~ r e aSlave t Lake (Kennedy, 1953) w a s somewhat above that of Paxson Lake. Growth ol: the round whitefish is depicted in Figure 13. Since growth of this s p e c i e s in Paxson and Siunmit Lakes i s t h e subject of a Masters t h e s i s TABLE 10. C A L C U L A T E D GROWTH I N F O R K L E N G T H ( c m . ) O F LAKE W H I T L F I S H T A K E N FROM P A X S O N L A K E , 1 9 6 2 A V E R A G E L E N G T H C A L C U L A T E D FROM E A C H A G E G R O U P A T E N D O F Y E A R O F LIFE Number of Fish I 2 5 11 23 17 20 28 11 3 2 I -I 0 1 to ' weighted Average II I11 IV V VI VII VIII IX X XI XI1 XI11 XIV xv XVI XVII F I G U R E 12. 1 2 3 GROWTH O F THE LAKE W H I T E F I S H ( C o r e q o n u s c l u p e a f o r m i s ) I N PAXSON LAKE BASED ON AVERAGE CALCULATED FORK LENGTH FOR EACH AGE GROUP 3 , 7 0 9 10 11 A g e group 12 13 14 15 16 17 18 FIGURE 13. CALCULATED GROWTH O F THE ROUND W H I T E F I S H (Prosopium c y l i n d r a c e u m ) I N PAXSON AND SUMMIT LAKES BASED ON AVERAGE CALCULATED FORK LENGTH FOR EACH AGE GROUP 4 Age group S i n g l e specimen by the junior author, discussion of round whitefish growth is only briefly mentioned in this paper. Figure 1 3 is based on sample s i z e s of 470 specfmens from Paxson Lake, and 572 specimens from Summit Lake. However, below a g e IV and above a g e X , sample sizes were too small t o provide accurate comparisons of growth. Sex Ratio In Paxson Lake s a m p l e s , the overall fema1e:male ratio for l a k e trout w a s 48:52. Among the l a k e trout s a m p l e s , males exceeded females only in a g e c l a s s e s IV-VII. Older a g e c l a s s e s contained a majority of females. In Summit Lake, the fema1e:male ratio of l a k e trout taken w a s 67:33. Females dominated a l l a g e c l a s s e s . Among t h e l a k e whitefish, males slightly dominated a11 a g e c l a s s e s . The fema1e:male ratio of iound whitefish w a s exactly the s a m e a s in Paxson Lake, 56-:44. Females dominated a l l a g e classes except a g e I V . The Paxson Lake fema1e:male ratio of t h e round whitefish was 56:44. Males dominated a g e c l a s s e s 111-VI; but older a g e c l a s s e s consisted predominately of females. Size and Age a t Maturity Most lake trout in Paxson Lake mature a t a g e s VIE-VIII. Almost a l l lake trout in Paxson Lake were mature when reaching 45 cm in length. Maturity i s defined here a s either having gonads developed t o s u c h a n extent t h a t spawning will t a k e place within the s a m e summer, or in t h a t gonads show evidence of prior spawning. By g r o s s examination, female l a k e trout were divided quite e a s i l y into f i s h containing large e g g s that were held loosely in t h e ovaries and were about t o spawn, and f i s h containing much smaller ova s t i l l held tightLy in the ovaries. Males could not be definitely separated into spawning males and mature males which would not spawn in the current year. I t i s unfortunate t h a t data w a s not compiled on the ratio of mature l a k e trout spawning e a c h year. Data similar t o t h a t obtained in Paxson Lake were collected in Summit Lake, e x c e p t that most l a k e trout matured between a g e s VIII-X instead of VIIVIIL. Numerous individuals were lmmature a t 50 cm, but a l l were mature a t 55 cm. Gonad development indicated t h a t spawning in Summit Lake would commence in mid-September and approximately 1 0 d a y s l a t e r in Paxson Lake. The above results compared favorably with a g e ana s i z e a t rnatu~ity of l a k e trout in Great Slave Lake. In this l a k e , Kennedy (1954) found that t h e first fish matured a t a g e VII and 4 3 cm in length, All fish were mature by a g e XI and 5 7 cm in length. Kennedy a l s o reported that Great Slave L a k e 'imbit apparently spawned once every two y e a r s after becoming mature. In contrast, Miller and Kennedy (1948) reported that in Great Bear Lake t h e first l a k e trout matured a t a g e XI11 a t 42 cm in Length, and a l l rake trout matured by a g e XVII a t approximately 55 crn in length. These authors reported that Great Bear Lake trout spawned once every three years after becoming mature. All l a k e whitefish from a g e VII were judged t o be sexually m a t w e . Not a l l individuals appear t o spawn e a c h y e a r , however, no data concerrling the ratio of ripe spawners t o unripe mature fish was compiled. Therefore, the frequency of l a k e whitefish spawning in Paxson Lake i s unkr~ovvrx. Aki individuals were mature a t 29 cm . Kennedy (1953) reported that in G-reat Slave Lake t h e lake whitefish first matured between a g e VLII and 30 crn with a l l fish mature by a g e XI11 and 4 5 cm. The same author reported l a k e whitefish in Great Slave Lake spawned once every two years after attaining maturity. In both Paxson and Summit Lakes, a11 observed round whiteiish shorter than 2 2 cm total length and younger than a g e IV were immature; whereas a l l f i s h were mature when longer than 29 cm and older than a g e VIII. In Paxson Lake, more than 50 percent of the males sampled were matfire a t a smaller s i z e ( 2 4 cm) than a comparable majority of females. However, in Summit Lake, t h e males attained 2 6 cm in length before more than 50 percent were mature, whereas the females were similar t o t h o s e from Paxson L,aLe. The round whitefish from both l a k e s had little difference in a g e at maturity, with more than 50 percent of both s e x e s maturing a t a g e VT, A greater percentage of males than females matured a t a g e IV and 11. In Paxson Lake, a slightly higher percentage of f i s h younger than a g e V were m a t ~ ~ ar te a n earlier a g e and a t a shorter length than for Summit L a k e f i s h . Food Habits A qualitative survey of the summer food habits of t h e iake t l = ~ ) ~ii'd t. round whitefish in Paxson a n d Summit L a k e s , and lake v~hltefrskl;r, PEA::^^, L ~ L P w a s conducted in conjunction with gill n e t sampling. In both l a k e s , lake trout under 40 cm fed principally on i n s e c t s and mollusks with some small f i s h being eaten by trout in the 30-40 cm range. Beyond 40 cm, the diet of lake trout w a s almost entirely f i s h . The slimy sculpin w a s common in stomachs of lake trout between 45 and 55 cm in Summit Lake. Sculpins were much l e s s common in the stomachs of comparable s i z e d Paxson Lake l a k e trout. Salmon fry were utilized by l a k e trout in Paxson Lake much more extensively than in Summit Lake. In both l a k e s , whitefish were intensively fed upon by large l a k e trout. In Paxson Lake, where both s p e c i e s of whitefish a r e present, fish remains were usually too badly digested t o allow identification a s t o s p e c i e s . Adult lake whitefish in Paxson Lake were observed feeding on salmon fry quite extensively during the spring and early summer. However, t h e predation ended a s the fry grew too big for t h e whitefish. It w a s a l s o observed that while some adult l a k e whitefish were feeding entirely on salmon fry, others in the same area would ignore the fry completely, feeding on the usual d i e t of molltrsks and other invertebrates. Round whitef i s h fed almost entirely on invertebrates. The principal food items were Trichoptera, Diptera larvae, , and Gyralus 9 . No fish remains were found; but 40 salmon eggs Lymnea were found in t h e stomach of one specimen from Paxson Lake. z. Parasites A total of 55 lake trout from both Lakes were examined for p a r a s i t e s . Specimens were selected randomly a s t o a g e and s e x and examined by the Department of Fish and Game parasitologist. The following parasites were found: Eubothrium salvelini. This tapeworm i s commonly found in many trouts and chars and w a s found in 3 6 of the 55 specimens. Nine individuals contained more than 30 of t h e p a r a s i t e s . Incidence was not correlated t o a g e or s e x . This parasite has a first larval s t a g e (procercoid) in copepods and a second larval s t a g e (pleroceroid) in prey f i s h and a r e not noted a s significant pathogens. Neoechinorhynchus rutili, a n acanthocephalan, h a s a world-wide d i s tribution and is commonly found in salmonoids, perches, and c a r p s . Twentyfive of t h e 5 5 specimens contained t h i s parasite, with 4 individuals carrying more than 30 of the p a r a s i t e s . Incidence w a s not correlated t o a g e or s e x . This parasite u s e s Ostracods or certain s p e c i e s of megalopteran i n s e c t s a s a host for t h e only larval s t a g e . Heavy infestations of this parasite have been reported to c a u s e mortality in trout. Crepidostomum farionis. This fluke i s a parasite of salmonoids throughout Europe and North America. Only 3 of the 55 specimens examlned were found t o be infected; none having more than 30 of the parasite. The 3 infected specimens were females and a l l were in the 1 2 year a g e c l a s s . The larval s t a g e of this parasite develops in s n a i l s and the a q u a t i c l a n ~ a eof some i n s e c t s . Heavy infestation h a s been suggested a s a source of trout mortality. Proteocephaius 9 . This tapeworm i s found in a wide variety g r i freshwater f i s h e s throughout the world. Copepods a r e the intermediate h o s t s for the only larval s t a g e . Twenty-three of the 55 specimens were infected, with 8 of the specimens containing more than 30 of the parasite. Generally, alder individuals were more heavily parasitized. Both s e x e s were equally infected. A l l parasitic infestations of l a k e trout were relatively mild. Ira no c a s e w a s t h e infestation judged t o be of sufficient severity t o appreciably impair the functions or performance of the iniected individual. SUMMAKY . 1 All limnological conditions, including water temperature, indicated Paxson Lake a s a significantly more productive l a k e than Summit Lake. Factors such a s elevation and surface-to-volume ratio were considered the main r e a s o n s for the difference in productivity. 2. A11 fish s p e c i e s present in Summit Lake were a l s o present in Paxson Lake. However, Paxson Lake p o s s e s s e d one s p e c i e s , the lake whitefish, nor found in Summit Lake. 3 . The c a t c h per unit effort of round whitefish in Summit Lake w a s more than twice t h a t of Paxson Lake, and even exceeded t h e combined c a t c h of both s p e c i e s of whitefish in Paxson Lake. No significant difference in c a t c h per anit effort w a s found for lake trout in the two- l a k e s . Lake trout in Paxson Lake grew f a s t e r than t h o s e in S.umira~tLake, Difference in growth rates t o a g e V w a s apparently correlated t o the reiatively greater invertebrate fauna in Paxson Lake. Differential growth r a t e s were especially pronounced between a g e V and a g e X , and were likely a restit: sf changes in diet from invertebrates t o small fish. 4. . The number of small f i s h suitable for l a k e trout food in Paxson Lake due chiefly t o the larger numbers of salmon fry and smolts greatly exceeded t h o s e in Summit Lake. Colder w a t e r temperatures in Summit L a k e during t h e ice-free period were a l s o partly responsible for slower growth. After age X , t h e samples were considered too small t o make valid conclusions. Growth of lake whitefish appeared t o be constant until a g e X; thereafter a d e c r e a s e in the growth rate w a s evident. ACKNOWLEDGEMENTS The author w i s h e s t o sincerely thank K. Neiland for compiling data on t y p e s and numbers of parasites; Dr. J . M . Morrow f o r u s e of laboratory equipment and a s s i s t a n c e ; and Dr. L. L. Hufman and S. Moore for a s s i s t a n c e and valuable information on the past history of t h e fishery; and Richard Haley , Dingle-Johnson Coordinator. , LITERATURE CITED KENNEDY, W . A , 1953. Growth, maturity, fecundity and mortality in t h e relatively unexploited whitefish, Coresonus clupeaformis , of Great Slave Lake. J. F i s h . R e s . Bd. Canada 1 0 (7): 413-441. KENNEDY, W. A . 1954, Growth, maturity and mortality in t h e relatively unexploited l a k e trout, Cristivomer namaycush, of Great Slave Lake. J. Fish. Res. Bd. Canada 1 1 (6): 827-852. MILLER, R, B. and W. A . KENNEDY. 1948. Observations on t h e l a k e trout of Great Bear L a k e . J . Fish. R e s . Bd. Canada 7 14): 176-1 8 9 , RAVVSON , D. S. 1950. T h e physical limnology of Great Slave Lake. R e s , Bd. Canada 8 (I): 1-66. J. Fish The Alaska Department of Fish and Game administers all programs and activities free from discrimination based on race, color, national origin, age, sex, religion, marital status, pregnancy, parenthood, or disability. The department administers all programs and activities in compliance with Title VI of the Civil Rights Act of 1964, Section 504 of the Rehabilitation Act of 1973, Title II of the Americans with Disabilities Act of 1990, the Age Discrimination Act of 1975, and Title IX of the Education Amendments of 1972. If you believe you have been discriminated against in any program, activity, or facility, or if you desire further information please write to ADF&G, P.O. Box 25526, Juneau, AK 99802-5526; U.S. Fish and Wildlife Service, 4040 N. Fairfax Drive, Suite 300 Webb, Arlington, VA 22203 or O.E.O., U.S. Department of the Interior, Washington DC 20240. For information on alternative formats for this and other department publications, please contact the department ADA Coordinator at (voice) 907-465-6077, (TDD) 907-465-3646, or (FAX) 907-465-6078.
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