Fishery Resources Theodore R. Merrell, Jr. Northwest Fisheries Center, Auke Bay Fisheries ~aboratory,National ~arineFisheries Sewice, of the National Ocear~icand Atmospl~ericAdministration, Vestern Aleutians Auke Bay, Alaska

Tlte fishery resources in the zuestent Aleutian Islnnds are zuill probably recover somezuhnt from tlieir preseat low diverse, nbtrnrlant, nrid heavily exploited, primarily by levels if average enuironniental conditions prevail and Japanese nnd Soviet fishermen. Seven groups make up the internationnl regulntions are successft!l in co,ttrolling the bulk of the crcrrent catch: snlmo~t (sockeye, chum, and high-seas cntch. King crnb harvests by U. S. fisher~ne,t pink), king crabs, Pacific hnlibut, Pncific ocean perch, shotrld grndrcally increase ns preuiotrsly atres~~loitedstocks sablefish, wnlleye pollock, mid Pacific cod. Three species of are fished. Pacific halibi~l,snblefislt, nnd Pncific ocean perch whales (syenn, fin, and sei) are also caplared. Tlre marine stoclrs nnd catcites will probnbly continue nt current low enuironmerrt is highly prodirctiue and is relaliuely trn- leuels. Il'alleye pollock a~tdPacific cod catches will prob- nffecterl by ,,ton's activities otlter fhai~fishing. nbly incrense somewhat. il'liole stocks will continue their Prospects for co,ttinaed or espanded fishery ltnruesls precipitous decline, nnd tlieir harvest will probnbly end vary nccording to species. Pink mid cltt~ersalmon stocks will within n few years nsn coaseq~~e~zceof co~rseruntion efforts probably remnin at about current levels. Sockeye stocks and internntionnl treaties.

The fishery resources around Amcliitka Island are the \vestern Aleutia~ls to fish virgin stocks of abulldailt and diverse, but rtiitil recently man Ims Pacific ocean perch, Sebnstes nlutus. During this used them to only a limited extent. Although the saliie period both nations expanded tlieir whaling original Aleut inhabitants of Amcliitka depended fleets tliroogliont tlic North Pacific Ocean. on the sea for most of their food, they had little Soviet or Japanese tra\vlers were frequently impact on the fish stocks because fishing n~ethods observed fishing off Amchitka during the Atomic were primitive. Their refuse middens, formed Energy Co~nmission (AEC) occupation of the during several milleni~uns of continuous occupa- island. Direct evidc~lceof the magnitude of these tion, consist almost entirely of the skeletal remains Japanese and Soviet fisheries is tllc treincndous of maine fishes, inanlmals, and in\'ertebrates accumulation of derelict fishing gear on (Desautels et ill., 1970). The size of the original Aiiichitka's beaches (Fig. 1). In 1972 I sitrveyed Aleut population is uiikrto~r~~~,but all cstiniates are 10,000 111 (33,000 ft) of these beaclies to provide a over 10,000, and nearly all the Aleutian Islands basis for estimating the total amount of fishing were inhabited; on Amchitka alone, 73 separate debris on tlte island. I estimated that at least occupation sites have bee11 identified (RlcCartncy, 22,000 items ~iladeof synthetic inaterials were on Chap. 5, this volume). After the Alc~~tianswerc the island beaches, including nearly 12 tons of discovered by tlic Western \Yorld in the eighteentli discarded scraps of tra\\rl web md 6600 salino~t ccntury, the Aleut populatioii d\vindlcd to less gill-net floats (Anonymous, 1973). than 1000 for the entire Aleutiail chain. As a By 1971 the Soviet and Japanese fislliilg effort result, for the 100-year period bet\\.een the mid- in Alelltians llad declined, and ollly tile nineteenth and tlie mid-tweiitietll centuries, the Japanese high-seas sal111on gill-net fishery was fishery resources \\,ere disturbed very little by man. contilluillg at a le\,el of effort colnparable to In the 1950s fish stocks in the western Aleu- peak years in the 1960s. The stocks of Pacific tians were exploited on a large scale on offshore or ocean perch and ~vhaleshad been so diininished by deepwater species that had not becn available to intensive fishing that they could no longer support the Aleuts. In 1952 Japan expanded the scope of the previous high levels of effort. its high-seas gil1.net fishing for salmon, O~ico~l~y?i- Although this relati\cely lo\$, fishing effort. is chlts spp., from Asian waters to the western continuing, the basic productivity of the seas in the Aleutians, ;md in tlie early 1960s Japan and the western Aleutians is unimpaired, and it seems Soviet Union sent enormous fleets of tra\vlers to ine\ritablc that additional species as well as renlna~~t 315 Alaska's continental shelf in competition with the foreign fleets. So far there has been little direct competition between the United States and foreign fleets in the western Aleutians. Several international agreements restrict the foreign fleets to species and waters not fished by U. S. fishermen, \vhose catches come mostly fro111 waters inside the 4.8-km (3-mile) territorial limit. Also, the United States has no distant-water trawl fleets, a reflection of the lack of ecollomic incentives to the U. S. fishing industry uilder institutional and market conditions that have prevailed. Discussion in this chapter is limited to the current status of commercially exploited fish and whale stocks around Amchitka. Other sectioils of this volume treat related topics, e.g., physical oceanography (h81cAlister and Favorite, Chap. 16), marine fish communities (Simenstad et al., Chap. 19), sea mammals (Abegglen, Chap. 20), and sea otters (Estes, Chap. 21). The area of coilcern is arbitrarily defined as those waters between longitude 170°E and 170°\\1 and latitude 50% and 55"1\7, which coincides with the area designated as Aleutian area 5 by the Itlterllational North Pacific Fisheries Commission Fig. 1-Japanese salmon giIl.net fragment washed up (INPFC) for reporting catch statistics. Amchitka on Amchitka Island beach. lies in about the center of this area (Figs. 2 and 3). The continei~tal shelf to the east of Amchitka, or recovered stocks of previously fished species \\rill including the Bering Sea and northeast Pacific again be exploited. Ocean, has very luge stocks of fish and intensive fisheries, but they are not considered here. The fishery resources in the wester11 Aleutians are now used only by Japan, the Soviet Union, and Only the major species currently sought within the United States. Other nations, however, may the area of concern by U. S. and foreign fishermen soon begin fishing there. Sout11 Korea has already are discussed here. These include sockeye salmon done some exploratory fishing, and Poland, (0.zerka), chum salmon (0.keta), pink salmon North Korea, \Vest Gerinanjl, wd the People's Re- (0.gorbusclta), coho salmon (0.kisutclz), chinook public of China have expressed interest in fishing in salmon (0. fshawytscha), Pacific ocean perch, the area. sablefish (Anoploponza filnbria), walleye pollock United States fisheries in the western Aleutians (Tlleragra clialcogmnt~~a),Pacific cod (Gadus mac- have been for Pacific halibut and crab, xvhich have rocephaltrs), Pacific halibut (Hippoglosstrs steno- a high unit value. These U. S. fisheries have been lepis), king crab (Pamlitllodes can~tscl~atica),sperm far exceeded it1 intensity, total value, and geo- whale (Pltyseter catodotz), sei whale (Balaenoptera graphic extent by the fisheries of Japan (Fig. 2) borealis), and fin whale (B. physahs). and the U.S.S.R. (Kg. 3), ~vhicl~concentrate 011 Other coini~~ercialspecies it1 this area, which species with a low unit value. In 1972, for are not included in the discussion because they example, over 1300 Japanese aud Soviet vessels make up only a minor portion of the current catch, \\?ere employed in fisheries off Alaska's coast include tanner crab (Clzionoecetes tnn~teri),Dun- (National Marine Fisheries Service, 1973). The geness crab (Cancer ntagister), arrowtooth flounder situation is changing, ho\ve\~er,because the target (Atherestltes stonzias), yello\vfin sole (Linlariada species sought by the foreign tralcrl fleets are being aspem), mck sole (Le/~ic/opsetta bilitieata), and depleted and U. S. fishing is expanding to include flathead sole (Hif>/>oglossoideselassodon). previously ni~exploited crab stocks in the western There is no single source of statistics for all Aleutians. Furthermore, as a result of changing species caught in the western Aleutians, but I have ecoiloinics and regulations, the U. S, fishing indus- included catches for the most recent year available try is exploring the feasibility of tra\vling on (usually 1970 or 1971). Fishevy Reso?~~cesof the ll'estern Alelctians 31 7 i

Fig. 2-Japanese fishing areas off Alaska, 1971 (excluding whales).

45' I I I I 170° 170' 1509 1 309

Fig. 3-Soviet fishing areas off i\laska, 1971 (excluding whales). DATA SOURCES foreign and U. S. fishc11nc11and discitss tlte geo- gaphic distribution, relative abundance, and biol- Detailed information on the fishery resources ogy of each species. A paper by Reeves (1973) in the immediate vicinity of Amchitka Islancl is provides similar but more current data for the Gulf scanty. Despite the limitations of available data, it of Alaska. Chitxvood (1969) provides a concise is important to include in this \~olumea risumi of history of Japanese, Soviet, and South ICorean what we kno\\, or can infer about the fishery Fisheries off Alaska through 1966 and describes resources of the western i-ileutians because they are fishing vessels, their numbers, ;u~dwhere they fish. potentially the nlost econo~llically\~aluable co111- I-Iitz (1970) gives detailed descriptions of vessels ponent of the Amchitka marine ecosystem. and fishing methods used by Soviet tra\vl fleets. The only assessment of local resources \\'as ICasahara (1972) reviews Japanese distant-\\rater limited sanlpling of nearshore fish stocks by the fisheries throughout the \vorld; he emphasizes the University of \\'ashington Fisheries Rcsearch Insti- role of international agreements and predicts that tute (FRI) during the AEC testing program. Al- the prospects for continued espansiort are not though Japanese and Soviet scientists investigated bright. Hart (1973) is one of the no st valuable some aspects of the fishery resources in the general references for information on distribution vest ern Aleutians during their hea\~csploitation and life history of Pacific Coast fishes. A3y discus- of demersal fish stocks in thc 1960s, most of the sions of individi~dspecies are drawn principally results of this research base not been published. In from these sources. a discussion such as this, the published statistics are adequate to describe the general characteristics of FISHERY RESOURCES the commercially exploited resources. The best published sources of catch data are A caveat that applies to any discussion of the INPFC, the National h'larine Fisheries Service abundance of ocean fish stocks is that it is seldom (NMFS), and the information exchanged annually possible to directly and precisely determine the since 1969 between the Soviet Union and the nu~nbersor biomass of any species. Usually com- United States at bilateral agreement meetings. mercial fishcry catch statistics provide the best Each year the INPFC publishes a statistical basis for estimating relative ab~undanceof a species yearbook that tabulates by area and gear thc in a particular area. Catches by Japrulese fleets weight of each species caught in the North Pacific supply thcs.: data for the western Aleutians. Soviet Ocean ,uld in the Bering Sca by Japan, the catch statistics are not so useful as the Japanese United States, and Canada. The INPFC also pub- statistics because they are not reported in as ~nuch lishes the proceedings of annual meetings and detail and include only recent years. In using catch bulletins describing research conducted by the statistics as a measure of relative abundance, we three treaty nations on North Pacific fisheries. must assume that the catch reflects actual abun- The Nh,IFS Division of Enforcement and Sur- dance. This is ob\riously an imperfect procedure, veillance in Alaska publishes an alnnal summary of but it is the best recourse available. It is also observations on patrols of foreign fishing fleets off important to remember that stocks of species not the Alaska coast. These patrols are made by NkIFS citrrently sought by fishermen may be abundant enforcc~ncntagents in cooperation with the U. S. but are not represented in proportion to their Coast Guard. Their purpose is to enforce numbers in catch records. United States fishery laws wd regulations, police The best single source of information for fisheries subject to international agreements, and assessing the relative abundance of commercially maintain surveillance of estra-treaty foreign fish- sought fish in the western Aleutians (escluding eries. 111 1971 the patrols covered about salmon, crabs, ancl \\'hales) is tlte series of INPFC 145,000 km (90,000 miles) by ship ru~d about Statistical Yearbooks, \vhich contain summaries of 380,000 km (236,000 miles) by aircraft. Japanese catch statistics. In 1969 a new system of Several publications describe the general reporting catches was initiated. This system identi- methods of operation of Japanese and Soviet fleets fied, for the first time, catches of each species for in the Gulf of Alaska and in the Bering Sea ru~dthe the western Aleutians specifically (Table 1).Before biology and life history of com~nerciallyesploited 1969 catches for the western Aleutians were species, although none of these pertain specifically combined with those from other areas to the east. to the western Alentian fisheries. The nlost com- Despite the scverc reduction of stocks, the prehensive single source of general information is catch of PaciCic ocean perch is still greater than the thc study by Al\~erson, Pruter, and Ronholt catch of dl other species co~nbined. (1964). These inr~estigatorsdescribe the history Esperimental research traxvling by FRI from and methods of fishing for denlcrsal fishes by 1967 to 1972 provides insight into the condition Fishery Resources of the ll'esteri~Alezrtiails 319 Table I-Con~mercial Catch of Fish (hlctric Tons) in tbe \Vesten1 Aleutians hy Japan and tbe U.S.S.R., 1969 to 1971

Nation Pacific Arrow- and ocean Sable- Pacific tootb Yellorv- Rock Platllead \\'alleye Pacific year pe~.ch fish halibut flounder fin sole sole sole pollock cod Other

Japan 1969% 15,597 1,673 330 65 20 2 2 512 222 2,558 1970* 13,650 1,247 351 288 9 2 11 179 284 2,002 1971t 14,664 2,766 387 44 1 1 16 622 432 3,928 U.S.S.R.$ 1969s 23,172 1,453 1970 53,274 9,490 9,490 1971 7,190 170 2,535 1,653 1,322

'International North Pacific Fisheries Commission (1972a; 1973). tFisliery Agency of Japan. $United States-U.S.S.R. Bilateral Scientific hleetings, 1970, 1971, and 1972. 8U.S.S.R. trawl fishery statistics for 1969, 1970, and 1971. Pmtial catch (does not include first cquarter). of fish stocks in the immediate vicinity of and that pink salmon are third-most abunda~lt. Amchitka and generally corroborates tile relative Coho and chinook sahno~lare present in insignifi- abundance of species deduced from Japanese cant nnmbers compared with the other three catches. Tlle narrative summary by FRI of their species. catches from 1967 to 1972 (Sinlenstad et al., Salmoll in the uvestem Aleutians originate from Chap. 19, this volume) states that bottom trawl streanls in both Asia and North America. Rlost catches ranged betweet1 200 and 1000 fish per cl~umsalmoll in the western Aleutians are from hour when dense schools of Pacific ocean perch, rivers in the U.S.S.R. and Japan (Shepard, Hartt, Pacific cod, or walleye pollock were encountered and Yonemori, 1968), and most sockeye sal~noll and that catch composition typically ranged from are from -rivers in Bristol Bay, Alaska (R,Iargolis 40 to 900 Pacific ocean perch, 30 to 55 nalleye et al., 1966; Bakkala, 1971). The abtuldance of pollock, 20 to 45 Pacific cod, 20 to 30 sculpins, 6 pink sallnoll fluctuates greatly. During some years to 12 arro\vtooth flounder, 6 to 7 rock sole, aud 2 pink salmon of Asian origin predominate; in other to 6 Pacific halibut per trawling hour. years pink sallnoll from Alaska streams are most abundant (Neave, Ishida, and Mttrai, 1967). Pink salmon are the predotniuant species spawning in Five species of Pacific salmon occur in the streams on A~nchitkaand other Aleutian Islands, western Aleutians as juveniles and maturing adults. but their nulnbers are relatively insignificant. They spawn in Asian and North American streams There is increasing evidence that many of the and intermingle in the western Aleutians in varying chiuook salmon caught by the Japanese are from degrees during the 1 to 4 years they spend at sea. western Alaska rivers, sucl~as the Y~rkonand The best source of infor~nationon the relative ICuskok~\.im. uu~nbersof the five species in the westem Aleu- Fisheries. Only Japan fishes for salmo~lin the tians is the catch records of Japanrse fishing fleets. \vestern Aleutian area. Kasahara (1972) revielvs the Table 2 sho\vs the mother-ship catches of the history and developlnellt of this fishery, which Japanese high-seas fleet for each salmon specics began in its present forin in 1952 with the signing from 1952 to 1970. Although the effort has of thc International North Pacific Fisheries Con- reinained constant since 1962, the catches fluctu- ventio~tby the United States, Canada, and Japan. ate widely because of annual variations in abun- This treaty requires Japan to abstain from fishiug dame of each species. The only data on salnlon for salmon east of longitude 175°1\' and was ab~uldancein the immediate vicinity of Amchitka believed to protect from the Japanese fishery ~llost are FRI samples for 1969 aud 1970 (International immature saln~onoriginating from North American North Pacific Fisheries Comn~ission, 1972b, streams (Fig. 2). Subsequent reseaxch has shorvn Tables 8, 9, aucl 10). ICondo et al. (1965) provide a that some North American stocks (mainly sockeye, comprehensi\~e review of ocean distribution and but also sonle chums, pinks, and chinooks) migrate migration of salmon based on tag returns. All three to the \vest of this abstention line, and since 1963 sources indicate that sockeye and chum sabnon the United States and Canada have attempted, predominate in most years in the western Aleutians without success, to lnove the liile farther \vest. Table 2-Japanese hlotl~er-ShipSalmon Fishery Catch* (Thousands of Fish), Longitude 160'~to 175~~

Year Sockeye Pink Churn Coho Chinook

*International North Pacific Fisheries Commission (1972c),

Froin 1962 through 1971 the Japanese fleet curtailed its ow11 catch. The present downward has consisted of 11 factory ships and 369 catcher trend of sockeye salmon abundance will probably boats. The salmon are caught in floating monofila- turn up~vadif environn~entalconditions during ment gill nets set off the stems of the catcher boats the fish's early sea life return to normal and (Fig. 4). The nets have a mesh size of about 120 international negotiations are successful in restrict- mm stretch measure atld are 5 m (16 ft) deep. ing catches on the high seas. When all the catcher boats are fishing, more than Chum and pink salmon seem to be maintaining 5500km (3400 miles) of net is fished daily their numbers better than sockeye. Barring the because each catcher boat sets 15 linear kilometers collapse of international fishery agreements, which of net each day. The catcher boats transfer each limit to some extent the scope of Japanese day's catch to a factory ship where the salmon are high-seas fishing, the abundance of pink and chum cleaned and frozen whole or canned. The processed salmon in the western Aleutians can be expected to catch is transported to Japan where it is marketed continue at more or less the same levels as at locally and throughout the world. present. The fishery, which lasts from mid-bhy until If other nations not party to the present July or August (Chitwood, 1969), starts south of fishery agreements between Japan, the United the Aleutian Islands and moves progressively north States, and Canada were to begin fishing, the toward Asia and North America to intercept unrestricted fishing could result in the decimation mature adult salmon migrating toward parent of present stocks of all salmon species. spawning streams. The thousands of net floats and huge wads of derelict gill nets that have aecumu- ICing Crab lated on Amchitka beaches provide striking evi- dence of this massive fishing effort. The king crab resource is the most significant Future Prospects. In 1973 and 1974 the present and potential fishery resource for U. S. valuable Bristol Bay sockeye salmon lull declined fishermen in the western Aleutians. The precise to very low levels, apparently as a result of both size of this resource is undetermined, but it is the Japanese high-seas catch and adverse environ- substantial in localized areas. ICing crabs are not inental conditions that caused unusually high common at Amchitka. National hflarine Fisheries mortality of juveniles in the Bristol Bay estoaly Service biologist-divers saw a few juvenile king and lakes. To reverse this decline, the United States crabs during extensive underwater observations at has aggressively attempted to persuade Japan to Amchitka over the 5-year period 1969 to 1973, curtail its high-seas fishing and has drastically and biologists from FRI captured a few adults Fishery Resources of the Western Alet~tiatts 321

Fig. 4-Japanese catcher boat hauling in salmon gill net. Note salmon on deck and small net astern of the gill net to retrieve salmon that fall out of gill net as it is being hauled aboard. (Photograph courtesy of NMFS Division of Enforcement and Surveillance.)

during the same period. Litlzodes aeq2rispina, a for about 11 months aiid hatch in the spring just species siinilar to king crab, was captured by FRI before llloltillg alld inatilig again. The larvae are in research trawrls near Amchitka at an average rate plallktoilic and may be transported long distances of 4 to 9 per hour (Simeiistad et al., Chap. 19, this by currents before settling to the bottom (Hebard, volume). 1959; IIaynes, 1974). Adults also migrate loilg distances by \\ralking on the bottom. After they Biology and Life History. Icing crabs occur along the etltirc North Pacific coast from northern reaclt maturity, king crabs have few natural enemies. Japan to British Columbia, usually 011 sand or inud bottoms in water depths to 400 m (1300 ft). The Fisheries. The U. S. king crab fishery in the commercial fisheries are limited by law and inter- western Aleutians is coilcelltrated east of Ainchitka national agrecinents to males only, which grow around Adak Island, but solne exploratory fishing larger than females. Icing crab males may reach a has been done around ISiska and Attu Islands. weight of 11 kg (25 1b) and an age of 16 years, Significant coillmercial catclies have bee11 made on although the averages are 111uch less. At Icodiak, Petrel Bank, northeast of Sc~ilisopochnoiIsland. Alaska, recent average ~reightsof individual crabs The Soviets and Japanese are prohibited from in the catch have been about 3 kg (6 ib) (Haynes fishing for crabs in the western Aleutians under and Lehma~i,1969). Both seses mature at about terms of bilateral agreements ~vith the United 5 years of age. The fecundity of females ranges States. from 25,000 to 390,000 eggs (Haynes, 1968). The The U. S. fisherinell catch king crabs in baited fertilized eggs are carried extenlally by the fe~nale pots in waters to about 200 in (650 ft) deep. The pots, wllicll are about 2 m (6 ft) square and 1 nl fish, and large females may produce 3 million eggs (3 ft) deep, we set on the bottom and attacbed by annually. Pacific halibut migrate long distances; line to two buoys at the surface. The pots weigh newly hatched young are sometimes carried thou- about 150 kg (300 lb) and are hauled with a po\ver sands of kilometers from the spawning locations by winch. Crab vessels are 15 to 55 m (50 to 180 ft) currents, and tagged adults have been recovered as long [average about 30 m (100 ft)]. The crabs are far as 3700 km (2300 miles) fro111 the point of kept alive in tanks \\'it11 circulating sealvater until tagging (Bell, 1973). Pacific halibut are omnivorous they arc taken to a shore plant !\,here they are but feed predominantly on othcr fish and crusta- butchered, cooked, and frozen. The U. S. catches ceans, inclitding tanner and king crab. \vest of longitude 17Z0\V for the years 1969 to Fisheries. Pacific halibut along the North 1971 were as follo~vs(Alaska Department of Fish American coast are exploited by the United States and Ganc, 1972): and Canada. The catch is regulated by the Intcma- tional Pacific Halibut Co~nmission (IPHC). In addition, large numbers of juvenile Pacific halibut arc incidentally captured off the Alaska coast by the Japanese and Soviets while trawling for othcr species. This catch has contributed to the recent declines in North American Pacific halibut stocks. Altho~~ghPacific halibut constitute only a small Fishing is regulated by season and catch-quota percentage of the total traxvl catch, in aggregate, restrictions, which vary from year to year. these tra\vl-caught Pacific halibut amount to a significant portion of the Pacific halibut resource Fnture Prospects. Crab fishing by U. S. fisher- in the Bering Sea. The IPHC has reduced the catch nlen on unexploited stocks in the western Aleu- linlits of the North American setline fishery, but tians is likely to increase. the expected benefits have been negated by the gro\ving incidental catch. Pacific I-Ialibat In the United States and Canadian fisheries, Pacific halibut occur on the continental shelf Pacific halibut we caught \\,it11 baited hooks along the entire North Pacific coast from Japan to fastened to 1.5-111 (5-ft) lengths of light line California in waters to 1100 111 (3600 ft) deep (Bell (gangions) attached at regular inte;.vals to a heavy and St.-Pierre, 1970). There is no reliable informa- groundline (skate) lying on the sea bottom. Each tion on their distribiction and abundance in the skate (a unit of gear) is about 550 m (1800 ft) long western Aleutians, but catches by research and and is anchored and marked by surface buoys at co~n~nercialvessels indicate that Pacific halibut are each end. The gear is pulled with a poxver winch present in lo\\, abundance throughout the area after fishing for periods of a few hours to Inore (Bernard E. Skud, Intenlational Pacific Halibut than a day. The fish are cleancd immediately and Commission, Seattle, \Vash., personal communica- transported to shore plants where they are frozen. tion), During the recent AEC occupation of Pacific halibut fro111 the \\restern Alcutiarts Amchitka, sport fishing for Pacific halibut in constitute only a small part of the total United Constantine Harbor xvas a pop~~larpastimc, and States-Canadian landings. The fishing intensity in large Pacific halibut were so~netimeshooked on this area varies greatly from year to year because of baited setlines. The FRI personnel also ci~icght its distance from processing plants. The area is sonle Pacific halibut near Amchitka \\,it11 research therefore attractive to fishermen only \\,hen areas fishing gear (Simenstad et al., Chap. 19, this vol- to the east are closed to fishing. During recent LII~e). years the catch in the western Aleutians ranged Biology and Life History. Pacific halibut arc from 1 to 69 metric tons (BenlardE. Skud, among the largest fishes in the \vorld. The North International Pacific Halibut Commission, Seattle, American recorcl Pacific halibut [225 kg (495 lb)] \\'A, personal communication): \\,as caught at Petersburg, Alaska. The average size of Pacific halibut in the U. S. commercial catch is Canada, U. S., Total, about 15 kg (33 Ib). Fe~nalesmatnre later, live Year ~netrictons metric tons metric tons longer, go\\r faster than nlalcs (Bcll and St.-Pielye, 1970). Most large Pacific halibut are females. Thc felnalcs reach an age of more than 40 ycars, and males, about 30 ycars. The number of eggs produced is proportional to the \\'eight of the Fishery Resonrces of the II'esten~rlleutiatls 323 Future Prospects. Because of the limited size American Pacific ocean perch resource, but it is of the stocks and the distance from process;ng reasonable to assiume that \vestcrn Aleutian stocks plants, it is rullikcly that Pacific halibnt fishing in of Pacific ocean perch have follo\ved the over.111 the \\restern Aleutians will expand much beyond its pattern of declinc. Published data are not available pesent relatively lo~vle\,el. for catchcs of perch by Japan and thc Soviet Union for specific areas over a series of years. The most Pacific Ocean Perch useful data illustrating the rise and decline of Pacific ocean perch have been (and may still Pacific ocean perch catches are Japanese statistics for the Bering Sea region, \vhich includes the be) the most abundant fishery resource in the western Aleutians, but intensive fishing by fleets of \vestern Aleutians (International North Pacific Soviet and Japanese tra\vlers during the 1960s Fisheries Commission, 1973, Table 32): caused a precipitous decline in abundance. Catcll, Catch, Chitwood (1969) s~~lnnlarizesthe development of Year metric tons tretric tons these fisheries. 1961 13,002 50,308 Biology and Life History. Pacific ocean perch 1962 12,449 38,517 adults occur in dense schools throughout Alaska's 1963 17,257 54,347 continental shelf south of the Bering Strait at 1964 43,750 30,892 depths of 150 m (490 ft) to over 400 m (1300 ft), 1965 47,648 22,772 generally over rocky bottonls (hslajor and Shippen, 1970). During the first few weeks of life, the larvae The Japanese and Soviets use the same fishing are planktonic and drift with the currents. Since no methods for Pacific ocean perch and other dcmer- verified collections of perch between the larval sal fish. Schools of fish are first located by stage and year-old juveniles have ever been re- electronic hydroacoustic instr~unents, and then ported (Carlson and Haight, in press), the balance traxvl nets are dragged through the water to of their first yea is unkno\\m. Juveniles over the intercept the schools. These schools contain pre- age of one year live in relatively shallo\\~\\rater [55 dominantly Pacific ocean perch, but slnall numbers to 130 m (180 to 425 ft) deep]. In the of other species are also caught. The catch is fifth or sixth year of life, they lnatltre and join hauled aboard ship either over the side or over the adult offshore stocks in deeper water. They are sten (Fig. 5) and cleaned immediately (Fig. G). long lived and slo\\' grolving, reaching a maximu!n Most are processed into frozen fillets. Nearly all age of about 30 years and a nlaxinlum size of fishing is no\v by large stern-ramp factory trawlers about 45 cltl (17.5 in.) and 2 kg (4.5 lb) (Alverson of 50 111 (160 ft) to more than 100 m (320 ft) long and \\'estrheim, 1961). (Figs. 7 and 8). These tra~vlers are co~npletely Juvenile Pacific ocean perch (1 to 5 ycars old) self-sufficient, i.e., they are equipped to catch, feed mainly on copepods and eupha~~siids.Young process, and store the finisl~edproduct until it is juveniles (1 to 2 years old) prefer copepods, and transferred to a refrigerated transport ship (Fig. 9) older juveniles (3 to 5 years old) prefer euphausiids or until they return to the parent conntry. In the (Carlson and Haight, in press). 1960s smaller tra\vlcrs were common, but the Adults also feed on copepods and euphausiids trend is to\vard larger ships. but show a prefercncc for euphausiids. Othcr larger Both Japan and the Soviet Union generally fish food items, e.g., shrimp and squid, make up a in sunllner (hzlay-No\~en~bcr)when Pacific ocean significant part of their diet (Shalkin, 1964; perch are more vulnerable to captnre by tra\vling; Paraketsov, 1963; Lyubimo\la, 1965). in winter the perch are in deeper water -~sl~eries.-. Pacific ocean perch in the western (Gunderson, 1974). In 1971 betxveen 1 and 6 Aleutians are fished exclusively by the Soviets and Soviet tratvlers and 2 and 10 Japanese tra\vlers Japanese. The history of the fishery encompasses a operated in the western Aleutians (National Marine period of only a few years, i.e., from the first Fisheries Service, 1973). explorations of Pacific ocean perch stocks in the Future Prospects. Bccause of the depleted late 1950s by Soviet research \'essels to one of the condition of the stocks and because Pacific ocean lnost massive fishing operations in history in the perch grors very slo\\,ly, the present relatively lox\, 1960s and lo the present reduced level of effort. level of fishing effort will probably continue. During that brief period the large virgin stocks of Pacific ocean pcrch in tltc Bering Sea and Gulf of Alaska \\,ere greatly reduced, and by 1968 only 39% of the original stock biolnass remained (Quast, Sablefish are distributed throughout the North 1972). Quast's estimate \\,as for the total North Pacific Ocean from California to Japan. In thc ('aa!uaS sauaqscj aqreK leu~!~~ (-aquas sauaqs!j augem puo!aeN 'o~our!qs!~or!f'jo Lsaamoa qd&o?oqd) 'oaotqqs!~or!rjo Lsaunoa qdex8o)oqd) .ia[nzea asauedef e jo dmr urals aq -1aIMeq dmr-was preoqe qaxad ueaao Burmala ueuuaqsg asaueder-g 1aAo U! pap9STW wadueaao 30 (q~000'~) SUO? 3qam nz JO yme>-? *s Fisltery Resources of the Western Aleutians 325

it

Fig. 7-Japanese stern-ramp trawler Taiyo Alaru No. 81 at dock in Shimonoseki, Japan, before leaving on fishing voyage to Alaska waters. (Photograph courtesy of Jiro Nishimoto, National Marine Fisheries Selvice.)

Japanese catch in the \vestem Aleutians, sablefish years of age and at 9 years are about 100 cm are second in ab~uxdanceto Pacific ocean perch. (39 in.) long and weigh about 18 kg (40 1b). They do not comprise as great a proportion of the Spawning takes placc in decp water during the late total Soviet catch, presumably because the Soviets winter, and thc eggs and larvae arc planktonic. do not use longline gear (the most effective Small immature fish fxequent shallow bays and sablefish gear).* Sablefish were not common in inlets during the late spring and early summer, catches by FRI in the immediate vicinity of often in the surface waters. Food items consist of Amchitka Island (Simenstad et al., Chap. 19, Pacific herring (Clzrpea harengns), Pacific sand lance Table 3, this volume), but Kobayashi (1957) (Am~nodytesilexapterzcs), aud crustaceans. Taggiiig caught larval sablefish in the western Aleutians in has shown that sablefish migrate long distances, July and August of 1955 and 1956. e.g., from Alaskan \\raters to the Asian coast and Biology aid Life History. Sablefish are known from Puget Sound, \Vash., to the Bering Sea. by the common name black cod in Alaska, but Sablefish apparently spawn in the western Aleu- they are not taxol~onlicallyrelated to the true cods tians because Icobayashi (1957) captured larval and bear only a superficial resemblance to them. sablefish at nine locatious in this area. The following summary of what is known of Fisheries. Sablefish are caught by Soviet aud sablefish distribution and biology is derived from Japanese tra~vlersin the western Aleutians with the Reeves (1973). Adult sablefish prefer deep water same gear and in the same arcas as Pacific ocean and are caught to depths of 530 fathoms (3200 ft); perch. The Japanese also havc a longlinc fishery juveniles often occur in shallo~verwaters (even specifically for sablefish. The longlinc gear is intertidally). They reach maturity between 6 and 8 similar to that used by thc U. S. and Canadian halibut fisheries (see the section on halibut). From - 1969 to 1971, bet\vecn 20 aud 50% of the *U. S.-U.S.S.R. Bilateral Scientific Meeting, 1972; Japanese catch of sablefish in the western Aleu- U.S.S.R. tra~vlfishery statistics for 1971. tians was by longline. Fig. 8-Stern-ramp trawler Tniyo ~\lorrr No. 81. Complex superstructure and rigging is used to set and haul the trarvl net and manipulate the gear after it comes aboard. (Photograph courtesy of Jko Nishituoto, National Marine Fisheries Service.) Fishery Resources of the IVesteni Alelrtians 327

Fig. 9-Soviet refrigerated transport anchored in Aleutian waters while transferring fish products from stern-ramp trawlers. The transport is about 130 m (430 ft) long, and the two trawlers are about 85 m (280 ft) long. (Photograph courtesy of NMFS Division of Enforcement and Surveillance.)

Sablefish are a highly esteemed food fish in prime target species (Ieasahara, 1972; National Japan. They are usitally marketed filleted and i\,larine Fisheries Service, 1973), althougll Pacific frozen, bitt their fine testltrc and high oil content ocean perch still predoininate in western Aleutian also make them a supcrior salted or smoked trawl catches (Table 1). This recent shift of empha- product. sis to pollock is primarily a consequence of Fatore Prospects. The catch of sablefish in depletion of other species and of develop- the western Aleutians rvill probably continue at ments in food technology, which have generated a about the same level. Most will continue to be heavy demand for tninced pollock to use in making caught incidentally by fishermen trawling for other surimi, a fish paste that is blended with othcr food species, such as Pacific ocean perch. products (IvIiyauchi, ICudo, and Patushnik, 1973). Previously the demand for pollock as human food \\'alleye Pollock was slight in both thc Soviet Union and Japan. \\'alleye pollock (also kno\\rn as Alaska pollock Biology and Life History. Temperature appar- and Pacific pollock) \\,ere, until recently, the only el+ determines where pollock occur (Icasahara, remaining large unexploited stocks of fish off the 1961). They are caught at all depths where the North Anlericatl continental shclf (ICasahara, xvater tempcrature is between 0 and 10°C. Pollock 1961). The Japanese catch of pollock in the mature when 3 or 4 ycars old and form dense eastern Bering Sea region increased from 24,000 schools during spalvning migrations in winter and metric tons in 1961 to more than 1,200,000 metric spring. The eggs are ptanktonic, and the young tons in 1970 (International North Pacific Fisheries hatcll in about 12 days. Pollock com~nonlyreach Commission, 1973, Table 32). Pollock are appar- an age of 8 years, but soine reach an age of ently abundant around Amchitka (Simenstad et a]., 15 years and a \\'eight of 18 kg (40 lb). The Chap. 19, Table 3, this volume). Sosict and primary foods are planktonic crustaceans, mostly Japa~icse trawlers are s\\ritching to pollock as a cuphausiids, althoug1:h shrimp and fish are also important. The diet varies in different geographic Fisheries. Pacific cod in the western Aleutians areas and is probably determined by the availa- are caught incidentally by Soviet and Japanese bility of food. trawl fleets and by Japanese longliilers fishing for Fisheries. \\'alleye pollock are caught mostly sablefisll. Soviet catches are geater than Japanese by large steni-ramp Japanese aild Soviet trawlers catches; e.g., in 1971 thc Soviets caught an with the same type of gear as is used for Pacific estimated 1653 metric tons of Pacific cod in the ocean perch. In 1971Japanese trawlers catiglit 622 rvestern Aleutians whereas the Japanese landed metric tons of pollock in the western Aleutians, only 432 metric tons. Pacific cod are filleted and and Soviet trawlers caught 2535 metric tons in the frozen and are considered a delicacy. same area (Table 1). Future Prospects. The catch of Pacific cod in Future Prospects. Future prospects are 11x1- the western Aleutians is likely to increase as other certain. Although the pollock catcli and demand target species decline in abundance because the are increasing, there is evidence in the eastern large stocks are still only lightly exploited. Bering Sea, where the largest catches off Alaska are made, that catch per unit of fisliing effort is dropping and the bulk of the catcli iucreasingly coiiiprises young and immature fish (Pruter, 1973). h,Iany aspects of the biology and distribution of The Bering Sea situation may be different from rcrhales arc poorly known despitc more than 100 that of the western Aleutians, however, and there years of worldwide exploitation. There is conse- appear to be substantial stocks of pollock that have quently little information about whale stocks only begun to be exploited. Therefore it is likely around Amcliitka. Behvecn 1968 aiid 1973 whales that catches near Amchitka will increase. were occasionally sighted fro111 shore by personnel at Amchitka, but their species was not verified. Pacific Cod Several dead sperm whales washed ashore on One of the earliest commercial fisheries in Amchitka during thc AEC occupation. Alaska \\.as for Pacific cod. Stocks in the Aleutian R,Iackintosh (1965) and Slijper (1962) are two Islands and Bering Sea were first esploited corn- of the best sources of general inforillation on inercially in 1864, and their great abundance Tvas whales and the whaling industry. Nishi~vaki(1966) first noted and recorded by a Russian navigator in describes the distribution and inigratioli of \\.hales 1765 (Cobb, 1916; 1927). The catcli by U. S. in the North Pacific Ocean on the basis of the kill fishermen reached its maximum in the early part of by the Japanese whaling industry between 1945 the twentieth century; in most years between 1905 and 1962. In the Rat Islands area (which inclitdes Amchitka), inore sperm whales (Pl~ysetercatodox) and 1925, the average catches were between 2.5 \\,ere taken th;ul any other species, but blue ~vliales and 3.9 million fish (icasahara, 1961). Pacific cod (Balae~~optera~n?~sculzis), humpback whales (fife- fishing by U. S. fisheniien has been at a low level off Alaska for many years because of the lack of gaptera novaea~lgline), fin whales (Bnlae~~optera U. S. market demands. Pacific cod are still widely pl~ysalirs),and sei whales (Balae~loptera borealis) were also taken. Presuniahly these five species are distributed and abunda~ton Alaska's contineiltal still present, although the killing of blue aud shelf, and tlie preseut stocks arc virtually unex- Iiccmpback svhales is 1101~prohibited, and their ploited. They are apparently abmidant around occurrence is no longer reflected in whaling sta- Anichitka in offshore waters (Isakson, Simenstad, tistics. and Burgner, 1971). Several dead Pacific cod were found washed ashore on Amchitka after tlie Biology aid Life History. Only the three Cannikin explosion. species of whales still hunted and occurriug in the Biology and Life History. Pacific cod live uear Aleutiaus (sperm, fin, and sei) arc discussed. the bottom at depths to 400 m (1300 ft) around The sperm whale is a toothed wliale that the entire North Pacific rim from California to reaches a length of 18 m (60 ft). Spertii \vhales are Honsltu, Japan (Icasahara, 1961). They apparently polygamous, and the ~ilalesarc much larger than migrate to shallow insliore areas in summer and to the females. Their distributioil is tvorldwide except deeper offshore areas in rvinter; these movements for polar regions. Several species of squid arc their are related to temperature. Females have between primary food. 1.4 and 6.4 lnillion eggs; they may reach a In tlie northern hemisphere the fin whale, maximum weight of about 18 kg (40 1b) after 15 a baleen wliale, reaches a length of 24 111 and is years of growth, although size, age, and fecundity second in size only to the ncarly extinct blue vary widely. They feed on a variety of inverte- \\,hale. Fin \vhales are monogamous, as are all brates and fish. baleetl whales. Their distribution is worldwide; in Fisfrery Resolrrces of the Hl/ester~tAletctio~ls 329 Alaska waters their distribution extends into the industrial products. Sperm whale oil is used only Arctic Ocean in summer. They are filter feeders, for industrial oils, whereas much of the oil of straining planktonic crustaceans and s~nallfishes baleeu whales is used for edible products, espe- from the water by horny bristled plates (baleen) cially margarine. The meat of all species is used for that are attached to the roof of the mouth. A large human and pet food; that of baleen \vhales is fin \\,bale may have as many as 400 of these plates similar to beef in palatability. spaced about half an inch apart on each side of the upper jaw (&lackintosh, 1965). Futnre Prospects. The numbers of whales Sei whales are relatively slnall baleen \vl~ales taken each year will probably continue to decline that reach a maximum length of 15 m (50 ft). because of restrictive regulatio~lsto prevent fitrtller Because of tlteir low oil content, they were not reduction of the stocks. Further, world opinion is harvested to any extent until recent years \vhen mobilizing against harvest of any whales. For these stocks of the larger species of whales were reduced reasons the high-seas whaling industry will proba- to the extent that hunting them became uneco- bly come to an end within a few years. nomical. They are widely distributed in subarctic waters throughout the world and are colnmon in [Author's Comments] Aleutian waters (Nishi\\.aki, 1966). Since the foregoing was prepared, the outlook for future foreign exploitatioll of fishery resources in the Fisl~eries. Japan and the U.S.S.R. are the only western Aleutians has been complicated by the probability nations tbat notv hunt whales com~nerciallyin the of unilateral action by the United States extending its Gulf of Alaska and in the Bering Sea. A severe fishery jurisdiction from 12 miles to 200 miles offsl~ore. worldwide decline in numbers of whales as a result Legislation has passed both houses of Congress, and the President is expected to sign the ultimate bill, wllich is of excessive killing and consequent restriction of likely to become effective by mid-1977. the kill by the International IVhaling Co~nmission Except for salmon and whales, most of the other are responsible for the present limited extent of corrently exploited fishery stocks of the western Aleutians the whalitlg industry. In 1971 the Japanese had 3 are rvitllin 200 miles of land. What action will be taken wit11 regard to these fishery resources and what the effect of this North Pacific Ocean fleets with 26 killer vessels; action will be is difficult to predict. the Soviets operated 2 factory ships aud 28 killer vessels (National Marine Fisheries Service, 1973). Table 3 shows the Japanese and Soviet pelagic REFERENCES whale kill for 1969 to 1971 in all North Pacific Ocean waters. Sperm ~vl~alesmake up the majority Alaska Department of Fish and Game, 1972, 1971 Alaska Catch and Production Commercial Fisheries Statistics, Table 3-Japanese and Soviet North Pacific Ocean Statistical Leaflet 23. \\'l~aleKill, 1969 to 1971" r\lverson, D. L., A. T. Pruter, and L. L. Ronholt, 1964, A Study of Demersal Fishes and Fisheries of the North Western Pacific Ocean, 1.1. R. hIachlillan Lectures in Nation Fisheries, University of British Columbia. and -, and S.J. \\'estrheim, 1961, A Review of the year Sperm Sei Fin Taxonomy and Biology of the Pacific Ocean Perch and Its Fishery, Rapp. P.-v Reuti., Cons. Int. Esplor. Mm., 150: 12-27. Anonymous, 1973, NMFS Finds Tons of Plastic Debris on Alaskan Island, NOAA Week, 4(March): 1. Bakkala, R. G., 1971, Distribution and Migration of lmma- ture Sockeye Salmon Taken by U. S. Research Vessels with Gillnets in Offshore Waters, 1956-1967, in International North Pacific Fisheries Commission Bulle- tin 27. Bell, F. H., 1973, The Pacific Halibut-A Managed *From National Marine Fisheries Service (1973). Resource, in A Review of the Oceanography and Renewable Resources of the Nortltern Gulf of Alaska, of the kill (Nishi~vaki, 1966). The \\,baling season D. H. Rosenberg ((Ed.),pp. 361-409, Institute of Marine occurs from hlay to October, and hunting takes Science, University of Alaska, Report R72-23 (Sea Grant Report 73-3). place far offshore. I11 1971 110 whales are believed -, and G. St.-Pierre, 1970, The Pacific Halibut, to have been taken near Alaska by tlle Soviets, and International Pacific Halibut Commission, Tecllnical only 622 whales, or 13%, of the Japanese kill was Report 6. from Alaska waters (National Marine Fisheries Carlson, H. Richard, and Richard E. Haight, in press, Service, 1973). Juvenile Life of Pacific Ocean I'erch, Sebastes nlut~ts,in Coastal Fiords of Southeastern Alaska: Their Environ- After the whale is hauled aboard the factory ment, Growth, Food Habits, and Schooliilg Behavior, ship, the entire whale is converted to food or Transactions of the American Fisheries Society. Chitrvoad, P. E., 1969, Jnpnaese, Souiet, nnd Sotrth Korean (Genus Oncorhy,rchus) Based on Tagging Studies Fisheries off Aloskn, Deuelopme,tt nnd History Tltroush (1958-1961), International North Pacific Fisheries 1966, U. S. Fish and \\'il:.il:I'e Service, Circular 310. Commission, Bulletin 171. Cobb, J. N., 1927, Pacific Cod Fisheries, Report of the Lyubimova, T. G., 1965, The Main Stages of the Life Cycle U. S. Commissioner of Fisheries, 1926, Appendix 7, of Pacific Ocean Perch (Sebostes alr~tusG.) in the Gulf pp. 385-499. (Doc. 1014). of Alaska, in Souiet Fisheries I>tuestigntion in the -, 1916, Pacific Cod Fisheries, Report of the U. S. ivortheast Pacific, Part IV, pp. 85-111. Commissioner of Fisheries, 1916, Appendix 4, Mackintosh. N. A,. 1965. The Stocks of iBhales. Fishine pp. 5-111. (Doc. 803). r\'err,s (Uooks), itd. Desautels, R. J., A. J. hlcCurdy, J. D. Flynn, and R. R. Maior. R. L.. axd H. H. Shiu~en. 1970. Svno~sisof Ellis, 1970, Arcllaeological Report, Amchitka Island, "~iolo~icalData on Pacific' 'Ocean Perch, ~ecnstodes Alaska, 1969-1970, USAEC Report TID-25481. nlattrs, FA0 Species Syilopsis No. 79, Circular 347, Gunderson, D. R., 1974, Availability, Size Composition, U. S. Department of Commerce, \\lashington, D. C. Age Conlposition, and Growth Characteristics of Pacific hfargolis, L., F. C. Cleaver, Y. Fukuda, and H. Godfrcy, Ocean Perch (Sebostes nlstes) off the Northern 1966, Salmon of tlze North Pacific Ocenn, Pt. 6, \\'ashington Coast During 1967-1972, j. Fish. RES. Sockeye Salmon in Offshore \jraters, International Board Ca~t.,31: 21-34. North Pacific Fisheries Commission, Bulletin 20. Hart, J.I.., 1973, Pacific Fishes of Cnttndn, Fisheries hliyauchi, D., G. Kudo, and h'l. Patushnik, 1973, Research Board of Canada, Bulletin 180. Surimi-A Semi-Processed \Yet Fish Protein. Alar. Fislt. Haynes, Evan B., 1974, Distribtction and Relntiue Abu,z- Reu., 35(12): 7-9. rlance of Lnrune of Kiq Crab, Paralithodes National hlarine Fisheries Service. 1973. Foreien Fishine camtschatica, in the Southenstern Beri~rg Sen, Activities, Bering Sea and Gulf ;f Alaska, 197c 1969-1970, U. S. National h'larine Fisheries Service, Enforcement and Su~veillanceDivision, Juneau, Alaska. Fishery Bulletin 72, pp. 804-812. Neave, F., T.Ishida, and S. Murai, 1967, Snlmoa of the -, and C. Lehman, 1969,hlinotes of the Second Alaskan North Pncific Ocenn, Pt. 7, Pink Salmon in Offshore Shellfish Conference, Alaska Department of Fish and JYaters, International North Pacific Fisheries Commis- Game, Information Leaflet 135. sion, Bulletin 22. , 1968, Relation of Fecundity and Egg Length to Nishiwaki, hl., 1966, Distribution and Migration of the Carapace Length in the King Crab, Pnrnlithodes Larrer Cetaceans in the North Pacific as S11ou.n bv cnmtschaticn, Pmc. ATnt. Shellfish Ass., 58: 60-62. ~a~anese\Vhaling Results, in IVlzales, Dolphins, nnb Hebard, J. F., 1959, Currents in Soetlreastenr Bering Sea Porooises, Kenneth S. Norris (Ed.),. . PP... 171-191, Uni- nttd Possible Effects Uf,on King Crab Lnrune, U. S. Fish versity of California Press, Berkeley. and JVildlifc Service, Special Scientific Report- Paraketsov, 1. \ 1963, On the Biology of Sebastodes Fisheries No. 293. nlutas in the Bering Sea, in Souiet Fislteries I,tuestign- Hitz, C. R., 1970, Operotion of the Souiet Trnwl Fleet off tion ill tlte Nortltenst Pacific, Part 1, pp. 319-327. the Iirosltingto!t Consts Dfrring 1966 nnd 1967, U. S. Translated by Israel Program for Scientific Translations Fish and \Vildlifc Service, Circular 332, pp. 53-75. (1968), Report 'rr 67-51203. International North Pacific Fishcrics Commission, 1973, Pruter, A. T., 1973, Development and Plwsent Status of Stntisticnl I'enrbook 1971. Bottomfish Resources in the Bering Sea, f. Fish. Res. -, 1972a, Stntistical Yearbook 1970. Board Can., 30: 2373-2385. -, 1972b, Annual report 1970. Quast, J. C., 1972, Reduction in Stocks of the Pacific -, 1972c, Annual report 1971. Ocean Perch, an Important Demersal Fislt off Alaska, Isakson, J. S., C. A. Simcnstad, and R. L. Burgner, 1971, Trmts. Ante,: Fish. Soc., 101: 64-74. Fish Communities and Food Chains in the i\mchitka Reeves, J. E., 1973, Groundfish of the Gulf of Alaska, in A Area, BioScietrce, 21: 666-670. Revier, of the Oceanography and Renervable Resources Kasahara, H., 1972, Japanese Distant-\Vater Fisheries: A of the Northern Gulf of Alaska, D. H. Rosenberg (Ed.), Review, Fislr. Bull., 70: 227-282. pp. 411-455, Institute of Marine Science, University of -, 1961, Fisheries Resources of the North Pacific Ocean, Alaska, Report 1172-23 (Sea Grant Report 73-3). Pt. 1, H. R. hlacMillan Lectures in Fisheties, pp. 1.135, Shepard, hI. P., A. C. Hartt, and T. Yoaemori, 1968, University of Britisll Columbia, Vancouver. Snbnon of the A'orth Pacific Ocean, Pt. 8, Chtlm Kobayashi, K., 1957, La~wne and Yoongs (Sic) of the Salmon in Offshore \\'aters, International North Pacific Sablefish, Anoplopo,nn fintbrin (Pallas), from the Sea Fisheries Commission, Bulletin 25. near the Aleutian Islands, Ball. jopnn. Soc. Sci. Fish., Skalkin, 17. A,, 1964, The Feedhg of Ocean Perch in the 23(8): 376 (abstract only). Bering Sea, in Soviet Fisheries I,cuestigntioa ill the Kondo, II., Y. ITirano, N. Nakayama, and AI. hliyake, 1965, ~\'orlheust Pacific, Part 11, pp. 159-174. Offshore Distribution ;u~dhligration of Pacific Salmon Slijper, E. J., 1962, llrltnles, Basic Books, Inc., New York. Oceanography W. Bruce McAlister Felis Fcrvorite United States Department of Commerce, National Marine Fisheries Sewice, North\\rest and Alaska Fisheries Center, Seattle, \\'ashitlgton

Circulation near Amcltitka Island is inflttenced by complex Dt~rittg spring and summer, constal turbzclence reduces patterns of exclta~rgennd mixing between tlte North Pacific thertnnl stratification in the surface layer, and ittsltore water Ocean and Bering Sea waters in tlre pnsses of the Aleutian tetnperatsres are 1 to 4'~ lozuer tlta,t in the offshore Islands, resulti~rgin wide unrintions in physical, chetttical, oceanic regimes. Zooplankton are typical stcbarctic Pacific and biological conditioi~sseaso,mlly, and also ouer short forltts. Zooplnsktoa uolunzes in tlte 0- to 150-m surfnce geopplric distances. Strong tuestwnrd flow in tlre A1ask;ft layer increased rorcghly one order of tnngrzitude from zuinter Stream south of A~ttchitknIslavtd brings tuater of 2 to 4 C to summer. A'utrie~rts do not nppear to be liatiting for into the area in tointer, nmeliornting sea and air tempern- phytopla~tktongrowth. ttcres and itrlribiting tlze deueloptnent of exte~tsiueshore ice.

-- The bulk of oceanographic researcll has taken place PHYSICAL OCEANOGRAPHY in those areas of tllc ocean rvl~ichpresent the most enticing lure to man. This attraction most often Subarctic Pacific Circulatio~l takes the form of a bountiful fishing ground or an The subarctic Pacific region of the North ecollo~nicalplactical sea route. Research data on Pacific Ocean is defined as bounded on the north the large circulations of water in and bet~veenthe by ~~~i~~~strait, lalld constriction at Bering Sea and the North Pacific Ocean lia\,e entrance to the Arctic Ocean, and on the south at acc~u~nulatcdmainly in co~lju~lctio~lwith tile ex- about 400~by a ill vertical water pansion of the area's fisheries and are only now ~l~~ southern edge, where marked temperature beginning to include detailed information on circu- inversions that are c]laracteristic of surface dillltion lation in the passes of tlle Aleutian Islands and in and elld alld marked salinity invex. the nearshore waters of the individual islands. sions that are characteristic of net surface evapora- The Aleutian-Commander Island arc divides tion begin, is generally referred to as the subarctic- the Bering Sea lrom the North Pacific Ocean along subtropic boundary and for convenience is a stretch of 1500 km. The arc is submerged in 39 specifically denoted by the 34 x0 isohaline in the- places, for~ningpasses and straits, but only 14 of surface layer (Fig. 1). The predominant oceanic these have an area greater than 1 km2 or a sill feature of the region is the large cyclonic subarctic depth deeper than 200 m. The series of passes g))re. Formed by a merging of cut~entsoff Japan among the Aleutian Islands is unique; the shdlour and driven by local winds, water in the gyre moves sills con~lecti~igmany of the islands block the free eastward across the North Pacific Ocean and exchange of intermediate and deep !\rater between sweeps around the Gulf of Alaska and northward the Bering Sea and thc North Pacific Ocean. Thc into and around the Bering Sea. Oceanographic deepest pass in the Aleutian portion of the island conditions in this region were extensively reviewed arc is Amchitka Pass, which has a sill depth grcater by Favorite, Dodimead, and Nasu (1976). Only a than 1000 nl. Because of exchange through this summary is presented llerc. pass, the oceanography of the Amcliitka Island Southward flow (Oyashio) along the north- area is related to offshore areas in both the North eastern coast of Japan and northward flow Pacific Ocean to the south and the Bering Sea to (Kuroshio) along the southeastern coast of Japan thc north. converge, mix, and turn east~vardacross thc North 331

Pacific Ocean, forming a transition zone between continental slope and then southwestward along the cold dilute subarctic waters and the warm the Siberian coast. At Cape Olyutorskiy a so~tth- saline subtropic waters. Although the demarcation wad branch forms an eddy in the norther11 Bering of these two flows is sharp and abrupt in the Sea, but the main flow continues southwestward western part of the ocean, it becomes gradually and completes the overall cyclonic circulation in more diffuse as stirring, advection, and diffusion the Bering Sea. The East I

Pig. 2-Vertical profile of all openings through the Aleutian-Commander Island arc and selective groupings of major ones.

Table 1-Depth and Area of the hlajor Openings in the Aleutia~l-Con~manderIsland Arc

General Depth, Area, opening PassIStrait m km2

East Aleutian Unimak 60 0.9 group Samalga 200 3.9 Chilginadak 210 1.0 Herbert 275 4.8 Yunaska 45 7 6.6 Amukta 430 19.3 Seguam 165 2.1 38.6 Central Aleutian Tanaga 235 3.6 group Amchitka 1155 45.7 49.3 \Vest Aleutian Kiska 110 6.8 group Buldir 640 28.0 Semichi 105 1.7 36.5 Commander-Near Near 2000 239.0 Strait Commander 105 3.5 242.5 Kamchatka Strait 4420 335.2 Total area 702.2

Alaskan Stream. This boundary current has speeds (northward) through my openings in the island of 50 to 100 cm/scc, and, although distributions of arc; however, this is not always the case. An water properties around the head of the Gulf of appreciable component of flow in the upper 200 to Alaska at times may indicate some lack of conti- 300 m is southward (the Aleutian Current), closing nuity in flolv in the surface layer, collti~luity circulatio~lin the surface layer of the Alaskan gyre. always exists belol\, 300 m. Normally the Coriolis At times inshore countercurrents with speeds in forces on the westxvard flow in the Alaskan Stream excess of 10 cm/sec occur along the continental would result in an elevation of the sea surface slope, but the continuity of such flows is poorly shoreward and a tendency for the water to not documented-these eastward flows may be associ- only hag the coastline but also to vcer to the right ated \\it11 large eddies in the Alaskan Stream. Tltomson (1972) described conditions dtat may in an attempt to ascertain the water exchange cause instabilities and separation of the flow from between the North Pacific Ocean and the Bering the coast. Sea. Arsen'cv (1967) summarized oceanograpliic Although the dominant flow in dte passes is conditions in the Bering Sea in an exhaustive tidal, there is increasing evidence dlat eddies exist analysis of station data prior to 1959. He consid- which encompass major island clusters and require ered the Commander-Near Strait as tlte most both north and south flows through tlte passes. significant source of flow into the Bering Sea Records of direct-current measuretnents in the (4.4 Sv),* whereas only 0.7 Sv occurred through Aleutian passes by the U. S. Coast and Geodetic the western groups; no net exchange was consid- Survey commencecl in 1934 with data collected ered to occur in the eastern Aleutian passes. These near Unimak Pass, and these data are of consider- transports appear derived from a reference depth able value in describing ~llaximumtidal velocities. of 3000 db, and it is also apparent that, except for However, estimates of nontidal components of a deep inflow in Icamchatka Strait, almost the these measurements are of questionable value entire flow illto the Bering Sea was derived from because of the limited time scale used in the the Alaskan Stream (Table 2). On the basis of measurements. Of the 18 records, 5 were of only continuity of the Alaskan Stream, Favorite (1974) 1-day duration; 13 other measurements taken from 1934 to 1946 covered periods of 2 to 8 days. Table 2-Flow (Sv) Through Aleutian-Commander Island Are Reed (1971) analyzed 25 surface tidal-flow records from radio current meters suspended from Transport out Transport buoys ancltorecl in various passes for intervals of of Bering Sea into Bering Sea over 4 days during 1949 to 1956. Northerly and Pass Arsen'ev Arser'ev Favorite southerly net floxvs were recorded at some loca- tions although at different times. Probletns ae Kamci~atka 18.4 encountered in such observatio~lswhen the tnove- Commander-Near 14.4 10.0 ment of tlte vessel or buoy may be greater than d~e !\'ertern Aleutian group 0.7 0 flow. This \\.as clearly demonstrated by obse~~a- Central Aleutian tions recorded aboard the U. S. Coast and Geodetic group 4.4 4.0 Su~veyvessel in Amchitka Pass (51°35'N, - - Sirrueyor Total 18.4* 19.5 14.0 180") in the late spring of 1963 when precise positioning at 15-min inte~valsindicated vessel "1.1 Sv loss through Bering Strait. speed as great as 85 cmlsec at anchor. The direct- current observations from the Amchitka area are summarized flow into the Bering Sea. Mean annual considered in detail in a later section. transport of 12.5 Sv in the Alaskan Stream soutlt Eight d~ift-bottleexperi~~tents \\,ere made be- of Adak Island was obtained ~vith4000 db (about txveen 1957 and 1972 in the Aleutian area 4000m) as a reference level. Because only (Favorite, 1974). Releases of bottles Isell south of Kamchatka Strait is deeper than 2000 m, 1.5 Sv of the Aleutian Islands from a merchant ship of this flow reaches this \vesternmost passage. opport~inityin tlte fall of 1964 implied a northerly Favorite found no net exchange in d~ewestern or drift from five of seven release locations southward eastern Aleutian passes. Of the llSv above of Adak Island, several recoveries being made on 2000 db, 4 Sv is lost through the central Aleutian Attu Island (the affinity of Attu Island for drift passes, 3 Sv is gained across the southern bound- bottles released from all directions .,\.as also re- ary, and 10 Sv flows northward through the flected in recoveries of bottles released from the Commander-Near Strait west of Attu Island. The research vessel George B. Kelez during ail oceano- rcsult is a net annual flow of 14 Sv into the Bering graphic cruise in the winter of 1966). One bottle Sea. This result falls bet\seen the transport values released south of Amchitka Island in 1964 was of 8.0 Sv proposed by Batalin (1964) and 19.5 Sv recovered on the Pribilof Islands. T\vo bottles proposed by Arsen'ev (1967). released north of Amchitka Pass in 1971 were recovered on the P~ibilof Islands (Favorite and Ingraham, 1972), but bottles released to d~enorth WINTER DATA 1968 and south of this location \\,ere recovered on Amchitka Island (Favorite and Ingraham, 1973). In Observations of temperature, salinity, and se- general, drift-bottle recoveries have reflected flow lected chemical properties were conducted in implied by the distribution of water properties. A number of authors have examined the density and \\rind-stress fields in the Aleutian area 336 Alcrilister and Favorite January-Ailarch 1968 in support of the AEC The coldest surface temperature (2.7OC) was ob- Supplemental Test Site Program on Amchitka served in the western portioll of the study area. Island (Fig. 3). These data permit a description of The wannest water (3.4 to 3.7OC) was also the marine environment in the vicinity of relatively lolv in salinity and characteristic of water Amchitka Island. transported westward by the Alaskan Stream. A brief drogue and dye current study was Water in the island passes, intermediate in tempera- conducted in February 1968 (IvIcAlister et al., ture (3.0 to 3.2"C), represents a mixture of the 1968). One 10-m parachute drogue was set in the warm Alaskan Stream water with cooler water channel north of Amchitka Island and followed for from the Bering Sea. Near the shore Alaskan 5 hr, during~vhichtime it moved co~u~terclock~vise Stream water occurred as a core of warm water at 25 to 40 cm/sec. Additional drogue studies were with a maximum temperature of 3.8'C between undertaken south of Adak Island in the summer of 150 and 400 m (see Fig. 6). Although inversions 1969 (Favorite, Ingraham, and Fisk, 1972). These and layering were frequent, continuity of the core studies again indicated the strong tidal component structure through Amchitka Pass indicated a net of current in this area and the great difficulty of north\vard flow of Alaskan Stream water. Below attempting to derive meaningful net current values about 500 m the temperatures decreased gradually from a limited program of disect current measure- with depth, and the vertical distribution of temper- ments. ature was similar in structure to salinity.

Temperature Salinity Surface temperatures in the North Pacific Collsiderable freshwater runoff enters the Ocean are normally lowest during March when coastal area of the Gulf of Alaska and is advected winter convective overturn forms an isotllermal westward south of the Aleutian Islands by the layer that may extend from the surface to 100 m. Alaskan Stream into the western North Pacific

Fig. 3-1,ocations of oceanog~aphicstations during Amchitka study, January-March 1968 (Research Vessel George B. Kelez). Fig. 4-Temperatoredistribution on sigma-t surface = 26.5 (100- and 1000-fathom isohatbs shoxvn). Ocean. Sharp fronts and eddies form at the nated by and generally similar to the distribution confluence of the Gulf of Alaska rvater ant1 the of salinity. The greatest stability occurred well Inore saline Bering Sea water that is usually present offshore ~vitlli~~ter~nediate stability in the Alaskan at the Aleutian Passes west of Unimak Island. This Stream and the xveak to neutral stability in the feature is present even during winter when runoff Aleutian passes. to the Gulf of Alaska is diminished by freezing and Water masses mix most easily along sigma-t \\,hen seasonal overturn causes mixing of surface (equal-density) surfaces. The distribution of tern- water with ]nore saline water to depths of about perature 011 a sigma-t surface indicates the extent 100 m. At the axis of the \\resttvard flow is a of horizontal nixing and suggests movement of tongue of dilute water (<32.8 Xo). Bering Sea water masses at various depths. South of Adak water (characteristically > 33.0 x0)was not fo~u~d Island the temperature distributions on the south of the entrance to Amchitka Pass. Offshore, sigma-t = 26.5 surfaces showed tongue-like exten- North Pacific water had a sligl~tlyhigher salinity sions westward toward Amchitka Island (Fig. 4). (-32.9 yo0)than Alaskan Stream \\rater. These extensions represented a westward advection Salinity increased with depth; the most rapid at depth of relatively warm water by the Alaskan change occurred in the well-developed halocline Stream from its source in the Gulf of Alaska. At between 100 and 300 m. South of the island arc, shallow depths, which are influenced by seaso~lal characteristic ridging of salinity and temperature overturn (0 to 100 tn), the coldest water was south was seen at all sections. The sharpest halocline was of Amchitka Island near 51°N, and water of well offshore at about 50°30'N. Although surface intermediate temperature occurred north of dilution in the Alaskan Stream would be expected Amchitka Pass in the Bering Sea. At depths greater to enhance a strong coastal halocline, the ltalocline than 150 m (sigma-t - 26.0 to 26.5), the coldest was less developed toward shore and weakest in the water was in the Bering Sea, and water of turbulent passes where high-salinity water of the intermediate temperature occurred south of Bering Sea was mixed in the surface layer. Amchitka Island. This reversal of temperature gradient ~vithdepth suggests that the warmer water Density of the Gulf of Alaska and the Alaskan Stream had mixed northward into the Bering Sea; at shallo~v The distribution of density, calculated from depth, belo\\' sill depths, however, Iiorizontal mix- the observed temperature and salinity, was domi- ing is restricted by the barrier at the island arc. i\icAlister attd Favorite

Fig. 5-Geopotential topogaphy (dynamic meters referred to 1500 decibars) indicating surface flow.

the Bering Sea. This transport can be compared Except for north-south tidal excursions with the average annual figure of 4 x lo6 Sv given through the shallo\v openings in the island arc (and by Favorite (1974). The flow was highly turbulent; probable shelf waves), westward flow south of the geostrophic velocities in eddies were calculated in Andreanof Islatlds follo\vs the general zonal trend excess of 40 cm/sec and may have been two or of isobaths from 172'\\' to Amchitka Pass where three times this value for periods of short duration. an irregular bathymetry is encountered. Consider- A series of direct-current obse~~ationswere able mixing results ivhich is due not only to flow made from the vessel Stt~veyor at 51°35'N, funneling through channels between the islands 179"57'E (about 40 km northeast of the eastern lining the eastern side of the pass but also to the end of Amchitka Island in about 1900 m of water) impingement of eastward and southxvard flow from on June 3 to 6, 1963. A plot of directions and the Bering Sea at the western side of the pass. speeds at 10,100, 200, and 300 m recorded each Roughly 20 to 30% of the flow in the Alaskan hour (commencing alternately at 10 m and at Stream moves northward through Amchitka Pass, 300 m) indicate the complexity of flow evident in and Amchitka Island appears to be near the critical a data series for June 5 (Fig. 7). Isolated, abrupt, focal point where this divergence occurs (Fig. 5). brief deviations from the direction of mean surface East of An~cllitka Island an eastward counter- flow from 0 to 300 m occ~uredat 10 m; at 10 and current occurred sea~varcl of the edge of the 100 In; at 100 and 200 In; and at 100 and 300 m at conti~lentalshelf. various inte~~rals.An extended period of flow Flow through Amchitka Pass is extremely (0400 to 1100 hr) with a consistently uniform complex at all depths. North-south vertical sec- easterly direction, opposite the offshore flow in tions through Amchitka Pass indicated a northerly the Alaskan Stream, was also indicated near this current on the east side of the pass and one flowing same location in the 1968 data (see Fig. 5). to the south 011 the !vest side at a depth as Velocities at all levels varied from about 25 to determined from the distribution of properties about 50 cm/sec, rarely in phase or with a specific (Fig. 6). Relative to 1500 db, a net northward relation to increasing rlepth; in fact, in several transport of 5 x lo6 SV was observed flo~vinginto instances maximum velocities occurred at 300 m. LATITUDE 52"N 5lQ30'N 50°30'N 52*N 51°30'N 51°N 61'30'N

TEMPERATURE,^^ SALINITY. %o

Pig. 6-Vertical sections of temperature and salinity from Semisopochnoi Island to 50'30'~.

GEOCHEMISTRY tions in the central Bering Sea are undersaturated by over 60 ppm. Although temperature and salinity data are Geochemical studies conducted in February reasonably complete, geochemistry data for the 1968 provided an information base on the concen- Amchitka area are fragmentary even though such tration of major and trace elements and aided, data could be informative not only as baseline where possible, in describing the general circulation information but also from a descriptive oceanogra- in the vicinity of Amchitka Island. I11 addition to phy standpoint. The equilibrium between the salinity and temperature, determinations were saturation of C02 in surface water with respect to made of pH and magnesium, calcium, silicate, atmospheric C02 has great potential for explaining phosphate, strontium, and heavier metal ions (zinc, variations in the surface environment that are not manganese, nickel, iron, and copper) (Table 3 evident from the distributions of temperature and and Fig. 8). The distributions are generally consis- salinity that reflect primarily large-scale physical tent with the circulation described on the basis of processes. However, the analytical methods are temperature and salinity distribution, and the complex, and only limited observations are avail- scatter of values in the passes north of Amchitka able (Park, Gordon, and Alvarez-Borrego, 1974). If (Fig. 8) suggests the strong mixing aud movement exchange with the atmosphere is restricted, cooling of water through the Aleutian passes; offshore produces a decrease in the partial pressure of CO2 values are relatively uniform. anrl warming the opposite effect. Although photo- synthesis by marine plants has the greatest effect on the reduction of C02, several factors can BIOLOGICAL OCEANOGRAPHY contribute to supersaturation compared to atmo- spheric CO,, such as biocl1emical oxidation, addi- Although relatively few biological observations tion of CO, -rich river waters, increase in the depth in the immediate area of Amchitka Island have of the surface mixed layer, upwelling of C02-rich been made, both the Bering Sea aud the North sitbsurface waters, and upward divergence of deep Pacific Ocean are regions of major fisheries and water in cyclonic gyres. Although considerably have an extended history of biological investiga- more information is required to determine the tions. individual effects of these processes, high vari- A'lcRoy, Goering, and Shiels (1972) discussed ability in relative carbon dioxide concentrations the work on primary productivity; Sanger (1974) between air and surface seawater exists in data reviewed the available data and obtained a value of from closely spaced stations in Amukta Pass near 415 mg C m-2 day-' as a best estimate of summer 171°1\' (I

Table 3-Chebnical Analysis of Water Samples 342 McAlister and Favorite

Pig. 8-Surface distributions of phosphate-phosphorus (pg-atlliter), pH (in situ tempera- ture), silicate silicon (pg-at/liter),. magnesium (Mg/Chl) x lo*, calcium (glliter) and (Ca/ChI) x lo*, strontium (Sr/Chl) x lo4, and metals reported as zinc @g/liter). FOOD TYPE AND ASSUMED AVERAGE PRIMARY PRODUCTIVITY RATE?* TROPHIC LEVEL REPRESENTATIVE SPECIES"

415 mg C m-2 day-' 100 mg C m-2 day-' 1. Primary

'nediscus cuwatulus 26% 26%

2. Herbivores 38.1 105 tons carbon 12% 6% tons carbon

634 x lo5 tons tons tons 26% ------

carnivores

4. Secondary carnivores Walleye pollock Squids (Conatidael Baleen whales Seabirds (murres, Uria spp.1

5. 3" Carnivores

Northern fur seal Turbot I True cod I I ------6. 4' Carnivores * Final carnivores Representative species listed here are representative of the eastern Bering Sea. "r9 t Area = 800,000 km2 for eastern Bering Sea and for Aleutian-Southeastern Alaska. w9 Killer whales * Note unit change from metric tonr of carbon per growing season to tons of biomass $ Large sharks on alternate assumptions that either 6%or 12%of biomass is carbon. 2 ?+

'r Fig. 9-Schematic of food chain and estimates ofproduction for the Bering Sea area, June-November. 2 344 McAlister and Favorite are also shown for an average daily primary noon or from local apparent noon to sunset. production rate of 100 mg C m ' day.', which is Chlorophyl a and nutrient concentration were also more representative of estimated productivity in measured at these stations (A,lcAlister et al., 1968). the North Pacific Ocean south of the island arc. Obse~ved concentrations of chloropllyl a As part of the investigations on North Pacific ranged between 0 and 0.44 1ng/m3. The wide Ocean aud Bering Sea fisheries, the U. S. National fluctuations of chlorophyl a and nutrients in the Marine Fisheries Sewice conducted a series of surface layer indicated that the phytoplankton investigations near Adak Island in the central were not evenly distributed in the surface layer and Aleutians starting in 1957 ~vhichprovided back- suggested large-scale turl~ulence.A single dominant ground data for investigations on primary produc- subsurface ~naximum of chlorophyl a occurred tivity and zooplankton distribution. Observations within the mixed layer when the layer did not in 1968 included measurements of primary produc- extend to 100 m. Where the mixed layer extended tivity (carbon isotope method), chlorophyl a for below 100 m, chlorophyl a displayed several maxi- standing phytoplankton stock, and other environ- mums and minimums, none of which were dis- mental variables, including incoming solar radia- tinctly dominant. The highest (14.5 mg/m2) and tion. Zooplankton samples were analyzed for lowest (3.8 mg/m2) values (summed through the standing stock and for relative importance and euphotic zone) of chlorophyl a were found at distribution of the major zooplanktonic groups. stations on the continental shelf south of Amchitka. The mean of chlorophyl a values was Pri~llaryProductivity 9.0 mg/m2. The values of chlorophyl a did not differ significantly between offshore stations and Rglotoda and I

Daily rate, Region mg C n17 day-' Dates Source

Bering Sea Bering Strait 4100 June 1969 McRoy et al. (1972) Eastern Bering Sea 2 1 February 1970 McRoy et at. (1972) Aleutian area Unimak Pass area 243 June1968and 1970 McRoy et al. (1972) 87 January-February 1967 McAlister et al. (1970) Amchitka Island area 38 to 45 February 1968 McAlister et al. (1968) Adak Island coast 686 June-July 1967 Larrance (1971) 581 August 1967 404 September 1966 Adak Bay 350 to 460 March 1966 Larrance (1971) 840 to 2400 Late spring-summer Central subarctic domain Subarctic waters south to Adak Island 133 February Larrance (1971, Fig. 5, p. 604) 325 AIarch 280 bfay 327 June 250 July 207 August 240 September

*Adapted from Sanger (1974). large catches were obtained preferentially in areas Zooplankton volume near Amchitka Island of high stability in the water column. ranged from 9.6 to 107 m1/1000 m3 with a mean Zooplankton samples were collected ~vith a of 40.7. Data from the U. S. National hclarine Pacific Oceanic Group (POG) net at 35 night Fisheries Service cruises in 1966 and 1967 indi- stations in February 1968. Two replicate tows, 5 cated volume values in the Alaskan Stream south to 10 mill apart, were made at each station. of Adak Island \vhich ranged from 10 to 70 Vertical tows from 150 to 0 m were made at each m1/1000 1n3 in winter to 400 to 500 m1/1000 m3 station, depth permitting, and were retrieved at a in summer. Thus zooplankton volume in the area rate of 2 mlsec. Total time required to retrieve south of the Aleutian Islands may change season- 150 m of cable was 75 ? 3 sec. Where the depth of ally by an order of magnitude. Values for displace- the bottom did not permit tows to 150 m, shallo~v ment volume of replicate samples varied as much as tows were made by lowering the net to approxi- 166%. This variation is probably due to a patchy mately 6 m above the bottom. The rate of retrieval distributioi~ of zooplankton and again represents for shallow tows was also 2 mlsec. Geographic effects of large- and medium-scale turbulence. position, date, time of tow, depth range, and meter Numerically, copepods dominated the zoo- revolutioits obtained from a flowmeter located at plankton, contributing 85.2% to the samples; other the side of the net mouth were recorded for all main zooplankton groups are shown in Table 6. collections. Samples were preserved at sea in 5% Analysis of variance showed a significant variation Formalin. (p < 0.05) in abundance between statious for all Displacement volumes were measured, and groups except ostracods. The maximum variation taxonomic groups were sorted and counted at the in biomass (9.6 to 107 m1/1000 m3) was observed Northwest Fisheries Center, National Aclarine Fish- across the Alaskan Stream south of Amchitka. eries Service, Seattle. Samples were snbdivided for Biomass was lowest near the center of the Alaskan taxonomic enumeration into subsamples of 300 to Stream but increased near its northern and south- 500 animals. A,Iost subsamples were '/8 splits but ern boundaries (Fig. 10). Southward of this flow, ranged from '/, to %, of a sample. The total the abundance of all organisms was relatively low numbersof organisms were counted in six samples except for euphausiids and amphipods. Total abun- (4, 28, 36, 38, 54, and 68) as a check on the dance of zooplankton organisms followed a distri- accuracy of subsampling. butional pattern similar to the total biomass except 346 AlcAlister and Favorite Table 5-Dominant Species of Zooplankton

Subclass: Copepoda Family: Oncaeidae Order: Calanoida Genus and Species: Family: Calanidae atcaea medk Genus and Species: Order: Harpacticoida Calanus plumchrus Family: Ectinosomidae Calonus cristatus Genus and Species: Calanus marshallae Microsetella noruegica Calanus glacialis Subclass: Malacostraca Family: Eucalanidae Order: Euphausiacea Genns and Species: Falily: Euphausiidae Eucalat~usbangii Genus and Species: Eucalanus califort~icus Euphausia pacifica Et4calanns elongatus hyalittus Euphausia gibboides Family: Paracalanidae Tltysanoessa gregaria Gen~isand Species: Tlrysanoessa i~termis Paracatanus pawus Tliysanoessa inspinata Family: Pseudocalanidae Thysanoessa longipes Genns and Species: Tliysanoessn spinifera Pse~rdocalannsniinuttrr Tessarabrachion oculatus Clausocalanus arcuicornis Clausocalanus pergens . . Ctenocalanus vanus Stylochelron maximum Order: Amphipoda Family: Aetideidae Suborder: Gammaridae Genus and Species: Family: Lysianassidae Aetideuspacificus Genus and Species: Gaetanus armiger Cyphocaris ciiallengeri Family: Eucl~aetidae Suborder: Hyporiidae Genus and Species: Tribe: Physoceplralata Pareuckaeta elot~gata Family: Vihiliidae Family: Scolecithricidae Genus and Specics: Genus and Species: Vibilia australis Scolecithricella ntittor Family: Ayperiidae Family: Metridiidae Genus and Species: Genus and Species: Hyperia medusarum Afetridia Iucens Hyperoche meditenanea Ple~~romammaabdominalis Parathemisto pacifica Family: Centropagidae Family: Anchylomeridae Genns and Species: Genus and Species: Centropages abdonlinalis Prin~noniacropa Family: Lucicutiidae Family: Lycaeidae Genus and Species: Genas and Species: Lucicutia flauicornis nyphaena malmii Family: Candaciidae Family: Osycephalidae Genus and Species: Genus and Species: Candacin colunibiae Streetsia challengeri Caridacia bipinnata Pllyl~~mChaetognatlla (Tokioka classification) Family: Acartiidae Class: Sagittoidea Genus and Species: Order: Phragmophora Acartia clatcri Family: Eukrohniidae Acartia lot~giremis Genus and Species: Order: Cyclopoida Eukrohnia homata Family: Oithonidae Order: Apltragmophora Genus and Species: Family: Sagittidae Oitl~onahelgolandica Genus and Species: Oithonn spiniroslris Parasagitto elegans Oithona sp. Flaccisagitta scrippsae

in the nearshore waters south of A~nchitkaIsland aries of the Alaskan Stream. Distribution patterns where biomass was high and total abundance low. for select zooplankton are shown in Figs. 11 to 18. Except for pteropods and amphipods, all taxo- The general north-south distributional pat- nomic groups were most abundant at the bound- terns of biomass may be summarized as follows: Oceanograplty 347 low north of Amchitka Island in waters of Bering eastward flow to the north of the island arc. Sea origin; high in nearshore waters south of Second, the island is adjacent to a major passage, Amchitka Island; high near the northern boundary Amchitka Pass, through which north-south excur- of the Alaskan Stream, decreasing in the axis of sions override and penetrate well into the east- flow and increasing near the southern boundary; west flows. Third, it is situated at the junction of and low south of the Alaskan Stream. the Alaskan and western subarctic gyres, the former having a predominant southward flow and Table 6-Percent Composition of Zooplankton Volume the latter a predominant northward flow in this Itern Percentage Item Percentage area. ~inall~,-itis at the southernmost part of the island arc; thus flows along bathymetric contours Copepods 85.2 Euphausiids 1.5 are subject to different geophysical effects east and Chaetognaths 5.2 Pteropods 1.0 west of the island. Amphipods 2.4 Ostracods 0.9 Cnidarians 1.8 Remaining There is no evidence of local upwelling; thus Larvaceans 1.6 zooplankton 0.4 chemical nutrients must be advected laterally into

Fig. 10-Zooplankton volume in oceanic and coastal waters -(mi per 1000 m3).

CONCLUSIONS the area. Nevertheless, concentrations are not limiting primarily because of turbulent mixing and Although broad easterly currents (I

OOt05W c::...;., :...... c::...;., 601 to 2000 $Ex4 >20W

60' I I I I I 50' 179- 180' 179* 178- 177-

Fig. 11-Estimated abundance of cnidarians per 1000 m3 of water, oceanic and coastal waters.

Fig. 12-Estimated abundance of ostracods per 1000 m3 of water, oceanic and coastal waters, Pig. 13-Estimated abundance of pteropods per 1000 m3 of water, oceanic and coastal waters.

Fig. 14-Estimated abundance of copepods per 1000 m3 of water, oceanic and coastal waters. McAlister and Favorite

Pig. 15-Estimated abundance of euphausiids per 1000 m3 of water, oceanic and coastal waters.

Fig. 16-Estimated abundance of chaetognaths per 1000 m3 of water, oceanic and coastal waters. Oceanography 351

0Oto 650 ;.;.> ...... ,. 651 to 1100 &?% >no0

Fig. 17-Estimated abundance of latvaceans per 1000 m3 of water, oceanic and coastal waters.

Fig. 18-Estimated abundance of amphipods per 1000 m3 of water, oceanic and coastal waters. 352 McAlister and Fauorite are associated with fronts between water masses, Gordon, L. I., J. J. Kelley, D. \V. Hood, and P. K. Park, lowering of temperature, and erosion of vertical 1973, Carbon Dioxide Partial Pressures in the North Pacific Surface Waters. 2: General Late Summer Distri- stability. Violent movements associated with the bution, dlar. Clrem., 1: 191-198. inshore turbiilent regime affect primary production Heinrich, A. K., 1961, Seasonal Phenomena in Plankton of directly as well as the moveinent and concentra- the \\'orld Ocean, Report TT 63-19929. (Translated tions of zooplankton herbivores. from Trans. Inst. Oceanol., 51: 57-81.) Kelley, J. J., L. L. Longerich, and D. W. Hood, 1971, Measurements of Carbon Dioxide in Northern Sea (Progress Report to Office of Naval Research, Washing- ton. D. C.\. Institute of Marine Sciences. Universitv of REFERENCES ~l&ka,Faiibanks. Koblents-Mishke. 0. I.. 1965. The Value of Primarv Pro- ductivity in ' the Pacific ocean, Ocea?:ology (USSR) Aron, IS., 1962, The Distribution of Animals in the Eastern (English Translation), 5(2): 104-116. North Pacific and Its Relationship to Physical and Larrance, J. D., 1971, Primary Production in the Mid- Chemical Conditions, J. Fish. Res. Board. Ca~r., 19: Subarctic Pacific Region, 1966-68, Fish. Bull. U. S., 271-314. 69(3): 595-613. Arsen'ev, V. S., 1967, Techeniya i vodnye masey Beringova McAlister, W.B., \S.J. Ingraham, Jr., D.Day, and morya (Cunents and Water Masses of the Bering Sea). J. Larrance, 1970, Oceanography, International North Izd. "Nauka," Moscow (in Russian, English summary). Pacific Fisheries Commission, Annual Report 1968, pp. (Translated in 1968 by National Marine Fisheries 90-101. Service, Northwest Fisheries Center, Seattle, \Sash.) -, C. Mahnken, R. C. Clark, Jr., \ J Ingraham, J. Batalin, A.M., 1964, On the Water Exchange Between the Larrance, and D. Day, 1968, Amchitka Bioenvironmen- Bering Sea and the Pacific Ocean (in Russia).. T?. tai Program. Oceanography and Marine Ecology in the VNIRO, 49: 7-16. (Translated in 1968 in Soviet Vicinity of Amchitka Island, USAEC Report Fisheries Investigations in the Northeastern Pacific, BMI-171-112, Battelle Memorial Institute. Part 2, pp. 1-12, Report 'IT 67-51024.) McRoy, C. P., J. J. Goering, and I\'.E. Shiels, 1972, Studies Bogorov, V. G., K. V. Beklenlishev, and h1. E. Vinogradov, of Primary Production in the Eastern Bering Sea, in 1961, The Distribution of Plankton in the Pacific Ocean Biological Oceanography of the Northern ~\'orth Pacific and Its Relation to Geographical Zonation, in Pacific Ocenn, 1. ', Takenouti et al. (Eds.), pp. 199-216, Basin Geography, J. L. Gressitt (Ed.), 10th Pacific Idemitsu Shoten, Tokyo. Science Congress, Honolulu, Hawaii, 1961, University Motoda, S., and T. Kawamura, 1961, Data on Phytoplank- of Hawaii Press. ton Photosynthetic Activity, in The "Oshoru Maru" -, and M.E. Vinogradov, 1968, Distribution of Cruise 46 to the Bering Sea and North Pacific in Zooplankton in the Kurile-Kamchatka Region of the June-August 1960, Data Rec. Oceanogr. Obseru. Pacific Ocean, U. S. Naval Oceanographic Office, Trans- Enplor. Fish (Hokkaido), 5: 142-165. lation 336. [Translated from Trans. Inst. Okeanol., NORPAC Committee, 1960, Oceanic Obseruntions of the Akad, Nauk SSSR, 34: 60-84 (1960).] Pacific, 1955, Vol. 2, The Norpac Data, University of Favorite, F., 1974, On Flow into Bering Sea Through California Press and University of Tokyo Press, Aleutian Islands Passes, in Oceanography of the Bering Berkeley and Tokyo. Sea, Institute of Marine Sciences Occasional Bull. No. 2, Park, P. K., L. I. Gordon, and S. Alvarez-Borrego, 1974, D. I\'. Hood and E. J. Kelley (Eds.), pp. 3-37, University The Carbon Dioxide System of the Bering Sea, in of Alaska, Fairbanks. Ocennography of the Bering Sea, D. \\I. Hood and E. J. -, 1967, The Alaskn?~Streanr, International North Pacific Kelley (Eds.), Institute of Marine Sciences Occasional Fisheries Commission, Bulletin 21,pp. 1-20. Publication No. 2, pp. 107-147, University of Alaska, Fairbanks. -, and Donald Fisk, 1971, Drift Bottle Experiments in Reed, R. K., 1971, Nontidal Flow in the Aleutian Island the North Pacific Ocean and Bering Sea: 1957-60, Passes, Deep-Sea Res., 18(3): 371-372. 1962, 1966 and 1970, U. S. Department of Commerce, Sanger, G. A,, 1974, A Preliminary Look at Marine Mam- National Oceanic and Atmospheric Administration, mal Food Chain Relationships in Alaskan IVaters, National Marine Fisheries Senrice, Data Report 67 (on Northwest Fisheries Research Center Processed Report, microfiche). National Marine Fisheries Service, Seattle. -, and IS. J. Ingraham, Jr., 1973, International North Steemann Nielsen, E., 1952, The Use of Radio-Active Pacific Fisheries Commission, Annual Report 1971, pp. Carbon (c") for Measuring Organic Production in the 89-97. Sea, J. Cons., CO?ls. Perma. Int. Explor. dfer., 18(2): -, and IS. J. Ingrabam, Jr., 1972, Influence of Bowers 117-140. Ridge on Circulation in Bering Sea and Influence of Taniguchi, A,, 1969, Regional Variations of Surface Pri- Amchitka Branch, Alaskan Stream on Migration Paths mary Production in the Bering Sea in Summer and of Sockeye Salmon, in Biological Oceanography of the Vertical Stability of Water Affecting the Production, Norther?: Nortl: Pacific Ocean, A. Y.Takenouti et al. Bull. fic. Fish., Hokkaido Uniu., 20: 169-179. (Eds.), pp. 13-29, Idemitsu Shoten, Tokyo. Thomson, R. E., 1972, On the Alaskan Stream; J. Plrys. -, A. J. Dodimead, and K. Nasu, 196,Oceanography of Oceanogr., 2(4): 363-371. the Subarctic Pacific Region, 1960-71, International Tsuruta, A,, 1963, Distribution of Plankton and Its North Pacific Fisheries Commission, Bulletin No. 33. Characteristics in the Oceanic Fishing Grounds, with -, \Y. J. Ingraham, Jr., and D. M. Fisk, 1972, Oceanogra- Special Reference to Their Relation to Fishery, J. phy, International North Pacific Fisheries Commission, Shi??~onosekiUniu. Fish., 12(1): 13-214 (in Japanese, Annual Report 1970, pp. 90-98. English summary). Ecology of Phillip A. Lebednikt John F. PalmisanoS Marine Algae* Fisheries Research Institute, University of \Vashington, Seattle, Washington

The rocky shores of Antchitha Island are densely corpeted The most conspicuous element of the sublittoralflora is by marine algal conrmuitities, which, beyond tlte coast Alaria fistulosa, which forms the extensive floating kelp exposed by the tides, are characterized by extensive beds. Down to 20-m (66-ft) depth, the bottom is densely floating kelp beds. An unusual feature of the intertidnl area covered with species in the genera Laminaria, Alaria, of the sorcthenstern third of tlte island is the occurrence of Cymathere, Agarum, and Thalassiophyllum. A third level of wide, often extrenrely flat, rock benches nt about mean tide vegetation is formed by populations of Ptilota, Hypophyl- level. lum, Cirulicarpus, and Constantinea, Fitay crustose The littoral region is characterized by a supralittoral coralline algae, particzclarly Clathromorphum nereostratom, fringe area with Prasiola, Rhodochorton, and Porphyra- and tlte green alga Codium ritteri dominate rocky bottoms Ulothrix zones and a midlittoral area with Fucns, Nedo- below 20 In (66 ft). The predation of the sea otter on phyllum, and Alaria zones. Afany species distribtctions are invertebrate herbivores is tntdoubtedly n major factor in the strongly i~tflue~tcedby the benches, being correlated with formntion of such dense nlgnl communities. horizontal distance perpe~tdicular to the shore. The pres- The Milrow nuclear test affected about 4 ha (10ncres) ence of Hedophyllum and Fucus zones indicates more of littoral vegetation when an uplift of about 12 cm (5 in.) protected conditio~rstitan might have been expected. It is occurred on an intertidal rock bench. Mortality was extensive suggested that the bench topography and offshore kelp in the upper portions of all zones 6 months after the distrcr- beds provide an exposure gradient across the intertidal area. bance, and siptificant changes were still occurring when last The sublittoral fringe is characterized by a zone of observed, 3% years after Alilrow. The Ca~t~tihintruclear test Laminaria longipes. Afost littoral algae occur throtcghout resulted in uplifting tip to a ~naximumof 1 m (3 ft). the year but exhibit growth in tlrespringairdsummer. The AJortnlity of littoral uegetation was severe along 1.9 hnr Porphyra-Ulothrix zone and the Halosaccion association (1.2 miles) of coast, including areas with 0.5 to 1 m of appear in spring and reach a peak of development in the uplift. Moderate mortality was observed along an additional sunrmer. Plots in the intertidal region denuded of all 1.5 km (I mile) of coast, and nrortality was detectable organisms exhibited colonization by a sequence of diatoms along a further 2.7 km (1.7 miles) of coast. Significant and filamentous algae, leafy green algae, and, last, larger red changes ill uegetation were reported to be occurring nearly and brown algae. Initial settlement was seasonal and 3 years after Cannikin. A normal littoral vegetation will occurred fastest in exposedareas at low levels. The rentoval eventrcally return to both the Milrow and Cannikin- of only the help canopy gave si~nilarresults, but coloniza- disturbed areas. The lifting of some rock benches aboue tire tion occurred ??tore qtrickly and was i~tflue~tcedby the midlittoral area as a result of the Cannikin test has resulted rentnining understory uegetation. in a significant permanent reduction in area available to most littoral species.

The benthic marine algae, commonly known as source of food for primary consuming organisms, seaweeds, grow luxuriantly around the rocky particularly marine invertebrates. coasts of Amchitka Island. These plants form a The history of phycological research in the conspicuous covering on rocks uncovered by low North Pacific Ocean and our studies of the physical tides. Floating kelp beds, consisting of plants that environment, the structure of marine algal com- are attached to the rocky bottom, occupy large munities, biological interactions, and the effects of areas of the ocean surface surrounding the island. underground nuclear testing on the algal commu- As primary producers these kelp plants are a major nities of Amchitka Island are reviewed in this chapter.

*All sections of this chapter except the section "Coloni- REVIEW zatioo Studies" were written bv Lebednik. ?Present address: Department of Botany, University of British Columbia, Vancouver, B. C., Canada. Until 1967, the starting point for studies $Present address: Auke Bay Fisheries Laboratory, reported in this chapter, the Aleutian marine algae National Marine Fisheries Service, Auke Bay, Alaska. were practically unknown. Therefore the floristic 354 Lebednilt and Pal?nisnno relationships of the Amchitka algal flora are best Bering Sea and Guryanova (1935), Kardakova- obtained by reviewing the literature of surrounding Prezhentsova (1938), and E. Zinova* (1940) on areas of the North Pacific Ocean. A comprehensive the marine algae of the Commander Islands. review of the Russian literature is given because The publication by Kjellman (1889) resulted most of the articles are practically unknown or from collections made on the Vega expedition in unavailable in North America. the Bering Sea from July 20 to Aug. 19, 1879. These collections were made at St. Lawrence Bay Aleutian Islands aild Konyam Bay on the Chukotski Peninsula, Union of Soviet Socialist Republics; Port Clarence, The first publication on marine algae from the on the Seward Peninsula, Alaska; and Bering Aleutian Islands (Mertens, 1829) resulted from the Island, one of the Commander Islands. This work is expedition of the Russian ships Seniavin and still an important taxonomic reference, and it is Moller, which were under the command of Liitke. the only work in which the distribution of the The famous "Illustrationes Algarum" (Postels and marine algae of the Bering Sea is treated compre- Ruprecht, 1840) presented the full treatment of hensively. the marine algae collected on this expedition by Guryanova (1935) discussed the history of Postels, Mertens, and ICastalsky. In the Aleutian scientific research, geology, climate, oceanography, Islands marine algae were collected at Unalaska and biology of the Commander Islands and made Island only. Further records of Aleutian marine extensive observations on the marine algal vegeta- algae, again limited to Unalaska Island, were tion of Bering Island from November 1930 to JUII~ published by Setchell and Gardner (1903). Algal 1931. The distribution of marine plants and specimens from the western Aleutian Islands, illcludi~lg the first collectio~~sfrom Atka, Am- animals on Bering Islatld, although treated in detail, was not studied with quantitative methods. chitka, and Agattu Islands, were gathered by A more detailed study of the distribution of the Kobayashi from May to August 1931 and were seaweeds of the Commander Islands was under- reported on by Okamura (1933). Brief comments taken by Kardakova-Prezhentsova from 1928 to on the marine and land plant communities of the 1932 (Kardakova-Prezhentsova, 1938). A compre- Aleutian Islands were made by Tate~vaki and hensive list, containing 174 species of marine algae Kobayashi (1934). Field work was conducted by collected at the Commander Islands from 1871 to Tatewaki in 1929 on Attu, Amchitka, and Atka 1932, was compiled by E. Zinova (1940). Islands and by Kobayashi in 1931 on Attu, Anlchitka, Atka, Umnak, and Unalaska Islands. Eastern Bering Sea The account of the marine vegetation, comprising 21 species of marine algae, includes no details on Several short papers have dealt with marine localities; thus it is impossible to determine on algae from the eastern Bering Sea. Setchell (1899) which islands each species was observed. Collec- published a short list of marine algae collected at tions of marine algae from Attu, Shemya, Adak, the Pribilof Islands and Chihara (1967)reported on and Unimak Islands were made in 1960 as part of a some marine algae collected at Cape Thompson, large study of the North Pacific algal distributions Alaska. From Izembek Lagoon, on the north coast by Scagel (1962; 1963). These collections, depos- of the Alaska Peninsula opposite Cold Bay, R.IcRoy ited in the Phycological Herbarium of the Univer- (1968) collected the Eurasian alga Fuczis inflatus f. sity of British Columbia, were later used by latifrons and Biebl (1970; 1972) studied the students and associates of Scagel in mollographic temperature resistance of a few local algal species. studies of the genera Hedopl~yllt~vz(IViddowson, 1965), Laminaria (Druehl, 1968),Alaria (IViddow- Pacific North America son, 1971a; 1971b), Porpl~yra (Conway et al., Research on marine algae east of the Alaska 1976), and Ulva (Tanner, in preparation). Druehl Peninsula, along the Pacific North American coast, (1970) discussed the distribution of the Laminar- has resulted in many publications. The importance iales in the North Pacific Ocean. Only three species of these studies to the Amchitka flora is generally of marine algae had been reported from Amchitka Island prior to our studies. Several other publications, although not deal- *Spelled E. S. Sinova as transliterated on the original ing directly with the Aleutian Islands, are impor- publications. This spelling is generally used in the phyco- tant either taxonomically or geographically in logical literature. To conform with transliteration standards now accepted, we have spelled her nalne with a "2" in this studies of the marine algae of Amchitka Island. paper. So that this author wilt not be confused with A. D. Probably the most pertinent of these are the Zinova, a niece, who also published on marine algae, the publications of Kjellman (1889)on the algae of the first initial is used in citations throughout this chapter. Ecology of A1nri11eAlgae 355 inversely related to their distance from the Aleu- the Kamchatka Expedition of F. P. Ryabushinski. tian Islands. The Harriman Alaska Expedition Savich" (1914) described the algal vegetation in collected 207 species of marine algae at Victoria, 'the bay. The taxonomy of Savich"s specimens was British Columbia, and at several localities along the determined by Voronikhin (1914). The area was southern coast of Alaska as far west as the studied again in 1929 and 1930 by a zoological Shumagin Islands (Saunders, 1901). Scagel's expedition, and the algal specimens were described allnotated list of marine algae of British Columbia by E. Zinova (1933). A few species in the red algal and northern Washington (Scagel, 1957) has family Delesseriaceae were also recently described recently been superseded by his monograph of the from the area (A. Zinova, 1965). 111 northeastern Chlorophyceae (Scagel, 1966) and revised lists and Kamchatka, eight marine algae were collected by keys for the Phaeophyceae (Widdowson, 1973) and Takayama in August 1923 at ICorf (Baron Korfe), Rhodophyceae (Widdowson, 1975) which cover on the northwest coast of Korf Zaliv (Okamura, the same geographical area. These publications 1928). The littoral and sublittoral algae from contain information on many species that occur in Anadyrskiy Bay were described by Vitlogradova the Aleutian Islands. A guide to the literature and (1973a; 1973b). A study of the littoral and distributions of Pacific benthic algae, including sublittoral algae of Karaginskiy Island was carried Alaska, was published by Dawson (1961). out in 1930 (ICongiser, 1933). Further studies of Johansen (1971) reported on the effects of the marine algae from southeastern ICamchatka in- Alaska earthquake of 1964 on intertidal benthic cluded an expanded list of species (E. Zinova, algae in Prince William Sound. Although the 1954a) and a detailed description of littoral zona- littoral marine algae were destroyed in some tion (Spasski;, 1961). localities, 120 taxa of macroalgae were collected, and 17 of these represented new records for the area. An extensive account in English of the marine Russian Studies algae of the ICurile Islands was published by the Japanese phycologist Nagai (1940; 1941). Subse- Bibliographies of the Russian literature on quently several Russian papers have been pub- marine algae published from 1900 to 1940 have lished. The distribution of the fauna and flora of been compiled by Ga'idukov (1901), Elenkin and the littoral region of the ICurile Islands was 01' (1929; 1935; 1950a; 1950b), Gollerbakh et al. described by Kusakin (1961). Several taxonomic (1966),and ICrasavina (1968). A taxonomic cross- papers have also appeared that are based on index to these bibliographies, spanning the years specimens of Laminariaceae (Gusarova and Petrov, 1737 to 1960, was compiled by Gollerbakh and 1970; 1972) and various red and brown algae Krasavina (1971). A short history of Pacific (Gusarova, 1972). Russian scientific investigations was given by Komarov (1962).Nearly all the Russian taxonomic Okhotsk Sea studies on marine algae have been carried out at the IComarov Botanical Institute in Leningrad (St. The first major paper on the marine algae of Petersburg in the older literature) by Elena S. the Okhotsk Sea was the classical phycological Zinova and later by her niece, Anna D. Zinova. A treatise "Tange des Ochotskischen Meeres" history of the Institute in English was recently (Ruprecht, 1851). This work was based on speci- published (Shetler, 1967). mens that had accumulated in the herbarium of the Imperial Academy of St. Petersburg since Gmelin's Eastern Kamchatka time. Most of these specimens had been collected in 1807 by Redovski (a few possibly also by The earliest records of marine algae from Steller) in the eastern Okhotsk Sea and later by I

Coloeizatio~~ Coastal studies Date T-1* T-2* T-3* IA-I* IA-2* IA-3* walk* (3areasl*

+March to April 1968 \Y I\' tMay 1968 0 0 August 1968 P P h,larch 1969 P P lvla\, 3 to 7. 1969 June 1969' P P Oct. 2, 1969 (i\lilro\v test) Apr. 21, 1970 Mav 21. 1970 ~ugust1970 September 1970 December 1970 P P March 1971 May 1971 July 1971 August 1971 September 1971 October 1971 Nov. 6, 1971 (Cannikin test) December 1971 February 1972 April 1972 Jine 1972 August 1972 October 1972 December 1972 A~rii1973

*P, photographic sample; I\', dry-weight sa~nple;and 0, visual observations. tSa~nplestaken by F. C. I\'einmann. $Samples taken by FRI staff; reported in Kirkwood (1974; 1975). the Cannikilt test were reported by Nakatani et al. coralline algae. IVeinma~incontinued his graduate (1973) and Nakatani and Burgltcr (1974). From career else~vhere.Extensive collectiolis were made August 1968 to August 1973, the preceding studies for this taxonomic study in the littoral and were conducted by Lebednik. Thc experimental sublittoral regions from 1967 througl~1973. denudation studics were conducted by Pallnisano In the first contributions to this study, IVynne (1975). Subseque~ltobservations of A,lilro\v and described the Dclesscriaceae (I\'ynnc, 1970a; Caltniki~teffects as reported in ICirk\vood (1974; 1975), Bomtemaisoniaceae (\\'yline, 1970b), and 1975) were made by other FRI investigators. The Porp1i)lra species of Amchitka (\\'ynnc, 1972), all littoral sampling program is summarized in Table 1. members of the red algae. An ecological study on From the beginning of the marine algal studies, algal canopy iliteractio~isin the sublittoral region it was apparent that many poorly kno\vn and \\.as reported by Dayton (1975). nndescribed species were common in tlte Amchitka A numbcr of later papers have beat bascd, in flora and that a comprehensive taxonomic study part, on collcctiolts made at i\mchitka. I\'ynne would produce a valuable contribution to our (1971) merged the broxvn algal genera A~zulipus knowledge of tlte marine algal flora of tlte North and Heterochordaria. Lebednik presented a mono- Pacific Ocean. In 1968, R. E. Norris, F. C. Wcin- graphic treatment of the coralline genera C1ntl~- mann, and M. J. IVynne agreed to undertake a ro~~zor/~hzpolychaete species (Fig. 3). In the identified to date. The tasonomic studies are winter this silt blanket may be eroded away, continuing, and we hope that the entire macro- exposing a rock surface sparsely occupied by the scopic algal flora will be described in future crustose coralline alga, Clatltromorpltum circunl- publications. scriptuna. Frequently this area is lower than the seaward portion of the bench; consequently sea- water may cover the rock bench at low tide. This PHYSICAL FEATURES OF THE depressed area is referred to as the moat. ENVIRONiVlENT The greatest fraction of the littoral region is occupied by the next seaward component, the The topography of the Amchitka littoral emergent intertidal rock benchflat (Figs. 1 and 2). region, {shich is important in the determination of The rnidlittoral vegetation reaches its greatest the distribution of littoral algae, is unusual and development here and is readily accessible; most of deserves description. An aerial vjew of a segment of our research was conducted in this area. The coastline at low tide (Fig. 1) sho~vstwo extensive benchflat lies approximately at mean sea level, rocky flats (B), dissected by large cl~annels(C), varying 2 to 3 m (7 to 10 ft) above and below. bordered landward by boulder strands and sepa- Powers et al. (1960) report the bench width to be rated by a sandy beach and the mouths of two about 15 m (50 ft) on the average, with an streams (S). Such stream mouths usually indicate a occasional segment as wide as 400 m (1300 ft). sand or gravel trough (T) projecting seaward This bench area is topographically diverse. It may through and beyond the littoral rock flats (Powers, be an extremely flat surface with a pure stand of a Coats, and Nelson, 1960; Lebednik, U'einmann, single algal species or it may be a surface undu- and Norris, 1971). The littoral rocky areas are lating a meter or Inore in height and with a diverse mainly characterized by a feature termed rock vegetation. Cobble-lined channels (Fig. I), usually benches (Po\\rers et al., 1960), intertidal platforms connected with the sea even at low tide, may (Morris, 1971), or strandflats (Everett, Chap. 8, dissect the surface, and local depressions may form this volume). At Bird Cape, at East Cape, at some tide pools (but tide pools are uncommon along the other promontories, and in the northwest third of whole coastline). Both channels and tide pools the island these benches are lacking, but, in the have avcgetation that is ~nostlysublittoral in compo- southeastern third of Amchitka, they are a domi- sition. A microtopographical feature of significance nant feature, occurring where there is a bedrock of is the occurrence of small or large masses of harder permeable fine-grained breccia (L. M. Gard, per- rock 5 to 100 cnl (2 to 40 in.) in diameter sonal communication; Powers et al., 1960). Sand, that project above the plane of the softer breccia. cobble, or boulder strands, which are mostly Often an isolated clump of filclts distichzts on the associated with stream mouths, occur for short bench will mark the presence of one of these intervals. protruding rocks. The film1 feature, cotnmon but Figure 2 is an idealized schematic section not present on all benches, is an elevated ridge or through a segment of coast where an intertidal rampart on the seaward edge of the bench (Fig. 2). bench is present. The landward nlargin is com- The rampart is typically 1 to 2 m (2 to 7 ft) highcr monly a sheer cliff 20 to 30 n~ (60 to 100 ft) high than the bench and tends to have an irregular surmounted by a mantle of tundra. At the foot of outline with small tide pools in depressions. \Vithin the cliff, huge blocks of cliff fragments and a small distance along its length, a ranlpart may sho~\ra \vide range of littoral algal species as well as a sharp difference between species on the exposed sealvard face and the protected landward face, the *The new genus dlikn,,~ieiln U'ynne has been proposed former being the most exposed habitat of the but not validly published (IVynne, 1975). bench region. Ecology of Alariile Algae 359 Table 2-hlarine Algde of Amchitka Island, Akska, as of June 1973

Phylum Chiorophycophyta Order Nemalialcs Order Ulotrichaler Pletrroblephnris japot>ico (Okam.) Wynne Enferontorpho sp. Rhodochorton prrrprtreurn (Lightfoot) Ilosenv. Adonortronzo orcticnm \Vittrock Order Cryptonemiales dlo,tortronrofrcscrc~n (P. et R.) Wittrack Borrielln cretoceo (P. et R.) Johansen Ulotbrix flncco (Dillwyn) Thuret CollophyNis flnbellulotn Harvey Ulvo loctuco L. Cirrtrlicarpur gmelhti (Grunow) Tokida et hfasaki Uluo sp. Clotkrontorpl~amcirczcntrcriptsm (Stroemf.) Fosl. Order Schizogoniales Clnthrontorphrcm cotnpnctum (Kjellm.) Fosl. Proriolo borealis Reed Clothromorplruat loculosutn (Kjellm.) Fosi. Rorenuingielln sp. Clnthromorpham nereortrotstn Lebednik Order Cladophorales Clothromorpkum reclbrotunz (Fosl.) Adey Clznefotnorpha ntelogo,ri,cnz (Weber et blohr) Kiitz. Co,,rtantineo rosa-ntorino (Gmelin) P. et R. Clodophoro sp. CoraNinn pilalifero P. et R. Lolo lr~bricn(S. et G.) A. et G. Eiantel CoraNinn un,rcouueriensir Yendo Spo~tgonzorplmspinescens Kiitz. Drtnontin rinzplex Cotton Order Chlorococcales E~tdoelndinmuricafn (P. et R.) J. i\g. Codiolutn petrocelidis Kuckuck Gloiopelfirfitrcoto (P. et R.) J. Ag. Order Codiafes Holynzenia ap. Codiunt ritferi Setch. et Gardn. Hildertbrondia prototypzrr Nardo Halicyrtir sp. Lithophyllunz spp. Lithoflrom,tinnr spp. Phylum Phaeophycophyta dlesophyllutn oleuticrrn~Lebednik Order Ectocarpales Ectocorper lonrentosur (Hudson) Lyngb. Order Gigartinales Pylniello littorolir (L.) Kjellinan Al~~rfellioplicoto(Hudson) Fries Rolfrinf~t?rgifornrir(Gutmerur) Setch. et Gardn. Gigortino pncifico (Kjellman) Tokida Order Chordariales lridoeo cornucopine P. et R. Anolipsrfiliforntir (Harvey) Papenf. Petroeelirfinncircn,ro S. et G. Anolipar joponicer (Harv.) Wyone Petroeelis nziddendorffii (Ruprecht) Kjellman Chordorin flagelliformir (bliill.) C. Ag. Plnlontn sp. Lenthesio difforotir (I..) Areschoug Rhodoglosrrrnz pulcl#m(Kiltsing) Setch. et Gardn. Order Desmarestiales Tznbraeoptera serrotn (P. et R.) Zinova dlernbm,toptera setchellii Gardner dle,nbranopfern rpinzrlorn (Rupr.) Zinosa dlicroclndin borenlis Ruprecht dlyriogmmnze kjellmaninnn,,, Zinova r\'ie,zbsrgia pr-rolifera Wynne Odo,rthniinfloccorn (Esper) Felkenberg Phylum Rhodophycophyta Pn,zlonesm jvergenrii 0. Ag.) Kylin Order Bangiales Pl~ycodrysornclritke~zrir Wynne Bnngiafrcrroperpurea (Dillw.) Lyngb. Pl~ycodrysriggii (Gardeer) Porphyra amplirri?nn (Kjellman) Sctchell et Hus Plotythomnion sp. Porphyro oclrolearir Nagai Polyriphonia pocifica Hollcnbcrg Porphyro perfornta J. r\g. Polyriphonin sp. Porplryro ps-reedolinenris Ueda Pteroriphonin gordceri Hollenberg Porpkymprlrpnren (Roth) C. Ag. Pterosiphonin sp. Porplzyro lorn (Yeedo) Ueda Ptilota arpletzioides (Espcr) C. Ag. Porpltym untbilicnlir (L.) J. Ag. Pfiloto filicipro (Farlow) J. Ag. Porphyro unriegatn (Kjellman) Hur Tokidodendron ballnto (Gardner) \Vynne Porphyra yezoenrir Ueda Yendonin crnrrifolio (Rupr.) Kylin Porphyrelln gordneri Smith et Hollenb. Ziuouaen ncaathocarpn i\'ynne

'This species war found only in drift on beaches and was never found attached to rocks in the Amchitka coastal area. Ecology of ~\.lari?zeA/gae 361

BIOLOGICAL SUPRALITTORAL SUBLITTORAL AREA FRINGE MIDLITTORAL FRINGE -1 I I TOPOGRAPHIC FEATURE BENCHFLAT RAMPART I- I-

n BEDROCK =SEAWATER GRAVEL n SILT HARD ROCK FRAGMENTS

Fig. 2-Schematic diagram of topographic features and biological regions of a typical intertidal bench.

Figure 4 shows schematically the relationship (National Ocean Survey, 1971). \\'eimnann (1969) of wave action (exposure*) to the bench configu- indicated that the lo\\, tide on ally day may be ration. At lo\\. tide \crave fronts (1) approaching the about 1 hr later than prcdicted. During our studies bench break solely along the sealvard face of the at fixed points in the intertidal area, \Ire found that rampart (2). As the sea level rises, water flows over the lo\vest tide \\lithi11 a spring tide cycle often the rampart onto the benchflat (3), resulting in occurred about 1 to 3 days later than predicted. currents shorexvard across the benchflat snd in the There are also large irregular deviations from the channels. Approaching the moat (4), these currents turn and ru~~parallelto the shore, finally emptying into sand ttoughs on either side of the bench (5). At high tide waves plunge over the tlten submerged rampart and dissipate most of their energy there. Ordinarily, only unidirectional flow occurs over the benchflat at high tide, and in solne localities the moat area appears to be a river of seawater. Only during storms do waves reach and break on the beach or on the cliff itself. Even under these conditions there is little or no plunging action over the benchflat because of the uniformity of water depth. Of all the components of the rock bench, the rampart has the greatest exposure. Tidal pllenomena are discussed elsewhere in this book. According to the tide tables, the tides altern;tte from diurnal in the spring phase to mixed seniidi- urnal in the ilcap phase, and the inasimum tidal range is just over 2 m (O'Clair, Chap. 18, this volu~ne).The lowest tides occur in June at about 1100 (BST) and in December at about 0100

*The term exposure as used in this chapter refers to Pig. 3-View of the moat area of an intertidal bench exposure to wave action. at Duck Cove (IA-1) on Aug. ZG, 1973. mean monthly wind speed of 13.8 knots occurs in July. Wind speeds and durations significantly greater than tliis occur in thc fall, blo\vi~lgfrom the southwest sector (Armstrong, 1971; Chap. 4, this volume). The Pacific Ocean coast receives s\velts of a greater \\ravelength than the Bering Sea coast owing to the greater fetch in that clirection, but no data on s\\rell frequency are available. The Berilig Sea coast has smaller s\\lells than the Pacific Ocean coast, and heavy wa\,e action is less often observed. In the evaluatioll of the degree of Fig. 4-Schen~atic diagram of xva\.e exposure in exposure, caution must be cxcrcised because bench yelation to an intertidal bench. (See text for explana- topography strongly i~lflue~lcesexposure of the tion of diagram.) [From P. A. Lebednik, F. C. \\'ein- mann, and R. E. Norris,BioScience, 21: 657(1971).] intertidal area. Seau~aterco~lditio~ls are of great importance to sublittoral communities. In the surrounding off- predicted tides from day to day. \ire have seen shore surface waters, winter conditions are about areas colllpletcly co\~erctt\\,it11 sca\vatcr thro~~ghout 3OC temperature ant1 33 Yo0 (parts per thousand) one day, and the next day they would be un- salinity (R'lcAlister, 1971 ; i\,lcAlister and Favorite, covered even though the predicted tide hcights Chap. 16, this volun~e);in summer they are about were identical and there was little n7a\.e action. G°C temperature and 30 %, salinity (Burgner et al., Nearshore currents are generally gentle except at 1968). '1'lierc is no sharp thermocline at any season Bird Cape and East Cape. above 30-111 (100-ft) depth, the limit of our Armstrong (1971; Chap. 4, this volume) de- collecti~tgactivities. In numerous profiles taken in scribes the island's climate in detail. We consider August 1967, neither salinity nor temperature here those factors of particular importa~lceto litto- changed more than one unit from the surface to ral algae. Record high and lo~vtemperatures re- 114-m (374-ft) depth (Burgner et al., 1968). At the corded at Amchitka during 1943 through 1948 were shore seawater temperatures ranged from a mini- +18.3OC and -lO.O°C, whereas extreme mean daily mum of 2'C in \\,inter to a maximum of about S°C high and low temperatures were +10.G0C and on the Bering Sea coast and 1O0C 011 the Pacific 2.Z0C, respectively. Perhaps most critical to litto- Ocean coast in summer. Thus the annual range of ral algae is the occasio~lalcoitlcide~lce of lo\\, tides temperature is only 8OC at ~naxlmum. .' with temperature estrcmes. In the cornparis011 of Turbidity of water is an importaut limiting maximum and minimum mean daily temperatures at factor in the detcr~lli~lationof the maximu~ndepth various times of the year with times of predicterl of sublittoral algal communities. Although no lo~vtides, t\vo such periods appear possible: an air measuren~entsare available, it can be said that the temperature of about +g°C near 1loo11 in August water surrounding Amchitka Island is one of the coillciding with water at the -61-cm (-2.0-ft) level least turbid waters in high northern latitudes. It is and an air tetllperaturc of -0.3'C near midnight in probably nearly comparable to the extremely clear January above \\rater at the -39.6-cm (-1.3-ft) waters reported for tropical areas. The best indica- level. The former is about equivalent to sca~vater tion of water clarity is the depth distributio~lof temperatures at that season and would not appear the algal communities themselves. to be critical, but the latter extreme, besides being The composition of offshore bottom substrates more than 4' belo\\, ambient sea temperatures, is varies widely around Amchitka. Sand, ledge, gravel, below the freezing point and may represcnt a and cobble bottoms, in approximate order of critical stress factor for littoral organisms. Indeed, frequency, may bc foulld on either side of the we have seen ice forming on emersed seaweed and island. In all cases, clown to depths reached by surface freezing of pools during nighttime winter scuba diving [about 30 m (100 ft)], these sub- lo\\, tides. strates are clean of fine detrital deposits. No tfue Desiccation as ~vellas temperature stress may mud bottoms nrere observeti in ~~umerousdives occur in summer months when lo\\, tides coi~icide arou~ldthe island. \\it11 clear skies and calm seas. Since clear skies and calm seas are rare at Amchitka (Burper et al., LITTORAL ZONA'TION 1969; Armstrong, 1971; Chap. 4, tliis volumc), these conditions occur infrequently. Along rocky coastlines the distribution of Wave exposure depends on local winds and the marine organisms utlco\~eretl at low tide is ex- iilcomillg swell from distant sources. The lowest tremely complex aixl depcilds on many factors Ecology of klarine Algae 363 (Lewis, 1964). A major factor influencitlg distribu- The boundary between the supralittoral fringe tion, aside from the tidal regime, is the degree of and ~nidlittoralareas is usually defined at the upper cxposurc (opposite of protected or sheltered). In limit of barnacles (Stephenson and Stephenson, highly exposed areas on the open coast, so-called 1949). Because barnacles are rare at Amchitka and intertidal marine organisms may exte~ld~vell above do not form a distinct zone there, we have assig~led the theoretical upper limits of the tides, i~ldicating the boundary between these two areas to the upper that the distribution of these organisms cannot be limit of Fzlcus disticlzz~s.In this chapter no attempt defined by a single physical factor, such as tidal is made to formally subdivide the sublittoral levels. Ho~vcver, the exte~lsiveliterature on this region. For convenience, the portion of the sub- subject (see Lewis, 1964, for a review of the littoral region exposed by the lo\\~cst tides is literature) indicates that marine organisn~son most referred to as the sublittoral fringe (infralittoral rocky coastlines of the world are observed in zones fringe of Stephenson and Stephenson, 1949). In or belts (". . . horizontally extensive, biotically the supralittoral fringe and ~nidlittoralareas, zones distinct. . ." groupings of organisms, Doty, 1957) of limited vertical extent, each usually dominated that are arranged one above the other in a pattern by a single species, are observed in a consistent of zonation. A'Iany systems of terminology have vertical sequence (zonation) over large horizontal been used to describe and classify the zonation of distances of coastline. In the lnidlittoral area, rocky shores. It is generally agreed that the where the greatest number of littoral species and organisms themselves best define the zones, and in the most complex zonation occur, a further break- this chapter we have adopted the schemes of do~vnof zones illto associatio/~sis possible. In this Stephctiso~land Stephenson (1949) and Le~vis chapter an association refers to an area that is (1964) \\!it11 slight modification (see Table 3). The locally dominated by a single species but docs not

Table 3-Classification Scheme and Approxi~nateElevations Above hlean Lo\ver Low Water of the Littoral Vegetation of Amchitka Island*

Region Area Zone Association Elevation, cm

Prasioln Supralittoral Rhodochorton fringe Porphyra-Ulo thrix

~ittoral 4 I I Analipas

Sublittoral Sublittoral Laminaria fringe

*Modified from \Veinmanu (1969). littoral region (littoral zone of Lewis) extends from ~leeessarily have horizo~ltal contiguity. Thus, the upper limit of the lichen (Verrucaria) zone within the Hedopl~yllzc?~zzone one call recognize do~vnto the upper limit of Lanlinaria (L. lo~zgipes several associations (including the Herlophylllo?z at Amchitka). The sublittoral region (sublittoral association), each of ivl~ichis named according to zone of Lexvis) has as its upper limit the top of the the domina~ltalga. La,ni/taria population, and its lower limit for the It is gerlerally agreed that the application of algae is defined as the greatest depth at \vhich these terms is often arbitrary and that the implica- benthic algae occur. The littoral region is usually tion of discrete zones and associatio~ls\\'ill not divided into an upper supralittoral fringe area hold up to close esa~lli~latio~lof littoral comm~uli- (Stephenson and Stephenson, 1949; littoral fringe ties. Despite these drarvbacks, apparently no better of Lewis, 1964) and a lo\ver midlittoral area system is available at present with \vl~iehone call (midlittoral zone of Stephenson and Stephenson, compare the general features of littoral zo~lationof 1949; eulittoral zone of Lewis, 1964). a particular locality with those of another area. 364 Lebed~likand Pnb~tisano The classification of zones and associations as used resampled. The sampling datcs of all marine algal in this chapter is intended solely as a general studies are given in Table 1. framework that we hope will enable the reader to visualize the major features of the marine vegeta- Description of Algal Zones and Associations tion at Amchitka. The zonation and floristic compositioli of the littoral region were studied by \\'eintnann (1969) at A,Iakarius Bay on the Pacific Ocean coast and \\'ei~imann's objective rvas to make an initial Square Bay on the Bering Sea coast. The follo\ving description of algal distribution in the littoral description is based on his study. region (\\'einmann, 1969). In March and April Seven zones, each denoted by the dominant 1968, he established one trailsect at h'lakarius Bay species in it, liar~ebeen characterized at Amchitka on the Pacific Ocean coast (T-1) and another at (Table 3). In areas at Amchitka with steep slopes, Square Bay on the Bering Sea coast (T-2). Species the general vertical relationship holds, but flat distributions in the supralittoral fringe were re- topography tends to blur the distinction between corded by visual observation and iu the midlittoral adjacent zones. Horizontally, the upper three zones area and the sublittoral fringe by the removal of are discontinuous, depending 011 the presellce of adjacent 90- by 20-cin samples of algae along the suitable substrate and exposure conditions. Some entire transect line. The abundance of each species of these zones are present only at certain seasons \\.as measured by dry-weight analysis of each of the year. sample. The abundance of crustose coralline algae Only qualitative data \\,ere obtained in the was estimated in percent cover valnes. Associatiorts supralittoral fringe area. Inconspicuous, but some- were studied by placing 35- by 35-cm or 20- by times occupying large areas, each zone of the 20-cm quadrats in the vicinity of the transect (but not \vithin the dry-\\,eight samples) and estimating the percent cover with the use of the Braun- Blanquet method (den Hartog, 1959). The trailsects established by \\'einmann were easily accessible yet sufficie~ltlydistant from the plailncd nuclear test that it \\.as thought possible to use them for two further purposes: (1) To study the seasolla1 changes in littoral species and (2) as controls for any test effects that might be detected in other areas. For these studies the sample areas could uot be disturbed; so a photogaphic salilpling frame (Fig. 5) was designed to sample a 0.25-m2 quadrat by means of an under~vatercamera acconi- panied by a strobe. A photograph and notes for each plot were taken in the field. In the laborato~y the 6- by 6-cm color transparencies were viexved with a hand magnifier over a light box with a clear plastic grid, and the perccilt cover of each species \\'as dctermiiled cluantitati\rcly. Since it was neces- sary that the plants remain undisturbed, the composition and percent cover of the understory vegetation could not be dctcrlnined with this technique. The plots were oliginally located by measuring distances fro~lla base-line point, but later, because of the problems in measuring, each plot \\.as marked by nailing polypropylene rope flags into the soft breccia bedrock. In an attempt to correlate the photographic sanlpliilg method with dry-weight values, a third trailsect (T-3) at Rifle Range Point \\,as sa~npledin May 1969 \\'it11 the use of both methods. This Fig. 5-Pftotographic sampling apparatus used for transect, rvhich ~vasstudied by Lebeclnik, \\,as not quantitati\.e intertidal studies. Ecology of Afarine rllgae 365 suplalittoral fringe consists of a very few species the top of boulders in the beach or moat areas and adapted to the stress conditions of the high littoral in the lot\rer zones on harder portions of the rock environment. Thus only one association can be that project from the benchflat (Fig. 7). The Fucus recognized in each of these zones. zone occupies a vertical range of about 70 cm The highest zone is characterized by the (28 in.). There are distinctly more species recorded lllacroscopic alga Prasiola borealis, a species found in this zone than in the Porpltyra-Ulothris zone throughout the cold waters of the North Pacific. It next above it (Fig. 8). Pylaiella littoralis, an occurs on the tops of large boulders, the sides of epiphyte, is commonly found on mature F. disti- sea stacks, and on high areas of the rampart on cltzcs plants around the island. As Table 4 indicates, rock benches. A luxuriant green carpet of this several species that occur in and characterize species may be conspicuous on promontories associations at lower levels also occur within the where sea birds alight and provide a habitat rich in Flccus zone, but they rarely form associations at nutrients. In places maritime lichens and small this level. Only Iridaea contucopiae, Ulua Iact~tca, nearly invisible blue-green algae may occur with and Gignrti~zapacifica occur with a percent con- this species. stancy greater than 50%. Rltodocltorton />tLrpzcreu?~t,the species charac- The second zone-characterizing canopy species terizing the next lower zone, occuls only in shaded of the midlittoral region is Hedophyllunz sessile. This or semishaded areas. This habitat preference lilnits is the major zone occurring on the benchflat aea. H. its distribution to sea cliffs that are exposed at sessile does not appcar to be adapted to exposed least intermittently to salt spray. Typically, this conditions. Its preferred habitat is on the very flat species can be found in caves or belo\\, oserhanging benches at the appropriate level where it may reach rock on vertical faces, such as at Rifle Range Point, biomass values of 12.2 kg/n12 wet ~veigllt. On but occasionally it may be found in the open, benchflats with a rolling topography, it occiurs in especially where suitable conditions occur with a depressions with little standing water (Fig. 9). The north or east aspect. The vertical distribution of Amchitka plants have blades that lack bullation, the species begins about 40 cm (16 in.) above, and but they have a large, continuous, more or less may extend to as much as 340 cm (134 in.) above, bowl-shaped morphology. In exposed areas, espe- the highest local Fucns distichlts plants. An un- cially on the Bering Sea coast and on \~ertieal determined species of Cladopliora someti~nesoc- ledges, there may be no Hedopliyflzr~~tbetween the cupies as much as 25% of the area in the Fzcczis and the lower lying illaria or Lantitlaria RItodochorton zone, and Ulotltris flncca occurs zones. In these situations the understory associa- sparsely in the lower portion of the zone. This tions are conspicuous. The Hedophyllzr?~tzone has zone seems to occur nlore frequently along the the greatest number of species of all the intertidal Bering Sea coastline than on the Pacific Ocean zones (Fig. 8). coastline, a fact that may be attributed to the Six associations can he recognized n'ithin the north aspect or to the less-~vell-developedbenches Hedopliyllzcn~ zone (Table 4): Uluci, A~taliptcs, and consequent increase in exposure on that side Coralli~~a,Halosaccio~t, Hedof~hyllzc?~~, and Iridaea. of the island. Itnmediately belo\\, the Rltodocltorto~tzone in elevation but much more widespread is the Porpliyra-Ulothrix zone. Species of these genera are dominant components of the vegetation on the boulder shores that are common on the Bering Sea coast (Fig. 6) as \\'ell as at high cliff, stack, and rampart locations. Battgia fitscopur/~vrea occurs sporadically in this zone. The Porpltym species usually have their lower limit solne~vhatabove the Fttczts zone, leaving a band spaxsely occupied by Ulotliris flncca. All but one of the Porphyra species (probably P. pse~tdoli~tearis)occur only in this zone. P. pse~cdoli~tearis,a species that attains lcngths of 50 to 80 cm (20 to 31 in.), is conllnonly found on cobbles in the moat area of rock benches as \\,ell as in the upper portion of the Fircus zone. The upper limit of the midlittoral region is defined as the upper limit at which Fitcns distichlts Fig. 6-Porphyra-(nothri,: zone on the Bering Sea is found. At Amchitka its preferred habitat is on coast, July 20, 1971. 366 Lebedfzik and Palf~lisano

LOCATION OF QUADRATS 1 to 28 29 30 31 32 33 34 35 36 37 38 39 40

PHYSICAL cmT3h r

LOW WATER1 TRANSECT

CANOPY

Fucus distichus

Hedophyllum sessile

Alaria crispa UNDERSTORY Analipus filiformis

Ralfsia fungiformis

0 10% Microcladla borealis OK - UNCOMMON SPECIES Foliose red X X X X X Gigartina pacifica X X X X Halosaccion glandiforme X X ,'MX X X X X Hildenbrandia sp. X X

Leathesia difformis X X X XXX

Odonthalia floccosa XXX X X X X X

Fig. 7-Species composition and percent cover at the T-3 transect Ecology of ~VJnriileillgne 367

ZONE

Fig. 8-Number of algal species (excluding blue-green and unicellular species) reported within the major algal zones at Amchitka.

Fig. 9-Shoreward HedophyNurn zone in Makarins Bay. Plot 2 of the T-1 transect is located on top of the boulder in the rear.

The Airnlipus association usually occurs on portion of the benchflat but forms an association boulders just below the Fztcrts zone and never (together wit11 Clntltro~ilorphzt~~~loczrlosum and C. extends out far along the benchflat. h3ost of the c~~cILII~scI.~~,~~,,~)only on extremely flat areas on year it is recogilizable only as an extensive holdfast the bench. Except for Clntlrroi~ror/~litt~~~locrtlosi~~n system, but in May the upright fertile fronds make and C, circt~nrscriptrim, only Hnlosc~ccio~lglnirdi- this species more noticeable (\\'ynne, 1971). Ulvn for~lre OCCUIS in this association with a percent Inctztca, Iriclnea cor~lrlcopine, and Hnlosaccio~z constancy greater than 50%. glri~d(for~~~eare coin~no~ily associated species. Halosnccio~l gla~zdt'for~~teis one of the i~iost Corallilrn piltilifera commonly occurs over the conspicuous of the association-cliaracterizi~spe- benchflat from belolv the Frccrrs zone to the outer cies. It occuis in ;dl the ;~ssociations of the 368 Lebedtiik and Pabliisano Table 4-Percent Constancy Values of Algal Species in Littoral Algal Zones and Associations of Amchitka*

Zone: Fucrrs Hedophyllum AIaria Lamittnria Corallina Halosaccion Association Ulva Analipus pilulifera glandiforme Hedophyllurn Iridaea

Species Ahnfeltia plicata 10 14 Alaria crispa 20 10 100 8 Analipus filiformir 10 100 40 50 Bossiella cretacea 8 Coilodesme bulligera 40 Corallina pilulifera 30 10 11 100 50 100 20 Corallina vancouveriensis 86 67 Cladophora sp. 10 Fucus distichrrs 100 22 30 10 Gigartina pacifica 60 10 20 30 10 20 28 ITalosnccion glandiforme 40 20 56 50 100 30 100 28 Holy menia 10 20 40 10 IIedophyllurn sessile 20 100 40 14 Hildenbrandia 20 Iridnea cornucopiae 70 50 56 40 80 70 100 7 1 Laininaria longipes 20 20 43 100 dlicrocladia borealis 20 10 40 20 20 Odontltalia floccosa 10 50 20 43 Petrocelis m iddendorffii 20 I1 20 20 I4 Ptilota asplenioides 17 Ptilota filicina 10 20 43 50 Ralfsia ftcngiformis 20 10 22 10 20 10 43 Rhodymenin palmata 11 40 10 Tokidadendroic bullata 20 14 8 Ulva sp. 50 100 89 60 30 50 Encrusting corallines 30 40 11 60 20 80 30 86 100 Total No. species in each association 12 10 10 12 16 14 Total No. species in each zone 13 22 14 8

midlittoral zones and has an average percent Iridaea association is second only to the Hedophyl- constancy of 53%. The most important of the lzcm association in the number of species present in associated genera are Corallirta, Iridaea, and Ulva. it (14 Since this species is dark or frequently The Hedopltyllz~massociation has the greatest black and since it grows low against the rock, I. number of species (16) in it, but outside its cor~izrcopiae is not nearly as collspicuous as II. association H. sessile occurs only in the Halosac- glandifornie, yet it occurs with a greater abundance cion and Iridaea associatiolls of the Hedophyllunt overall. zone, and in no case does it occur with greater than The Ulva association is found primarily in the 40% constancy. Halosaccioil gla~tdifor~iteis often upper part of the Hedopl~yllzrmzone. \\leintnann associated wit11 I;ledophyllzcnt, and encrusting cor- (1969) found U. lnctzsca common during h'lay alline algae and Iridnea also occur frequently in this through September in shallo\v tide pools just below association. The Hedoplzyllum association occurs the FZICZLSzone. Scattered plants of U. lactzlca can over a small vertical range (Weinmann, 1969), but, be found in ally part of the benchflat at most times since this frequently coincides with the level of the of the year, but the species appears to be most benchflat, it covers extensive areas (Fig. 7). abundant in the moat area (Burgner et al., 1969; Iridaea coriizscopiae is the most frequent spe- 1971) (Figs. 3 and 7). The species most commo~lly cies tl~rougl~outall the associatiolls of the intertidal found with U. lactz~cain this associati011 are I. region (average percent constancy of 67%), and the cornzrcopiae and Odo~~tltaliafloccosa. Ecology of ~VIari~feAlgae 369

The Pacific coast intertidal benches of Am- The lo\\rest zone of the intertidal area is that chitka demonstlate that some of these species, characterized by Lantiuaria lottgipes. This sublit- given appropriate conditions, can form nearly pure toral fringe species occupies large areas of rock stands on particular benchflats. Figure 7 sho\vs an down to -3 or -5 m (-10 or -15 ft), thus consti- abundance profile for a transect at Rifle Range tuting a transition species between the littoral and Point indicating extensi\re dominance by H. sessile. sublittoral regions. Most of the species associated Coralli~~apilulifera also sometimes occupies exten- with this zone are sirblittoral species at the tipper sive benchflat areas nearly to the exclusion of all limit of their ranges, and the number of species other algal species. found in this zone (8) is fewer t11a11 that in the The presence of a distinct Alaria zone along the three previous zones. Regardless of exposure or Beling Sea coast may be conelated with the lack of be~lchtopo~aphy, L. longipes appears to be fairly Hedophyllic~i~.Wlen Alaria crispa and Hedopltyl- consistent in occupying ledges from about mean ltctn sessile occur together, A. clispa generally lower low water down\vard. Unlike all the other occupies the more seaward portion but may grade canopy-formitlg kelps at Amchitka, which form a slo\vly into the Hedopl~yllut~tzone. The association single stipe from a limited holdfast, L. lo~lgipes a~~alysis(Table 4) sho\vs that A. crispa occurs in gro\vs by means of a prostrate branching stipe that insignificant frequencies in other associations, but produces numerous upright stipes and blades a high number of species (14) occur in association (Markham, 1972). with it. A. crispa \\'as not present on the Pacific Sublittoral fringe vegetation is found in chan- coast transect in 1968 (\\'einmann, 1969), but we nels (Fig. 1) cutting through the bench and in deep found this species there from 1969 to our final tide pools (which are rare) on the bench. In one visit in 1973. sense much of the bellcliflat could be considered a On the outer face of the rampart, plants of tide pool since at low tide seawater stands up to a various Alaria species form an association in the few centimeters deep in many undrained depres- most exposed area of the bench (Fig. 10). A,lorpho- sions. However, only littoral species are present in logically these plants differ from those of the tbese shallo~sdepressions. The rare deep tide pools benchflat in that they have tufts of thick linear lra\re an abundance of Tlfalassiophyllz~?11clatlfrtls, sporophylls; from each tuft protrudes a short Ptilota spp., Cymatltere tviplicata, and other sub- ragged main lamina, the whole being reminiscent of littoral species in them. L. longipes is only occa- small Postelsia paltifaeforn$is plants seen in similar sionally seen in tide pools. habitats along the west coast of North America. Tatewaki and I

Feb. April June Aug. Oct. Apr. Aug. Species 1972 1972 1972 1972 1972 1973 1973

Alaria crispa 6 7 7 Analipus filiformis 3 3 Clathrotnorphum circtttnscriptun~ 1 2 Corallinn pilulifera (live) 4 4 3 CoraNina pilulifera (dead) 1 3 1 Fucus distichus 4 5 4 Hnlosaccion glandiforn~e 5 5 5 Hedophyllum sessile 12 11 12 Iridaea cor~~ucopdae 6 5 4 Latninaria longipes 4 4 3 Ulua sp. 3 4 3

Table 6-Frequency of Occurre~lceof Conm~on Species in 24 0.24-n12 Plots at T-2 from February 1972 Tllrough August 1973

Feb. April Jirne Aug. Oct. Apr. Aug. 1972 1972 1972 1972 1972 1973 1973

Alnria crirpa A~ralipusfiliformis Clathromorphrrm circumscript~rm(live) Clathromorphum circumscriptrlm (dead) Coralli~rapilulifera (live) Fucus distichus Hnlosaccion glandiforme Hedophyllum sessile Iridaea cor~racopine La~ninarialoirgipes Ulua sp.

*Dead plants rvere found with a frequency of4 in this sample.

in ~vinter,probably February-i\,larch. In agreement great deal of variability in frequency of occurrence. with IVeinmani~ (1969), wc observed that the This species is a slow-groxsing elustose coralline illcrease in cover \\.as due to the growth of new alga, and seasonal variability in cover would not be up:.ight thalli in early summer, altliough a few expected. Holvevcr, the percent cover on four plots plants survi\'ed for two summers. at T-2 sho~\.cd definite seasonal variation and Iridnea conzzlcopiae has a perennial crustose die-off (Fig. 15). This undoubtedly resulted from base that produces upright blades each year in the removal of silt by stornls and uncoveri~lgof early spring. The frequency of this species (Tables plants in winter and buildup of silt and burying of 5 and 6) \\,as remarkably constant throughout the plants in summer. As shown by the peak in dead year. Figure 14 shows that the cover of the species cover in Fig. 18, seine summer mortality occurred (based on the upright blades because the crusts are in the population, but some plants survived and i~npossible to distinguish in the photographs) colltinued gro~vingthe followi~lgwinter. peaked in early summer and declined thereafter as Alaricc crispn sporeliilgs appear in the spring, the blades became fertile and disintegrated in the and growth continues throughout the summer. A process of releasing spores. slight increase in frequency \\,as seen in the spring Ulva lactuca sho~veda great deal of irregularity of 1972. An even greater increase in frequency was in its frequency of occurrence. This is a species recorded in the spring of 1973, ~rrhenA. crisf~nIvas characteristic of the moat area of the benchflat, an also esccptioilally abundant in other midlittoral area of annual deposition and removal of silt. This areas we visited. This Isas the oilly instance we factor coiltributes to a strong seasolla1 variatioil in recognized of a definite annual variation in this cover. species' abundance at Amehitka. Because illaria Another species commoil ill illoat areas, plants are strap-shaped, attached at one end and Clatltrontorpltz~,nt circlctitscri[~tir~tt,also SIIO\VS a usually much larger than a 0.25-in2 quadrat, FEB. APR. JUNE AUG. OCT. APR. AUG. 72 72 72 72 72 73 73 MONTH OF SAMPLING

Fig. 14-Percent cover of Iridaea cornucopiae on plot 3, T-1 transect, from February 1972 to August 1973.

Fig. 12-Growth and reproduction of Fucus dis- tichtrs plants on plot 7 of the T-l (blakarius Hay) transect. A, Feb. 22, 1972; 8, Apr. 15, 1972; C, June 10, 1972; D, Aug. 20, 1972; E, Oct. 22, 1972; F, Apr. 5, 1973; G, Aug. 25, 1973. Stippled portions are receptacles; the dashed line in "A" indicates the outline of a hard volcanic rack fragment on \vhich the plants were growing. n4 0 L2/.%I OCT. FEB. APR. JUNE AUG.OCT. APR. AUG. 71 72 72 72 72 12 13 13 I MONTH OF SAMPLE Fig. 15-Percent cover of live and dead Clathro- morphum circnn~scriptu,n on the T-2 transect from October 1971 to i\ugust 1973. a, plot 2. X, plot 3. o, plot 4. 0,plot 9. Species cover too small to measure accurately is assigned a value of 0.1%.

abundance data from the trailsects are poor mea- A 0 sures of growth and recruitment. Generally Lami~zarialotzgipes exhibits no sea- X-X sonal change in its populations. There were occa- OCT. FEB. APR. JUNE AUG. OCT. APR. AUG. 71 72 12 12 12 72 13 73 sional dead patches, someti~nesover a meter in MONTH OF SAhlPLlNG diameter, xvithin othencrise healthy areas of the species (Fig. 16). This phenome~lon caused a Fig. 13-Percent cover of Holosaccion glandiforme on the T-2 transect from October 1971 to August decline in frequency on T-2 as seen in 1973, 1973. a, plot 3. X, plot 4. 0, plot 8. 0,plot 9. A, plot although visual observations indicated that in the 11. Species cover too small to measure accurately is zone as a whole die-off was more extensive in June assigned a value of 0.1%. 1972 0. Isakson, personal communication). The Ecolog), of 11larirte Algae 373 holdfasts, stipes, and nleriste~naticareas dcgcneratc and the blades are lost con~pletely.

Conlpariso~lwith Other Regions There are many inherent problems in the comparison of studies of zonation from different parts of the North Pacific Ocean with our studies at Amchitka. Aside from any real differences that may exist, different application of vegetation terms may be found in different countries and even between x\~orkerswithin one country. Misidentifi- cation of species or other taxonomic proble~nsmay strongly influence the apparent similarities or differenccs between two studies (particularly in the older literature \ellere recent taxonomic changes might make comparison with present species names very misleading). Finally, one must ask how representative of an entire geographic region a particular study is. These problems can be reduced somewhat by concentrating on specics that are Fig. 16-Dead patches of Laminnria longipes at conspicuously and consistently dominant either as Duck Cove near IA-1, Aug. 26, 1973 (meter stick in a zone or as an association, arc easily identified, one patch). and/or are known to be characteristic of particular environmental conditions. Kardakova-Prezhentsova (1938) made a detailed study of the littoral region of the Commander Islands. She found a zonation pattern very similar, if not identical, to that at Amchitka. The supralit- toral zone was characterized by Urospora perticil- lifontiis, Baicgia $iscoptr~tcrea, Porpl~yra sp. (all winter species), and Prasiola borealis. The principal species of the upper littoral zone (comparable to the midlittoral Fttczts zone at Atnchitka) were FZLCNSdtiticfcus (as F. eua,cescerts*), Gloiopeltis firrcata, hlortost~omagreuillei, and ilrtalipus japoit- icus. t The occurrence in this zone of Clcordaria flagellifon7cis, an arctic species, suggests a slightly colder environment than at Amchitka. The me- dium littoral zone of ICardakova-Prezhentsova is equivalent to both the Hedopltyllzrtrt and Alaria zones at Amchitka. This difference probably re- flects a matter of opinion rather than a significallt difference in zonation because fllaria is scaso~~al and does not form a zone every~vhereat Amchitka. The major specics of the medium littoral zone at Fig. 17-Reproductive Lnminaria longipes on plot the Co~nmanderIslands were Alaria larcceolata, 22 of the T-2 transect, April 1972. The photograph is Hedophyllzori sessile [as If. st~bsessile,see wid do^\.- of a 0.5- by 0.5-m (1 %- by I %-ft)area. Reproductive son (1965)], Rl~odomelaIarix, and Fttctcs disti- areas that have released spores are conspicuous as chus. The lolver littoral zone at the Connnander white margins on blades. Islands (comparable to the sublittoral friuge Lami- ?,aria zone at Amchitka) was dominated by Lanti- ?!aria lo~tgipes, L, der~tigern, l'halassioplr)~lhtm release of zoospores results in ~narginal linear colorless patches on the fronds (Fig. 17), wvl~ichare 'Ftccus eunnescens is now recognized as a subspecies of most conspicuous in April each year. This repro- F. distichus (Powell, 1957). ductive process is apparently not related to the ?For synonyms used in the literature (Chor[llnria abie- mortality mentioned because in the later case tina and IIeterochordaria abietitta), see TVynne (1971). 374 Lebedt~iltand Palntisa~to clathrtts, Codiztn~adlterews (probably Codilcnz rit- The littoral zo~latio~lof Sakhalin Island in the teri), and Pterosiphonia Dipitz~~ata.Except for the Okhotsk Sea was recently studied by Vozzhinskaya absence of the Halosacciotz glandiforme and Iridaen (1964b). On the southern coasts of the island, the couttccopiae associations, the preceding description supralittoral area was dominated by Gloiopeltis of zonation at the Commander Islands agees very capillaris, which was often associated with Pop well with the results of our studies at Amchitka. pltyrn sp. and Urosporn sp. The littoral region was characterized by three horizo~ls(comparable to the spasski; (1961) reported on the littoral zona- midlittoral zones of Amchitka), which were, from tion of southeaster^^ Icamchatka. In the supralit- top to bottom: (2) toral region, two "horizons" \\,ere distinguished: (1)Aual@us ja/~otticus, Coral- litfa pilulifera and Iridaea conttccopiae (Fucus in (1) An upper first holizon of Rltodochorton protected localities), and (3) Cltordarin n~agellanica pt~rp~cretct~zand Battgia fitscopttrpurea and (2) a lo\ver second horizon of Rltizoclouiu~~t~if>ari~~?tt. and Lantittaria japonica. The occurrence of Sargns- on the southern coasts once again indicates a These horizons are probably comparable to the sunt Rhodoclzortott and Porpltyra-Ulothrix zones, re- transition to a warmer water flora. From the comparison of the above studies with spectively, of the sup~alittoralfringe at Amcl~itka. Spasski; recognized three horizons in the littoral our study of Amchitka, one can conclude that the region which were, from top to bottom: (1) Gloio- littoral zonation of the Commander Islands is peltis colifor~tlis,(2) A~~alipusja/>o~zictrs, Chordaria nearly identical with that of Amchitka. The vegeta- flagellifomtis, and (3) Corallitla and various Rhodo- tion of the Kurile Islands is generally similar, but pltyceae and Laminariales. Ftccus distichus, Por- that of southeastern Kamchatka is less similar. phyra perforata, Ulua lactnca, Lantinaria lottgipes, Finally, the islands of Hokkaido and Sakl~alin and three species of Alaria were also present in the bridge the transition to a distinctly warmer water littoral region but were apparently not sufficiently flora. abundant to establish zones or associations. The Practically no studies have been made of geog~aphical distribution of Hedopltylltcm sessile littoral zonation in the northenl Bering Sea, which does not extend beyond the Commander Islands precludes a review of the arctic affinities of the (Icardakova-Prezhentsova, 1938; Widdomson, Amchitka flora. 1965). Along the west coast of North America, sur- prisingly few descriptive studies of littoral zonation The littoral zonation of the ICurile Islands was have been done. Rigg and Miller (1949) reported studied by Nagai (1940; 1941). In the northern on studies of littoral zonation at Ned1 Bay, on the Icurile Islands, he recognized a cold-water flora Strait of Juan de Fuca. They distinguished seven that he na~nedthe "upper boreal district." Within zones in the littoral region, \\rhich were, fro111 top this district three fo~mationswere recognized in to bottom: (1) Ralfsia-Pmsiola, (2) Et~docladia- the littoral vegetation: (1) An upper Fucus disti- Gigartiua, (3) Postelsin, (4) Halosaccio~t glmtdi- chtts formation (including Halosacciott saccatfcnt, fornte, (5) Alaria, (6) Lessoniopsis, and (7) Lnlni- Ulua fe~testrata, Porpltyra spp., Iridaea contfcco- ?laria anderso~tii.The first zone occurred in their piae, Ulothrix psetidoflacca, Analipus japonictis, "splash zone" (comparable to the supralittoral and Corallitta piltclifera), (2) a loxver Alaria angtts- fringe area of Amchitka). The second zone strad- tatn formation, and (3) a Lan~it~arialo~tgipes for- dled the splash zone and the "upper intertidal mation. Thus it \vould appear that the zonation of zone," the latter including zones 3 and 4 (com- littoral algae in the Icurile Isla~ldsis more similar to parable to the Fucus and Hedophyllzcnt zones of that of the Commander and Aleutian Islands than Amchitka). The "lower intertidal zone" of Rigg that of the southeaster~lcoast of Icamchatka. Only and Miller included zones 5 and 6, and zone 7 was the midlittoral Iledoplzyllzr~~zzone was not ob- classified the "demersal zone." Zones 6 and 7 served in the ICurile Islands. appear to be comparable to the Latttinaria lolzgipes Considerably less affinity in littoral zonation is zone at Amcllitka. Both FILCI~Sdistichus and seen on the east coast of Hokkaido. Thc main Hedoplzyllzr~tzsessile occurred in abundance in the "belts" recognized by Taniguti (1962) were, from more protected areas studied by Rigg and h,filler. top to bottom: Porpltym pseudocrassa, Peluetia The above review indicates the variation of zoriglttii, Fzccus clisticltzts, and Lrotzinaria longiee- classification schemes as used by different workers dalis. In exposed localities Peluetia and Fttczts were in various pats of the North Pacific Ocean. The replaced by Attalipus japo~tic~isand Iridaea cortzzc- presence or absence of the large kelp species in the copiae (as Iridopl~ycus).Coralli~~a pilzcliferct was midlittoral area appears to have the greatest in- also present. The presence of a distinctly soutl~ern fluence 011 littoral zonation schemes adopted by element, Srtrgass~~t~t,signals a transition to a various authors. In contrast, the supralittoral fringe warmer water flora else\vhere on Hokkaido. area can be recognized in most localities by the Ecolog)~of ~VlarineAlgae 3 75 presence of characteristic genera (Prasiola, Pop these factor? and to aid in predicting the cffects of phyra, Uloti~~is,and Bangia) of widc geographical nuclear testing on littoral algal comnlunities. In distribution and with a strong seasonal difference these experiments either the entire community or in abundance. The sitblittoral fringe area is charac- the canopy only mas removed from pertnanently terized in exposed localities by a dominance of marked plots, and the subsequent colonization was kelps wit11 massive holdfasts and strap-shapcd recorded. blades. According to some authors (Taniguti, 1962; Methods Vozzl~inskaya, 1964b) Fzrctrs occurs in abundance only in relatively protected areas. The same distri- So that differences in colonization due to bution appears to characterize Hedophyllrcm sessile exposure could be acco~unted for, plots wcre (Rigg and Miller, 1949). The n~orphology of established at sheltered (R'lakarius Bay), inter- Hedopl~yllut~tplants at Amchitka (smooth entire n~edi.ate (Duck Cove), and exposcd (Riflc Range blades) also indicates a relatively protected cn- Pt.) areas. These plots were completely indepen- vironment (see \Viddowson, 1965). These facts dent of the T-1, T-2, and T-3 transects. Local correlate with the idea presented above that the differences in colonization in the midlittoral area rock-benclt topography provides an exposure were sti~diedby establishing plots in pure stands gradient so that zonation of an exposed type is (associations) of the follo\\~ing species: Alaria obscrved on the pro~nontories and outer bench crisps, Hedopliyllu?~~sessile, Coralli~za uancou- areas, whereas a more protected zo~latiotlpattern, veriensis, and Halosaccioil glaln,~diforn~e.In the including extensive ITedophyllrrm and Frrcrcs zones sublittoral fringe arca, plots \\rere establisl~cdin the and a diverse flora, characterizes the benchflat Lanti~rarinlo~~gipes association. Not all associatiotls area. were sampled at each locality (see Table 7). Sea-

Table 7-Summxieed Descriptions of Plots Established for Denudation and Canopy Reinoval Studies at An~cliitkafrom hlarcl~1971 to April 1973

Number Species of plots Zone Locntion Rlanilipk~latioa

Lamitrnria losgipes Laminaria Makarius Bay Rock denuded Laminnria lollgipes Laminaria h'lakarius Bay Canopy removal Hedophyllun~ssessile Hedoplryllum blakarius Bay Rock denuded Halosaccio~tglandiforme HedophyN~im Makarius Bay Rock denuded CoraNiitn un~rcorcuerienris Alorin Rifle Range Point Rock denuded Alnrin crispn Alnrin Duck Cove Rock denuded Alnrin crispn Alnria Duck Cove Canopy removal IIedophylls~~rsessile Hedophyllum Duck Cove Canopy removal Hedophgllum sessile Hedophyllum Makarius Bay Canopy re~noval Corallinn unncorcuerie~rsis Alaria Duck Cove Rock denuded

* 1-m2 plots (1 by I m). t0.25-m2 plots (0.5 by 0.5 m).

COLONIZATION STUDIES solla1 differences \sere studied by clearing plots in some areas at nearly bilnonthly itlter\gals from Denudation experiu~~cntsaltd studies of subse- klarch 197 1 to December 1972 and in April 1973 quent colollization hase been suggested as a mcans (see Table 1). Two types of n~allipulatiolt(each on of understandillg the Factors influencing the zona- separate plots) were used: (1)Total denudation of tion of littoral algae (Chapman, 1943). Such rock surface by the remo\'al of all organisnls and studies have bee11 conclucted in 1-Ialvaii (Neal, (2) rc~novalof the canopy species only by selective 1930), California (Northcraft, 1948), \\'ashington cutting of stipcs. (Dayton, 1971), England (I(itching, 1937; Kecs, For a study of colonization on newly denudcd 1940), Soutlt Africa (Bokenllam, 1938), and elsc- rock, 1-mz areas of L. longipes, A. cris/m, and \shere. No si~cllexperinlerlts had been performed Hedopizyllu~nsessile and 0.25-m2 areas of Coral- in the Alcutian Islallds prior to our studies. linn va~tcozrveriensis and Halosacciotr gla~rdiforn~e Starting in 1971 Palmisallo (1975) cond~~cted \\,ere cleared svith a shovel, putty knife, and wire denudation esperitnents at Anlchitka to delitleate brush. Each area was burned twice with a 1 : 1 mixture of fuel oil and white (unleaded) gasoline. toms and filamentous brown aud green algae; For an evaluation of the effectsof canopy re~noval (2)ulvoids, i.e., small leafy green algae (Ulua on colonization, the blades and stipes of L. Iuctlrca and A!lo~tostronta spp.); and (3)macro- lo?tgi)es, A. crispa, and Hedophyllum sessile were phytic red and brown algae (Iridaea corttucopiae, removed with a wire cutter in 1-m2 areas (Table 7). Halosaccion glattdifor~~te,Hedophyllunt sessile, All plants and animals were removed from a Alaria crispo, Fucus disticltus, Coralli~lavancotc- 50-cm(20-in.)-wide strip around each plot to mini- uerieitsis, Lnmitzaria longipes, etc.) (see Table 8). mize physical disturbance from adjacent algae Succession observed in areas where the canopy was during surge at high tide and from grazing by removed was similar to that indicated earlier, but benthic invertebrate herbivores (sea urchins, lim- the rate was faster (Table 9), primarily because pets, and chitons). One disturbed plot and one they received a lower level of experimental distur- undisturbed control plot were established for each bance. In the canopy-removal studies, only Lnmi- study area. nuria exhibited vegetative growth from its hold- So that study areas could be located and fasts, a site of meristematic tissue in this kelp. identified, one corner of each was marked with a Alaria and I~edopltyllzc?~tplauts did not regenerate 0.3-nl length of cord secured with a concrete nail because their meristelnatic region occurs in their and a numbered washer. Each 1-m2 area was blades. Hedophyllu~ttsettled in all areas where the divided into four contiguous 0.25-m2 plots that canopy was removed, but Alaria settled only in the were delineated with concrete nails and identified Ln?tti?laria zone. For Alaria to settle, bare substra- with a numbered piece of survey flagging. Each tum, wllich was more plentiful in the Lnmi~zaria 0.25-m2 area \\!as further marked by an additional zone than in the Hedophyllunt zone or its own, is concrete nail and numbered washer placed in the required. corner diagonally away from the cord. Others have noted similar successional patterns Data were collected from all study areas (\\'ifson, 1925; Bokenham, 1938; Northcraft, (experimentally disturbed and control plots) at 1948; Southward, 1956; Dayton, 1971). Boken- nearly bimonthly intervals from March 1971 to ham, Northcraft, and Dayton have described December 1972, in April 1973, and in August various groups of colonizing algae. In general, these 1973. At each time each 0.25-m2 plot was photo- are (1)pioneer species (diatoms); (2)fast-growing graphed during lo\\, tide, and species compositio~l annuals (ulvoids and Halosaccion); (3)fast-growing and percent cover were determined for algae on perennials (Hedopltyllzr?n and Alaria); (4)slow- that plot. The photographic sampling frame de- growing perennials (La~ni?taria and Corallinn); scribed in the section Littoral Zonation \\,as used, (5)canopy-producing species, i.e., those which are but the photographs were taken with a 35-mm nlost successf~~lin competition for light (Hedo- camera. pltylltc~nand La,~tittaria);(6) obligatory understory species, i.e., those which require a canopy to Results and Discussion survive or are more often subtidal (coralline algae); The sequence of algal species observed on and (7)fugitive species, i.e., those which exist in denuded rock benches at Amchitka %%'as(1) dia- disturbed areas or on the fringes of communities

Table 8-Optimum Algal Colonization Scl~eduleObserved on Denuded Rock Surfaces at Alncllitka Island, Alaska, from hlarcl~1971 to August 1973

Optimum clearing Period of time Time to attain Season of time for rapid before alga at least 75% Alga settlement recolonization is observed coyeraee

Diatoms* AIL All \\'ithin 1 month 1 month Ulvoids* All All 2 to 4 months 2 to 4 months Iridaea * Pall and winter Summer and fall 2 to 4 months t Halosaccio~~ Summer and fall Fall 4 months 6 months Hedopl~yllum Spring and summer Pall and winter 4 to 6 months 1 year Alnrin \\'inter Summer and fall 4 to 6 months 4 to 6 months Fucus* Summer and fall Spring and summer 6 to 8 months 14 to 16 months Corallina Spring and summer$ Fall and winter$ 6 to 8 months$ t Laininaria Fall and winter$ \\'interS 8 to 10 months$ t *Settlement did not occur in its own zone. tCover never exceeded 18% (see Table 10). $Little information is available. Ecolog)! of hlari~zeAlgae 377

Table 9-Optimom Algal Colonization Sclledule Observed in Canopy-Removal Areas at Alncliitka Island, Alaska, fro~nhlarch 1971 to August 1973

Optimum clearing Period of time Time to attain Season of time for rapid before alga at least 75% Alga settlement colonization is observed coveraee

Diatoms* All All \\'ithio 1 n~onth t Ulvoids* 1\11 All 2 months 2 to 4 montbs Iridaea * All 1\11 2 months t Halosaccion* Spring, summer, Spring, summer, 1 month t and fall and fall HedophyNum Spring, summer, Summer, fall, 2 to 4 months 11 months and fall and winter Alaria* Fall and winter Summer and fall 2 to 4 months 10 to 12 months Fucus* Summer, fall, Sp~ing,summer, 4 to G months t and winter and fall Corailina $ $ $ t Laminaria Fall and winter 5 Fall and winters 2 months 3 7 to 9 months8

'Settlement did not occur in its own zone. tCo\.er never exceeded 62% (see Table 11). $Little information is available. sColonization \\-\.asby vegetative growth.

(Porpilyra and ulvoids). This last category, sug- ulvoids occurred in summer on areas disturbed in gested by Dayton (1971), is so broadly defined winter. that it could include diatoms, ulvoids, Halosaccio,l, Species whose stands were denuded started to or even illaria at Amchitka. colonize their original areas immediately after Most algae that colonized disturbed study areas diatoms and ul\,oids disappeared, but only if the at Amchitka settled in broader vertical expanses area was cleared just before the season nrhen their aild in more diverse exposures than they normally spores were released and settled, ~vhichwas in- occupy (Tables 10 and 11). At the indicated ferred fro111 the subsequent appearance of exposures to wave action wd heights above sea sporelings (see Table 8 for thc optimuin time level, the follo\ving algae were the most abundant: period). If a species was cleared too early or too Diatoms, in all exposures and at all levels; ulvoids, late relative to the time its propagules settled, then in all areas except those most exposed to \vave any species \vliose propagules \\.ere settling ~vlten action; Halosaccion, at higher levels in areas of the diatonls and ulvoids disappeared occupied the little or intermediate exposure; Hedoplzyll~~nz,in area. Thus a Hedophyllz~ntplot cleared in Decem- protected areas at lo\ver and intermediate levels; ber (early winter) \\.as replaced by Hedophyllz~n~. Alaria, at lower levels and in exposed areas; and a Halosacciora plot cleared then recovered to Laminaria, at lolver levels at all exposures; and Halosaccioia. Ho~vever,a Lamittaria plot cleared in Coralli~~a,in exposed areas at intermediate and the fall contained Alaria by spring, and a Halosac- lo\ver levels. In general, initial settlement rate was ciotl plot cleared in spring contained FILCUSthe faster in exposed areas at lower levels than in follo\ving year. protected areas at higher levels. Three of the five species whose stands nrere Diatoms and ulvoids settled on denuded rock denuded and ttvo of the three whose canopies were surfaces during all seasons. Their year-round repro- removed persisted and increased their percent ductive ability enabled them to occur whenever cover when thcy settled in their o\vn zones. The space \\.as available. All other algae settled on areas that did not contain at least 80% of the denuded rock during specified seasons (Table 8) original algal species at the end of our study in \\.hen their spores were available. In aeas where April 1973 were the Laminnria aud Coralli~la the canopy was removed, settlement of most denuded areas and the Alaria canopy-removal species occurred almost througliout the entire year areas. A low reproductive potential may have becn (Table 9). Again this was correlated xvith the lo\\. responsible for the sparse settling of Laminaria level of esperime~ltaldisturbance. loizg$>es. Laboratory esperiinents (Markham, Diatoms persisted for 1 to 8 months after 1972) indicated little sexual reproduction by this settling, and ulvoids persisted for 5 to 15 months kelp. At Amchitka coloiiization by L. lorzgi/>es after replacing the diatoms. Peak densities of the occurred primalily by vegetative gro~vth.11s stated above, the lack of bare substratnlt probably For example, lo\\, temperatures, with little prevented settlelncnt of illa~iain areas x\ll~erethe seasonal fluctuation, high humidity, and lo\\. le\~cls canopy was remo\wl. rllaria and Hedopl~yllzt?~~did of direct insolation, provide favorable conditions invade other areas, especially the La?,~i~lnriaplots, for desiccation-sensitive littoral algae. Other physi- and large percent cover values were recorded for cal stresses, LC., \vave shock and abrasion, also these species at the end of tlte study. Algae that appear to be reduced, as sho\\rn by the abundance settled outside their normal area tvill not persist of species characteristic of protected en\'~~rolllnents. there indefinitely, but it is not known ho\v long it This protection is due to a reduction in \va\,e \\fill take the original climas vegetation to establish action causcd by tlte sheltering effect of offshore itself. kelp beds and by tlte topography of the intertidal rock bencltes. Herbivory, an important biological Causes of Intertidal Zonation factor in littoral communities, is redltccd otving to the reduction of the ntajor herbivores by intensive The rcsults of the colonization studies can noxv sea otter predation (Palmisano and Estes, Chap. 22, be related to the physical and biological features of this volume). The relatively low levels of physical the Atnchitka exlvirolllncnt and can be used to and biological stresses result in a relatively simple understand the dynamic aspects of the present system in wlticlt the important illteractioils are littoral zonation. Competitive success may be probably confined to direct competition for space influeltced by physical featnres of thc habitat and anlong algal colonizers. by biological interactions. Some of the physical Each of the species studied in the denudation stresses typical of littoral communities appear to experiments is a dominant in the Amchitka littoral be relatively ameliorated at Amchitka. region and colttpriscs either a zone or an associa-

Table 10-Maxin~am Percent Cover Attained at Any Time by Intertidal Algae ie Five Different Denuded Rock Study Areas at A~ncllitkaIsland, Alaska, fron~hlarcl~ 1971 to August 1973

Percent cover in denuded rock areas

Laminnria Alaria HedoPhvllunt. , Halosaccion at at at at at Rifle at Algae Makarius Bay Dnck Cove hlakarius flay Alakarios Bay Range Point Duck Cove Diatoms 100 100 90 Ulun lactrccn 100 20 82 Iridnea con~ucopiae 18 T Halosaccioit glar~rlifor~ne 29 5 95 Hedophyllum sessile 100 Alaria crispa 98 Fucus distichus 89 Cornllina uancouverie~~sis Laminaria longipes Filan~entousbrown Filamentous green Spongon~orphasp. Gigartinn pncifica Porphyra spp. Xhody ntcnia palmnta illicrocladia borealis Tokidndendron ballata Durnontia sinlplex Foliose red* Crustose coralline Anniipus spp. Scytoriphoft lomen taria Leathesin diffortnis Petnlo~tinfascia Cymathere triplicata

*T, trace, < 1%. +Circled values indicate the dominant species prior to denudation. f Unidentified red alga. $Upper sublittoral alga. Ecolog~of hlarit~eAlgae 377

Table 11-Maximum Percent Cover Attained at Any Time by Intertidal Algae in Three Different Canopy-Removal Study Areas at Amcl~itkaIsland, Alaska, from hlarcll 1971 to Aurmst 1973

Percent cover in canopy-removal area

Laminnrin HedoPhvllum. , Alaria at at at - hlakarius Bav Duck Cove Duck Cove Diatoms 14 56 UIua lacttcca 78 55 79 Iridaea cornacopiae 5 49 13 Halosnccion glandiforme 39 62 2 Hedophyllum sessile 100 @* 36 Alnria crispa 100 3 Qt Fucts disticl~u~ T 35 T CoraNinn pilulifern 5 5 Lantinaria longipes* 6 Filamentous brown T Spoltgomorpha sp. T -Gigartinn pncifica T T T Porphyra spp. 1 T 2 Odon thalia floccosa T Rhodynte~~iapalmafa 9 1 T iVficrocladia borealis 35 T Tokidadendron bullata T 1 Dumontia simplex 14 4 Foliose red$ 7 14 14 Crustose coralline 5 9 Analipus spp. T T Scytosiphon lomenfaria T Leatltesin difformis T Cymntltere triplicatan T

*Circled values indicate the dominant species prior to denudation. tT,trace, < 1%. $Vegetative growth. $Unidentified red alga. gupper sublittoral alga. tiou. The attributes of these species, which may withstand wave exposure as well as the round give them a competitive advantage, can nolv be stipes and thin blades of Lanlinavia spp. Alaria spp. collsidered in view of the results of the coloniza- nor~nallyoccur in areas of slowly receding tides tion studies. and where there is wave splash. They settle during Lanti~zariaspecies, principally L. lo~agipes,char- winter (a season when few species settle) and grow acterize a zone in the sublittoral fringe area. A rapidly, both of which characteristics appear to be secure holdfast system, round stipes, and long competitively advantageous. Alaria's dense canopy narrow blades may allo~rthcm to occupy more (and possibly the \\hiplash effect of their blades) exposed areas, but their apparent inability to may also help in the competition for space. withstand desiccation probably restricts them to Corallitla species, particularly C. pilulifera, this region of extensive spray. Their dense canopy occur throughout the tIedop/tyllt~lnzone, at times excludes ~llostother algac except crustose coralli~le it1 sufficient density to characterize at1 associatioll algac from occupying their zone as understo~y within that zone. Their small size and secure species. The colonizatioll experime~lts indicate holdfasts make them usually an understory species. that, when disturbed, Lnl~bblnrialol~gipes is slos\r to Their environme~ltaltolerance is apparently quite returu u~llessholdfasts remain from whiclt vegeta- wide, but they are slow to return when disturbed. tive growth can take place. Heclopltyllz~m sessile is a canopy species of Alaria species, principally A. crispa, character- sufficie~ltdominance to characterize an associatio~l ize the next higher zone. Their disk-like firmly and a zone of its own, the zone that covers most of attached holdfasts and lol~gflat stipcs and blades the middle intertidal bench (Figs. 7 and 11). A probably enable thcm to occupy exposed areas, weak holdfast, the lack of a stipe, and a large but the flatness of these blades and stipes may not inflexible blade probably exclude 13. sessile from 380 Lebed~tikand Pnlnrisnt~o the exposed outer edge of the bench but are likely not disadvantageous on the protected middle re- gions. Tlie colonizatioii experiments indicate that it is a vigorous colnpetitor that quickly recovers fro111 disturbance. Hnlosnccioiz gla~tdifornte is an understory species of the ntiddle bench. Its ability to retain seawater in its thallits during lo\\' tide possibly hclps it to tolerate dcsiccatioll and to occur at high intertidal levels. It timy bc helped in coinpetition for space by tlie fact that its spores settle at all times of the year except \\'inter. Ividnea cor~tztcopine also characterizes an as- sociation within the IIedophyllltnt zone. A peren- nial alga, it outcompetes H. glnttdifornte (an annual) at higher levels by perma~ientlyoccupyilig space aild presumably prevents H. gla~~difornle spores from surviving. I, corstucopine occors in slight deprcssioiis that tend to accuin~~latesilt, and thus this tolerance to silt may enable it to outcompete other algae in this area. The ulvoids, Ulvn lnct~rccc and ~\lot~ostvo~tla Fig. 18-Underwater photograph of the kelp bed at spp., are abundant probably as a result of their -10 m (-33 ft) near Kirilof Point on Apr. 23, 1972. spores being a\,ailable and settling year round. I\ dense lower canopy of Lanlitzaria pmenlandicn Fzrctrs rlisticitzcs characterizes tlie highest of the (broad blades, lower foreground) and a few plants of Aforia fistl~losn floating torvard the surface (upper midlittoral zones. It not only withstands desicca- background) are visible. tion well but also it inay actually require it. Johnsoti et al. (1974) showed that I;. disticlztcs rcaclies maxinium photosynthesis after some de- gree of drying. It may be absent from lolver zones in width. We made no measurements of the length because of increased exposure there (Southward, of plants owing to collection problems, but we 1956) and because it requires high light iiltcnsities observed that the densest bcds occur in water of 5- (Gail,1918) that would be absent under the kelp to 15-m (16- to 46-ft) depth, where by late canopies of lolver zones. summer blades of plants often are over 10 m (33 Not all species of littoral algae at Amchitka ft) long along the surface. Tlie reports of total occur in distinct vertical zones. Some species occur blade lengths up to 25 ln (82 ft) (Setchell and in seemingly random places on the intertidal bench Gardner, 1903) should be considered minimum for or outside their normal location. This soggests the large plants in late summer. At the lo\vest tides of existence of favorable microhabitats or of naturally the year, a felv of tlie shallowest A. fistztlosa plants occurring disturbailces that permit their temporary may be eineised on seaward-sloping rock benches presence. not dominated by other kelps, but it cannot be considered an intertidal species. The deepest plants were observed at about 20-m (66-ft) water depth. SUBLITTORAL COhIhIUNITIES On the shallow Pacific coast, bcds may be seen several kilometers offshore. These beds have a great As emphasized by \Vynnc (1970a), the state- dalnping effect on \\raves. In Illoderate seas, there is ment by Ruprecht (1851, p. 97) that coasts of the usually only gentle surging on a coast behind these Bering Sea are depanperate of algae is "grossly beds. This is probably a major reason for the inaccurate." Nowhere is this more evident than in relatively protected aspect of the littoral coiiiniu- the sublittoral communities. Po\\.crs et al. (1960) nities at Amcliitka. conimeilted that the floating kelp beds are dcnse As seen at the surface, f1. fistttlosa beds arc enough to impede transit of small boats. Thesc small in winter, but by late summer they reach a beds are formed by a single species, Alnria fistulosa peak of density, and much of the nearshore area is (Fig. 18). This is the largest species in the genus covered. This change results froin the gro\\~tliof and is distributed from the southeni Icurile Islands individual plants, at least some of which persist to southeast Alaska (\Viddowson, 1971a). At tlirougll the \\,inter. It is also brought about by an Amcliitka we saw plants mith blades over a meter increase in population size in the spting. Since \\re Ecology of Marine Algae 381 I did not conduct a tagging study, the longevity and I population turnovcr of these plants were not dctcrmined. Most of the population may be annual with few plants remaining throughout the ~vinter; ho\\,e\rer, winter storms may break off the floating fronds but leave the holdfasts intact to produce ncxv blades the follo~\~ingspring. Other species of ~llariaare perennial and have a pcak of summer gro\sth (\Viddo\uson, 197la). Specimens of i\'ereocystis ltcetlteaifn, the well- known floating kelp of the West Coast of the United States, arc coinmonly found drifted onto the beaches of Amchitka and a few have been found tangled in offshore kelp beds, but no attached ~Vereocystirplants have been observed at Amchitka. The same situation occurs in the Com- lnander Islands (Kardakova-Prezhentsova, 1938). These plants must drift along the Alaska Stream current from the \vesternmost point of its known distribution, Unalaska Island (Druehl, 1970), or from points still farther east.* Beginning with the Lanzi~faiialottgipes popula- Fig. 19-Under

*Asterocolnx hypophyllophila, Hypophyllum dentn- Milrow-Affected Areas turn, Laingia oleulica, Neinburgia prolifern, Phycodrys a~nclzitke~~sis,and Zhovoen aca~~thocnrpo(see also foot- Lebednik (1973) reported the effects of the note, p.358). nuclear test Milrow, which caused a fault shift Ecology of A'laritte Algae 383 resulting in an uplift of about 12 cm (5 in.) of an intertidal area (Fig. 20) of about 4 ha (10 acres) on the Pacific Oceau coast. The uplift caused die-off in certain algal populations, especially those of the Hedophyllt~~ttzone. Twenty-five species were ob- served in plots on the uplifted portion of the bench. Seventeen of these species were so infre- quent or sporadic that my possible effects could not he deduced in their populations. Of the eight most frequent species for which effects could be observed, five showed mortality attributable to the uplifting, one (fitctls distichus) increased siguifi- cautly in frequency, and the remaining species exhibited little change in frequeucy (Fig. 21). Most die-off occurred in the first 6 months after the test. Lebednik (1973) concluded: "The rather rapid aud extensive die-off suggests that the populations affected were vertically close to their upper physi- Fig. 20-View of the IA-1 area at Duck Cove. The ological limits before the uplifting. . . ." Con- fault runs across the center of the intertidal bench. tinued sampling of the area indicates a gradual shift To the left the bench was uplifted; to the right no elevation change occurred. Photograph was taken on of algal species. Three and one-half years after the Mar. 29, 1971, at a 30-cm (-12-in.) low tide. test (in early 1973), ~~~ajorincreases in the FZLCZLS and Alaria crisp populations were still occurring in 64% of the plots (Nakatani and Burgner,l974). algae that had settled since Cannikin were limited This confirms Lebednik's (1973) predictiou that to areas belo\\' the 1970 low-water line, and the the die-off in the Hedopl~yllzr?nzone would he only other vegetation visible Isas on the offshore followed by forn~ationof a Fuclcs zone. There is reefs behind the sea stack. no doubt that a stable algal community will Following the uplifting, Iridaea cor?atrcopine cvcntually be established. was the first species to sho\v conspicuous mortal- ity. By April 1973 all the 20 surviving plots (5 Cannikin-Affected Areas more had been buried by rockfalls), which preshot were representative of the entire vertical range of The 1971 Cannikin test caused au uplift of as intertidal vegetation, were devoid of algae except much as a meter dong the Bering Sea coast. Two for a fervPorphyra sp. and geen algal plants. Total areas with fixed plots (IA-2 and IA-3, see Fig. 26) destruction of littoral vegetation also occurred in were located in the area most affected by the test. other areas with co~nparableupliftiiig (Nakatani An average uplift of 91 cm (36 in.) occu~redin the and Bmgner, 1974). IA-2 study area (Nakatani et al., 1973; 1974). An At IA-2 the sublittoral region, including the indication of the damage caused is given in Fig. 22, sublittoral fringe, was lifted to a new level, and a which sho\\.s a portion of the IA-2 area. The upper new littoral vegetation is expected to develop. The photograph was taken in August 1970 (15 months shallow suhlittoral fringe is one of the most before Cannikin), at which time the rock bench in difficult areas to sample because it is mostly the foreground xvas co~npletelycovered by Halo- underwater even during low tides, aud kno~vledge sncciott ghttdiforttte, Rhody?netfia pal7nata, a~td of the suhlittoral communities occurring there has Corallitla spp. The surge cliannel at the center was been limited (Weinmann, 1969). The uplifting of dominated by Alaria crispa aud Lanzi?tnria lotagipes, Cannikin allowed observations of these communi- and, betxveen this area and the sea stack, a varied ties. At the IA-2 area a large portion of the kelp vegetation of Heclophyllzt~nsessile and Lantittatia comrnui~ity was occupied by three Laminaria lo~~gipesnras dominant in low pools. The lower species: (1) L. lo?fgipes, (2) L. groettlattdica, and photograph \\'as taken in April 1973 duriug an (3) L. yezoetfsis. Tlzalassiophyllu?~ clatlir~is was equivalent lo\\, tide. h4ore rock was exposed both psesent in some localities, and Alarin fistzrlosn, in the surge channel and in reefs behind the stack. which forms the extensive offshore floating kelp A great deal of rockfall from both the stack and beds fringing the island, was abundant on one rock the adjacent sea cliff had buried some intertidal bench at the lowest level exposed by the uplift. areas, including three study plots. Of greater These species and their associated flora, except for siguificance was the conlplete lack of algal vegeta- Lanti?faria longipes, wherever they had been lifted tion on the rock benches. In the surge channcl the into the intertidal area, were dead by April 1973. 384 Lebednik and Palnzisa~to 18 I x-x . Corallioa I I I I is lage boulders. In both plots the pre-Cannikin vegetation was dominated by a tall dense canopy of Lantinal-ia yezoettsis, beneath which was a dense stand of L. lolzgipes, covering most of the rock. Although the entire L. yezoensis population died,

I Z A M S TIME, months Fig. 21-Summary of frequencies of the eight most frequent species (except Atzalipus filiforntis) on the uplifted plots at IA-l in 1970. 2, deduced pretest pattern. A, April. hi, hIay. S, September. [From P. A. Lebednik, dlnritre Biology, 20: 201 (1973).]

New plots were established at IA-2 after Catlllikin in four types of habitat: (1)High outer bench, (2) surge chan~lelbasin, (3) newly exposed Fig. 22-View eastward at IA-2 in August 1970 rock, and (4) lo\\, outer bench. (upper) pre-Cannikin and April 1973 (loxser) post- Two plots in the first habitat were located on Cannikin. the outer portion of the intertidal bench at a relatively high level. This was an area exposed to heavy wave action; the pretest vegetation was characteristic of the lower levels. The reaction of Lnnti~zaria longipes at a relatively protected plot (Fig. 23) is compared with that of a more exposed population (Fig. 24). In vegetation the plots were identical initially and followed generally the same pattern of decrease in cover. However, some plants remained alive 2 months longer on the more exposed plot, and the remains of dead plants were still visible 17 months after Cannikin. The expo- sure conditions have changed because the outer vertical face of the bench is well above mean lower lo\\. water and does not atlow wave splash on the bench top as it did pre-Cannikin except duriug the highest tides. Should any new vegetation develop Fig. 23-Percent cover of Lamittorin longipes versus on these plots, there should be a significant time at IA-2 (dead cover of L. longipes indicated by difference between the two. shading), December 1971 to April 1973. Preshot T\\ro new plots were established in the bottom elevation, 34.3 cm (14 in.); postshot, 122.1 cm of a surge channel basin where the stable substrate (48 in.); uplift, 87.8 cm (35 in.). Ecology of nilari~leAlgae 385 months after Cannikin, two species were present: (1) A large population of Cladophora sp. and (2) numerous plants of Po,plryra sp. (P. pseltdo- lineatis?). Two months after that, in April 1973, both species had decreased in cover whereas Halosaccion glandifovtne had become dominant in two plots and Rhodyntenia palmata in one. Alnfia crispa had appeared in two of the plots; this was the first indication of the appearance of a kelp canopy. The four plots established in the final habitat were on a lo\\, rock bench with a uniform seaward slope that was subtidal pre-Cannikin but became intertidal post-Cannikin. Observations after the uplifting indicated a threefold preshot zonation from landward to seaward dominated by Lamiflaria Fig. 24-Percent cover of Lamirraria longipes versus lottgipes, T/zalassiopl~yllt~mclatlzrzts, and Alaria time at IA-2 (dead cover of L. longipes indicated by fistulosa-Lanti?ta,.inn groenlandica. The first two shading), December 1971 to April 1973. Preshot elevation, +58.3 cm (23 in.); postshot, +153.2 cm domitla~ltswere represented by one plot each and (60 in.); uplift, 94.9 crn (37 in.). the last one by two plots that reached to the outerrnost portion of the bench emersed at low tide. By April 1973 the L. longt$es population had L. longipes, which initially showed some die-off, died in the plot originally dominated by it. During began to recover in August 1972 (9 months after 1972 Rltodyntenia palntata appeared and reached a Cannikin), as illustrated in Fig. 25. Laminnria maximum in October 1972 but still was abundant longipes normally occurs in the sublittoral fringe, (64%) in April 1973. The appearance of Fl'zccus and its survival here is consistent with the present distichus and Alaria ccrispa in April 1973 signaled elevation of these plots. the reestablishment of a kelp canopy in the plot. ~uccessiohon newly exposed rock surfaces \\'as At the plot where T. clatl~rttswas the dominant studied in three plots. Two of them were on the species pre-Cannikin, Rhodymertia pal~natainitially vertical surface that formed the outer face of the showed an increase but was soon covered by a pre-Cannikin intertidal bench and one was on the burgeoning Alaria crispa population that first horizotltal surface of a large rock boulder immedi- appeared in June 1972 and occupied 100% of the ately seaward of the other two. All these plots plot by April 1973. At the outermost plots showed a similar pattern of colonization. Three Lantinaria longipes settled and dominated one (75%) and Alaria crispa dominated the other (80%). Rltodynzenia palnlata was fairly abundant in these two plots in June and August 1972 before its populatiotls were covered by the growing kelp plants. The seco~ldarea, IA-3 (Fig. 26), established to study effects of Cannikin, had only 13 plots sulvive after Cannikin since 2 of the origi~lalplots were buried by rockfall. Although the 1A-3 area underwent an average uplift of only 50 cm (20 in.) as compared with 91 cm (36 in.) at IA-2, mortality \\.as nearly as complete. As of April 1973, five plots bad no live algae; eight had one or both of two Porplt)wa species; Ulva lnctttca occurred on three plots; and Rlbodyntenia palmata, Analt$zts filiformis, and Halosaccion glandifornte each oc- TIME, months port-Cannikin curred on one plot in insignificant amounts. Only at one plot, where a small FZLC~CSdisticItzts popula- Fig. 25-Percent cover of Lomi~torinlongipes versus tion had settled and started to develop, was there time at IA-2 (dead cover of L. longipes indicated by shading), February 1972 to April 1973. Preshot an indication of permanent colonization. Except elevation, -93.2 cm (-37 in.); postshot, 1.7 cm for this last plot, no midlittoral vegetation colo- (1 in.); uplift, 94.9 cm (37 in.). nized the IA-3 plots after the Canuikin test. New 386 Lebedttik nttd Palmisatto

BERING SEA I iTREL

ITALICE CREEK

IA-3 BANJO POINT i CANNlKlN SGZ @

Fig. 26-Location of points discussed in the Bering Sea coast survey.

plots were not established at IA-3 to docnment and crustose coralline algae showed die-off. East- colonization, but \ve did make general obser\~a- ward from point P8 to point 96B, only I. cowttr- tions. By April 1973 there was heavy Ft1cics copiae and L. longipes mortality was observed, and disticlttcs colo~lizatio~ljust below the original plots. beyond point 96B no test-related die-off was seen Alaria crispa appeared to be newly abundant and at all. domiuallt near low !\rater levels [at the 0.5-nl Severe damage was observed along 1.9 km (1 (-2-ft) tidal level] . There was little evidence of mile) of shore, including the IA-2 and IA-3 areas. Lat?zinaria lotzgi/~esor Hedophylltrtn sessile. Moderate damage was seen on 1.5 km (1 mile) of So that the data taken at the IA-2 and IA-3 shore and damage detectable along 2.7 km (2 plots could be generalized, observations were made miles) of shore. Altogether, 6.1 km (4 miles) of at low tide along the 15.4-km (10-mile) stretch of shore \\.as affected. shore between Sea Otter Point and Crown Reefer In April 1973, except for a slight increase in Point (see Geological Survey map in the pocket at mortality around Petrel Point, the distribution of the back of this volume). !\'alks were made in mortality was similar to that in April 1972. Dead August 1971, 2Yz months before Cannikin, and in plants not yet removed by wave action made it April 1972 and 1973, 5 and 17 months after\vard difficult to detect colonization in the moderately (Table 1). affected areas. In the severely affected areas, The obseivations made in April 1972 on algal colonization occurred as reported for IA-2. die-off on this stretch of shore are summarized in Fig. 27. Coastal landslides were visible west of Petrel Point, but algal die-off did not become SEVERE a apparent until just east of Petrel Point. From there m to point 9B, die-off was light and patchy. Between ? MODERATE -", points 9B and 10B, die-off was more extensive aild 0 involved more species. From just west of point 10B to the mouth of White Alice Creek, die-off \eras extensive wd comparable to that at IA-2 except lllllllllllllll lllllllllllllllllllll that near where the creek flowed over the bench LOCATION E PETREL 98 108 WHITE ALICE P8 968 Iridaea contztcof~ineappeared to be less affected. 0.25 East of White Alice Creek to point P8, die-off \vas DISTANCE, krn ~1.1~1.9-t1.25+1.61 illoderate and discontinuous. At Banjo Point only the Alatia zone showed die-off. Elselvltere portions Fig. 27-Alongshore die-off of littoral algae due to of the populatioils of I. cort~trcopiae,L. long+es, Cannikin, April 1972. Ecology of hlari?te fllgae 387 Followi~lgthe field studies of Lebednik, obser- of an abundant and diverse marine algal flora. A vations of the IA-2 and IA-3 areas were made in review of phycological research in the North &Pay and August 1974 by other FRI staff menlbers Pacific Ocean, in particular that of Russian work- and were reported by ICirkwood (1974; 1975). No ers, indicates a dearth of knowledge of the marine quantitative data on cover values of the species algal flora of the Aleutians. were presented; however, significant changes in A somewhat unusual topographic feature, ex- species con~positionappear to have occurred since tremely flat rock benches, occurs at about mean August 1973 in 8 of the 11 plots discussed tide level around the southeastern third of the previously. It is cstitnated that the total area of island. These benches provide an exceptionally intertidal bench uplifted into the supralittoral large area for the tle\relopment of littoral vegeta- fringe by Cannikin was 5.8 ha (O'Clair, Chap. 18, tion. The seatcwd edge of the bench, \\,here waves this volume). * are often breaking, is the most exposed portion, The effects of uplifting from underground xvhercas the land~vard edge is usually quite pro- testing at Amchitka and those of uplifting from the tected. Cloud cover and humidity are very high Alaska earthquake of 1964 as reported by Johan- througllout the year, and thus the littoral algae do sen (1971) were similar. In both localities littoral not experience much desiccation. Air tenlperatures populations did not tolerate the environmental are generally about the same as seawater tempera- conditions at higher elevations and extensi\.e mor- tures except that in the winter subfreezing condi- tality resulted. A subsequent succession of species tions prevail for short periods. Heavy wave action in the disturbed areas led eventually to a return to is characteristic of Amchitka. Seawater tempera- the original pattern of zonation, which was consis- tures range from 2 to 10°C at the shore, and the tent with the new elevations and changes in water is exceptionally clear. The extensive rocky exposure. areas, cool nloist climate, and clear water condi- Uplifting at Amchitka ill probably have a tioils provide an ideal environ~nent for marine significant long-term effect because of the presence algae. of flat rock benches in the nlidlittoral region. The littoral marine vegetation was studied by These benches were lifted above the nlidlittoral means of two transects, one each on the Pacific region at IA-2 and IA-3 folloxving the Cannikin Ocean and Bering Sea coasts. The littoral region test. A new midlittoral vegetation in those locali- can be classified into the supralittoral fringe and ties will have far less area on \vhich to develop. midlittoral areas. The supralittoral fringe is com- Because the supralittoral fringe populations depend posed of three zones, from top to bottom: on wave splash, which occnrs only at the edge of (1) Prasiola, (2)Rltodocl1orto1~, and (3)Por- the benches, only a small portion of the bench area phyra-Ulotltrix. The last zone is practically absent will be colonized by these species. in the fall and winter and reaches its highest lLIortality may occur over a period of months development in late spring and summer. The upper in uplifted populations, depending on the habitat limit of the midlittoral area, usually taken as the and amount of uplifting. The early stages of upper limit of the barnacle zone, \\.as assigned to colonization appear to be strongly influenced by the upper limit of the Ftlclls zone owing to the seasonal spore production. A disturbed plot tnay lack of a barnacle zone at Amchitka. The next remain nearly devoid of algae or with unchanged lo\\.er zone is a Hedopltyllum zone that usually vegetation during the winter lnonths but may occupies the greatest area because its occurrence exhibit rapid changes in species composition and coincides \\'it11 the level of most of the rock abundance in the spring. The return to a normal benches. Six associations can be recognized within community may require years, as exetnplified by the Hedopltyllzcnt zone: (1) Ulua, (2) A?tal@tts, the marked changes in the Ftrclis populations at (3) Co,ulli~~a,(4) lIalosaccion, (5) Hedo/dtyllt~?1t, IA-1 3% years after the Milrow test and the and (6) Iridaea. The geatest number of species and changes observed in Cannikitl-disturbed areas the most complex vegetation were observed in this nearly 3 years after that test. zone. The third zone of the midlittoral area, the Alarin zone, occurs primarily near the outer edge SUMMARY of the benches. The sublittoral fringe consists of the Lanti~zaria longipes zone, in which se\rcral The climate, sea conditions, and topography of sublittoral species have their upper limit. Tide Amchitka Island are favorable for the development pools are rare, but channels running across the benches and conncctcd with the sea at lolv tide have a sublittoral flora. The bench topography *Previously reported as 57.6 ha (Kirktvood, 1975). influences littoral vegetation in that the distribu- tions of many species were correlated xvith hori- red algae, such as Ptilota asple~tioi~les,P. filiciiln, zontal distance perperidicular to the shore regard- Hypophy llunt ~tcprecl~tia~izcnt, Cit.rzclicar/~trs less of elevation. Seasonal photographic sampling gmelini, and Constantinen ~osn-iiiariiza.The fourth indicated that most littoral species occur through- level of vegetation, coilsisting of crustose forms out the year. Most species appeared to release and extending well below 30 m (100 ft), included spores in late winter-ealy spring, and the appear- the green alga Codi~cnt ritteri and the corallille ance of sporelings follo\vs this period. Growth of genera Clathromol;ohzcm, Litl~otltant~tizen~,and sporelitlgs as \\,ell as of older plants reached a peak A.lesophyllzci~~.The richness and density of marine in late spring-early summer. The Porphyra- algae at Amchitka might be due to the removal of Ulothrix zone and the Halosaccio~t association invertebrate herbivores by sea otters. A strong were annual, occurring from early spring to late relationship between the Amchitka flora and the summer. The littoral zonation \\'as nearly identical western Pacific, eastern Pacific, and Arctic floras to that described at the Commailder Isla~ldsanct \\.as found by comparing species lists. The initial was similar to that described at the northern ICurile results of a long-term taxonomic study have Islatlds. The zonation at Amchitka \\'as also com- resulted in the descriptio~tof three new genera and pared with that of southeastern Icamchatka, eight new species from Amchitka. Sakhalin Island, Hokkaido, Japan, and the Strait of The R3ilro\\. nuclear test caused a 12-cin (5-in.) Juan de Fuca. uplifting of a portion of intertidal bench totaling Total relnoval of organisms from rocky inter- about 4 ha (10 acres). h.(ortality ensued in the tidal plots resulted in a colonization sequeilce of upper portions of all zones, especially the diatoms and filamentous algae, leafy green algae, Hedopltyllzei~~zone, within 6 months after the test. and larger red and brown algae. In plots where only Although new vegetation characteristic of the the kelp canopy had been remo\fed, the sequence higher levels had begun to develop, significant was similar but faster. Initial settlemeilt occurred changes \\'ere still observed in some plots 3%years most quickly in exposed areas at low levels. after the test. The Cannikin test caused a maxi- Colonizatio~lwas strongly influenced by the length mum uplift at the shore of about 1 m (3 ft), which of time between the clearing of plots a11d the next occurrect in one of t\vo study areas. Total mortality anilual spore release of the species originally of all algal species ensaed in both study areas with dominant in that plot. Three of the five species uplifting of 0.5 to 1 n~ (1.5 to 3 ft) and along whose stands were dent~dedand two of the three 1.9 kt11 (1 mile) of coastline. Moderate mortality wvl~ose canopies bad been remo\led attained a was observed along an additional 1.5 km (1 mile) of coastline, and solne mortality \\,as detected for a coverage of at least 80% in their OISII zone by the end of the study. Those which did not attain this further 2.7 hn (2 miles). A total of 6.1 km degree of coverage were Lamiitaria lottgipes and (4 miles) was affected. New plots were established Corallina ua~zcouuerie~tsisin denuded areas and in formerly sublittoral areas, and coloilizatio~l\\,as Alaria crispa in areas rvhere the canopy had been follo~ved. Significant changes were occurriitg in removed. The lo\\, recovery of L. longipes was these plots \\,hen last obseived, nearly 3 years after attributed to its low degree of sexual reproduction. the test. As was the case with the Milroxv-disturbed The scarcity of A. crispa \\,as attributed to the lack areas, new vegetatiou was for~ningin coilsollance of bare substratum needed for settlement. The with the ne\v elevations and exposure of the establishment and tnaintetla~lceof algal zones aud disturbed areas. I11 contrast to the Milrow distur- associations, at times in almost pure stands, are bance, which will have no long-term effects, the enhanccd by the physical liomogeileity of the Ca~mikin disturbance has resulted in a marked intertidal bench, the relatively few species of reductioil of area at midlittoral levels by perma- dominant algae, aud the lack of excessive physical nent lifting of the intertidal bench into the and biological disturbances. supralittoral fiinge. Supralittoral fringe species, \vhicb require constant wave splash, will be able to The sublittoral commu~lities are extremely colonize only the outer edges of the uplifted dense. Alaria fistulosa, which forms huge floating benches. kelp beds, mas observed do\vi~to about 20 m (66 ft). These kelp beds have a considerable damp- ing effect on wave action. A dense covering of kelp occurs on the bottom from the intertidal area down to about 20 m (66 ft) and rvas formed by The research reported in this chapter \\,as sup- species in the genera Laminaria, Tltalassiophyllutn, ported by the Atomic Energy Commission contract Cymatltere, Alaria, and Agancin. A third level of AT(26-1)-171 to Battelle Columbus Laboratories vegetation is made up of filamentous and foliose and the Fisheries Research Institute, University of \\'ashitlgton. \\re express our thanks to Drs. R. L. -, J. S. Isakson, and P. I\. Lebednik, 1970, Ohse~vations Burgner and R. Nakatani for their encouragement, on the Effect of the Milrow Nuclew Test on Marine Organisms at ,\mchitka, BioScience, 21: 671-673. to J. Isakson, C. O'Clair, C. Simetlstad, P. Slattery, -, et al., 1971, Amchitka Bioenvironmental Program, and G. Tutmark for help with the field work, and to Research Progranl on Marine Ecology, Amchitka Island, Drs. I<. Chelv and R. E. Norris for their guidance. Alaska, Annual Progress Report, July 1, 1969-June 30, Additional equiplllent and support were provided 1970, USAEC Report BM1-171.137, Battelle hlemorial to Lebedltik by the faculty of the Department of Institute. -, et at., 1969, ~\mchitka Bioenvironmental Program, Botany, University of \\'ashington. Annual Progress Report, July 1, 1968-June 30, 1969, Preparation of the manuscript was accom- Research Program on Marine Ecology and Oceanog- plished with the support of a Postdoctoral Re- raphy, Amchitka Island, Alaska, USAEC Report Bh,lI- search Fello\\~sliip(to Lebednik) through National 171-128, Battelle AIelnorial Institute. Research Co~~llcilof Canada grant A-4471 to Dr. -, et al., 1968, Amchitka Bioenvironmental Program. Annual Progress Report, July 1, 1967LJune 30, 1968, R. F. Scagel and x\rith the support of the Auke Bay Research Program on Marine Ecology and Oceanog- Fisheries Laboratory of the National Bllarine raphy, Amchitka, USAEC Report BAII-171-114, Hat- Fisheries Service (to Palmisano). \\re thank Dr. P. telle Memorial Institute. Silva for providing access to Gtnelitl's paper. The Chapman, 1'. J., 1943, Zonation of hlarine Algae on the help of G. Lehednik in preparing the final manu- Sea-Shore, Trans. Lin~t.Soc. Londo., 1941: 239-253. Chiliara, B1., 1967, Some, Marine Algae Collected at Cape script is gratefully ackno\vledged. Thompson of the Alaskan Arctic, Bull. Natl. Sci. r\Ias., Tokyo, lO(2): 183-200, plates 1-4. Conr\,ay, E. C., T. F. hlumford, Jr., and R. F. Scagel, 1976, The Genus Porphym in British Columbia and \\'ashing- REFERENCES ton, Syesis, 8: 185-244. Dawson, E. Y., 1961, A Guide to the Literature and ,\dey, \ H., T. Masaki, and H. Akioka, 1976, The Distributions of Pacific Benthic Algae from Alaska to Distribution of Crustose Corallines in Eastern Hokkaido the Galapagos Islands, Pac. Sci., 15: 370.461, and the Biogeographic Relationships of the Flora, Bull. Dayton, P. K., 1975, Expel-imental Studies of Algal Canopy Foc. Fislr., Hokknido Uniu., 26: 303-313. Interactions in a Sea-Otter Dominated Community at Armstrong, Robert H., 1971, Physical Climatology of Amchitka Island, Alaska, Fis11.Bull., 73: 230.237, Amchitka Island, Alaska, BioScience, 21(12): 607-609. -, 1971, Competition, Disturbance, and Community Biebl, R., 1972, Temperatore Resistance of Marine Algae, Organization: The Provision and Subsequent Utiliza- in Proceedings of the Seuenth Inter~~ntio~~nlSeaweed tion of Space in a Rocky Intertidal Community, Ecol. Symposiufn, Aug. 8-12, 1971, pp. 23-28, University of ~\~oIIogT.,41: 351-389. Tokyo Press. DeCew, T. C., and J. \\lest, in press, Culture Studies on T\vo -, 1970, Vergleichende Untersuchungen zur Temperatur- blarine Red Algae, llildele,zbmndin occidentalis Setchell resistenz \,on Meeresalgen entlang der pazifischen Kiiste and H. prototypus Nardo (Cryptonemiales, Hilden- Nordamerikas, Protoplasnta, 69: 61-83. brandiaceae), Bull. Jap. Soc. Phycol. Blinova, E. I., 1968, Morskie vodorosli severe-vostochno.i Doty, hl. S., 1957, Rocky Intertidal Surfaces, in Treatise OII AIarine Ecology nnd Pnleoecology, \'ol. 1, Ecology, chasti Okhotskogo ~norya[h'larine Algae of the Nortlt- east Part of the Sea of Okhotsk] , Arouosti sistenlatiki J. \\'. Hedgpeth (Ed.), pp. 535-585, Geological Society of l\merica, New York, N. Y. nizshiklt msteni; Akad. Nauk SSSR, Bot. Inst., 5: 33-38. -, 1946, Critical Tide Factors that are Correlated xvith -, and I. S. Gusarova (Subhotilia), 1970, ivlorskie the Vertical Distribution of hlwine Algae and Other vodorosli, notye dlya poherezh'ya \~ostochoo; Kam- Organisms Along the Pacific Coast, Ecolo.~, 27: chatki [New Marine Algae from the Coast of Eastern 315-328. Kamchatka] , Novosti siste~nntiki r~irshikh msteni; Druelil, L. D., 1970, The Pattern of Laminariales Distriba- Aknd. l\'ouk SSSR, Dot. Inst., 7: 68-71. tion in the Northeast Pacific, Pkycologin, 9: 237-248. - 1968, Taxonomy and Distribution of Northeast -, and 11. D. Zinova, 1967, Nov),; vid Rhodymenia iz Pacific Species of I.nminarin, Can. J. Bot., 46: 539-547, severo-vostochno; chasti Okhotskogo morya [A Nen, plates I-VII. Species of Rkodymenia from the Eastern Part of the Elenkin, A. A,, and L. OI', 1950a, Bihliografiya al'golo- Sea of Okhotsk] ~\buosti sistemntiki nizst~ikhrasteni; , gicheskikh trttdov v predelakh SSSR s 1931 po 1935 g. Akad. A'auk SSSR, Bot. Inst., 4: 107-109. vklyuchitel'no [Bibliography of Algological Studies in Bokenhani, N. A. H., 1938, The Colonization of Denuded the USSR from 1931 to 1935 Inclusive], Tr. Bot. Inst., Rock Surfaces in the Intertidal Region of the Cape Akad. ,\'auk SSSR, Ser. 2: Sporouye rosteniyn, No. 5: Peninsula,A~tn.Natnl. dl~cs.,9: 47-81. 95-115. Buxgner, R. L., and R. E. Nakatani, 1972, Amchitka -, and L. Ol', 1950b, Dopolnitel'nye dannye po biblio- Bioenvironmental Program, Research Program on Ma- grafii algologicheskikh wudov v predelakh SSSR. rine Ecology, Amchitka Island, Alaska, Annual Progress Dopolnenie 11. [Supplementary Data for the Bibliog- Report, July 1, 1970-June 30, 1971, with contribu- raphy of Algological Studies in the USSR. Supplement tions by J. S. Isakson, L. G. Gilbertson, P. A. Lehednik, 111, Tr. But. Inst., Aknrl. Nnuk SSSR, Ser. 2: Sporouye C. E. O'Clair, J. F. Palmisano, C. A. Simenstad, and rastet~iya,No. 6: 5-22. G. J. Tutmark, of the Fisheries Research Institute, -, and L. Ol', 1935, Bihliografiya al'gologicheskikh University of !\lashington, USAEC Report BA1I- b'udov v predelakh SSSR s 1926 g. po 1930 g. 171-144, Battelle Memorial Institute. vklyuchitel'no [Bibliography of Algological Studies in the USSR from 1926 to 1930 Inclusive], Tr. Bot. Inst., Tikhookena. Na~tchito-Issled. Inst. Rybn. Khoz. Akad. Nauk SSSR, Ser. 2: Sl>orovye rasteaiya, No. 2: Okeatrogr., 14: 77-108. 171-255. Kireesa, &,I. S., 1965, Syr'evye resursy vodorosley more; -, and L. OI', 1929, Bibliografiya al'gologicheskikh sovetskogo soyuza [Natural Resources of Marine Algae trudov \. predelakb SSSll s 1900 po 1925 gg. vklyu- of the Soviet Union], Okennologiyn, 5(1): 14. chitel'no [Bibliography of Algological Studies in the Kirkwood. 1. B. (Comu.). 1975. Bioenvironmental and USSR from 1900 to 1925 Inclusive], Tr. Bot. Sada ~ydroiogicstudies, khchitka island, Alaska. Fall 1974 (Actn Horti Petrop.), 42(1): 1-139 (not seen by Task Force Report, ERDA Report Bh'iI-171-158, Bat- authors). telle h,Iemorial institute. Ga;dukov, N. hi., 1901, Literat~rrnyeistochniki k russko; (Comp.), 11974, Amchitka Biaenvironmental Program. flore vodorosle; [Literary Sources for Russian Flora and Bioenvironmental Safety Studies, Amchitka Island, Algae], Tr. Bot. Sndn (Scripta Bot. HortiInlp. Petrop.), Alaska, Spring, 1974 Task Force Report, USAEC 17: 1-126 (not seen by authors). Report BhI1-171-157, Battelle i\lemorial Institute. Gail, F I. 1918, Some Experiments with Fucrts to Kitching, J. A,, 1937, Studies in Sublittoral Ecology. 11. Determine the Factors Controlling Its \'ertical Distribu- Recolonizatio~lat the Upper Margin of the Sublittoral tion, Pttbl. Puget Sound Biol. Sta., Univ. Il'nsh., 2: Region; with a Note on the Denudation of La~ninnrin 139-151. Forest by Storms,]. Ecol., 25: 482-495. Gmelin, S. G., 1768,Historinft1cortr111,[lo] + 239 + 6 pp., Kjellman, F. I<., 1889, Om Beringhafvets algfloran, Kgl. plates lA, lB, ?A, 2B, 3-33, Typographia Academiae Suetrska Vetettskapsnkad. Hnndl., 23: 1-58. Scientiarom. Petropoli. Komarov, F., 1962, Botany, in The Pncific Rsssinn Scien- Gollerbakh, M. M., ,\. A. Elenkin, L. K. Krasavina, and tific Investigatio~~s,pp. 126-136 (English Edition; Orig- L. A. OI', 1966, Bibliografiya sovetskoy literatu~ypo inal Published in 1926, not seen by author), Greenrvood r,odoroslyam za 1936-1940 gg. (s dopelnenicm za Press Inc., Westport, Conn. predydushchie gody). [Bibliography of Soviet 1,itera- Kongiser, R. A,, 1933, Gidrobiologicheskie raboty s Bering- ture on Algae for the Years 1936-1940 (with a ovom more u severo-vostochnogo poberezh'ya Supplement for the Last Year)], Akademiya Nauk Kamchatki [Hydrobiological Studies in the Bering Sea SSSIl, Botanical institute (not seen by authors). Near the Northeast Coast of Kamchatka], Issled. -, and L. K. Krasavina, 1971, Vodorosli. svodn); Dnl't~evost.illore; SSSR, 19: 115-124. ukazatel' k otechestvennym hibliografiyam po vodoro- Krasavina, L. K., 1968, Bibliografiya sovetsko; literatory po slyam za 1737-1960 gg. [~\lgae.A Consolidated Guide vodoroslyam za 1941-1960 gg [Bibliography of Soviet to Native (Russian) Bibliographies on Algae for the Literature on Algae for the Years 1941-19601, Years 1737-19601, \'. L. Komnrov, Akademiya Nauk Akndemiya Nauk SSSR, Bota~ricnlInstittrte (not seen SSSR, Botanical Institute. by authors). Gur)~aaova,E. F., 1935, The Commander Islands and Their Kusakio, 0. G., 1961, Nekotoqre zakonomernosti raspre- hlarine Littoral Fauna and Flora, Prirodo, 11: 64-72. deleniya fauay i flory v osachno~zone yuzhnykh [Translation by L. Sagen, The Soviet Fishery I'erti~mlia. Kulil'skykh ostrovov [Certain Regularities of the Dis- No. 16 (Mar. 25, 1974), pp. 1-17, College of Fisheries, tribution of the Fauna and Flora in the Intertidal Zone University of IYashington, Seattle.] of the Southern Kurile Islands] , Isslerl. Dnlheuost. Gosarova, I. S., 1972, Noqre vodorosli dlya nekotorykh More; SSSR. VII: 312-343. ostrovov bol'sho; ~uril'skoy g~yady [New Algae from Lebednik, P. A,, in press, a, Postfertilization Developn~ent Some Islands of the Big Kurile Chain], A'ouosti sis- and Evolution in the Rhodo~hvta... .. I. Pltycol.. temntiki nizsltikh rasterzit Aknd. Nask SSSR, Bot. Inst., 13(Suppl.): (abstract). 9: 32-34. -, inpress, b, Algal Communities of the Aleutian Islands, -, and Yu. E. Petrov, 1972, Noyi rod i vid J. Plrycol., 13(Soppl.): (abstract). laminariev)~kh vodorosle; c 0. Simushir (Kuril'skie -, in press, c, Postfertilization Development in Clnthro- Ostrova) [*\ New Genus and Species of Laminaria11 azorplzum, wel lobe sin and ~\Iesophyllaaz, \\fit11 Com- Algae from Simushir Island (Kurile islands)] , Nouosti ments on the Evolution of the Corallinaceae and the sistemaliki rrizshikh rastettii; Akad. Nnuk SSSR, Bot. Crgptonemiales, Pl~ycologin,16: 171st., 9: 39-44. - 1977, The Corallinaceae of North\vesterrt North -, and Yo. E. Petrov, 1970, Novyi rod i vid America. I. Clathromorahum Foslie emend. Adev.,. la~ninarievykllvodorosle; s Kuril'skikh ostrovov [A Nerv Syesis, 9: 59-112. Genus and Species of Laminarian Algae from the Kurile -. 1975. Bios\~stematicStudies of ClathromorDhao~and Islands] , A'ovosti sistenlntiki nizshikh rosteni; Aknd. A'auk SSSR, Bot. Inst., 7: 87-90. ...~.... den llartog, C., 1959, The Epilithic Algal Communities -, 1974, The Genera Clntlrromorphum arid dlesophyllat~r Occurring Along the Coast of the Netherlands, Ilre,ztia, (Corallinaceae) from the Northeast North Pacific 1: 1-241. Ocean, Ph. D. Thesis, University of i\'ashington. [Diss. Johansen, H., 1971, Effects of Elevation Changes on Abstr. Int., 36(Sect. B): 2596-B (1975).] Benthic Algae in Prince I\'illiam Sound, in Tlre Great -, 1973, Ecological Effects of Intertidal Uplifting from Alaska Enrthqrrnke of 1964, pp. 35-68, Biology, Nation- Nuclear Testing, ~\Inr. Biol., 20: 197-207. al Acadenly of Sciences, \\'ashington, D.C. -, F. C. Weinmann, and R. E. No~ris,1971, Spatial and Johnson, I\'.S., 11. Gigon, S. L. Gttlman, and H. A. hlooney, Seasonal Distributions of hlarine Algal Communities at 1974, Comparative Photosynthetic Capacities of Inter- Amcllitka Island, Alaska, BioScience, 21: 656-660. tidal Algae Under Exposed and Submerged Conditions, Lewis, J. R., 1964, ?%e Ecology of Rocky Shores, English Ecology, 55: 450-453. Universities Press, London. Kardakova-Prezlle~~tsova,E. A,, 1938, Vodoroslevaya B'iarkham, J. IV., 1972, Distribution and Taxonomy of rastitel'nost' Komandorskikl~ Ostrovov [The Algal Lnmitrnrin sh~clnirii and L. longipes (Phaeopltyceae, \regetation of the Komandorski Islands], Ir. Laminariales), Phycologin, ll(2): 147-157. Ecolog)~ofMarine Algae 391 McAlister, I\'. B., 1971, Oceanography in the Vicinity of sistonntiki eizsl~iklt msteni< Aknd. A'auk SSSR, Bot. Amchitka Island, Alaska, BioScie~~ce,21: 646-651. It~st.,7: 81.87. h4cRov. C. P.. 1968. A Eurasian Alea" in Alaska. Pnc. Sci.. Polanshek, A. R., 1975, Hybridization Studies on Cultured ai(i): 138. Petrocelis Species from California and the i\leutians, J. hlertens, Heinrich, 1829, ljber serschiedene Focus-Arten, Pl~ycol.,1 I(Suppl.): 17 (abstract). Lbinnen, 4: 43-58. -, and J. A. \\lest, 1976, Culture and Hybridization BIokievski;, 0. B., 1967, lzmeneniya litoral'noi bioty ot Studies on Petrocelis (Rhodophyta) from Alaska and Beringova morya do zondskogo proliva [\'xiations of California, J. Phgcol., 11: 434-439. Littoral Biota from the Bering Sea to the Zond Straits], Postels, A., and F. Ruprecht, 1840, lllustrationes Algarum Okea,~ologiyn,2: 321-324. in itinere circa Orbem. . . Executo in Oceano Pacifico Morris, R. H., 1971, Marine Terraces of the \\'estem Imprimis Septentrionali ad iittora Rossica ~\siatico- Aleutian Islands. Alaska. USAEC Reuort USGS-474- Americana Collectaru~n,St. Petersburg. 139, U. S. Geological ~uniey. Pourell, H. T., 1957, Studies in the Genus F~icusL. I. Fucas Naeai.-. Evl.. . 1941. Marine Aleae- of the Kurite Islai~ds.11. J. distichus L. emend. Porvell, J. lllnr. Biol. Ass. UK, 36: Fnc. Agric. Hokknido Uniu., 46: 139-310, plates 407-432. 77, I,, Porvers, FI. A,, R. R. Coats, and \\I. H. Nelson, 1960, -, 1940, hlarine Algae of the Kurile Islands. I. J. Fac. Geology and Subtnnrine Pltysiogmphy of Amclzitkn Ayic. Hokkaido Univ., 46: 1-137, plates 1-111. Islnt~d. Alnska. U. S. Ceoloeical Sulvev., . Bulletin Nakatani, R. E., and R. L. Burgner, 1974, Amchitka 1028-P, pp. 521.554. Rioenuironmental Program, Research Program on hla- Rees, T., 1940, Algal Colonization at hlunlbles Head, J. rine Ecology, r\mchitka Island, r\laska, Annual Progress Ecol., 28: 403-437. Report, July 1972 Through September 1973, and Riee. G. B.. and R. C. hliller. 1949. Intertidal Plant and Summary Report, -1967 Through 1973, with contribu- Animal zonation in the vi;inity of Ned, Bay, Washing- tions by J. S. Isakson, P. A. Lebednik, C. E. O'Clair, ton,Proc. Cnlif. Acnrl. Sci., 26: 323-351. J. F. Palmisano, and C. A. Simenstad, USAEC Report Iluprecht, F. J., 1851, Tange des Ochotskischen Meeres, in Bhll-171-156, Battelle blemorial institute. A. T. \.on bliddendorff, Reise in den aiissersten Norden -, J. S. Isakson, and R. L. Burgner, 1973, Amchitka und Osten Sibiricns rvillrend dm Jallre 1843 und 1844, Bioenvironmental Program, Research Program on hla- Vol. l(2): 191-435, Botanik, plates 9-18, St. Peters- line Ecology, Atnchitka Island, t\laska, Annual Progress burg. Report, July 1, 1971-June 30, 1972, USAEC Report Saundcrs, Def\., 1901, Papers from the Harri~nanAlaska BMI-171-150, Battelle hleinorial Institute. Expedition, SS\r. The r\lgae, Proc. 1Vnsl1.Acad. Sci., 3: National Ocean Suwey, 1971, Tide Tables, Il'est Const of 391-486. r\'orth nnd Soutlr Bn~erica, 1972, Soperintendent of Sa\7ichM, 1'. P., 1914, ~l'~ola~icheski~ob"ezd"Avachinskoe Documents, U. S. Government Printing Office, \\'ashing- guby v" hlae 1909 g [r\lgological Circuit of the Gulf of ton, D. C. Avacha in htay 19091, Kanchntskaya Ekspeditsiyn, 2: Neal, Marie C., 1930, Hawaiian Marine Algae, Btcll. Berrfice 449-472, 593-594. Patralri Bislroy ~\luseuw~,67: 1-84. Scagel, R. F., 1966, hlarine Algac of British Columbia and Northcraft, R. D., 1948, hlarine Algal Colonization on the Northern \\'ashington, Part I: Chlorophyceae (Green h'lonterey Peninsula, California, Anier. J. Bot., 35: Algae), A'nt. dlas. Cmt. Bttll., 207. 396-404. -, 1963, Distribution of Attached Marine Algae in Okamura, K., 1933, On the Algae from i\laska Collected by Relation to Oceanographic Conditions in the Northeast Y. Koba)~asBi,Rec. Ocea~logr.Il'orks Jay., 5: 85-97. Pacific, in iV10ri.e Distributiot~s, kl. J. Dunbar (Ed.), -, 1928, Algae fiom Kamtschatka, Rec. Oceanogr. IIrorks Royal Society of Canada, Special Publication 5, Jay., 1 : 52-55, plates xiii-xv. pp. 37-50, University of Toronto Press. Pahnisano, J. F., 1975, Sea Otter Predation: Its Role in -, 1962, Coastal Studies of Marine Algae in British Rocky Intertidal Community Stracture at Atnchitka Columbia and the North Pacific, in Proceedings of the and Other Aleutian Islands, PB. D. Thesis, Universit). of A'ational Coastal and Shallolv Water Research Confer- \\'ashington. [Diss. Abstr. hi., 37(2): 595-B.] ence, D. S, Gorslioe (Ed.), pp 748-750, National Perestenko, L. P., 1967, O dvukh vidakh vodorosleiiz roda Science Foundatio~l-Office of Naval Research, Talla- Rhodoglossum J. Ag., obitayushchiiih v moryakl~ hassee, Ha. dal'nego vostoka [Concerning T~voSpecies of Algae of - 1957, An Annotated List of the hlarine Algae of the Genus Rhodo~lossumJ. t\g., Occurring in the Scas British Columbia and Northern t\'ashington, ~\'nt. dlits. of the Extreme East), Nouosti sistelltatiki nizshikl~ Can. Bell., 150. rasteiri< Akad. Natik SSSR, Bot. Inst., 4: 150-152. Setchell, I\'., 1899, SSIV, Algae of the Pribilof Islands, in Petrov, Yu. E., 1968, Rod Sargassum C. r\g. r dal'nevosto- The Fur Seals and Fur-Seal Islands of the North Pacific cllnykh moryakh SSSR [The Genus Sorgnssu?if C. Ag, in Ocean, D. S. Jordan, Part 111, pp. 589-596, Special the Extreme Eastern Seas of the USSR], i\'ouosti Papers Relating to the Fur Seal and to the Natural sistemntiki nizshikh mste,zic Aknd. A'azck SSSR, Bot. History of the Pribilof Islands, Washington, D. C. Inst., 5: 42-53. -, and N. L. Gardner, 1903, Algae of Northwestern -, 1966, Rod Cystoseira C. Ag. v dal'nevostochnykh America, Publ. Botany, U,liu. CnliJ., Berkeley, 1: moryakll SSSR [The genus Cystoseira C. Ag. in the 165-418, plates 17-27. Extreme Eastern Seas of the USSR], r\'ouosti sis- Shchapova, T. F., 1948, Geograficheskoe rasprostranenie tematiki nizshikh mste~~icAkad. i\'nuk SSSR, Bot. Inst., predstavitele; poryadka La~ninarialesv severnol chasti 3: 96-99. tikhogo okeana [Geographical Distribution of Repre- -, and V. B. \'ozzhinskaya, 1970, Nov).e vidy rada sentatives of the Family Larninariales in the North Laminaria iz Okhotskogo nlorya [Nerv Species of the Pacific Ocean], Tr. Inst. Okemzol., ilknd. Araak SSSR, Genus I.nminnria from the Sea of Okhotsk] , l\'ovosti 2: 89-138. -, 1946, On the Amphipacific Distribution of Certain -, 1964a, Novye vodorosli dlya Sakhalina, 11 [New Algae Species of Phaeophyceae, Con~pt.Rend. (Doklady) from Sakhalin, 11] Arouostisistentntiki~~izshikhrosteni; Acnd. Sci. URSS, Nouu. Ser., 52(2): 171-173. Aknd. ~\'nuk SSSR, Bot. Inst., 1: 151-153. -, and V. B. \'ozzinskaya, 1960, Vodorosli Litorali - 1964b, hlakrofity morskikh paberezl~ii Sakhalina Zapadnogo Poberezh'ya Sakhalina [Algae of the Lit- [hiarine blacrophytes of the Coast of Sakhalin], Tr. toral of the Vest Coast of Sakhalin] , Tr. Inst. Okennol., Inst. Okeaitol., 69: 330-441. (English Translation: Aknd. Arauk SSSR, 34: 123.146. hiacrophytes of the Sakhalin Sea Coast, U. S. Naval -, 0. B. hIokicvskii, and F. A. Pasternak, 1957, Flora i Oceanographic Office, Translation 462.) Fauna Litorali Zapadnogo Sakhalina (Predvaritel'noe -, 1960a, Novye vodorosli dlya Sakhalina [New Algae soobshchenie) [Flora and Fauna of the Littoral of from Sakhalin], Botanicheskie materialy, Aknd. Armck I\'estern Sakhalin (Preliminay Report)] , Tr. 111st. SSSR, Bot. Inst., 13: 119-128. Okeaaol., Akad. ~VarckSSSR, 23: 102-111. - 1960b, Nekotorye endofit], Sakhalinskikh r.odoroslei, 1. [Some Endophvtes of Aleae from Sakhalio. I1 Shetter, S. G., 1967, Tlze Komarov Botmticol Institute, . >.. Smithsonian Institution Press, \\'ashington, D. C. ~oianicheskiemate;.ialy, Akad-~~aakSSSR, Dot. Inrt., South\rrad, A. J., 1956, The Population Balance Between 13: 128-130. Limpets and Seaweeds on It'ave-Beaten Rocky Shores, -, and E. I. Blinova, 1970, Materialy raspredeleniyu i Rep. dlm. Biol. Sta., Port Erbt, 68: 20-29. sostavu vodoroslei Kamchatki (Okhotskoe more) [Mate- Spasski;, N. N. 1961, Litoral' yugo-vostochnogo rials on the Distribution and Composition of Algae of poberezh'ya Kamchatki [The Littoral Zone of the Kamchatka (Sea of Okhotsk)] , Tr. Iirst. Okeariol., Southeast Shore of Kamchatka] , Issled. Dal'i~evost. Aknd. A'nuk SSSR, 88: 298-307. I\IOIZ~ SSSR, \Ill: 261-311. -, and N. hI. Selitskaya (T'ishnevskaya), 1970a, Vidovo; Stephenson, T. A,, and I\. Stephenson, 1949, Universal sostav, raspredelenie i zapasy fukoidov \, Okhotsko~n Features of Zonation Between Tide Marks on Rocky more [The Composition, Distribution, and Quantities Coasts, J. Ecol., 37: 289-305. of Fncoid t\lgae in the Sea of Okhotsk], Tr. Inst. Tanignti, hl., 1962, Phytosociologicnl Study of hrbte Okeaf~ol.,Akad. A'ouk SSSR, 88: 281-287. Algae in Jnpnn, 1-111, Inoue and Company, Ltd., -, and N. i\I. Selitskaya (Vishnevskaya), 1970b, Tokyo. Tvlorskaya ratitel'nost' pribrezh'ya 0-va bol'shoi Sha~tar Tanner, C., in preparation, A Study of the Northeast Pacific (Okhotskoe more) [hIarine Vegetation of the Coast of Species of Ulvn, Ph. D. Thesis, University of British Big Shantar Island (Sea of Okhotsk)], 7i.. Inst. Columbia. Okennol., Aknrl. ~VaukSSSR, 88: 288-297. Tatewaki, h.I., and 1'. Kobayashi, 1934, A Contribution to -, and N. M. I'ishnevskaya (Selitskaya), 1968, Redkie i the Flora of the Aleutian Islands, J. Fac. Aqic. novyc vidy vadorosle; dlya severnykh i zapadnykh Hokkairlo U,~iu.,36(1): 1-118. poherezhi; Okhotskogo morya [Rare and Neu, Species Tokida, J., 1954, The klxine Algae of Southern Saghalien, of Algae from the Northern and I\'estern Shores of the dle??t. Fnc. Fish. Hokknido U~riu., 2(1): 264 pp., 14 Sea of Okhotsk] , A'ovosti sirfemntiki ~~izsliikkrosteai< plates. Akad. Ararck SSSR, Bot. Inst., 5: 53-56. -, and H. Hirose, 1975, Arlvance of Pl~ycologyin Japan, I\'einmann, F. C., 1969, Amchitka Bioenvironmental P1.o. I\'. Junk, The Ilague. gram. Aspects of Benthic Marine Algal Ecology at Ushakov, P. V., 1951, Litoral' Okhotskogo morya [The Amchitka Island, Alaska, USAEC Report BMI-171-115, Littoral Zone of the Sea OF Okhotsk), Dokl. Aknd. Battelle Memorial Institute. ~\Tnuk SSSR, l\'ouayn Ser., 76(1): 127-130, -, 1968, Aspects of Benthic Maine Algal Ecology at Vinogradova, K. L., 1973a, 0 notlykh vidakh Rhodomela Amchitka Island, r\laska, &I. Sc. Thesis, University of Ag. i Polycerea J. Ag. iz Beringova morya [Concerning I\'ashington. Next, Species of Rhodomelne ,\g. and Polycerene from I\'iddo\r.son, T., 1971a, A Taxonomic Revision of the the Bering Sea], il'ovosti sistematiki ,tizshikk mstenit Genus lllaria Greville, Syesis, 4: 11-50. Akad. Arauk SSSR, Bot. Irrst., 10: 22-28. , 1971b, A Statistical Analysis of Irat.iation in the Brown Alga Alarin, Syesis, 4: 125.144. -, 1973b, \ridovoi sostav vodoroslei na litorali i sub- , 1965, A Taxonomic Study of the Genus Hedoplrylltiin litorali severo-zapadnoi chasti Beringova morya [Com- - position of Species of Algae in the Littoral and Setchell, Con. J. Bot., 43: 1409-1420. Sublittoral Zones of the Northwestern Part of the I\'iddowson, T. B., 1975, The Marine Algae of British Bering Sea], iVouosti sisteeratiki r~irsl~ikhraste~ric Columbia and Northern IYashington: Revised List and Aknd. ,\'oak SSSR, Bot. Iust., 10: 32-44. Keys. Part 11. Rhodophyceae (Red Algae), Sgesis, 7: \'oronikhin", N. N., 1914, Morskiga vodorosli Kamchatki 143-186. IThe h'larine Aleae of Kamchatkal. . Kmi~chatskiva - 1973. The Marine Algae. of British Cohtmhia and .kkspeditsiyn, 2: 473-524, 592-5113. Northern \\lashington: Revised List and Keys. Part I. Phaeophyceae (Brown hlgae), Syesis, 6: 81-96. \~ozzhinskava.. . 1'. B.. 1967. Novve vodorosli dlva serernvkh . . -. . poberezl~i; Okhotskogo mo& [New Algae from thc \\'ilson, 0. T., 1925, Some Experin~ental~bservations of Northern Shore of the Sea of Okhotsk] , ~Vouosti hlarine Algal Succession, Ecology, 6: 303-311. sistematiki nizshikh rasterzit Akad. Nnuk SSSR, Dot. \\'ynne, hl. J., 1975, Further Studics of the Delesseriaceae Inst., 4: 138-140. of Amchitka Island (,\leutian Islands), J. Phycol., -, 1965a, hlorskie rodorosli zapadnogo pobcrzh'ya ll(Supp1.): 17 (abstract). Knmchatki [hlarine Algae of the \\lest Coast of -, 1972, The Genus Porplrym at hnlchitka Island, Kamchatka] , A'ouosti sistenzafiki r~izslriktr rastenir, Aleutians, in Proceedings of the Seuerith Irrterrrotional Aknd. Nnuk SSSR, Bot. Inst., 2: 73-78. Seaweed Symposirrm, Aug. 8-12, 1971, pp. 100-104, -. 1965h. Raspredelenie vodoroslei u beregov zapadno: University of Tokyo Press. ~amchatki[~istribution of Algae Off the coasts of -, 1971, Concerning the Phaeophycean Genera Atnlipris Western Kamchatka] , Okeatrologiyn, 5: 348-353. andHeterochordnrin, Plzycologia, 10: 169-175. Ecology of lblnrble Algae 393

-, 1970a, Alarine Algae of Amchitka Island (Aleutian Novostisistentatiki nizslzikh mste,zi; Aknrl. A'aak SSSR, Islands). I. Delesseriaceae, Syesis, 3: 94-144. Bot. last., 1: 125-126. -, 1970b. hiarine Algae of Amchitka island (Aleutian Zinova, E. S., 1954a, Morskie vodorosli yugo-vostochno; Islands). 11. Bonnernaisoniaceae, Pnc. Sci., 24: 433-438. Kamchatki [AIarine Algae of Southeastern Kamchatka] , Tr. Bot. Inst., Akarl. Nnuk SSSR, Ser. 2: Sporovye Zinova, A. D., 1970, Dopolnenie k stat'e o novom vide Rnsteniya, 9: 365-400. lalninarii c 0. Sakhalin [An Addition to the Comments -, 1954b, Vodorosli Okllotskogo morya [Algae of the an the Neb,. Species of Laminnrin from Sakhalin Sea of Okhotsk] , li.. Bot. hst., Aknd. Nouk SSSR, Ser. Island], i\'ovosti sirtentntiki ltizshikh rastenii, Aknri. 2: Sporovye Rasteniyn, 9: 259-310. Nnuk SSSR, Bot. Inst., 6: 65-68. -, 1940, AIorskie vodorosli Komandorskikh ostrovorr [Blaine Algae of the Kon~andorsk Islands] , Tr. -, 1965, Predstarriteli sem. Delesseriaceae (Rhodophyta) Tikhookea,~.Kom., V:166-243. v serernoi chasti tikhogo okeana [The Family Deles- -, 1933, Vodorosli Kamchatki [~\lgaeof Kamchatka] , seriaceae (Khodophyta) in the North Pacific Ocean], Isslerl. Alore; SSSR, 17: 7-42. Novosti sistematiki nizshikh msteiiii: Aknd. A'aak SSSR, -, 1930, \'odorosli Okhotskogo mo13.a s poberezhi; Bot. Inst., 2: 78-97. bol'shogo Shantarskogo ostrova [Algae of the Sea of -, 1964, Not7< vid Laminaria u beregov Sakhalina [A Okhotsk Off tl~eShores of Big Sbantar Island], Tr. New Species of Lantinnrio Near the Coasts of Sakhalin] , Leni~igmrl.Obshch. Estest., 60(3): 81-125. This page intentionally left blank

Marine Invertebrates Charles E. O'Clair* Fisheries Research Institute, University of \Vashing- in Rocky intertidal ton, Seattle, \\'ashington J Communities

Seue~rzoogeograpAica1 elements were recognized i~rthe Previous ktrowledge of Alerrtian marifze bentlzic inverte- shallow-water marine fnuna of Amchilka. The greatest brates was based 011 few obseruatioas "ad collecliorts, the proportion of species ure North Pacific or North Anzerican Iristory of which is srrntmarized. In this study tiwee belt in their distribution. Tzuo oceanographic features (the tronsects (one sampler1 once a~rdtruo sampled seuen times) Alasknn Stream aud the Kamclmtka Current) and one anrl two intertidal arrays (sampled tzuice each) of 0.25-m2 geologtc feature (theflleutiaa "stepping-stone" isla~rdsbe- quadrats were used for descriptive studies of intertidal tween Amchitka and the Asian and A'ortk Bnrericnn conzmu,~ities at Amchilka Island Tlrese com~~rrcnitiesore ~~taiirla~tds)htcrmse tlre immigration rates of North Ameri- dominnted by algae nt all lidnl leuels. Three comnrnitities can species over Asiatic sf,ecies. designated nccordit~g to their dominnr~t mncrophytes nre The effecfs of Cnn~rikinwere deternri~zedzuifh the use the Laminaria community, the Alaria-Hedophyllum com of two irztertidnl arrays offixed 0.25-m2 qr~adrals(totaling munity, and the Ilalosaccion-Fucus com~xunity.Inucrte- 40 plots) ~xnntincdtwice preruent nnd ten times posteuent Orates in these co~nmu,rities are tnostly inconspicuous. (over 33 nfontlfs)anrl an iirtertirlnl grid (control) sampled Despite tlte large proportiotr of North Anrerica~rspecies in ra~~dotrrlyonce preeuent nud tzuice (ouer 9 inortths) the fauna of Anrchitkn, species that pkzy key roles in posteuent. Tlte rate of die-off and enrigration of iittertirlnl straclurirrg i~rtertirfolcon~manities elsezuhere on the west species in masimnlly trplifted (as ~ntjch as 1 111) areas const of Nortlr America are absent or in low ~bunrlaucent depended on exposure to open ocenn waves, Alost preeuent Amchitka. Inuertebmtes in subtidal commrr~titiesare rlis- i~zterfidalspecies were replaced by supralittoral frilf.re cussed briefly. An nanotnted list of over 365 littoral and species. sublittoral inuertebrate species u nppended

The marine invertebrate fauna of the Aleutian some of the salient differcnccs in community Islands has received little attention beyond the structitre bet~\.ecn these communities and other initial taxonomic descriptions that ltave follo\ved conlparablc North Pacific intertidal commttnities. mainly the collccti~lgactivities of E. \\'osnesenski Descripti\'e studies alone provide little insight into and W. H. Dall. Except for the pioneering work of commiwity organization, and tlley have largely Gurj,anova (1935; 1966) on Bering Island (Ostrov been replaced by approaclics of a con~parativeor Bering?) and the brief description by Barabash- experiniental nature. I justify a descriptive ap- Nikiforov (1947) of the benthic fauna at Copper proach hcre 1))' pointing out the absence of a Island (Ostrov Mednyi) in the Commander Islands previous detailed study of Aleutian intertidal cum- (I

0 - 0 0

( KOMANDORSKIE 0 5 Kodiok

LEUTIAN -

50°N - 50"N -

."'l~~~lllll~lrlIIII'IaI'sL 160' 170-E 180" 170"W 160'

Pig. I---Map of the northern North Pacific Ocean and Bering Sea sho\sing the location of Amchitka Island, Alaska, the Alaskan Stream (information from Favorite, 1967), and tlie Kamchatka Currcnt (information from Dodimead, Favorite, and Hirano, 1963). describe the most destructive and lo~lg-lasting trlrrlcatn. Steller and those early ~~aturalists\vho effects on sonie rocky intertidal co~nmuiiitiesdue catne after hi111 (Merck, Sauer, Tilesius, Chamisso, to uplift of the iiltertidal benches by the under- Eschsclioltz, and L~lertens)into this region during ground ~iucleartest, Cannikin. the late eighteenth and early nineteenth centuries participated in expeditions ~vhoscprimary mission was to discover and explore new lands. Tbe PREVIOUS INVESTIGt\TIONS IN THE opportunities for naturalists to make observations ALEUTIANS on the shores of tlis region were limited. \\'hen it was possible to go ashore, their attc~ltio~lwas Georg \\'ilhelm Steller's importailt contribu- drawn to the more obvious flora and fauna, such as tions marked the beginning of zoological investiga- nlarine mammals, birds, and the more co~~spicuous tions in the North Pacific. \\re do not know nrlietlier Steller collected or described invertebrates terrestrial plants. The oi~lypublications on Aleu- from Bering Island, the only Aleutian island on tiail invertebrates that eniergcd from these early \vhich he set foot, because most of his collectiolls expeditions were very brief descriptive accouiits of from Bering Island and his ma~lttscriptson inverte- the fauna of one or two islands (Sauer, 1802; Chamisso, 1821) and some taxonon~ic work brates \Irere lost (Stejneger, 1936). \\'e do know (Chamisso aid Eysenhardt, 1821; Eschscltoltz, that hc collected and described invertebrates from 1833). Icamchatka, e.g., Cry/>toc/zito?rste//erih and 1V1ya The first real scientific docume~ltationof Aleu- tian ill\rertebratcs came about the middle of the nineteenth ceiitury in the taxonomic works of *Dall (1884) states that specimens of Cryptochiton h,liddendorff (1846; 1849a; 184911; 1849~;1851), slelleri, described by Stelter under its IEamchntkan name Brandt (18.19; 1850; 1851), and Grube (1855), "1Ceru," were collected by Steller on Bcring island. Dall which describe several species collected in the cites as his rcfcret~ceSteller's Beschreiblrng uon dent Lflitde Ko,nsclratkn; howetrer, I could find no reference to Bering Aleutians prin~arily by E. \\'osnesenski. Prior to Island in Stellcr's description, and hliddendorff (1846) lists I\'osiiesenski's excursions, \vIiich extended to Atka only I'eter and Pnul (Petropaslovsk) Harbor in Kamchatks and At~uIslands, invertebrate collecti~lghad been as the collection localit). of C. slelleri. limited to the elids of the Aleutian chain, espe- Ii~vertebrntesbz Rocky Ii~tertidalCo?i&ti~l~lities 397 cially around Unalaska in the east and the Com- PHYSICAL ENVIRONiVIENT ~nariderIslands in the west. Tides The greatest contribution to our kxio\vledge of Aleutian invertebrates has come froin \\'illiam H. Tlie most importaut phenomenon ultimately Dall, \\~hoscassignmeilts in the Aleutia~lsunder the controlling the lives of iilost littoral invertebrates is U. S. Coast Survey from 1871 to 1874 ellabled him the tidal cycle. The amplitude and frequency of to make collections on a number of islands, tidal fluctuations deterl~linetlie length of time a iiicluding Amchitka, from which illvertebrates had particular level on tlie shore is either subnierged or not previously beet1 collected. Dall subseque~ltly exposed to atmosphelic conditions, xvhich, in turn, described most of the species he had collected, ge~leraily determines, directly or indirectly, especially the mollusks (see Boss, Roselvater, and whether a species call maintain a population at that Ruhoff, 1968). Collectively, these publicatio~lson level. For example, the upper limit of distributioii tlie systematics and zoogeograpliy of Aleutian of a particular species inay be deter~niueddirectly ~iiolluskscoiistitute the definitive source of infor- by the ainount of time it is subject to the stress of mation 011 this the best known group of Aleutian desiccation; at the salile time its lower limit may be invertebrates. In addition, Dall's collections have set indirectly at that level below which it is provided most of the Aleutian littoral material for excluded by ~narinepredators or competitors more a number of publications on other invertebrate susceptible to the stresses of emersion. [There are groups: hydroids (Clark, 1876), barnacles (Pilsbry, exceptions to this generality, e.g., R. T. Paine 1916), isopods (Richardson, 19G5), starfish (personal communication) finds that the mussel (Fisher, 1911; 1928; 1930), and others. d1)~tilllscnlifortliatltls sets the upper limit of the After Dall, collectiilg activity was again mostly alga Alnrin i~~nrgit~ntu.] restricted to the eastern aud western ends of the Unlike the semidiicrnal tides that predominate Aleutiau chain. Investigations near the eastern end throughout the ~vorld,the tides in the vicinity of of the Aleutians have included the trawling atid the Aleutian Islands ale irregular diurnal (Plate 1 of clredging activities of the U. S. Fish Commission Doty, 1957), being primarily diurnal hut tending steamer /Ilbatross in the \\raters from U~lilnakto tolvard mixed durilig neap tides (for definition of U~iiliakIslands in 1888, 1890, aud 1893 a~idthe terms, see U. S. Department of Commerce, 1949). Harriman Alaska Expedition in the vicinity of Figure 2 shows representative tidal curves for Unalaska in 1899. At the western end of the chain, March atid June 1969 at Amcliitka. the Comlnauder Isla~ldswere visited by the Vega Tlie tidal rauge at Amchitka is narro\cr com- expedition under Nordenskiold in August 1879; by pared \\,it11 those at other locatio~lswhere descrip- Stejneger under the auspices of the Smithsonian tive studies of littoral co~ntiiurtities have been Institutioil and the U. S. Signal Service in 1882, performed. For examplc, the diurnal range, the 1883, and 1895; by the Albatross in 1892, 1895, difference between meall higlier high water 1896, and 1906; and by several Russian llaturalists (h,lHH\\') and mean lower loxv \\later (MLL\V), at (see Gurjano\ra, 1935). However, some collections South Bight, Amchitka, is 1.1111, \shercas it is 2.5 n~ wcrc ~ilade near Amchitka. They illclirde those at Three Saints Bay, Kodiak Island, Alaska, 2.4 iii made by iilvestigators on board the illbatross at at Neah Bay, \\lash., and 1.6 111 (5.2 ft) at blonterey IGska Islaud in 1894 a~idnear Sen~isopoclinoi Bay, Calif. (U. S. Department of Commerce, Island in 1906 and by 1'. Scheffer aboard the 1969). The ~nasiniu~iitidal range at Amchitka is Brozu11 Beur in 1937 at Amchitka, hlore recently, just over 2 111 (2.1 In in 1969 at South Bight). Newell (1950; 1951) and I'amada (1955) have (Computed from hourly tidal hcight tlieasureli~e~lts published on collectio~isfroill this area. at Sweeper Cove, Adak, Alaska, 1969, \vhicli \sere co~rectedfor Amcliitka \vith the correctious pub- Despite all the collecting activity, very little has been published on the ecology of Aleutian benthic lished in the tide tables.) invertebrates. Dall (1884) and Annenkova (1934) have provided sollie i~lforlnationon the natural Tc~iiperat~~reand Salinity history oC the littoral mollusks and polychaetes of McAlister (1971) has discussed several aspects Bering Island, but their publications were primarily of the offshore lilari~iephysical en\ironment in the taxoiioiiiic in nature. The studies on the distribu- vici~iityof Amchitka. Two aspects of the illshore tion of algae and invertebrates at Bering and lilarille environment, water temperature and salin- klcdnyi Isla~ldsby Gurjanova (1935; 1966) and ity, \\,ere iiloliitored during the course of our Barabash-Nikiforov (1947), respectively, represent studies. the only published attempts to deal \\.it11 the Water telilperature \vas lllollitored in Constan- benthic cotntiiuilities of any Aleutian island. tine Harbor ~vitli45-day recording thermographs 4.00 SMHW -- CMHW r-' I2.00 SMTL w CMTL w I ' I I v 0.00 I '"V w \I ' ll ll 1 ll I/ " ' u I1 1 V \I V II \IV V\ MLLW V\IIVV T T' t -2.00 I I I I I I I I I I I I I I I 0.00 2.00 4.00 6.00 8.00 10.W 12.00 14.00 16.00 18.00 20.W' 22.00 24.00 26.00 28.00 30.04 32.00 DAYS MARCH I969

4.00 SMHW

% CMHW r-' I 200 SMTL w CMTL w I 0.00 MLLW

-2.00 0.00 2.00 4.00 6.00 8.00 10.00 12.00 14.00 16.00 18.00 20.00 22.00 24.W 26.00 28.00 30.00 32.W DAYS JUNE 1969

Fig. 2-Representative tidal curves for March and June 1969. The curves were constructed from hourly tidal height measurements at Sweeper Cove, Adak, Alaska, 1969, obtained from the National Ocean Survey. The hourly tidal heights were corrected for Amchitka with the use of the Constantine Harbor (c) and South Bight (s) corrections published in the tide tahles. The corrections are mean values of range and tidal planes computed from automatic tide gauge records. The value for Constantine Harbor on the Bering Sea side of Amchitka is based on a 6-month record by automatic tide gauge, August 1946 through January 1947. The value for South Bight on the Pacific Ocean side is based on a 6-day record by automatic gauge from June 29 to July 5, 1945.

hung approxiinately 6 m (20 ft) belo\\, mean tide the U. S. Naval Oceanographic Office data. Surface level beneath a dock. Figure 3 shows the mean, temperatures of sea water over the intertidal rocky maximum, and ininimum inonthly temperature benches sho\\, a geater variability than those variability for Constantine Harbor based or! ther- recorded in Constantine Harbor. Temperatures mograph data (32 tapes) collected intermittently taken (178) of the \crater over the bench show a fronl 1968 to 1973. low of 1.l0C in January 1970 at Square Bay and a The curve inset in Fig. 3 shotvs tlte same water high of 12.1°C in June 1970 at h,lakarius Bay. temperature characteristics as the main figure but Sa~nples of surface sea water for salinity it is dmxvn from sea-surface isotherms shown for analysis xuerc taken in citrate of magnesia bottles the vicinity of rlmcl~itkain U. S. Naval Oceano- concurre~ltlywith surface temperature readings at gaphic Office Publication on Sea Surface Temper- our study sites. Samples were usually taken on a11 atures of the North Pacific Ocean (1969). The inconling tide at a water levcl bet\veen mean lo~rrer cnrve of ~ncanwater temperatures for Constantine lo\v water (MLLIV) and mean tide level (VITL). The Harbor differs very little from that based on Naval analysis was done on a laboratory salino~neterat Oceanographic Office data except that mean sum- the Department of Oceanograplty, University of mer temperatures were lotver. The position of tlte \\lashington. Salinities at the study sites remained, tllermograph 6 nl (20 ft) belo\\, the surface may for the tuost part, between 32 %, and 34 %, . have influenced the readings. Variability in the Extremely lo\v salinities were often recorded at extreme water temperatures at Constantine Har- Makarios Bay on days with onshore winds. The bor, for the most part, falls within the range for study sitc at Makarius Bay is less than 100 m from ~llnri~zeI?tuertebrates in Rocky Intertidal Conrnrn~tities 399

Fig. 3-Water temperatures at Constantine Harbor, Amchitka Island. Records are from 32 recording thermograph tapes taken intermittently from April 1968 through January 1972. Inflections in the cuwe of means reflect variability resulting front few tapes. hlinimum and maximum values are plotted opposite the time of month at \vhich tltey were recorded. Inset shows the same temperature characteristics based on sea-surface isotherms shorvn for the vicinity of Amchitka in U. S. Naval Oceanographic Office Special Publication 123. the mouth of a short stream draining a 10-ha nay be as great as the annual range in mean (25-acre) lake (Clevenger Lake). Dye releases at the air temperature at Amchitka. Glynn (1965) mouth of the stream indicate that, when onshore recorded temperatures in the E~tdocladia-Balanus winds prevail, much of the stream's effluent is association at Pacific Grove, Calif., for an 8-lir swept south nearshore over the bench. period in September 1961 and found that the range bet\veen the highest and lowest of his in situ Climate measurenients on exposed rock surface was greater than the range bet\veen the mean monthly maxi- The stresses of the physical environment are ~iiuni(September) and mini~nuln(December) air most severe on nlarine organisms when they are temperature for tlie year at that locality (see also exposed during low tides. Aspects of the atmo- Hedgpeth and Gonor, 1969). spheric climate \vhich are i~nportantto these organ- isms are air temperature, precipitation, insolation, Severe osmotic stress occurs when e~iiersed relative humidity, and \\rind. Tlic foilo\\~ingdiscus- intertidal organistns are exposed to heavy rain. sion is based on tlie climatological data of However, heavy rains are infrequent at Amchitka; Armstrong (1971; Chap. 4, this \'oltutne), \\rho has the mean anni~alprecipitation is only 83 cm, and summarized tlie surface weather observations of the average number of days with greater than 1 cm the U. S. Department of tlie Air Force arm)^ Air precipitation is only 16. Light rain benefits inter- Corps before 1947) at Amchitka during tlie period tidal organisms by reducing desiccation rates. Fcbruary 1943 through June 1948. Other aspects of the maritime climate that As is characteristic of ~nariti~neenvironments, promote low rates of desiccation in the intertidal tlie lnean daily and illcan annual ranges in air area ;we a cloudy sky, high relative humidity, and teiiiperilture (3.9"C or less and 9.4OC, respectively) onshore wind. The total freque~lcyof broken and are relati\rely narro\v at t\mchitka. However, my overcast skies at Amchitka is at least 70% every records from thermographs placed intertidally month of the year. Also, heavy cloitd cover and fog sho\v that the range in temperature (from subnier- are most frequent in the sunlmer (90 to 95% and sion to emersion) to tvhich intertidal organistns on 30 to 52%, respectively) when lo\\' tides occur in Amchitka are exposed during a single tidal cycle daylight hours (Armstrong, Chap. 4, this volume). There is no published summary of relative humid- collections from over 300 additional plots at these ity at Amchitka, but at Shelnya Island, 180 miles and 12 other intertidal sites (Fig. 4). Most of the (290 km) wcst of Amchitka, the incan inontltly time I spent at Amchitka (432 days) \\'as from relative humidity is 80% or more for all months June to September, but I have made observations (U. S. Department of Commerce, 1971). there in every month except No\reinber. Winds increase desiccation rates by increasing Sal~~plii~gProcedures. Each of the three tran- evaporation; however, onshore winds increase \va\'e sects rvas divided along its length into sections on splash aiid sea spray and, if strong enough, may the basis of substrate or doininant algal cover. A even prevent complete tidal ebbing. Winds from minimum of trvo 0.25-in2 plots judged to be the south~vest prevail during the summer; thus typical of each scction were selected for sampling. desiccation rates of intertidal organisms along the The algal canopy of each plot was photographed Pacific coast of Ainchitka with its southerly aspect and then removed. As the fronds were removed, are alnelioratcd during daylight low tides. individuals of each species of invertebrate epifaunal on them \sere identified and co~unted.The plot was Substrate photographed again. This second photograph \\'as Po~vers,Coats, and Nelson (1960) and Gard used to deterinine the pcrcent cover of algal (Chap. 2, this volume) describe the geology of understory and the relative abundance of such Amchitka. The general map of the geology of invertebrates as Halicho~~driapatlicea,* an encrust- Ainchitka in Polvers et al. (1960) indicates that the ing sponge. The number of individuals of each of so-called Banjo Point formation, described as "bed- the larger macroinvertebrate species was then ded marine sandstone, conglomerate, tuffaceous recorded. This number \\'as coinbined with the shale aiid some lapilli tuff of basaltic composi- individual counts of species epifaunal on the fronds tion," forms the substrat~unof all the study areas of the algal canopy, Invertebrates that were too to be discussed in this chapter. I-Iowever, in some small and too numerous to count over the whole areas, especially at Rifle Range Point, the snbstrate plot nrere subsainpled with a 5-cm(2-in.)-diameter is highly indurated. circuls ring. Everything within the ring \\,as removed do\vii to bare rock. The nnlnber of s~rbsanlplestaken varied with the heterogeneity of METHODS the algal understory and substrate cover. It was sonletiines necessary to extrapolate counts of large Trailsects and Intertidal Arrays invertebrates froin subareas of a plot when the plot Study Sites. Trvo intertidal belt transects pre- \\,as threatened \\,it11 i~ninineilttidal inuudation or viously established in i\,karch 1968 by F. C. contained one or two species too numerous to IVeinniann at A~IakariusBay (T,) and Square Bay count coinpletely in the time a\railable. Subsamples (T2) for studies of algal ecology xverc selected as and representatives of tunidentified species were initial benthic invertebrate study sites (Fig. 4). In removed to the laboratory \\'here they were fixed in 10% neutral-buffered Formalin. The subsamples July 1969 I established a third transect at Rifle \\,ere sieved through a 1-mm sieve, sorted, and the Range Point (T4) (T4 is not the T-3 of Lebednik animals identified. and Palniisano, Chap. 17, this volume), and in h,Iost invertebrates \irere initially identified August 1970 P. A. Lebednik established intertidal from the literature. When possible, representative arrays (IA-2 aiid IA-3) of 0.25-m2 quadrats on the specimens were coinpsed with niuseuln spcciinens Bering Sea side of the island. These five study sites or sent to specialists for identification or verifica- provided data on invertebrate distribution and tion (see the appendix). abundance from sheltered and exposed areas on Transects TI and T2 were sampled in \\,inter, both coasts. 011 the Pacific Ocean sidc, TI is in a early spring, late spring, summer, and early fall sheltered area and T4 is an exposed site. On the (Table 1). I did not sainple the same quadrats on Bering Sea side, T, is sheltered, IA-3 is slightly less different dates (Figs. 7, 10, and 12 sho~vthe sheltered, and IA-2 is exposed. The azi~n~ithsand locations of quadrat centers sampled on dates profiles of the transects and the elevations of the other than August-October 1969 at Square Bay). plots in the arrays \\'ere fixed by standard survey- Because of the difficnlty of identifying and count- ing techniques. 1\11 measurements of elevations \Irere subsequently related to incan lo\ver lolv water (MLLIV) lor Amchitka by a professioiial surveying crc\v. The true azimuth and length of each transect *'l'he appendix contains a partial list of invertebrates are given in Tablc 1. In addition to sampling these collected at An~chitkaincludine " the authoritv and date of five main study sites, I made obsel-\rations of and the original description for each scientific name. A.lari~zeI?zuevtebrates bt Rocky I~ttertidalCo71~1izunities 401

Fig. $-Maps of r\nlchitka island showing intertidal collecting areas (@, upper ]nap) and major study sites, Cannikin surface zero (e) and area of Chionoecetes bairrli catches (shaded area, lower map). ing snlall invertebrates in situ in the dark in early belo\\.); thc transects were abandoned after blarch fall and winter, data from T, and T, are inadc- 1970. quate for detcrmilli~lgseasonal variability in in- In August 1970 two intertidal arrays (IA-2 and vertebrate abundance. Seasonal data xverc nsed IA-3) of 0.25-m2 quadrats were establisbcd by oilly to refine the distributioilal limits of inverte- Lebednik (Lebednik and Palmisano, Chap. 17, this brates. [If a species \\,as recorded at a position volume) on thc Bering Sea coast of Amchitka alotlg the tratlsect other that1 one of those sampled (Fig. 4). Forty quadrats (25 at I/\-2 and 15 at in August-October 1969 (but \\~ithinthe same 1.4-3) \Irere placed in patches of more or less community) in a different season and there was no uniform algal covcr and distributed over the a priori reason for not espectii~git at that positio~l surface of the intertidal bench so that all lnajor in August-October 1969, its distribution was algal zones were rcprcscuted. Each plot was extended accordingly.] I found it iillpossible to marked in the ccilter and at t~vocorners with flags contini~esatnpli~lg more than one control ~vlhenI of polypropylellc rope ilailed to the bench surface began the stady of the effects of Ca~lnikin(see with concrete nails. \Vhen data collection on the Table 1-Azinlutlls, Lengtlls (or Size), and Sampling Dates of 'l'bvee Transects and One Intertidal Grid and Sampling Dates for Two Arrays at Alncbitka Island

Number Study site Azin~uth Sainpling dates of plots

Length, m

Makarins Bay Aug. 7-10, 1968* (TI) Ian. 1-2. 1969 Mar. 24-26,1969 June 12-15, 1969 Square Bay 098~58' Aog. 25 to Oct. 17, 1969 (T2 True Ian. 3-12. 1970 (kr. 12-16, 1970t Rifle Range July 30 to Aug. 24, 1969 Point (4114)

IA-2 and It\-3 Aug. 15-18, 1970 Sept. 6-12, 1971 Dec. 2-7, 1971 Feb. 21-25,1972 Apr. 15-26,1972 June 14-19, 1972 Aus 2-12,1972 OC; 22-24, 1972 Apr. 13-20,1973 Aug. 22 to Sept. 2, 1973 May 22-27, 1974 Aug. 27-28, 1974 Size, m' h4akarius Ray x 192'55') 3717.5 intertidal grid (61-m square) Feb. 20-24,1972 \, 282'55' Aog. 18-22, 1972

*Both TI and T:, were visited during each time span throogh hlarch 1970. $General obse~vationsand physical sumeying \\,ere done during this period; no plots were sampled.

algae in each quadrat was completed (see Lebednik transects. Ho~vever, because of the effect on and Palmisano, Chap. 17, this volume), the number subsequent observations of sampling without re- of indi\,iduals of the larger macrofau~lalinverte- placement on the same plots, subsampling was brate species (those retained by a 2-111111 sieve) were stopped after Augnst 1970. Thereafter only semi- recorded, and the percent coverage of all colollial quantitative data urere collected on small abundant species lvithin each plot was estimated by visually species. Counts of individuals per species were bisecting the plot into equal-sized rectangles and made twice (August 1970 and September 197 1) then subdividing these rectallgles into even smaller prior to Cannikin. Postevent cotunts begall 1 month equal-sized units until an estimate could he gained. after the test and continued on a I~imonthly It was impossible to obtain collsistently accurate schcclule for 10 months. Thereafter cou~ltswere coul~tsof individuals of the smaller macrofauna made at 17, 22, 30, and 33 months postevent. (the majority of whose adults would be retained by Rockfalls due to Cannikin buried seven plots (five a 1-111111 sieve) o\ving to the dense algal cover on at IA-2 and two at IA-3). So that the loss of these most plots, especially in the H(l1osnccio11-Fuctrs plots could be compensated for, t\\w stations \\.ere commimity. In tlugust 1970 plots that contained added at IA-2 in February 1972. Table 1 gives almost exclusively these tiny abundarlt species saml~lingdates ;old the number of plots sampled were subsampled ~vith an 8-cm-diameter, for each d;ite for IA-2 and 1A-3. A11 intertidal grid 1.4-cm-deep circular ring in a manner similar to established at h3akarius Bay for another study that used to subsample the quadrats on the acted as the control. I~luertebratesin Rocky I~ltertidalConzn~rr)~ities 403 Intertidal Grid algal zones \\,as met. If the qoota of one algal zonc \\.as filled before the others, coordinates that A stratified random sampling scheme with subsequently fell in that zone were ignored. major algal zones as strata \\.as adopted for sampling populations of invertebrates at h4akarius Quadrats were l~ositioned in the field by Bay. A 200-ft(61-~n)-squaregrid was laid out with measuring the distance of one coordinate along one standard surveying equipment to fix the azimuths asis (usually that axis parallel to the shore) and of the axes. Each axis was then divided along its then stretching a surveyor's tape in a pcrpendicr~lar length into 10-ft (3.1-m) intervals, each of ~vhich direction to a length cqual to the othcr coordinate. was marked with a flag of polypropylene rope or Perpendicularity tvas determined with a Bruntot~ surveyor's flagging nailed to the bench surface xeith pocket transit. The length on the surveyor's tape a concrete nail and \\rasher. Thcse m;~rkcrs facili- corresponding to the second coordinate marked tated the procedure for mapping the major algal the center of the quadrat to be sampled. zones \vitItin the grids and helped in locating the positions of the plots to be sampled. Quadrats \\,it11 a surface area of %, m2 \\,err used to sanlple the grid. As each quadrat was Mapping of Algal Zones. The procedure for positioned, a photograph xvas taken to record the mapping the nlajor algal zones was as follo\vs: A percent cover of the canopy-forming algae. The surveyor's tape leas stretched out the width of the fronds of these algae falling \vitllin the quadrat grid in a direction perpendicular to one of the grid \\'ere then removed and placed in a polyethylene axes from one of the nlarked points along the axis. bag, and the plot was rephotographed to record the The algal zone within \vhich the tape intersected percent cover of understory algae, I~oldfastsof the axis was recorded. I then walked along the tape canopy algae, and colonial invertebrates, such as and recorded the sequence and \\~idthsof the ntajor encrusting sponges. The plot xvas then stripped to algal zones \\,here they were intersected by the bare rock wit11 paint scrapers, stone chisels, and tape. The tape \\,as stretched tight before each modified spoons, the contents \\,ere placed in the measurement was taken. This procedure \\,as re- polyethylene bag with the canopy algal fronds, and peated for the rcnlaindcr of the marked points the bag was labeled. The plot was thcn marked at along the same axis and all the points along the one corner wit11 a concrete nail, washer, and perpendicular axis. Forty runs \\,ere made on the surveyor's flagging for future reference. \Vhen all grid. The runs \sere reproduced to scale on graph plots had been sanlpled in each season, their paper as lines ~shosesegments represented seg- elevations were related (using standard surveying ments of the surveyor's tape \\,hich passed over a techniques) to points \vhose elevations \vcrc detcr- particular algal zone. By connecting the points that nlined by a professional surveying crew. The marked the boiundaries of the algal zones along the samples were removed to the laboratory for pro- cessing. Time did not permit f~~llprocessing of the line segments, I obtained a rnap of the distribution of the major algal zones \vithin the grid. \\'hen samples at the field station; so most samples were therc \\,as doubt about the position of the bound- fixed in 10% neutral-buffered Formalin and shipped aries, as in areas of irrcgtular topography whcre to Seattlc. A4akarius Bay was sampled in the fall algal zones changed rapidly and irregtlarly, the 1 month prior to Cannikin and in late winter and map \\,as compared directly \\,it11 the algal distribu- late summer following Cannikin (Table 1). No tion in the field. obvious changes (LC., no uplifting) had occurred on the grid after Cannikin, and therefore it was not The algal map was used to detern~ine the remapped. allocation of sample size within strata (algal zones). After the map had been drawn up, the relative Each sanlple was screened successively through areas of each algal zone rvithin the grid were standard screens \\,it11 mesh openings of 4, 2, and deter~nincdwith a compensating polar planimeter. 0.991 mm and the invertebrates were separated for The number of quadrats allotted to each stratum species abundance co~tnts.All invertebrates \\,ere \\,as proportional to thc relati\re area of that counted and preserved in 70% isopropyl alcohol stratum in the grid. containing 2% glycerin. Invertebrates retained by the 0.991-nlm screen were counted under a dissect- The coordinates of each plot were taken from a ing microscope. Invertebrates of the nlajor taxo- random-number table and plotted on the map of nomic groups klollt~sca, Ecltinodernlata, and thc grid. The numbers taken front the table Crustacea, excluding Amphipoda, \\?ere identified represented distances (in meters) from the grid to species, and the individuals of each of these origin along the ases. Coordinates \\,ere taken in species were counted. Invertebrates of minor phyla this manner until thc sample size quota for all the \\.ere identified to species \\rhcn possible. Sources of Bias and Error. The grid at emersion and thus reduces the necessity for orgall- Makarius Bay was established priniarily to act as a isms to move to lo\ver levels \\'it11 the receding tide. control for another study. Because the sample The spatial relationships among sessile organisms, surfaces (Yl6 m2 versus '/q m2) and the salnpling such as barnacles, of course, do not change \sit11 schemes for Makarius Bay and IA-2 and IA-3, the tide. respectively, were different, the iVIakarius Bay grid 5. Because of the dense algal cover on the IA-2 was not a proper control for the plots at IA-2 and and IA-3 plots prior to Cannikin, it \\,as often IA-3. Ho\vevcr, limits on field time and resources difficult to get complete counts of even the larger precluded thc maintenance of two controls. In macrofauna. As the algal cover died and sloughed view of the striking changes that occurred at IA-2 off, bleu. away, or was xvashed a\tray in stornt and 1A-3 follo\ving Cannikin, the results would \vaves after Cannikin, invertebrates became much probably not have been different had an additional easier to see, and species counts could be made control been added at h~lakarius Bay using a easily. Therefore the probability of undcrestimat- sampling scheme and sample surface identical to ing the preshot abundances of species in the IA-2 that at IA-2 and IA-3. Both sample surfaces and and IA-3 plots was much greater than that for the associated data-collection procedures were used at postshot counts. h~iakariusBay, and those used on the intertidal grid Three sources of error ntay have affected my always turned up nlore species and indi\~idualsin a given plot despite the reduced sample surface and zoogeographic analyses. The first results from the \vould be expected to be at least as sensitive to tendency of early taxoltontists to describe more changes in species abundance patterns as that used species than are valid. Thus sonle species of at IA-2 and IA-3 even with the introduced spatial mollusks described by Dall from the Aleutians may \'ariabiIity in abundanccs due to random sampling. prove to be synonymons with previously described fornls as have, for example, sonle of the tellinids he Other sources of error are as follows: described (Coan, 1971). This type of error \\rould 1. The abundanccs of certain highly nlotile bias my zoogeograpllic analysis by erroneously forms, e.g., gantmarids, capable of withdralving inflating the ltuntber of endemic species. deeply into the holdfast systems of algae whcn ti second and opposite source of error results enlersed or of escaping unnoticed from quadrats from the tendency of students newly initiated to a during sa~nplingwere probably often underesti- faluxla to force their specinlens into existing keys, mated as compared \\,it11 the less active species \\,it11 thereby niissing ne\v species and \vrongly assigning the use of both of the sampling schenics described specime~ts to more \videly distributed species. I above. have made every cffort to have my identifications 2. The abtuldances of the smaller fornts of both verified by specialists (see the appendix), therefore the large (retained by a 2-mm or larger mesh the above sources of error are probably nealigible. screen) and small (retained only by a 1-mm screen) Another source of error resitlts from an incom- size classes \\.ere probably icnderestimated as corn- plete knowledge of the distributions of some pared \vitb the larger forms, depending on the species. Because of the relative inaccessibility of sa~nplingprocedure. the Russian and Japanese literature, my deterniina- tion of the relative contributions of Asiatic and 3. Small species about the same sizc may be North American faunas to the benthic invertebrate under- or overrepresented because different growth fauna of Anlchitka may be biased. The recent fornls have different probabilities of retention by papers of Spassky (1961) and Golikov and the sieves. Ies at all rank and greater dominance than thc other three tlcgrecs of esposurc. It ranges from -3 or -5 111 species. However, because Al. azitiazttis is ~nuch (Lebednik and Palmisano, Chap. 17, this \.o1111ne) smaller than an average individual of the above to as higll as t0.61 111 in exposed situ;ttions. species, it is clearly not dominant in biomass. Of the 142 spccies of i~lvcrtcbratcs(excluding Statistics based on abu~ldanccwere not calcnlated those not identified to species) found at the five for the sponge Sc)lplta cotzt[)ressa because abun- main study sites, 110 were found in the Lnzniztaria dance \\,as measured as percent cover, not numbers community. The average number of species per of individuals. 0.25 m2 anlong the large species was also greater in The assignment of characteristic spccics status the Laiztittarin community (13 species) than in the to the five species of the small-size class is tentative nest richest commutlity (/llnria-HedophylItrzzt, 11 because the lumber of plots sampled for the small species). Considering data from transects T2 and species in tllc Ln~ztiz~nricicon~~nu~~ity (3) is cx- T4 only and including s~llallspecies, the difference tremely small. Three species, Dentoztnx sp., Atrtol- bctu~een these t\vo communities is more pro- yttts Deriztginzzus, and Cercops contpncttrs, \\,ere nounced (42 vs. 32 species per s;imple). The foinld in only one sample, but that sample oc. number of spccics that were faithful (f greater than curred in the Lnittiztaria community, making their 0.5) to the Laz~tiztarin commtcnity (73 spccics) far fidelity 1.0; so their rank (R) \\'as consequently escceded that for the other two communities (19 l-iigll. Several other species, including Bnlcis ran- species and 17 species for the illnrin-Hedoplzjrllzczz~ dol/~lti,Syllis (T)~pos)dlis)sp., and Desiosf,irn sp., and Ha1osnccioz~-Ftrcus communities, respectively). occurred frequc~ltlyin the Lamiztnria com~nu~lity Within the Lnzitiztnrin comnlunity species at Makarius Bay and IA-2, but data from these t\vo counts were greatest among the Polychaeta (30 localities are not complete for all species and \\'ere species) and the Gammaridca and Gastropoda (19 not included in the calc~~lationsfor the s~nall species each). Figures 5 and 6 sI~o\vthe distribu- species. If these data are considered, Bnlcis, cspc- tions and relative abundances of species in the cially, shows both high coilstancy in (0.73) and Lazztittnrin community for two transects representa- high fidelity for (0.80) the Lniniz~orincommunity. tive of an esposcd location [Rifle Ranze Point The sea urchin Strottgj~loceztbrott~spoljt- (T4)] (Fig. 7j and a sheltered one [square Bay ncnzttl~t~shas potentially the most important role in (T,)] (Fig. 8). structuring the Lnzitirtnrin community of all the The ten most characteristic invertebrates of the invertebrates found there. Descriptive- La~ztittnrin comm~unityare listed in Table 2 along comparative studies (Estes and Palmisano, 1974; with several statistics suggcstivc of their role in Chap. 22, this volun~e)of islands \\,it11 lolv (Am- commullity structure (see Fager, 1963). Species arc chitka and Adak) and high (Shemya and Attu) listed in order of decrcasi~lgrncatl rank (R) for urchin biomass indicate that this species call have a each size class (large-size class on top). They profound effect on populatio~lsof lorver intertidal include a sponge, four polychaetes, t~voechino- and shallow subtidal algae and probably other derms, ttvo gastropods, and an amphipod. invertebrates as well. These authors argue that Other species anlong the large-size class with o\~eresploitationof urchins by sea otters (Eztltydrn large rank (R) included O/~hiopholis nctrlecitci 11~tris)reduces considerably thc impact of the sea (10.5), Ilezzricin leuitcsculn (10.5), Aletcticrster urchin at Amchitka. They contelld that, in the sclzefferi (lo), i\'crlicn clnzisa (9.5), and Hnli- presence of otter predation, urchin individual size cltottdricr pniticea (8). The first four species shelved and popi~latio~lbiolnass arc reduced, permitting high fidelity for the Lroztiztnrin community (f = 1.0, benthic algae and associated invertebrates to flour- 1.0, 0.72, and 0.86, respectively), but they were ish. rare. The sponge Halichoztclrin p(rnicea sho\ved a Other species that may be important to the Iligh constancy (c = 10/15) for the Ln~ztiztcrria structure of the Lntzziztaria community are the community, but it also occitrrcd frequently in the starfish Le/>tcrsterins nlnslieztsis and L. alezrticcc (see Alcrrin-fIedo/~l~~~II~riztcommunity; therefore f was tasonomic note in the appendix). Preliminary small (0.53). feeding observations on these species in the field Three of the large-size-class species in Tablc 2 indicate that they are ge~leralistpredators that feed (Stroztg)~loce~ttrottrs pol)fcrcnzttl~us, i\'otocto~ten mostly on gastropods and peracaridan crustaceans scuirott, and Leptcrsterias alet~ticn)had comparable (personal observations; Palmisano, 1975). These abundances and did not differ greatly in average two spccics were not distinguished daring these ~blnt'ilref~luet'trb,ute$ ill Rocky hltotirlal Co~~llliii?~ities407 4 A Fucus d;$tishor W' Hoioasccionglondiforme tr iridaeo cornucopiac os Cobble 10.000 8 Hcdophylium scnilc ;-, Silt md rand '+10 0 3 Lominoria iongipcs *12,io c,iwo 6 - L- SPECIES 3'

Scypha compressat Acliniidaerpp. Sertubrella pinnata Ereooe fisvr, Syllir lTv~orvlli~l - rp.' Lum&inee;d ;nn>r, P CB~iierielir)>lam maculara - - Cimfulus Type IV - Asabeiiides sibirica Dexiospira rp.. - Tubficidaerp. - Phaseoiosomz ugmizii - - Ectopioma rpp.$ - Ophiophoiis aevieara Henncia tumida Alcuoasr6.i schefferi LwrarTeri~~o!eotico Srrongylocenrrorur ~D~~BCJJI~US Noroaemea xurom A1"inru aurivill;;' - Mirre!/a amianr;~ Neboiia rp. - Mvnna rrephenreni lschyrmeror Type I I - lschyios~rusType I Jam sp. Ponlogeneia rp. Mesomeropo rp. 0) - Oh$LD. - Caprelie kinc~idi Caprclia rp. cmops COmp8CIUS - Acarina sp. - swei* coiisced Srvoia 7 rp. Aplousobrvnchia rp. Aplid;um Type It - Aplidivm Type I1 t -

MLLW ------25 -3021 - - $5; '[ Ourdmt No. 26 23 28 27 24 16 19 17 22 29 20 18 Ousdrat mrni . - " . - -.. - . - .. . . ., . -. . .. .:...... 2 CC Yl L"= 0 > 123 I~~I1~1111~11~11~1~111111~~11(1~1111~111~~11i11~1i111~l~~1111~1~111~111~~~111111 m -30 -20 -10 0 10 20 30 40 50 HORIZONTAL DISTANCE DISTANCE. m Fig. 6-Distributions and abundances of species that were most abundant in the Laminaria community at Square Bay (T,), Amchitka Island, in August and September 1969. Abundances are numbcrs of individuals per 0.25 m2 except as noted. Centers of quadrats sampled on dates other than August and September 1969 are shown below the quadrat numbers of Auyst and September 1969. *, Estimates based on two or more subsamples of 19.6 cm2. t, Scale is in units of percent cover. :I,Presence data only. Illarii~e17tverte6rates in Rocky Intertidal Conti~cz~~lities409

I

Fig. 7-Rifle Rang-e Point transect (T4). (Photogaph taken by C. A. Silnenstadin hlay 1974.)

feeding obser\rations, so we do not know ho\v their Ceritl~iopsisstei,~egeri was rare at the five ilrai~l diets differ. Our observations are scanty, but study sites, but, because of its s~nallsize, it can be neither species appears to feed on barnacles as does easily overlooked in the field. Since it \\'as rare it the closely related L. hesactis (h'lauzey, Birkeland, did not sho\v high affinity with Hcrlicho~~clricr. and Dayton, 1968; Menge, 1972). Presu~l~ablythis However, C. stej~legeri\\,as foiind only in quadrats is because of the lo\\, abu~ida~lceof Bn1a111rs containitlg H. pa~ticecr;\\~llen seen in the field, the glnttdrtla and B. cnriosus at Amcltitka. snail \\'as almost invariably on or under the sponge. A fourth species that may be an important Another North Pacific snail, C. stef~lre~~sae,has occupier of space is Halicl~o~~driapnnicea, because been observed in association with II. pa~~icea of its e~lcrustinggrourth form, pote~ltiallylarge size (Bakus, 1966). In the British Isles C. t116erctrlnrisis (mean and range of percent cover \\'ere 12.6%, 1 to kno~vn to eat IIalicho~tdria aild other spo~lgcs 55% for all those 0.25-m2 quadrats in which it was (Fretter, 1951). found), and relative comn~o~lllessin the lower 1lri1-HeclopIl~tI Community. 'I'he intertidal area (c, 33/60), Itlteractiotls iilvolving Alcrrin-Iledopl~y~llo,, community ranged in vertical Haliclto~~clriainclude possible predation on it by distributio~l from -53.4 to +9.1 cm (-21 to the starfish Henricia ttrl~liclaand parasitism by the +3.6 in.) in extreme shelter (tlmnsect TI) to 0 to snail Ceritl~iopsisstejl~egeri. I-lo~ricia has been seen +lGO.l cm (0 to 63in.) in extreme exposure pressed against H. pn~~icecr;\\,hen thc starfish \\.as (transect T4). Hedophylltt~l~sessile usually domi- rcn~o\red,the sponge tissue beneath it \\.as found to nates this community in sheltered situations, but, be discolored. Mauzey et al. (1968) discuss the as exposure increases, Aluricr crisps becomes domi- possibility that another North Pacific congener, H. nant (Lebednik and Palmisano, Chap. 17, this Ieviuscrtla, may also prey on e~~crusti~lgsponges. volume). 111 exposed localities IIerlopl~)~l/u~~~is Fig. 8--Square Bay transect (T2).(Photograp11 taken by Charles E. O'Clair on May 26, 1974.) usually absent except well back from the sca~sard Bnln~z~rscnriostrs, x\ritll increasing esposure and edge of the benches in clcpressions, title pools, and tllerefore xvitlt increasing dominance of illaria. drainage chanilels (Fig. 9). illorin can be found in iIo\eever, tlicse differences did not justify the patclics at h2Iakarius Bay, the lllost sheltered establishment of two separate communities, one locality studied (see L.ebccl~iik and Palmisano, dominated by Ilerloph)~llrr~~tand one dominated by Chan. 17. this volt~ulle). fllnrin. Despite the fact that thc species of algae The total specics comnt in the Alnrirr- dominating this commutlit)~ changed 1vit11 the Hedopl~~~llrrnrcommunity \vas great (11.1 specics), degree of exposure, i~ivcrtcbmtespecics composi- but fc\v species (19) were faitlifttl to it. As in the tion was silllilar regardless of \\~Itichspecics of algae Ln~r~i~rnrincom111unity, specics counts were grcatest dominated. Only two specics of invertebrate prcs- ;Itlions the Polychaeta (35 species), follo~vedby cnt in lllore tllatl one quatlrat were restricted to the Gastropods (23 spccics) and Gammaridea (18 Ilerlo/~lz)~llzr111-d0111i11atedquadrats, and none \\,ere species). Tlte ciistribations and relative abundances restricted to /llnri(t-domi~iittcdquadrats. If differ- of species in tlic illnrin-Hrdophyllrr~~rcommunity ences in thc relative ab~u~dancesof spccics of are sho\\w in Figs. 9 and 10 for representative invertebrates bet\eccn Herloplr)rlln~~r-dominated localities, T, and T, , respccti\rely. c~uadratsand illorin-dominated ouadl.ats ;Ire consid- ered, there does appear to be ai increase in tlic Table 3 shows the ten lllost characteristic ahund;ulces of ccrtain suspension-feeding species, invertebrates of thc illrrricr-llerloplr)~llrr~~rcomma- such as mussels, ~\f)lti/rrs edrtlis, and barnacles, nit). and their statistics. This table cont;liits all the hlari~leInvertebrates in Rocky Intertidal Con~il~u?lities 411

Table 2-Statistics for the Five hlort Characteristic Species of ltwertebntcs of Two Size Classes froru tBe l.o,,,irtnrin Commueity* Fidclity Constancy - Average Species Taxollt (I.)$ (c.)P KF Abu~tdasee'* rank (i)tt Dominancrii Dispersio~~gp Vitality

I.urger Species Counted in 0.25.in2 Quadrats

Slro,~gyloce,~lrofer K 0.84 10 16 2, 1-10 1.4 1/15 kt = 0.6 Sires up to polyocnnfhtcrg (, (I, 1-10) Aggregated 30-mm test diameter present. dlilr~~lln G 0.73 13/15 15 12, 1-124 4.8 7/15 K, = 1.2 All stages nnzinnrir (16, 1-49) Aggregated present. ~Votoa~~~~co G 0.92 G/l5 14 1, 1-2 0.47 0115 Prcseet year SCILIIO1I (I. 1.2) r0"lltl. Scypho P 0.78 9/15 13.5 2.5%, 1.5'79 Present year con~prossn (3.2%. 1.9%) round. I.rplnrfcrirrr A 0.62 13/15 12.5 3.2,l-21 2.73 2/15 K, = 2.4 Ju\scnile and nk~sfint*** (2.5, 1-9) Slightly adult stages aggregated present.

Slnall Species \\'hose Ai,u~~dances\\'ere Estln>utcd from 50-8~1n1-Di;t~reterSuhsaznplrs of Quadrats PC 0.89 213 7 69,38-90 8.3 113 k, = 1.2 Adults present. (80,69-90) 3gregated PC 0.80 213 6 38. 1-167 7.0 113 KI = 0.9 ,\dulls prescnt. (98. 23-54) A~gregated PC 1.O 113 6 6 1.33 013 K, = 0.4 ,\dulls present. 9gregated l'c 1.O 113 6 2 0.67 013 K, = 0.68 ,\dulls present. Aggregated C 1.11 I I3 6 1 0.33 013 Random ,\dulls present.

*Format ewdificd after 'rable 2 of 1:;xger and hlcGowan (1963). ti\. i\nteruidea; E, Erhieoidc;t; C, Caprel1ide;i;G. Gar~ralloda;P, Porifera; PC, Polychaeta. i l'rvbich tl~cspecics war found and (in parentl~eser) there same statistics for qoadrats from thin community in which the specics war found. Abuodance of S. comprcrsn is the mean and mnse of percent rover per quadrat; other statkticr 1,;trrd on :~bundirnccare not included far ScgpAa. ttllanks for eaclb species \rere averaged ovrr 15 (large species) and 3 (snlall species) quadrats. The average ranks of the most ;~bonrlaotIcwge ;tnd sm;tll species in tine l.n,,rhtnrin qaitdr;ttr were 6.53 ;md 12.66, respectively (1, least abundant). i$Proportion of qe;gdr;tts fn,m tltir colnm~tnityie wl~ich tlre species was antong those making up 50% of tile individuals. Summ;ttiun is r;trlt quadrat brg;te wit11 tl~eamst ;tbuod;tet species. 3 \' K, . tbc first ;tpproaie~;ttior>of the cspoerot K of tltc negative binoniinl distribution. Overdispersion increarcs as K gocs to 0. 1)irpersion \v:.;ts jadged i-.todom if tile quotient of the sample \,zriitsce divider1 by thc sample mean war not sig~lificnntlydifferent from one; the relation ?;' = ~\s~I~\\.~~susPL~COT tests of ~i~nili~itnc~. (;(Seet;trotarmic anole ondrr S. polyrrrsr,fh!uin ;%ppenrlix. "*See t;taonosnic note attder I.. alt.atie-nin ;q~pendir. largc specics that are faithful to this coln~ni~liity Stephenson, 1972), frequelltly to tlie exclusiorr of and sliglitly inore than one-third of tlie sillall other species, such as barnacles and limpets (I.e\vis, species o\ving to the s~iiallnumber of faithful 1964, p. 95). U'hetlier 1\1. edtrlis conipetiti\~ely species. exclitdes these species has yet to be demonstrated. Spccies that \\re might predict to have impor- 11l)~tiltrsccrlifornia~~~rs has bee11 sho\vn to displace tant roles (as successful competitors for space) in other upper intertidal species \\,lien it is released structuring tlie Alaria-Hedophyllti~n comiiitnlity from predation by experimcnt;tl remo\~alof its Ltrc ~\lytillrsednlis and Bala~litscnrioslrs, although primary predator, the starfish, Pisaster ochraceus this proved not to be trite at Amcliitk;~.11t)~tilrrs (Paine, 1966; 1974). For at least oiic species, ecl~rlis is a \\vidcly distributed species in north fl~~t/ropleuraelegn~ztissi~~~u, Paine has observed teluperate and sitbarctic regio~isasid forms conspic- competitive exclusion by 11lytil11scnlifor~liu~tlrs (see uous belts in the midlittoral zone (Stephenson and Dayton, 1971). SPECIES

Haii~hondriapaniccat Plafyhelminfhes SPP. Nemerfined rpp. Nematode rpp.' Poiychaera rp.': Pholoe minum' Ereone ilava' syiiis rp. Syiiis ormilhiir' s~liiraltern~rj

C;rratulvr rp.' Ca~ll~rie~ladata ~JCUI~IJ Sedentaria rp.% Upiteiia opirsrs' Chone eincm' DemOnJx rp. Oligochaer8 rp. 1 I' C,i& 5o.t

Alioishc~fesIP. - - Paraiiorcheries ochorensis Amphichw rp.' - Ponrogeoeia maknrovi - Paraphoxus spinosur ' Oichorncne 7 rp. - - Parapleusm pugemensis' _C - - Sryeia 1 rp. Aseidiaeo ril. I - -

.- .. Fig. 9-Distributions and abundances of species that were most abundant in the Alario- Hedophyllum community at Rifle Range Point (T4),Amchitka Island, in August 1969. Abundances are numbers of individuals per 0.25 m2 except as noted *, Estimates based on two or more subsamples of 19.6 cm2. t, Scale is in units of percent cover. $, Specimens incomplete and unidentifiable. 5, Presence data only. .ran03 luaarad jo sl!un u! s! a1e3s '4 9'61 $0 salduresqns arour ro OM? uo paseq salemps3 ', .6961 raquraldas pue isn8nv $0 sraqurnu lerpenb aql MO[aq UMO~Sare 6967 raqwaldas pue isn;inv ueql raylo saiep uo pa[dmes slerpenb $0 saalua3 .palou se lda~xa,m 92.0 'ad sFnp!A!pu! $0 siaqurnu are sa3uepunq.b. '6961 raqura~daspue asn3nv u! 'puep~eyl!q~mv '(ZJ,) Legi arenbs le Lz!unucmo3 un?lXydopaJj -uuqv aql u! Iuepunqe lsom aiaM ley3 sapads 30 samzpunqe pue suo!lnq!rls!a--oT 33NVlSIO LU '33NVJSlO 1VINOZI~OH LU 05 00 OE Oz 01 n 01- 02- Lt11 ~~1~~~~'~~'~~~~~~~~~~~""~"~'~"""'""'""'1"111"'111"'1 ...... Y. -. . ."... . . -...... - s_l -62 44 -61 --12 OE -52 9' -vz -LZ , .. . , Ml1W

ill adAl ioP!!O!ldo!liQ - - gsuullJ6nbsJlSnuidO>Od - WOJO!l!9"03 OU!ON j2a-ii.S O,iod02uwOJOd 'dB sa>JnJldUAS .S!AW wn,qaoi02 .'ds aoq2!qOuV Z!IIJIOqW SO~ilr~~IDli~~Od - 'dr SOJwq>JOllV mso!~orro rnuoleg IrWOJ!U/~hS"l"JIi)MI - - !O$"C,Y e!uo,*OPO -- wnuc!qJJOW !Jam u"U!JJJg !ia63o!ars s!rdo!ul!,a2 __L. .'a5 O!isU!d,!A .8 '6s NP!OWU .!omcu oin6u!3 .os!>a,,oa!o 01n6613 ,qd/opuor ?>lW - s!su~~u!J~~w>lia6a6B1V 09!>!,EpA* OJaullalqord - ",,>"",,m,,,m Table 3-Statistics far the Five hlost Characteristic Species of Invertebrates of Two Size Classes front the illorin-Hedophylla,?~Coe>manily"

Fidelity Constancy Average Species 'Taxoltt (f.)? (c.)B R5 Abandunce** rank (gtt Dominance$? Dispersions 8 Vilality

Larger Species Cotlnted ill 0.25-tn2 Quahats

B 0.7 6/44 3.5 4, 1-40 0.50 1/44 kt = 0.66 Prcrent ycar (5, 1-40) Stroogiy round. a gegated Ci 0.67 10/44 3.5 4, 1-24 0.65 1/44 a, = 0.1 P,,,,,,, year (4, 1-24) Strongly round. aggregated hi1 benthic stayes. .prercnt. Juveniles and (1) adults present. I3 0.66 3/44 2.0 2, 1-3 0.07 0144 Kt = 0.2 Juveniles (1, 1-2) Strongly present. aggregated Ga 0.57 3/44 1.5 11. 1-26 0.18 0144 kl = 0.08 Jevet>ilesand (3, I-'!) Strongly adults present. aggregated

Small Species IYbore Abundat>cesWere Estimated fro11150-mm-Diameter Subsan,ples of Quadrats G a 0.62 5/12 3.5 412, 1-3937 2.92 0!12 k, = 0.16 (190. 1-2201) Strongly a~gregated G 1.O 2/12 3.5 6, 1-11 0.33 0112 KI =0.1 Jesenilcs and (6. 1-11) Strongly adu1:s present. aggregated Lt~o~bricillnrsp. 0 0.57 12/12 3 328, 11-2661 9.67 1/12 Kt = 1.3 Jureniien and (372, 100-1698) r\ggregated adults present. G 1.O 1/12 3 64 0.25 0112 Kg = 0.08 Adultr preresl. (64) Strongly aggregated PC 0.66 2/12 3 54, 1-64 1.08 0112 Kt = 0.2 Immature stager (59,5444) Strongly present. .ayyrer;tted

*Format tnodified after 'Table 2 of Faser and r\lcGowi$o(1963). t Am, hmphincura; B, Biualvia; Ci, Cirripedis: G,Gastropods: Ga, Gamolaridea; 0. Oiiguchaeta; PC, Polychaeta. $Proporlion of quadrats containitlg the rpecier that was in this cotnmunity corrected for differences it, san,ple sire between communities. §Proportion of quadrats from this community that contaieed the rpecier. q Average of ranks of fidelity and constancy for each specier. '*hledian and range of numbers of individuals per 0.25 nl' in ail quadrats in which tile species was found and (in ptrentherer) litere same statistics for quadrats from this comrnoeily in x\~hici>the species was found. it Rankr for each species were averzgcd over 44 (large species) and 12 (rmaii species) qoitdritts. 'The average moks of the most abundant large and small species in the Alnria-IleduphyNan quadrats \rere 7.02 and 13.67, respectively (I, least abuntiznt). t?Proportion of quadrats from this community in svhicix the specicr war itmoag those making up 50% of the individuals. Sarnma!ion in each quadrat began with the most abundant specicr. S B K, , the first approximation of the caponen1 K of the negative biz>omiai rlirtrii>utiun. Overdispersion itwrearer as K goes to 0. Dispersion was judged random if the quotient of the ramplc variance divkird by the sample mean war not rigeificaotly different from one; the relation x2 = nr2/kmas used for tests of significance.

At Amchitka s~nallto large (from about 15 mm nance is not suggestive of a successful ~i~onopolizer to about 40 mm* in shell lcngth) 111. ediilis showed of space. Holvcver, very sinall individuals of a high fidelity for the /Ilaria-tledoph~~lli~t~zcan- ~Vlytilits werc found to be quite numerous (see munity but occurred there relatively infrequently discussion of Haloscrccio,~-Flrcits community). and in low abundance. Its lox\, numerical domi- Dayton (1971) has sho\sn that, in the absence of predation by the snails, Thais spp., and the starfish, Pisaster oclrracetts, and of disturbance by *Shell length was measured from beak to posterior edge limpets, Collisella spp., Balntlirs cariostts compcti- of sllell. lively csclitdes other species of barnacles and I~tuertebrntesill Rocky Ilzterticlal Col~bt~ttrtbities 415 dominates space in the lowcr middle intertidal tropoda (20 sl~ecies)antl the Ganiliiaritlea (15 area. Indeed, B. cnriostls may escape the cffects of species). The distributions and relative abundanees liiiipets and Tllnis and eventually occupy 100% of of species in tlie iInlosaccio~t-Ftrcus co~ii~iiunity the space if tlie probability of being removed by arc slio\vn in Figs. 11 and 12 for Rifle Range Point drift logs is lo~v. atid Square Bay, respectively. Pisnster is absent from timcliitka, and only tlie Table 4 shows ten characteristic invertebrates sea otter is close to the starfish's ecological of tlie Ilnlosciccio~~-F~tc~fscommunity and their cqi~ivalcnt.ICenyon (1969) reports on tlie esamina- statistics. All the large species are l~crbivorous tion of sto~iiaclls fro111 475 otters killed at Am- gastropods. Two other species, Bnln~tttsgln~~dttln cliitka in 1962 aid 1963. Bnlr~tnscnrioslcs was not and Colliselln stri&ztelln, had tlie same rank (R) as reported from any of them. Norvever, Barabasli- Litto~i~lnnletcticn, but tlicir median abundaoces Nikiforov (1947) foiuid that B, cnriostcs remains were less. were occasionally present in sea otter fcces, and sea Bnln~tlcs glni~dr~lamnges from tlic Aleutian ottcrs havc been seen feeding on B, cccriosus in the Islands to Enscnada, Baja Calif., aiid is normally Aleutians (J. F. Palmisano, personal communica- one of the ~iiostabiuidant animals in the upper tion). The populatioii statistics for B. cnrioslis intertidal area (Ricketts et al., 1968), forming indicate that it is uncomnion, loxv in abundance, along \\,it11 other barnacles a distinct zone and rarely numerically dominant at Amcliitka (Stephenson and Steplienson, 1972). Yet, at the (Table 3). Yet at Shemya Island, approsi~iiately five ~iiainstudy sites on Amcliitka, the median [I 180 miles (300 k111) to tlie \vest, B. cnrioszrs is use the median here; it is a better statistic of more frecluent (60% of Alnria-Hedo/~l~)~lI~~i~tplots) location than is the mean ~vlien spccies Iiavc and more abiuidant (median abundance, 17; range clumped distributions (e.g., Fager, 1963)l number 1 to 77) per %, xii2 and forms a conspicuous belt of individuals of B. glntld~cln pcr 0.25 11i2 in in tlie lo\ver ~iiidtlleintertitlal area. Considering the quadrats in tvliicli it \\,as found was very low (3), data of i

'em3 9'61 30 sajdmesqns arom ro OMI uo paseq saletu!asa ', '6961 xaquraldas pm isn8nv 30 slaqurnu lerpenb aql Molaq u~oqsare 6961: ~aquralda~pue asn2nv ueql raqlo salep uo paldmes slerpenb 30 sraaua3 .palou se 3dasr.a Zm 5z.0 lad s.(enp!n!pu! 30 sraqurnu are sasuepunqv '6961 raqwa?da~pmls&nv u! 'puels~e~l!qsmv '(2~) iies arenb~le iil!unurmos sn3nd-uon -3vsolw~aqa u! luepunqe lsom araM ley1 sapads 30 sasuepunqe pm suo!lnq!ns!(l-21 ...... ~VLNOZIUOH 0s 0s OZ 01 0 01- 02- OE- YI k l"l'l',,'l,l',lll,II,'"1"',l,~,,~,,,,I,,,,I,,,,I,,~,ltl,,\,,,,~i,l,\,II,~II,IJ r: .$ t ...... %-..-. -..... - .. .. r . . rrazuw rmrpew A 81 62 ZZ Ll 61 LZOC -SZ 91 02 Lt 'ON LPJPCnD ------$2 g 2 ...... MllYY

-a, .-c .=4

- -- -

- oa!~~O!JOIV ~~~~~UOIOI~OY~'YD~- pur, lIIS :- all- UnllA~dOPH 0001 2 ulqq03 90 1P!d030~103 DDDPUl 11 DWJOjlpYBj6 UD!1m$OIBH 8- SOFlPSnlnjy Table 4Statistics for the Five Most Cl~aracteristicSpecies of Invertebrates of Two Size Classes from the IInlosnccion Ftlcus Conlnn~nity*

Fidelity Constancy - Average Species Taxol~t (f.)p (c.) S R(i Abut~dance** rank (P)tt Dominance$ $ Dispersion$ g Vitality

Lwger Species Coonted in 0.25-n1~Quadrats G 0.67 41/53 6.5 6, 1-844 2.79 11/53 IZ, = 0.1 Present year (6, 1-844) Strongly roond. aggregated Sipl~o,tnrin I<, = 0.3 All stages thersites Strongly present. aggregated KI = 0.4 Present year Aggregated round. Random Present year round. Present year round.

S~nallSpecies \Vhose Abi~ndances\\'ere Estin~atedfro~n 50.111111.Diamete Sobsan~ples of Quachats A Ci~rgulo G 0.57 7/11 3.8 234,2-3810 3.27 1/11 K, = 0.3 All benthic martyni (14,2-3810) Strongly stages aggregated present. Pnrn~fnis 0 1 2/11 3.5 190, 127-254 0.91 011 1 K, = 0.2 Adults littornlis (190, 127-254) Strongly present. a6gregatcd Cnl/iopiello(?) Ga 0.8 4/11 3.4 85, 24-345 2.91 1/11 KI = 0.2 Adults and SP. (89, 51-345) Strongly )loong aggregated present. r\l),selln B 1 1/11 3 127 0.27 0/11 KI =0.1 Adults nleuticn Strongly prescnt. azgregated Cirratulss PC 1 0111 3 64 0.36 011 1 K1 = 0.1 'l'ype 11 Strongly aggregated

*Format rnodificd after Table 2 of Fager and hlcGowan (1963). tU, Uivalvia; G, Gastropoda; Ga, Gammaridea; 0, Oligocl~aeta;Pc, I'olychaeta. $Proportion of quadrats containing the species that r\?as in this commonity corrected for differences in sample size between communities. 8 Proportion of quadrats from this community that contained the species. (I Average of ranks of fidelity and constancy for each species. **i\ledian and range of numbers of i~tdividualsper 0.25 1n2 in all quadrats in rvhich the species \\.as found and (in parentheses) these same statistics for quadrats from this community in rvl~iclltllc species was found. ttRanks for each species were averaged over 53 (large species) and 11 (slnall species) quadrats. 'l'lte average ranks of the most abundant large and small species in the Hnlosnccioii-Fucus quadrats \\,ere 6.02 and 8.91, respectively (I, least abundant). $$Proportion of quadrats from this community in rvhich the species was among those making up 50% of the individuals. Summation in each quadrat began with the most abundant species. 5 g k,,the first approximation of the exponcnt K of the negative binomial distribution. Overdispersion increases as K goes to 0. 1)ispersion \\!as judged random if the quotient of the sample variance divided by the sample mean \\-as not significantly different from one; the relation ,yZ = nsz/z was used for tests of significance.

benches. Sea stacks and vertical rock cliffs repre- \vard of aid above the benches. Even \\,hell nearly sent a smaller area of tlic intertidal region and \trcre optillla1 conditions for B. glo~dtrlrr arc sought out not salllpled c~uantitati\~ely.'Sherefore my sanlpliilg on the island, such as on vertical silrfaces in 111iry have underestimated the species abundance, csposed localities, a distinct barnacle zone is rarely but this is unlikely. Bnlnlftts glanduln is rare on the found, and even then individuals are scattered and cobble and boulder beaches, ~vhichexteild shore- do not form dense shcets as elsewhere in the I~~vertebrntesit1 Rocky Itttertidal Com~ttu~~ities417 eastern North Pacific. There seems little doubt tliat energetically feasible for Tltnis to begin preying on lo\ver B, glat~dulaabunda~ices are a real phenome- B.glat~duln, may not be very high since barnacles non at Amchitka. re probably easier to capture and handle than Why are B, glat~dttlaabundances low at Am- li ttorines. eltitka? One might expect that, since the i\lcutians . Dayton (1971) concludes from his studies that are near the ~iorthernli~iiit of B, glandtcla, popula- limpet disturbance alone cannot completely limit tions at Amchitka may frequently suffer \\,inter kill barnacle populations but does reduce barnacle or poor reproductive success in spring oxving to low recruitment. In the presence of both limpets and temperatures. Ho\vever, if we compare the long- Tlrais, recruitment and survival of B. glattdttla are term data on water temperature in tlie vicinity of virtually nil. At Amcliitk;~two species of limpets, A~ncliitka (inset, Fig. 3) wit11 that for Kodiak Collisellr digitalis and C. striyntella, sho\v high Isla~d,Alaska (Nybakken, 1969, Fig. 5), ~vhereB. fidelity for the Hrrlosaccio~t-Fttctis commnnity. glattduln maintains populations, we see that the Collisellc d~itnlis\\,as rarc in the IInlosacciott- minimum water temperature recorded in the vicin- Fttctts con~munitybecause it rvas near its lo\ser ity of Amchitka is higher than the mean water limit of vertical distribution. Colliselln strigatella temperature 'in the coldest ~iionthof the year at occurred somewhat more frequently in tlie Ilalo- Woman's Bay, ICocliak Island. Similarly, comparing saccio~t-Fttc!rs community (e = 19/53) than in the air temperatures between tlie two islands, we find Alarict-IIedopl~)~llur,t comnicmity (c = 1 1/44), but tliat the average monthly temperatures recorded at it \vas relati\rely uncommon in both com~iiunities. the ICodiak Na\d Station in winter and early spring A third species, C. pelpa, also occurred in the from 1942 through 1956 are about the same (29 to IIa1osctcciot1-Flcclts conimunity, but it teas more 3G°F) (Kieger and H1underlich, 19GO) as those common and more abundant in tlie Alaria-Hedo- recorded at Amchitka from 1943 through 1948 1ll1community. Dayton's exclusion esperi- (31 to 3G°F) (Armstrong, 1971; Chap. 4, this ments included the above three species as rvell as a volume), but the record lows at Kodiak (-5' to fourth, i\'otoacttzect sctctnnl. At Anichitka I\'. 10°F) are much lower than those at Amchitka (4" to sc~itttt~trarely occurred in the Halosncciotz-Fucns 20°F). Further, large populations of B. glat~dtclaare community. Densities for these four species ;it present on islands to the east and \vest of t\m- Amchitka are, for the most part, much lower than chitka (Adak and Shemya). The alq>arcnt tolerance those recorded by Dayton. Nevertheless, C. digi- of B. gla~tdtrlato a wide range of environmental talis, C, strigatelln, and C, pelta may play an conditions indicates that it is imlikely that such important role in liniiting barnacle recritit~i~entat factors as Ion, salinities or loxi. oxygen content in Amcliitka. the water are limiting Balantts populations at Amchitka. Sublittoral Invertebr;~tes Btclnnus cariostts populations are also in low A variety of semiquantitatise sa~nplingnieth- abundance at Anichitka; so conlpetitive exclusion ods rvcre used to collect sublittoral invertebrates of B. glat~d~tlaby B. cctriosus is ruled out. The and to assess their populations. These tilethods results of previous studies of the effect of other have included 215 shrimp trap sets, 37 crab trap species on B. glnndula populations (Conncll, 1970; sets, 26 ~ninnowtrap sets, and 11 ampliipod trap Dayton, 1971) suggest tli;\t predation by Tltais sets. In addition, I tnacle a limited number of direct lit and/or disturbance by limpets of the genus observations (23 scuba dives) of sublittora I mverte-' Collisella and perhaps other herbivorous gastropods brate communities to a depth of 34 111 (112 ft). nlay be liniititig B. gl~tt~dtcl~cpop~~l;ttions at Am- Sampling of conniiunitics below 34 111 (1 12 ft) to a chitka. Palmisano (1975) has found that 93.6% depth of 183 m (600 ft) was acconiplished by (N = 106) of the diet of 1: littta on Amchitka dredging \\~itIirock and pipe dredges (10 hat~ls). consists of Liltorha spp.; only 2.8% consists of Sublittoral and neritic invertebrates \\,ere also barnacles (B. glattdttlc and B, cariostrs). I have collected incidentally during sampling for fish \\,it11 madc very fe\v feeding observations of T. Iittza mid\sater and otter tralvls, bottolii gill nets, tram- (N = 7), but on only one occasion did I see Tllnis tnel nets, and long lines by Fisheries Research feeding on a barnacle (;I young B, cariosus). Tl~nis Institute biologists. Bottom longlines xvere particu- may inaintain its popt~lation size by preying on larly effective in capturing Ptilosarctts sp. Plank- littorilles and switch to barnacles then B, glrttdala tonic organis~nswere collected with a plankton recruitment makes it energetically tvortli\vhile for pump. The appendix lists those sublittoral and finis to cio so. The thresliolcl of barnacle density, ~ieriticinvertebrates which have been identified. above ~vhichthe time required for Tlzrris to search Sampling of sublittoral invertebrates with traps for its prey is significantly reduced and it becomes \\,as patchy. Traps were set in the depth range 1 to 61 in (3 to 200 ft) and almost exclusi\~ely in 1\111scrilusver~~icosrts, He~zricicr sp., and Pteraster sp. Constantine Harbor 011 the Bering Sea side of the Occasionally the sinall crabs Cailcer orego~~e~lsis island and in Makarius Bay on the Pacific Ocean and Der?lrcctzrr~tsriln~ldtii could he see11 in pits and side; most \\.ere set in Constantine Harbor. The crevices in the rock surface. At the base of these traps \\.ere both species and size selective, presum- blocks, shell and rock rubble usually accumulated, ably selecting species the foraging behavior of anlong which the tubes of Clmetopterlrs variopedn- which includcs scavenging and which are in the size ~IIScould be seen. range that can enter yet be retained by the traps. All the dredge hauls were made on the Bering The species that was most numerous and most Sea side of the island. Of these, one-half were nlade frequent in aud on the shrimp and crab traps was in the outer sublittoral zone on the insular shelf Stro~tg)~loce,~t~ot~tspolyacu~~tl~ns. Other species aild one-half \\,ere made in the upper bathyal zone. frequently collected included Fusitrito~lo~ego~teil- The appendix lists those species froin the dredge sis, Leptasterias spp., and BL~CC~IIILI)Ispp. hauls which have been identified along with the Traps set near the nliddle of Constantine depths at \vhich these species were caught. Harbor on a sandy substrate (Fig. 4) consisteiltly caught Cltio~~oecetesbairdi. A total of 78 crabs Zoogeography ranging in carapace \\idth from 24 to 77 mill \\,ere The Aleutian Island archipelago falls \\'ithill the recordcd from this area. Thc size of thcse crabs Aleutian province, a nlarine biogeographic subdivi- indicated that they were young; nevertheless, 20% sion of the Pacific boreal region. The Aleutian of the fenlales were gravid. The presence of adults province extends from Dixon Entrance off British in Constantine Harbor is not precluded by these Columbia to Nunivak Islaud at about 60°N latitude trap data, hut the largest of these crabs was \veil (Briggs, 1974), but its \vestem liinit has never been belo\v the size selecti\~ityof the traps. Relatively precisely established. It has long been recognized sl~allow bays, such as Constantine Harbor, may that the Itunicates. The fi~vally (7) the "endemic" clc~nentconsisting of stalked tunicate Sl)lela clavata and various spe- those species restricted to the Aleutian (and .cies of Actiniaria, including ~bletridian~sp., \irere Commander) Islands. This definition of "endemic" often munerous. Common motile species included refers to a much more linlited geographic area than ~Vlargarites sp., Dicrdora sp., dorid nudibranchs, previously considered (O'Clair and Chew, 1972), ~\lrrriizeI~ivertebrates ill Rocky I~lte~ticlalCo1~~1,iiinities 421 Table 5-Number and Percent of Species of Intertidal and Sl~allosrSublittoral* Invertebrates of Tllree hlajor Taxonon~icGro~ips at Amchitka Island and T\vo Groups in tllc Conlalander Islands Distributed According to Their Zoogeographical Affinities

Zoogeographical elenlentst North Nortll kctic- l\'idely Ende~nic American Asiatic Pacific Boreal Antphiboreal distributed Total Si: % St % St % sg % St % s; % St % -s;

Amchitka Island Polycbaeta 1 5.9 4 23.5 2 11.8 2 11.8 3 17.6 0 0 5 29.4 17 hfoll~~sca 3 7.5 10 25.0 5 12.5 15 37.5 6 15.0 0 0 1 2.5 40 Crustacea 1 3.3 10 33.3 6 20.0 12 40.0 0 0 1 3.3 0 0 30 Commander Islands Polychaeta p 4 14.8 2 7.4 2 7.4 1 3.7 10 37.0 1 3.7 7 25.9 27 h~lollascaq 1 3.1 2 6.2 7 219 14 43.8 8 25.0 0 0 0 0 32

*Restricted to species that are not found below a depth of 60 m. tSee text for geographical areas inclitded in zoogeographical elements. i: S, number of species. g Compiled from Annenkova (1934), who esamined material from Bering Island only. q Compiled from Da11 (1884), Barabash-Nikiforov (1947), Skarlato (1960), and Golikov and Kussakin (1962). reducing the endemic element considerably. For that have been identified \vhether or not species the purposes of discussiou of the relative contribu- with broad depth distributions are included. tions of nlajor faunal source areas to Amchitka, the The anlphiboreal clemeut of the invertebrate present definition is preferable because it restricts fauna of Amchitka is either very small or nonesis- "endemic" to "insular" otlly. tent, depending on the tasonomic group (Table 5). Table 5 sho\vs the zoogeographical affinities Except for one crustacean, all species found on of three major tasonomic groups of Amchitkan North Atlantic and North Pacific shores also range invertebrates. These three tasa leere selected for into the Arctic. These arctic-boreal species consti- illustration because they \\.ere the most abundantly tute fro111 0 (crustaceans) to 18% (polychaetes) of represented macrofautla. The distributions of all the shallow-water fauna of Amchitka. It is note- the identified intertidal and shallo~\rsubtidal (to a \vorthy that, for those groups for \vhich \ve have depth of 60 m) species of these three groups xvere comparable species lists from the Commander used to construct Table 5. Only intertidal and Islands, the arctic-boreal element on the Corn- shallolv subtidal species are considered here for the mander Islands is consistently greater (37% for purposes of the discussioxl of the relative contribu- polychaetes and 25% for mollusks) than that on tions of Asia and North America to the fauna of Amchitka. Amchitka. Species that do not "vie\\," the Aleutian For thc three taxonomic groups included in Islands as islands, LC., those with very broad depth this discussion, bet~vecn12 and 40% (depending on distributions, are cscluded from the discussion. the group) of the species restricted to thc North Thc depth liinit !\,as arbitrarily set at 60 m because Pacific arc found on both Asiatic ;111cl Nort!? this depth is shallotc~erthan all the passes separat- Anlerican shores. Of greater interest are the re ing major island groups in the Aleutians as well as mailling North Pacific species, whose ranges esteila many of the passes separating islands xvithin these either to Asiatic or North American shores but not groups. The distributions of spccies included in to both. Table 5 are noted follolving the species name in the appendix. The follo\ving sources were used to Asian 1,s. North American Faunal Contributions compile lists of species from the Commanrler Islands to obtain the percentages used in Table 5: Amchitka lies mid\vay bct~seenthe tip of the Annenkova (1934) (polychaetes) and Dall (1884), Alaskan peninsula and the easternmost point on Barabash-Nikiforov (1947), Scarlato (1960), and larthropods in the \vestem Aleutian Islands (Fig. 1). As Dall (1884) and A1euti;tns (Lindroth, 1963), ;md in tlie terrestrial Ekman (1953, p. 156) have suggested, this current flo1.a of the islands in tlie vicinity of Amchitka: of probably acts as a b.~111icr -.' to faunal eschange the 33 species of vascular plants occurring in the behveen nortlin~estemiin~crica and northern Asia. Kat Isl;~ndgroup (of 1\4lich Amchitka is a member) [Briggs (1974) surlllises that the "deep" betrveen \vllose rangcs estend to continental Alaska or Kamchatka and the Aleutians is a barrier, but it is Iwsis never completely excluded except tain populations on Amchitka when we consider perhaps in very sheltered areas. \Vhether 111. cnli- the follo\ving? First, for marine benthic organisms fortzinrntcs at the edge of its geographic range could oceanic islands like Amchitka can be thought of as be excluded by 1l.I. edztlis from an island that tnainland islands (MacArthur, 1972) in the sense contains no habitats as exposed as those on the that dispersal over water is much less hazardous for outer coasts of California is a matter only for them than it is for such organisms as birds or speculation. insects. Second, all the species above have bee11 The low abundance of Bnlarttis cariosus and recorded from mainland Alaska and on large B. glnnd~~lnon Amchitka may support but one islands near the Alaskan peninsula (Pilsbry, 1916; species of predatory snail. Why this snail should be Oldroyd, 1924; 1927; Rathbun, 1930) and could Thais 1i11za and not T. 1a111ellosamay have to do presnmably benefit from the Alaskan Stream and with tlte former's adaptations. Thnis liwzn is very stepping-stone islands east of Amchitka to increase similar morphologically to T. errlnrgi~tatn;in fact, their immigration rates to Amchitka [Cryptocltito~~ Icincaid (1964) considered the two species to be stelleri, Clathn~rtnltcs dnlli, and Tel~rlessusclieirn- gorzus have been reported from Icamchatka (Spassky, 1961)j. Finally, all the species above *This latter assun~ptionis almost certainly true for the except T. lnntellosn have planktotrophic larval intertidal species !\I. cnliforninsas, T.ln~sellosn, and C. dnlli. development and consequently good dispersal \Ire cannot be completely certain that it ltolds for the polver. These species ~vouldtherefore be expected sublittoral species C, magisfar, T. clreiragonss, and C. to have rather high Aleutian immigration rates. In sfelleri because our sublittoral sampling was not as exten- sive as our littoral work (see tbe appendix). If populations fact, we have collected one i~ldividualeach of C. of these species do exist on Amchitka, their population rnctgister and M, cnlifor~tictr~ztson Amchitka, ~vhich levels ae probably so low that tbey run a high risk of indicates that immi~rationis occurring, but exten- extinction fro111 random pop~rlationfluctuations. synonymous. Tlrnis lil~rcc and T. el~tnrgi~zatnarc selves (limpets) and ~\rouldbe subject to starvation, small con~pareclto other inembers of the genus, desiccation, and predation. and both possess a comparatively thin sltcll, a large 2. Allii~ralsxvere probably crushed by falling opeiling, and a short spire. Connell (1970) suggests boulders and rubble overturned or propelled tip- that these adaptations allow T. e~~rn~gi~ratccto ward from the bellch surface \vhen the shock \\rave persist at upper shore levels provided that its main passed. food supply, B. glntrcl~clcc, is vcry dependable. At 3. Immersed organisms \\,ere subjected to low shore levels the thin shell of T. e~l~nrgi~rntn overpressures that may have been as high as 100 psi makes it \rulnerable to predation by crabs that live (Arlerritt, 1972) follo\ved by underpressures and there, \vhercas T. la~~tellosa,\vhich has a thicker cavitation. shell, avoids this predation (Co~lnell, 1970). On Amchitka, crabs inhabiting the intertidal zone are These immediate effects alone \sould not have Itad De~~~tnttcrttsatii Hnl,nloguster grebnitrkii, a profound influence on co~nmtinitystructure. Pag~~rrtshirsrltircscrclns, Cn~tcer orego~te~rsis,and A more important source of mortality to Pupttin g~ncilis.All thesc species are small and in illvertebrate populatioils on those parts of the lo\\' abiindance* and it is unlikely that ally of them intertidal benches near the bases of cliffs md sea prey on Tl~nis.Tlzccis linrn may therefore be free of stacks was burial by rockfalls and turf slides. crab predation at lo\\, shore levels on Amchitka. If, Althouglt the total volume of these falls and slides in addition, T. lir~rn,because of its sinaller size and has been estimated by the U. S. Geological Survey thinner shell, can subsist on a lo\ver rate of intake to be 35,000 m3 (ICirk\\~ood,1974), most of this of food than T, la~~zellosn,as Col~ncll(1970) has material was deposited on the beach shorelvard of suggested for T. enrnrgi~rntn1,s. T. lnn~ellosn, then the intertidal benches and affected intertidal organ- T. li111cc may be able to esclude T. lan~ellosn from isms only in localized areas (Lebednik and Palmi- lolver shore levels. (Ho\vever, there is no docri- sano, Chap. 17, this volume, and personal observa- mented e\ridence of competitive esclusion among tions). At the edges of the benches, bench spa11 thaids.) resulted in large chunks of rock and accompanying Anotl~er possible cause for estinction of organisms pluilging from the edge of the bench inariile invertebrates on Amchitka is predaticn, into the sea (ho\ve\~er, some of this material especially by sea otters (Eirlryrlrn lntris) (see probably \vashed onto the bench surface). The fate Palmisallo and Estes, Chap. 22, this volume). Size of animals not killed in these rockfalls is utlknown. selective predation by otters on M. ccrlifo~~zia~tus, hlost of the evidence acc~umulatedso far indicates \vhich grows larger and matures later than ~bl.ediclis that intertidal ailimals are excluded from the (Wargcr, 1972), may contribute to the former sublittoral area by predators and competitors species being excluded fro111 Amchitka. lio\\,evcr, (Connell, 1972). Intertidal in\~ertebratesthat sur- ilI. cnlifo~~rin~~usis also absent from Shemya Island vived the plunge into the sublittoral area probably where sea otter predation is negligible. Sea otter \\rill not be able to maintain popitlatiolls there. predation inay also be responsible for the cstinc- Long-Term Effects. All the disturbances dis- tion or near estinction of C. ~lrngiste,;T. clreirrl- cussed above, although by no meails inconsequcn- golrns, aid C. stelleri populations on ilmchitka. tial to intertidal invertebrate communities, are relatively trailsitor)? in ecological time; i.e., though Effects of Cannikin mortality inay be high as a result of these Immediate Effects. The til~derground disturbances, recoloilizatio~l is rapid compared nuclear test, Canilikin (No\,. 6, 1971), produced ~viththe generation time of most of the species several effects traumatic to intertidal in\rcrtebratcs. involved unless the habitat is made ~ulsuitablefor Innnediatc effects associated nrith the passage of recolonization. The most disruptive and lasting the ground shock wave (Merritt, 1972) could have disturbance to intertidal communities resulting caused mortality in the follonfing \\rays: from Cannikin was the pcrmanellt uplifting of the 1. Some invertebrates dislodged or over- Bering Sea coast near the site of Cantlikin. Detect- tunled by the shock wave xvould bc unable to able uplift in the range of 3.4 cin (1.3 in.) to 1.1 m reattach (barnacles and ascidians) or right them- (3.6 ft) Isas recorded along at least 7.8 km (4.8 miles) of coastline (I

SEVERE Figures 14 and 15 sho\\. the changes in mean species coullts with time following Cannikin for three intertidal communities at IA-2 and IA-3, MODERATE respectively, compared \\(it11 R,lakarius Bay (con- "I- 0 trol). The plots in the H~~1osnccio11-Ft~cttscommu- 3 nity at IA-2 \\,ere divided into t\vo groups. The first 5 DETECTABLE group contains plots nearer the sea\vard edge of the 4 bench (outer plots). The second group contains iplots nearer the shorex\~ardedge of thc bcnch a'ivay from the area of frequent \vetting by wa\re splash and salt spray \\'hen etnersed (inner plots). ALONGSHORE DISTANCE. krn IA-3 and AIakarius Bay arc relatively shcltcred, and all plots at these sites \\'ere subject to the (a) same degree of exposure; therefore outer and inner plots \\,ere not distinguished. All species on the plots that could be distinguished arc inclncled in Figs. 14 and 15. Species of Platyl~elminthes, Nemertea, Oligochaeta, Nematoda, and Ectoprocta \\'ere not distinguished. Poiychaetes, except ereis is pelagica, Desios/>irn sei~tide~~tatn?,and Desios/)irn sp., and gammarids, escept Parallorcl~estesocllo- te~tsisand Orcllestia sp., \irere lulllpcd into Poly- cllaeta and Gammaridca, respectively. Actiniaria \\,ere di\rided into species that \\,ere and tliat were not lnembcrs of the Family Actiniidae. Specics counts at IA-2 began declining in all communities immediately after Cannikin (Fig. 14). From the beginning the greatest source of mortal- ity of invertebrates not crushed or buried by rock and soil \\,as prolollged elnersion due to uplift, especially for the sessile species, such as sponges, sedentary polychaetes, bi\ral\res, and ascidians. Solne motile specics at the seaward edges of tlte benches \vhich \\,ere not thro\vn into the water Fig. 13-(a) Distribution and relative magnitude of with the passage of tlte shock may have algal die-off due to Cannikin along the section of migrated to lonrcr le\rels soon after icplift. Preda coastline shown on the map in part (c). Detectable, tion by gulls and oyster catchers may also have from one to a few species showing die-off. hloderate, bcen an important source of mortality in the first a few species shorving extensive die-off (more than few days follo\ving uplift. During this period large about one-third of the total population). Sesere, all abundant species showing extensive to complete flocks of gulls (Lartrs glaucesceits) and oyster die-off. (Data from Lebednik and Palmisano, Chap. catchers (flaeinntop~rsDncl~11ra11i) wcre seen at IA-2 17.) (C. Simcnstad, personal communication). Also, a (b) Vertical displacement at several locations large number of overturned aucl empty linipet along the same section of coastline as part (a). (Data shells (CoNisella spp.) \\,ere observed on the bench. from Kirkr\.oad and Fuller, 1972, Table A-5.) (c) Map of a section of Bering Sea coastline IIow ~iluchof the activities of these birds can be adjacent to Cannikin surface zero (SZ). Points along attributed to predation and how much to sca~~eng- the shore that are depicted in parts (a) and (b) are ing of dcad and dying anitnals is not kno\vn. directly above the appropriate locations on map. The fiftcctl plots established at In-3 wcre uplifted from 47 to 56 cm, depending on the plot (Table 6). Only one plot isas uplifted above cs- The 25 plots initially established at 1A-2 \\,ere trenie high \\rater of spring tides. However, prob- uplifted from 75 clli to 1.1 111 (3.6 ft), depending ably bccause 1A-3 is more sheltered than hi-2, on the plot, as a result of Cannikill (Table 6). mean species counts there declined similarly to but Sixtcen were uplifted above extreme high water of less drastically than at 1A-2 (Fig. 15). The sole spring tides for the Bering Sea coast of Anlchitka csception to tllis trend \vas plot 1. (146 cm in 1969); four were lifted above mean Plot 1 (Table 7) was established in the Ln~lti- high water. Five plots were buried by rockfalls. ~iariacolnlnunity pre-Cannikin. It had been up- Table 6-Physical Survey Data for 0.25.m2 Plots at IA-2 and IA-3 Before and After Cannikin*

Elevation, Elevation, Upward Upward Clll Clll displacement, displacement, Plot Before After c tn Plot Uefore After CI~

IA-2 1 40.7 150.7 110.0 2 57.0 157.7 100.7 3 51.4 151.0 99.6 4 41.8 139.6 97.8 5 139.1 234.0 94.9 6 68.8 Buried 7 15.4 Buried Buried 145.6 229.5 177.7 99.1 138.5 146.5 -. .. .- 171.8...... 16 69.0 Buried 8 43.0 90.6 47.6 17 33.4 117.5 84.1 9 53.3 100.5 47.2 18 66.6 153.5 86.9 10 44.8 92.2 47.4 19 99.8 189.5 89.7 11 59.1 Buried 20 66.0 156.4 90.4 21 20.0 Buried 22 47.5 122.2 74.7 23 47.5 125.6 78.1 "Compiled by P. A. Lebednik from Holmes and Na~verand Fisheries Research Institute survey data. tPreevent elevations of plots 26 and 27 \\.ere back-calculated assuming that each plot \\.as uplifted equally with the nearest origiual IA-2 plot.

lifted 54 cm to an elevation of 70 an. Species The total tlunlber of species seen preevent in connts on this plot at first declined after Cannikin each comnlunity decreased steadily after Cannikin but increased again by 3 lliollths post-Cannikin. to relatively stable lescls within 1 year (Fig. 16). Part of this increase was an artifact in that species, Conversely, the total number of new species not such as Actiniaria spp., Desios/)ircr sp., Plzascolo- previously recorded on plots in each of the three so11in agassizii, and Bolcis rcrlzdolplii (Table 7), were commtunitics increased. The ~ic~vspecies consisted probably present pre-Cannikin but \\,ere missed in chiefly of species from upper intertidal levels, the dense La?~ihznrinholdfast cover. Also contribut- especially the supralittoral fringe, and included ing to the increased species count on plot 1 at Collembola, Diptera (Salo~derinn~o~inus and Porn- 3 months was the addition, to the rctnllallt popula- clu~zio aloske~isis),Acaritta (~Veo~?tolgcislittorcrlis, tions of species of the Lo~~ziiiariacomnlunity, of Pa~ositztssp., ctc.), Ligici pallnsii, and the gammarid species comlnon to higher levels in tlie intertidal Orcl~estia sp. The only intertidal species that area, such as Litto~i?iaspp., ~vhicllhad migrated persisted throughout the year on the uplifted plots into the plot t\~hcn their fornier habitat was \Irere tlie gastropods Litto~i~laspp. uplifted above the intertidal area (Table 7). By The control at Makarius Bay sho\sed a slight 7 months postevcnt, the character of tlie fauna on decrease in mean species count per plot in Febrn- plot 1 had changed from Lomilintin conlmunity to ary 1972 (3 months postevent) and then a return a fauna contai~lillgspecies \vitli broader vertical nearly to October 1971 le\rels by t\ugust 1972 ranges and species of higlter intertidal levels. As \ve (9 months postevent) (Figs. 14 and 15). Ho\\.ever, \\dl see later, IA-3 plot 1 is the only plot at either the decline was only 23 to 24% of the October IA-2 or IA-3 to shale signs of recolonization by 1971 levels, depending on the community. Tlte intertidal organisms after 33 months postevent. decrease by February 1972 at IA-2 varied from 3 11kriite I~~vertebrcrtesit? Rocky Iiltertidcrl Contiit~c~tities 427

Hai~s~cia,~-F~~u~inner, L...... ""..*...... Hslosccion-Fuces outer. b-4 Alar-Hd110- - -O La",inaria.

70171 72173 73 174 CANNlKlN TIME. months

Fig. 14-Mean species counts for each algal zone before and after Cannikin at IA-2 and Makarius Bay (control). Asterisk indicates a sample size of 1. All macroscopic species at I,\-2 with distinguishable individuals are included. Anemones, polgchaetes, and gammarids, which could not he distinguished to species in the field, were lumped into their respective taxonomic groups. At Makarius Bay all identifiable species with distinguishable individuals retained by a 1-mm sieve are included.

to 100% of the September 1971 levels, depending Hulosnccioir-Ftrctts aiid the Alciria-Hedopl~ylltrii~ on the community. The decrease at IA-3 was 53% comn~unitiesshowed increases in the mean i~umber and 58% in the Halosciccioit-Fuczrs atld /Ilaria- of individuals per plot, especially in April and June Hedo/~hylluntcommunities, respectively. The only 1972. At IA-3 the Laiitiitaricr and /llaria-Hedo- Laii~irznria community plot (IA-3 plot 1) sho~ved phylluiit plots showed increases in the mean an increased species count of 36%. I attribute the ili~nlberof individuals of comparable magnitude in decrease in illen11 spccies count in February 1972 August 1972. Both of these increases are ac- at A,kikarius Bay to seasonal changes in species counted for by increases in the abundances of composition. The I\,lakarius Bay grid was not Littoriitn spp., especially I>. crtkaita. Further, litto- sampled at 17 and 30 and 33 n~onthspostevent rines too sinall to be assigned to species in the field because of inadequate field time and poor tides, also sho~vedillcreases in April, June, aiid August respectively. The grid was sampled at 22 months, 1972 (Figs. 19 and 20). but the data are not incl?rded here. General The drop in the mean number of indi\iduals observation of the grid at these sampling times did per plot in the H(r1osacciott-Fucns communit)t at not reveal obvious departures from the normal h,Iakarius Bay is probably seasonal (Figs. 17 and pattern of seasonal change in the intertidal ~011111111- 18). Halosaccioil gla~tclifor~itedominates this corn- iiities at h'lakarias Bay. munity at R,Iakarius Bay and sllon~sseasonal die-off The inean niunber of indivitluals per plot in all in late fall and xvinter \\,llich extends to late spring communities at IA-2 and IA-3 generally decreased (Lebednik and Palinisano, Chap. 17, this voliun~e, throughout the ycar follox\~ingCannikin (Figs. 17 and personal observations). The drop in the ~ILI~I- and 18, respectively). However, the decline was ber of individuals may be clue to the loss of the erratic in some communities. At IA-2 the outer moderating effect of the algae on the iilicroclinlate - - - - 25 30 Cammunity - "'..~...o" Hdlosa~~io,?-Fu~o~.&...... A - 20 Afarib-Hedophvllunl.D- - -0 - Laminaria, - 10 MAKARIUS BAY (CONTROL1 20 - - t- - Z 3 - - 0 - - 0 - - - ", - - IA-3 - 5 - 10 - 'n* - b' b' - 5 - - - -a- a - - ...... - - &.. ..A -

0 1 *" -15 -2 1 0 2 4 G 8 10 12 17 22 30 33 70171 71172 72 173 73174 CANNlKlN TIME. months

Pig. 15-Mean species counts for each algal zone before and after Cannikin at IA-3 and h'iakarius Bay (control). Asterisk indicat~sa sample size of 1. All macroscopic species at 1A-3 with distinguisltable individuals are included. Anemones, polychaetes, and gammarids, \vhich could not be distinguished to species in the field, were lumped into their respective taxonomic groups. At Makarius Bay all identifiable species with distinguishable individuals retained by a 1-mrn sieve are included.

of the invertebrates and perhaps to a reduction in F~cclts con~~llunitiesat IA-3 in h'lay 1974. Large tlie spatial heterogeneity in this co~ll~llu~lityin numbers of littoriiles, especially L. ntltn~~a \vin ter. (Fig. 20), on sotlie plots kept the meart ~lu~liberof After 1 year post-Cannikin, tilea~l species individoals relatively high in the fornler Ala~in- cou~ltsper plot at I/\-2 ruld 111-3 remained in the Hedophyll~oztconnnunity at IA-3 in August 1974. range 0 to 4 (Figs. 14 and 15, rcspecti\rely). The i\,Iost of the invertebrates observed on the 1A-2and total uu~nberof species for~llerlypresent in each IA-3 plots after 1 year postevent were in rnoist community before Calnliki~iat both IA-2 and IA-3 depressio~lsand crcvices or beneath pieces of dead never csceeded sevelt in those com~~lu~iitiesduring kelp rvhich \\,ere tossed up on the benches during the period of 12 to 33 1110ntlis posteveut (Fig. 16). storms and n'hich had bccome hung up on the nails Counts of i~idi\,idoalsalso remained low duriug this markirtg the plots. Other species, such as the flies pcriod, seldonl excccditlg a mean of 10 indi\kiuals S(t~o~rlel.ici~~~ctri~l~rs and mites ~\~eo~i~olg~rslittoralis, per plot at IA-2 (Fig. 17). In May 1974 (30 months mo\red through the plots very rapidly. postevent), for~iierH(t1osctccioz1-FIICIIS community 1A-3 plot 1 \\,as the only plot at either 114.2 or piots at thc sea~varcledges of the benches at IA-2 1.4-3 that showed sollle signs of recovery by the showed large numbers of tiny pink ~ilites(Acarina end of the study. Species cou~ltsthcre reached a sp.), but these mites \\,ere prescnt in large llu~nbers ~lli~lin~umsolneti~lle between 12 and 22 months on only one plot (formcrly in the Alnria-Hedo- postevent but then beg.ln showing some signs of p/z//~icomm~mity) in i\ugust 1974. These tiny recovery. By 17 montI>s postevent a few young abundant mites also accounted for no st of the E'ztctts disticlz~rs plants were seen on the plot incrcasc ill the incall ~iumberof inclividuals in plots (Lebednik aud Palmisano, Chap. 17, this rolunie), in tlic ~11~trirc-l1e~1opl1)~1111111and Hrclosnccio~l- and by 33 11lo11thsthe perccltt cover of I;: distichzts Alarb~eIilvertebrntes in Rocky Intertidal Co~niiztolities 427

Table 7-Species Abundances per 0.25 m2 011 IA-3 Plot 1 Before and After Caanikin

hlontl~s before Cannikint Alontl~salter Cannikir Species Taxon* 2 13 5 7 9 11 17 22 30 33

Scypha compressat Ilolichondria panicea t Actiniidae spp. Desinspirn senlirleatnfo I.epidochitona nlenlicus Hnloconchn mittor Cinguln spp.Q i\lifrelln mnian tis Buccinlrm bneri ~norchim~s~~z i\largnrites beringensis Idotea wosnessenskii PnrnNorclrestes ochotensis Leplnsterins nlerr fica Cfrcunfariauegoe ?Styela coriacen Actiniaria sp. Nemertea spp. Oligochaeta sp. Plrascolosomn agnssizii Schizoplns brnndtii Colliselln peltn ~\'otoncmen scirlunf Oolcis rondolphi /.ittorinn aleuticn Litforinn atkana Littorina sitko,m Aluinia nzrriuillii i\Issculas uenzicosus dlytiltrs edlclis Tzo.to,ria nlbrtcfn dlnco~,fnsp. Exos/>hneroma anfplicn~crlo Po~ilogo~ein,aoknroui Anisogo~~rmarusconferuicolt~s Gammaridea spp. I,iporoce/>hnh~sDreuipe,znis Sa!c,tderin mnrinus Pnraclfcrrio nlnskensis Diptera sp. Insecta spp. ~\'eon~olguslitlornlis Acarina spp.~. *A, asteroidea; Ac, acarina; Am, ampliineura; As, ascidiacca; U, bivalvia; C, cnidaria; G, gastropods; Ga, ga~mnaridea;H, holothuroidca; I, isopoda; In, insccta; P, porifera; Pc, polyclraeta; S, sipuncula. tAbundance is percent cover. $Plus signs indicate relative abundance: +, present. 5 Numbr:rs in parentheses indicate nu~nberof dead individuals. (1 Cingule alettti~oand C. ,,fnrtyni not distinguished. **Letters accompanying abundances refer to life stages: I., larva; A, adult. 30 Former rpecior. New species,

20

10

0

."

30 Lanlinaria Community 20

10

0 I I I I I I I I I I I I I I -15 -2 4 1 3 5 7 9 I1 17 22 30 33 CANNlKlN TIME. months

Fig. 16-Replacement following Cannikin of orisinat species by species not previously recorded on plots in three communities at 1A-2 and IA-3 combined. The numbers directly above the vertical bars indicate the number of plots sampled.

had increased to 40% (C. Simenstad, personal 2. The estimate excludes areas of offshore islets communication). Io\~ertebrate populations in- that could not be rcached. creased in size .ivitIi the increase in Fztcirs cover 3. The estimate does not include uplifted (Fig. 18 and Table 7), but the species count was bench arca east of Baujo Point or \vest of Sand \\.ell belo\\, the mean species colunt recorded for Beach Cove. Although the length of coastline undisturbed Hnlosnccio~t-Fzrcrrs plots by included in this estimate is only half that \\!herein 33 months postevent. Apparently plot1 at algal damage was detectable (Lebednik and Palmi- 33 inoiitlls postevent !\,as near the upper limit of sano, Chap. 17, this \'olume), the area of bench the fIa1osnccio1~-Fztctrs community (thc highest outside this length of coastline that was uplifted intertidal community) for the degree of exposure out of the intertidal zone is probably small by at IA-3. comparison. IIOIVmuch of the pre-Cannikin intertidal zone was uplifted into the supralittoral fringe? Thirty- Before Cannikin the intertidal zone \\,as a three months after Cannikin was detonated, I made nearly horizontal plane bct\\reen Banjo Point and an estimate of this arca in the field by superinipos- Sand Beacli Cove. In this stretch of coastline, the ing plane geometric figures on the bench surface. new intertidal area is a steep slope at the outer The areas included within the figures were essen- edge of the uplifted bench and occupies a much tially bare rock containing scattered supralittoral smaller area than before. 12inally, intertidal recolo- in\rertebratcs and no macrophytes. The estimate nization of the foriner bench surface may not be arrived at was 5.8 ha (14.3 acres). This estimate is coinplete; there is sonle evidence that the uplifted conservative because: bench is being eroded. By spring 1974 mechanical 1. The method treats the bench as a plane and cl~cinicalweathering appeared to have reduced surface and iguores irregularities. the level of the upliftcd bench by 1 to 2.5 cm, ~\larineIitvertebrates in Roclcy I~ttertidalCoi?t~ici~icities 431

Hal~~~cion-Fu~~~souter, b. 4 Alaria-Hedaph~llt~m,D - -13 Laminaria, 0------0

CANNtKlN TIME. months

Fig. 17--Mean number of individuals in plots for each algal zone before and after Cannikin at IA-2 and for the control, Makarius Uay (circled symbols). Asterisk indicates a sample size of 1. Species not retained by a 2-mm sieve, such as oligochaetes, Turtonio mhn~ta,andCinguln spp., gammarids, except ParaNorchestes ochotensis and Orcltestia spp., polgchaetes, except Nereis pelagica, and the holothurian Cuormaria vegae are excluded. All gammarids and polycl~aetesat Makarius Bay are excluded. dependiilg on bedrock lithology (I

typing se\~eral drafts of this chapter. She has Barabash-Nikiforov, 1947, The Sea Otter, Council of supported me morally and financially during most AIinisters of the RSFSK. Main Administration of of the ~siitingof this paper. Reserves. (Translated from Russian by Israel Program for Scientific Tz-anslations, Jerusalem, No. 621, 1962.) Barnard, J. 1 1962, Benthic hlarine Amphipoda of Southern California: Families Amphilochidae, Leoco- REFERENCES thoidae, Stenothoidae, Argissidae, Hyalidae, Pac. ~\'nturaIist, 3: 116-163. Abbott, R. T., 1974, Americntr Seashells; the dfarine -, 1960, The ,\mphipod Family Phoxocephalidae in the dfolluskn of the Atlnatic and Pncific Consts of A'orth Eastern Pacific Ocean, \\pith Analyses of Other Species Bnzwica, Van Nostrand Reinhpld Co., New York. and Notes for A Revision of the Family, Allan Hancack Annenkova, N. P., 1934, Kurze Ubersicht dcr Polychaeten Pacific Expeditions, \'ol. 18, pp. 175.375, der Litoralzonc der Bering-lnsel (Konunandar-Inseln),, Barnes, K. D., 1974, Inuertebrnte Zoology, 3rd ed., I\'.B. nebst Beschrcihume~ ~ neuer Arten. Zool. 2.106: 322-331. Saunders Company, Philadelphia. Armstrong, R. H., 1971, Physical Clin>atology of Amchitka Barr, L., 1971, Studies of Populations of Sea Urchins, Island, Alaska, BioScience, 21: 607-609. Stronp3~locentrot!crsp., in Relatiorl to Underground Bakos, G. J., 1966, A,larine Poeciloscleridan Sponges of the Nuclear Testing at Amchitka Island, Alaska, BioScieace, San Juan Archipelago, \Yasbington, J. Zool., 149: 21: 614-617. 415-531. Barr, N., 1973, Extension of the Known Range of the Crab, Ball, I. R., 1974, Contributions to a Revision of the Rlarine Cryptolitltodes typic~rs Brandt, to Amchitka Island, Triclads of North America: The hlonotypic Genera Alaska (Decapoda, Anoinura, Lithodidae), Crustocaana ~\'esiIis, i\'esion, and Fouiello (Turbellaria: Tricladida), (Leiden), 25: 320. Can. J. Zool., 53: 395-407. Banse, K., and K. D. Robson, 1974, Benthic Enn,ttiate Boss, K. J., J. Rosewater, and F. i\. Ruhoff, 1968, Tlte Po1)tchaetrs of British Cols~nbin nnd Il'nslti~~gfo,r, Zoologic01 Tosa of ll'illinm Healey DON, U. S. National Fisheries Research Board of Canada, Bulletin 185. Museum, Bulletin 287. ~Vlari~teI~tuertebrates in Rocky I~ttertidalCo?~lntzrnities 433

Fig. 19-Mean abundances of littorines on 0.25-m2 plots in three communities before and after Cannikin at IA-2. H-F;, Ha1osnccioi1-Fucus community inner plots; H-F,, Halosnccio,r- Fucus community outer plots; A-H, Alarin-Hedophyllzzm community; L, Lambraria commu- nity.

Rrandt, J. F., 1851, Krebse, in Reire in rlerf dicsserste~t Burton, M., 1963, A Revision of the Classification of the Norden und Osfen Sibiriens, A. Th. V. Middendorff, Calcareous Sponges, British i\luseum of Natural History, \fol. 2, pp. 77-148. London. -, 1850, \'orliiufige Bemerkungen iiber eine neue aus Carlquist, S., 1973, Islnnd Biology, Columbia University zwei ,loch unbeschriebenen Gattungen und Arten Press, Nexv York. gebildete Unterabteilung (Hapnlogastrica) der Tribus Chamisso, A. V., 1821, Remarks and Opinions of the Litl~odinn,begleitet von einer Characteristik der eben Naturalist of the Expedition, in A Voyage of Discovery genannten Tribas der Anomuren, Bfzll. CI. Phys.-dfatlt. into the Soutlr Sen rind Beering's Straits, Otto von Acnd. Imp. Sci., St. Petersboarg, 8: 266-269. Kotzebue, Bibliotheca i\ostraliana, No. 18, \'ol. 2, pp. -, 1849, Die Gattung Lithodes Latreille nebst vier neuen 351-433, No. 19, Vol. 3, pp. 1-318, Da Capo Press, Nerv ihr serwandten von 1Yosnessenski entdeckten, als Typen Vork. einer besondern Unterabteilung (Tribus Lithodea) der ,and G. G. Eysenhardt, 1821, De animalibus quid- Edrsads'schen Anomuren, BUN. CI. P1tys.-illnth. Acod. busdam e classe vermium Linneana, in circumnavi- Imp. Sci., Sf. Pk.fersborrrg, 7:171-176. gation~terrae, auspicante Comite N. Romanzoff, duce Rriggs, J. C., 1974, hIari?~eZoogeogrophy, h,lcGra\\'-Hill Ottone de Kotzebue, annis 1815-1818, Fasc. 2, ~\'oun Book Company, New York. Acta Pltysico-illedicn, Acad. Cnesarene Leopoldhzo- Bolycheva, A. I., 1957, The Sea Fleas of the USSR and Cnroli~zneBonn. ~\'atrtme Crtriosorem, 10: 343-374. Adjoining M'aters (Amphipoda-Talitroidea), Acad. Sci. Clark, H. L., 1911, Arorfh Pacific Opttiurntts in the USSR Zool. Inst., 65: 7-185. (Translated from Russian CoNecfion of fhe U. S. i\'ntionnl dIfcseu,,r, U. S. National by The Bureau for Translations of the Department of Museum, Bulletin 75. the Secretary of the State of Canada.) Clark, S. F., 1876, Report on the Hydroids Collected on Burgner, R. L., K. K. Chew, J. S. Isakson, and 0. A. hlathi- the Coast of Alaska and the Aleutian Islands by \Ir. H. sen, 1969, i\mchitka Bioenviron~nental Program. Dall, U. S. Coast Survey, and Party, from 1871 to 1874 i\nnual Progress Report, July 1, 1968-June 30, 1969, Inclusive, Proc. llcnrl. '\'a tur. Sci., Plfilndelpl~ia, 3 : Research Program on h'larine Ecology and Oceanog- 209-235. raphy, Amchitka Island, Alaska, USAEC Report BMI- Coan, E. V., 1971, The Northwest American Tellinidae, 171-128, pp. 45-46, Battelle Memorial Institute. Veliger, 14(Suppl.): 1-63. Fig. 20-Mean abundances of littorines on 0.25-m2 plots in two communities [Halosaccion- Fucus (H-F) and Alarin-Herlophyllt~n~(A-H)] and abundances on one plot in the Lanliilaria (L) com~nunitybefore and after Cannikin at It\-3. Data collected in the Ln~~tinnriacommunity 15 months before Cannikin were lost at sea.

Connell, J. H., 1972, Community Interactions on Marine J. I\'.Hedgpeth (Ed.), pp. 535-585, Geological Society Rocky Intertidal Shores, An?:. Reu. Ecol. Syst., 3: of America, Memoir 67, New York, N. Y. 169-192. D'yakonov, A. M., 1954, Opbiuroids of the U.S.S.R. Seas, -, 1970, A Predator-Prey System in the Marine Intertidal Opred Faaile SSSR, Izdau. Zool. Ifist. Akad. rVnuk. Region, I.Balanss glattdula and Several Predatory SSSR, 55. (Translated from Russian by Israel Program Species of Thais, Ecol. ~\Iottogr.,40: 49-78. for Scientific Translations, Jerusalem, No. 1880, 1967.) -, 1961a, The Influence of Interspecific Competition and -, 1950, Sea Stars (Asteroids) of the USSR Seas, Opred. Other Factors on the Distribution of the Barnacle Fatli~eSSSR, Zzrlav. Zool. Irrst. Aknd. ,\'ask. SSSR, 34. Cl~thnmnlusstellatus,Ecologj*, 42: 710-723. (Translated from llussian by Israel Program for Scien- -, 1961b, Effects of Competition, Predation by Thais tific Translations, Jerusalen~,No. 1922, 1968.) lapillus, and Other Factors on Natural Populations of Ekman, S., 1953, Zoogeography of the Sea, Sidgwick & the Barnacle Enlanus bnlnnoirles, Ecol. i\Io~?ogr., 31: Jackson, Ltd., London. 61-104. \V. F. Dall, \\I. H., 1884, Report on the r\lollusca of the Emison, B., S. L. \\'illiamson, and C. hl. \Vhite, 1971, Commander Islands, Bering Sea; Collected by Leonhard Geographical Affinities and Migratio~~sof the Avifauna Stcjneger in 1882 and 1883, Proc. U. S. ,\'at. i\Jtrs., 7: on Amchitka Island, Alaska, BioScience, 21: 627-631. 340-349. Eschscholtz, J. F., 1829.1833, Zoologischer Atlas, Dayton, P. K., 1975, Esperimental Evaluation of Ecological G. Reiner, Berlin. Dominance in a llocky Intertidal Algal Community, Estes, J. I\., and J. F. Palmisano, 1974, Sea Otters: Their Ecol. dfoltogr., 45: 137.159. -, 1971, Competition, Disturbance, and Commu~>itg Role in Structuring Nearshore Communities, Science, 185: 1058-1060. Organization: the Provision and Subsequent Utiliaation of Space in a Rocky Intertidal Community, Ecol. Fager, E. \Y., 1963, Communities of Organisms, in The Sea, i\Jonoq., 41: 351-389. Ideas and Obseruatio~~sOIL Progress bt the Study of the Dodimead, A. J., F. Favorite, andT. Hirano, 1963, Salmon Sens, Vol. 2, The Co,nposition of Sea-\\'ater Compara- of the North Pacific Ocean, Part 11. Reviev of Oceanog- tive and Descriptive Oceanography, M. N. Hill (Ed.), raphy of the Subarctic Pacific Region, International pp. 415-437, Interscience Publishers, Inc., now John North Pacific Fisheries Commission, Bulletin 13. \\'iley & Sons, Inc., New York. Doty, hi. S., 1957, Rocky lntertidal Surfaces, in Treatise on -,and J. A. b'icGowan, 1963, Zooplankton Species ilfarine Ecology and Paleoecology, Vol. 1, Ecology, Groups in the North Pacific, Science, 140: 453-460. Inuertebrntes b~Rocky Intertidal Conin~unities 435

Favorite, F., 1967, The Aloskan Stream, International Hobson, K. O., and K. Banse, Benthic Sedentariate Poly- North Pacific Fisheries Commission. Bulletin 21. chaetes of British Colilmhia and \\'ashington, in prepara- pp. 1-20. tion. Fisher, W. K., 1938, Hydracorals of the North Pacific Hulthn, E., 1960, Florn of the Alentiair Isln~rds,2nd ed., Ocean, Proc. U. S. Nat. dfirs., 84: 493-554. J. Cranter, \Yeinheim. -, 1930, Asteroiden of the North Pncific and lldjncent Kenyon, K. \V., 1969, Tlie Sea Otter in the Eastern Pocific Il'aters, Part 3, Forcipulata (concluded), U. S. Natioital Ocenll, U. S. Department of the Interior, Bureau of Illuseum, Bulletin 76, pp. 1-356. Sport Fisheries and \\'ildlife, North American Fauna, -, 1928, Asteroidea of the North Pncific and Adjaceitt No. 68. Il'aters, Part 2, Forcipulata (part), U. S. National Kincaid, T., 1964, Notes on Tlrais (Nucella) linra (Gmelin), h,luseum, Bulletin 76, pp. 1-245. a hlarine Gastropod Inhabiting Areas in tbc North -, 1911, Asteroiden of the Nortlr Pacific and Adjacent Pacific Ocean, 'Die Calliostoma Company, Seattle, Il'aters, Part 1, Phanerozonia and Spinulosa, U. S. Na- Wash. tional Museum, Bulletin 76, pp. 1-419. Kirkwood, J. B., 1974, A~nchitkaBioenvironmental Pro- Praser, C. M., 1937, Hydroids of the Pacific Coast of gram. Bioenvironmental Safety Studies, Amchitka Cannda mid the United States, University of Toronto Island, Alaska, Spring, 1974 Task Force Report, Press, Toronto. USAEC Report Bi\lI-171-157, Battelle L'Iemorial Insti- Fretter, V., 1951, Obsen'ations on tlte Life History and tute. Functional Morpllology of Cerithiopsis tuberculnris -, and R. G. Fuller (Comps.), 1972, Uioenvironmental (h,lontagu) and Triphorn peruersa (L.), J. illnr. Biol. Safety Studies, Amchitka Island, Alaska, Cannikin Dt2 Ass. U. K., 29: 567-586. AIonths Report, USAEC Report BMI-171-147, Battelle Garth, J. S., 1958, Brachyura of tile Pacific Coast of hfemorial Institute. America. Os\~rh\,ncBa,. , Allan Hancock Pacific Ex~edi- Latrbitz, D. R., 1970, Studies on the Caprellidae (Crustacea, tions, Vol. 2 1. Amphipoda) of the American North Pacific, Publica- Glynn, P. W., 1965, Community Composition, Structure, tions in Biological Ocennograpliy, Nation01 dluseunr of and Interrelationships in tlte Maine Intertidal Endo- Natural ScieTrces, Ottazun, No. 1. cladin nruricata-Balantrs alanrlirla Association in Lewis, J. R., 1964, The Ecolog). of Rocky Shores, English h4onterey Bay, California, en-ufortia, 12: 1-198. Universities Press Ltd., London. Golikor,, A. N., and 0. G. Kussakin, 1972, Sur la Biologie Lindroth, C. H., 1963, The Aleutian Islands as a Route for de la Reproduction des Patelles de la Famille Tecturidae Dispersal Across the North Pacific, in Pacific Basin (Gastropoda: Docoglossa) et sur la Position Sys- Biogeography, Tettth Pncific Science Congress, Hono- tematique de ses Subdivisions, dfalacologin, 11: lulu, J. L. Gressitt (Ed.), pp. 121-131, Bishop hluseum --.9R7.944 -" .. Press. -.and 0. G. Kussakin. 1962. Fauna and Ecoloes of r\lacArthur, R. H., 1972, Geographicnl Ecology, Harper & ~astropod ~rosobra!;chian i\lollusks (~astroKoda, Row Publislters, Inc., New I'ork. Prosobranchia) of the Littoral of Kurile Islands, Issled -,and E. 0. %\'ilson, 1967, Tlte Tlieory of Island Dnlheuosf. SSSR, 8: 248-346. (In Russian.) Biogeography, Princeton University Press, Princeton, 7 Grube, A. E., 1855, Beschreibungen neuer oder \venig n... J. hekannter Anneliden,Arclt. ~\'atargesclr., 21: 81-136. MacGinitie, N. 1959, blarine Alollosca of Point Barro\v, Gurjanova, E. F., 1966, Comparative Research of Biology Alaska,Proc. U. S. Not. i\lus., 109: 59-208. of the Littoral in the Far Eastern Seas, Proc. Pac. Sci Makarov, 1'. V., 1938, Crustacea: r\nomura, Opred. Fatn~e Co,igr., Sth, 19: 75-86. SSSR, Izdau. Zool. Inst. Akad. Nauk. SSSR, I\'. S., 16. - 9 Bokoplavy morei SSSR i Sopredel'nykh vod (Translated from Russian by Israel Program for Scien- (Amphipoda, Gammaridea), Opred. Faune SSSR, Izdnu. tific Translations, Jerusalem, No. 225, 1962.) Zool. Inst Akad Nauk. SSSR, 41: 1-1029. Mauze)., K. P., C. Birkeland, and P. IC. Dayton, 1968, - 1935, Komandorskie Ostrova i ikh Morskaya Pribezh- Feeding Behavior of Asteroids and Escape Responses of rtaya Fauna i Flora (The Commander Islands: Their Their Prey in tlte I'uget Sound Region, Ecology, 49: Littoral Fauna and Flora), Prirodn, 11: 64-72. (Trans- 603-619. lated from Russian by Leda V. Sagen.) McAlister, B. \\I., 1971, Oceanograpl~yin the Vicinity of Gustus, R. hl., 1972, A Species of the Genus Eirnice (Poly- Amcltitka Island, Alaska, BioScience, 21: 646-651. cltaeta) from the Pacific Northwest Coast, ~\'ortlrwest r\lcLaughlin, P. A,, 1974, The Hermit Crabs (Crustacea Sci., 46: 25 7-269. Decapoda, Paguridae) of North~vesternNorth America, Harger, J. R., 1972, Variation and Relative "Niche" Size in Zool. l7erh. (Leiden), No. 130. the Sea Ivlussel i\fytilas edirlis in Association with hlcLean, J. H., 1969, h'larine Shells of Southern California, ~\lytiIascalifortiinni~s~ikliger, 14: 275-283. Los Angeles County A'luseum of Natural History, Hartman, O., 1969, htlas of the Scdentariate Polycbaetous Science Series 24, Zoology, No. 11. Annelids from California, Allan Hancock Fonndation, -, 1966, \Yest American Prosobranch Gastropoda: Super- Los Angeles, Calif. families Patellacea, Pleurotomariacea and Fissurellacea, -, 1959, Catalogue of the Polycbaetous Annelids of the Ph. D. Thesis, Stanford University. \old Pxts I and 11, Allan Hancock I:oundation, Menge, B. A,, 1972, Foraging Stratepy of a Starfish in Occasiond Paper No. 23. Relation to Actual Prey Availability and Environmental Hedgpcth, J. W., and J. J. Gonor, 1969, Aspects of the Predictability, Ecol. illonogr., 42: 25-50. Potential Effect of Thern~alAlteration on hlarine and hfenzies, K. J., 1952, Some Marine Asellote lsopods from Estuarine Benthos, in Biological Aspects of Thermol Northern California, rvith Descriptions of Nine New Pollutiorr, Proceedings of the National Svm~osiumon Species, Proc. U. S. Nat. dl~rs.,102: 117-159. Thermal Pollution, ;p. 80-118, Vanderbjlt -University hlerritt, &,I. L., 1972, Cannikin Ground Alotion and Water Press. Pressures, in Amchitka Bioenvironmental Program, Bio- environmental Safety Studies, Amchitka Island, Alaska, Richards, B. R., and C. I. Belmorc, 1976, Occurrence of the Cannikin D+2 hlonths Report, USAEC Report BhII- \\'oodborer, Limnaria ligaorum (Rathke) at Amchitka 171-147, Battelle Memorial Institute. Island, Alaska, J. Fish. Res. Bonrrl Can., 33: 1642.1644. Middendorff, I\. Th., 1851, hlollusken, in Reise in den Richardson, H., 1905, ~\lonogroph or1 the Isopods of ~Vortlr itssersten iVorden und Osten Sibiriens, A. Th. v. America, U. S. National h'luseurn, Bulletin54, bliddendorff, Vol. 2, No. 1, pp. 163-508. pp. 1-727. - 1849a, Beitrage zu einer hIalacozoologia Rossica. Ricketts, E. F., J. Calvin, and J. 1%'. Iledgpeth, 1968, I. Beschreibung und Anatomie ganz neuer, oder fur Between Pacific Tides, 4th ed., Stanford Universit), Rusrland neuel- Chitonen, i\lAnt. Acad. Imp. Sci., St. Press, Stanford, Calif. PAtersbourg, Ser. 6, Sci. ,Vat., 6: 67-215. Rieger, S., and R. E. \\'underlich, 1960, Soil Survey and -, 1849b, Beitfige zu einer Malacozoologia Rossica. 11. Vegetation of Northeastern Kodiak Island Area, Alaska, Aofz'ihlung und Beschreibung der zur Meeresfauna U. S. Department of the Interior, Bureau of Land Russlands gehijrigen Einscltalcr, ill&n~.Acad. 11np. SCi., hlanagement, Soil Conservation Service, Soil Sulvey St. P&tersbourg, Ser. 6, Sci. A'at., 6: 329-516. Series 1956, No. 17. -, 1849c, Beitfige zo einer Malacozoologia Rossica. Ssuer, I 1802, A,r Account of a Geographical and 111. Aofi'ihlung und Beschreibuna der zur i\leeresfauna Astro,tomicnl Experlition to the A'ortlzern Ports of Russlands gellarigen Z\veischaler, fildm. Acad. Imp. Sci., Russia, for Ascertninitlg the Degrees of Latitude and St. P6tersboarg, Ser, 6, Sci i\'at., 6: 517-596. Lojtgigitllde of the AIouth of the Hiuer Kouin~a;of the -, 1846, \'orlHufige Anzeige bisher unbekannter hlollas- Il'l~ole Const of the Tshetski, to Enst Cape; and of the ken, als Vorarbeit zu einer hIalacozoologia Rossica, IslanrLr in the Eastern Oceon, Stretching to the Anreri- verijffentlicht durch, Bttll. CI. Phys.-i\lnth., Acnd. Imp. ca,z Const, by Cotnnrodore Joseph Billings, in the Years Sci, St. P6ter?botcrg, 6: 113-122. 1785 to 1794, A. Strahan, London. Scarlato, 0. A,, 1960, The Bivalved Molloscs of the Far hlortensen, T., 1943, A dlonograph of the Echinoirlea, East Seas of USSR (Order D~sadonta),Opred. Farcfie Carnorodonto II, Echinidne, Strongylocentrotidae, SSSR, Izrlau. Zool. I~asf.Aknd. ~Vask.SSSR, 71: 1-150. Parosnle~liidne, Echh~ometridne, Vol. 3, Part 3, C. A. (In Russian.) Rejtzels, Forlag, Copenhagen. Schultz, G. A., 1969, How to Know the i\farine Isopod Newell, I. M., 1951, Nerv Species of Agnue and Thalass- Crustncenns, \iIlim C. Brorvn Co., Publishers, arach,m fsom the Aleutians, Amer. ~lles.~\'ouit., 1489: Dubuque, Io\va. 1-19. Shelford, V. E., A. 0. Weese, L. A. Rice, D. I. Rasmussen, -, 1950, Ne\v Species of Copidog~zathusfrom the /\leu- and A. hlacLean, 1935, Some Marine Biotic Communi- tians, An~er.illus. i\'ouit., 1476: 1-19. ties of the Pacific Coast of North America. Part I. Nybakken, J. I\'.,1969, Pre-Earthquake Intertidal Ecology General Smvey of the Communities-Their Extent and of Three Saints Bay, Kodiak Island, t\laska, Biol. Pap. Dynamics, Ecol. Monogr., 5: 249-332. Uniu. Alaska, 9: 1.123. Soot-Ryen, T., 1955, I\ Report on the Family hlytilidae fPetecvuoda). t\llan Hancock Pacific Expeditions, O'Clair, C. E., and K. K. Chew, 1971, Transect Studies of v0l. 2b; pp.'i-i74. Littoral I\lacrofauna. Amchitka Island. Alaska. S~asskv.N. N., 1961. The Intertidal Zone of the South- ~iscience,21: 661-664. Eas; shore of ~amchatka,Issled. Dnl'neuost. Morei Oldroyd, I. S., 1924 and 1927, hlarine Shells of the \\lest SSSR, 7: 261-311. (In Russian.) Coast of North America, Staitford Uni\rersity l'ublica- Steineaer, L., 1936, Georv Il'illtelm Steller, The Pioseer of tion, University Series, Geological Science, Vols. I and -Al&kan r\'atural ~i;ory, Hanfard University Press, 11, Parts 1, 2, and 3. Cambridge, Xlass. Paine, R. T., 1974, Intertidal Community Structure, Ex- Stephen, A C and S. J. Edmonds, 1972, The Phyla perimental Studies on thc Relationship Between a Sipuncula and Echiura, British hluseiim of Natural Dominant Competitor and Its Principal Predator, Oeco- History. logio, 15: 93-120. Stephenson, T. A,, and A. Stephenso:,, 1972, Life Between -, 1966, Food \\'eb Complexity and Species Diversity, Tiden~nrkson Rockv Sllores. 1V. H. Freeman & Co., San Amer. ~Vntur.,100: 65-75. Francisco, Calif. Palmisano, J. I:., 1975, Sea Otter Predation: Its Kole in -.and A. Steuhenson. 1949. Universal Features of Rocky Intertidal Community Structure at Amchitka Zonation Bet;veen ide em arks on Rocky Coasts, j. and Other Aleutian Islands, Ph. D. 'lllesis, University of Ecol., 37: 289-305. \trashington. Stephenson, \V., 1973, The Validity of the Community Pilsbry, H. I. 1916, Tlte Sessile Barnacles (CirripedinJ Concept in Marine Biology, Proc. R. Soc. (QueenslnndJ, Confained in tlzc Collectio~?~of the (I. S. r\'ational 84: 73-86. ~\luseum, Incl~rding n ~\lo~togmphof the America11 Tzvetkova, N L., 1975, Geographical Distribution of Species, U. S. National hluseum, Bi~lletin93. Species of the Genus Anisogan~marus Derzhavin Powers, H. \ R. R. Coats, and \\'.H. Nelson, 1960, (Amphipoda, Gammaridae), Crusfacennn (I,eirlenJ, 28: Geology and Subv~arine Physiography of Amcl~itka 191-199. Island, Allnska, U. S. Geological Survey, Bulletin 1028-P, pp. 521-554. Ushakov, P. V., 1955, Polychaeta of the Far Eastern Seas of Rathbun, hi. J., 1930, The Cnncroid Crnbs of America of the U.S.S.R., Oprerl. Faeae SSSR, Irdau. 2001. Inst. the Fantilies Eurynlidoe, Portuaidne, Atelecyclidae, Akad. Nauk. SSSR, No. 56. (Translated from Russian Concridae and Xa,zthidna, U. S. National hluseum, by Israel Progran~for Scientific Translations, Jerusalem, Bulletin 152, pp. 1-609. No. 1259, 1965.) Reid, J. L., 1973, Northwest Pacific Oceml Il'aters ill U. S. Department of Commerce, 1971, National Oceanic Il'inter, Johns Hopkios Oceanographic Studies No. 5, and Atmospheric Administration, Locnl Climatological Johns Hopkins University Press, Baltimore, Md. Datn, Ai~nual Summary with Comparatiue Dafa, 1lla~i11eI?tuerteltrates in Rocky Intertidal Cotnmtcltities 437

Sheaiyn, Alaska, Supcrirttendent of Documents, U. S. Dr. J. kliltotl Campbell, Research Bninch, Government Printing Office, \\'ashington, D. C. Biosystematics Research Institutc, Agricul- -, Knvironmental Sciences Services Administration, 1969, Tide Tables, West Coast ~Vortlt and South ture Canada, Ottawa, Ontario, Canada. America, Superintendent of Documents, U. S. Govero- Coleoptera. ment Printing Office, \\'ashington, D. C. Dr. David G. Cook, Dcpartlnent of the En- -,Coast and Geodetic Survey, 1949, Ede nnd Current vironment, Fisheries and Marine Service, Glossary, Special Publication No. 228, Superintendent Ottawa, Ontario, Canada. Oligochaeta. of Documents, U. S. Government Printing Office, Wash- ington, D. C. hpIr. Matthew Dick, 1902 State St., illamosa, U. S. Naval Oceanographic Office, 1969, BIontltly Charts of Colo. Bryozoa. Mean, BIinintum and blaximum Sea Surface Tempera- Dr. A,Iaureen E. Doxvncy, Dcpartment of In- ture of the Nortll Pacific Ocean, Special Publication vertebrate Zoology, National h,Iuscum of No. SP-123. Natural History, Washington, D. C. Van Name, \Y. G., 1945, The North and South American Ascidians, Ball. At~ter.Uus. at. Hist., 84. Asteroidea. Vermeij, G. J., 1972, Endemism and Environment: Some Dr. Arthur R. Fontainc, Department of Biol- Shore blolluscs of the Tropical Atlantic, Amer. ~\'atur., ogy, University of Victoria, Victoria, British 106: 89-101. Columbia, Canada. Bo~telliopsis. \\'ilimovsky, N. J. 1964, Inshore Fish Fauna of the Ms. A~Iadeline Green, College of Fisheries, Aleutian Archipelago, in Proceedhtgs of the Fourteenth University of \Vashington, Seattle, \\'ash. Alaska Science Conference, pp. 172-190. Yamada, hl., 1955, Some Hydroids from Agattu, in the Japetelln Iteatlti. Aleutian Islands,A~tnot.Zool. Jap., 28: 121-125. Dr. Olga Hartntan, deceased, formerly at the Allan Hancock Foundation, University of Soutltern California, Los Angeles, Calif. APPENDIX: t\NNOTKrED SPECIES LIST C;rratzilus. OF THE IN\IERTEBRATES AT Dr. Melville H. Hatch, Burke A,Iemorial h,Iu- AMCHITKA ISLAND, ilLASIaroce/~hah~s. The follon'ing species list is a partial list of tlte spccies of itl\,ertebratcs collected by Fisheries Dr. Charlotte ~Iolntquist,Sekt. Evertebratzool, Research I~tstitute personnel in the vicinity of Naturhistoriska Riksmuseet, Stockholm, Alnchitka Island, Alaska. A nutttber of specialists Sweden. Pnrnca~ltlto~~tysisk~~rile~tsis. have contributed their expertise to the preparation Dr. Paul L. Illg, Dcpartll~ent of Zoology, of this list. These specialists are listed below and University of \Vashington, Seattle, \\'ash. are acluto~vledgcdpreceding the taxonomic groups Blakeartus. or species they identified. Dr. A3argit Jensen, Zoolo$cal A~lusettn~,Univer- Dr. Donald P. Abbott, I-Iopkins h8karineStation sity of Copenhagen, Copenhagen, Denmark. of Stanford University, Pacific Grove, Calif. Strottg)~loce~ttrotf~s. Styela clavata. Dr. A. htyrit Keen, Department of Geology, h,Is. Suzan~teAllyson, Research Bratlch, Biosys- Stanford University, Stanford, Calif, h4ol- tematics Research Institute, Agriculture lusca. Canada, Ottawa, Ontario, Canada. Lepidop- tera. Dr. Allan J. Icohn, Department of Zoology, Dr. Gerald J. Bakus, University of Southern University of \\'ashington, Scattle, \\'ash. California, Los Angeles, Calif. Halicho~tdria Pltascoloso~~ta, pa~~icea. Dr. Eugene N. Kozloff, Department of Zool- Dr. Karl Banse, Departlnent of Oceanography, ogy, U~tiversityof \\'ashington, Seattle, \\'ash. University of \\lashington, Seattle, \,\lash. l'etrastel~t~~ta. Syllis. Is Diana R. Laubitz, National h~luseum of Mr. Louis Barr, National h~larine Fisherics Watural Sciences, Ottawa, Ontario, Canada. Service, Biological Laboratory, Auke Bay, Cerco/>s. Alaska. Rhittolitl~odeszuos~tesse~~skii. Dr. Ulf Lie, Biological Station Espegrend, Dr. Ed~vardC. Becker, Research Branch, Bio- Blomsterdalen, Nor~vay.Gammaritlca. systematics Research Institute, Agrici~lture Dr. Evert E. Lindquist, Research Branch, Bio- Canada, Ottalva, Ontario, Canada. Coleop- systenlatics Research Institute, Agriculture tera. Canada, Ottalva, Ontario, Canada. Acarina. Dr. Edward L. Bousfield, National h,luseum of Dr. Jaltles H. A~IcLean, Los Angeles Coiulty Natural Sciences, Ottawa, Ontario, Canada. h,Iuseu~n of Natural &tory, Los Angeles, Gammaridea. Calif. h4ollusca. Dr. Carl F. Nyblade, Depart~llellt of Zoology, insertebrates occurring at the five main study sites University of \\lashington, Seattle, Wash. iilclttdes the population statistics fidelity (f.), Paguridae. constancy (c.), and abu~ldance(a.). Classification Dr. Rita hl. O'Clair, Friday Harbor Labora- generally follo\vs Barnes (1974). Specitne~lshave tories, Friday Harbor, Washington. Poly- been depositcd in the invertebrate collections of chaeta. the National h,luseum of Natural Sciences, Ottawa, 1.Gordon \$I. O'Connell, Department of aud the College of Fisheries, University of \\'ash- Biology, University of Victoria, Victoria, ington. Abbreviations used in this species list are British Colutnl)ia, Canada. Pycnogonida. listed below. Dr. Do~~aldR. Oliver, Research Branch, Biosys- n (abundance), median and range of numbers tematics Research Institute, Agriculture of illdividuals per 0.25 m2 in quadrats in which the Canada, Ottawa, Ontario, Canada. Diptera. species mas found. Dr. Bruce Ott, Institute of hk~rineSciences, c (constancy), proportion of quadrats from the A4cGill University, A'lontreal, Quebec, Can- community to which the species was assigned that ada. Porifera. contained the species. Dr. David I.. Pawson, Department of Itlverte- f (fidelity), proportion of all quadrats contain- brate Zoology, National h'luseum of Natural ing the species that was in the community to \vhich History, \Yasliington, D. C. Echinodermata. the species was assigned. A species was assigned to Dr. Stephen Prudhoe, Department of Zoology, that comnlut~itywhere the value off: \cras greatest. British Museum (Natural History), London, NA,lNS, National Museum of Natural Sciences, England. Turbellaria. Ottawa, Canada. hfls. Fanlida Rafi, National bluseum of Natural USNkI, National bluseuin of Natural History, Sciences, Ottalva, Ontario, Canada. Idotea. \,\'ashington, D. C. Dr. Gordon A. Robilliard, Woodward-Envicon, Phylum Porifera (Unless otherwise noted, all Porif- hlc., Sa11 Dicgo, Calif. Nudibranchia. era identified by B. Ott and Dr. h'lary E. Roussel, Research Bratlch, Biosys- deposited in the NMNS.) tematics Research Institute, Agriculture Can- Class Calcarea ada, Ottaxva, Ontario, Canada. Diptcra. Scypha contpnctutn (Lambe, 1894). h4r. Kenneth Sebens, Department of Zoology, Sublittoral, 6 In. University of Washington, Seattle, Wash. Scyplzn contp~essa (Fabricius, 1780). Actiniaria. Cottlpared with NMNS specimens of Mr. Ronald Shirnek, Department of Zoology, Gra~ttinnloltstncosa (Breitfuss, 1898) University of \Vashington, Seattle, Wash. [= S. conipressa (see Burton, 1963)l Sztavorl~illia. from Bering Island. Verified by B. Mr. Peter N. Slattery, Moss Landing h4arine Ott. See Table 2 for population statis- Laboratories, h,loss Landing, Calif. Gamma- tics. ridca. Lezccnird~a taylori Lambe, 1900. Dr. Herbert J. Teskey, Research Branch, Bio- Alnria-Hedopltyllz~t~z community to systematics Research Institute, Agriculture 91 m. Canada, Ottawa, Ontario, Canada. Diptera. Leicco~tin pyrifOrt~ziS (Lambe, 1894). Mr. Gregory J. Tutmark, 2200 Northeast 75th Ala~ia-Hedoph)~llzam coinmu~lityto Street, Seattle, Wash. Ccphalopoda and 6 111. Crustacea. Leuco~tiniteathi (Urban, 1905). Ln~iti- Dr. Charles W. Thayer, Department of Geol- tzarin community to 183 n~. ogy, University of I'ennsylvania, Philadelphia, Leucoltin sp. A. Sublittoral, 6 to 24 111. Pa. Brachiopoda. Rha6dodernzella ttztttirigi Urban, 1905. Ulllcss otherwise noted, I identified the rcst of Sublittoral, 91 in. the species included in the list from the literature, Class Hyalospollgea and references are included follo\ving the names, KltaDrlocal)~/~trtsdazosotri (Lambe, authors, and dates of thcse species. Additiollal 1893). Sublittoral, 183 m. annotatiolls include the vertical distribution of the Hyalospongea sp. Sublittoral, 183 111. species in the vicinity of Amchitka, and, for those Class Demospongiac species covered in Table 5, the geographical dis- Haliclona pernt 011;s (Bolverbank, tribution. (Species in those groups covered whose 1866). Sublittoral, 6 111. ranges estelld to a depth greater than 60 nl were Hnliclo~rasp. Sublittoral, 91 m. not included in Table 5.) Inforlnatioil on intertidal Gellitts sp. Sublittoral, 183 m. hlnrit~eInuertebrntes iii Rocky Intertido1 Conzm~oiities 439

Signindocia edn/)lt~ts de Laubenfels, Asbesto/~l~~n~alyco/>odiun~ (Le\' 'IIISCII, 1930. Sublittoral, 55 m. 1886). Sublittoral, 91 m. Isodictyn qrcntsi~toeiisis(Lambe, 1893). Awirtelln sp. A. Sublittoral, 183 m. Sublittoral, 6 to 24 m. Syringelln ntn/>his/~iclclnde Laubenfels, ~\Jycale hispicla (Lambe, 1894). Sub- 1961. Sublittoral, 183 m. littoral, 6 to 24 m. Phylum Coeletlterata iV1. litlgiia (Bowerbank, 1866). Sub- Class Hydrozoa (Fraser, 1937). littoral, 18 to 24 111. Order Hydroida A{. loveni (Fristedt, 1887). Sublittoral, Syitcoryi~esp. Loininnria community; f, 91 m. 1.0; c, 0.06; a, 1. M. helios (Fristedt, 1887). Sublittoral, Sertulnrelln pinnntn Clark, 1876. Lanti- 91 In. imrin community; f, 1.0; c, 0.06; a, 1. Mycnle sp. Sublittoral, 91 m. Order Stylasteritla Atitl~onrc~catagrncene Bakus, 1966. ?Allo/>ola canlpylecn (Fisher, 1938). Sublittoral, 6 to 24 m. Sublittoral, 183 111. ~Vlicrocioi~npritnitivn IColtun, 1955. Class Anthozoa Sublittoral, 91 to 183 111. Order Pcinlatulacea Iotrocl~otamngna Lambe, 1895. Speci- Ptilosnrci~s sp. Subtidal, 30 to 35 m. tnen collected by R. Martin. Sublit- One very young specime~lcollected in toral, 15 to 23 m. Lainitfarin com~nunityat IA-2. Lissodeitdo,yx nn~nkt~akensis(Lambe, Order Actiiliaria (All anemones \\.ere identi- 1895). Sublittoral, 6 111. fied by I<. Sebens except L. firitla (Lambe, 1895). Laminorin kletriclitoii sp.) community to 91 In. Family Actiniidae Stelodoryx alnske~tsis (Lambe, 1895). Cnirlopus ritteri (Torrey, 1902). Lam- Sublittoral, 6 to 55 m. inaria comrnni~ity. IlJ(ygb1sin zu~giitside Laubenfels, 1953. Epinctis sp. Lnininarin community. Sublittoral, 183 m. Several specimens bearing young. iVl)~~illnlncnnosn Lambe, 1893. Sublit- Tenlin sp. Lnininarin community. toral, 6 m. Unidentified sp. B. Lnn~iimria com- Halicliondrin panicen (Pallas, 1 766). munity; the most common anemone Identification by G. J. Bakus of speci- in the lorver midlittoral zone and mens collected by J. F. Pahnisano. inf~alittoralfringe. Unidentified sp. C. Lniiiinaria com- Lniititiaria community; f, 0.53; c, nluuity. 0.66; a, 12.6%, 1 to 55%. Unidentified sp. D. Lniiziiznria com- H. Inntbei Brdt~sted,1933. Sublittoral, munity. 6 to 183 in. Family Ho~mathiidae Sziberites domzotcltln doiizuncicla (Olivi, Unidentified sp. A. Lnminnrin corn- 1792). Sublittoral, 12 to 128 nl. muni ty. S. d. ficus (Olivi, 1792). Sublittoral, 55 Family Metridiidae to 183 m. Afetridinnt sp. Subtidal. S. n~o~ttinigerCarter, 1880. Sublittoral, Phylum Platyhelminthes 55 to 183 in. Class Turbeilaria (All Turbellaria identified by S. Polyntnstia pnclry~nnstinde Laubenfels, PI udhoe.) 1932. Sublittoral, 6 to 274 m. Order Alioeocoela P. pncificn Lambe, 1894. Sublittoral, Plagiostomidae sp. Hnlosnccio~t-Fuc~is 91 to 183 In. community: f, 0.66; c, 0.18; a, 1, 1-2. Lntvitc~~lintricinctn Hentchel, 1929. Order Tricladida Sublittoral, 183 m. ~Vexilis epichitoni~is Holleman and Tetilln uillosa (Lambe, 1894). Sublit- Hand, 1962. Alnrin-Hedoph31llzcnz toral, 6 to 183 In. community: f, 0.6; c, 0.25, a, 9, T. s/)iitosn (Lambe, 1894). Sublittoral, 2- 133. Not previously recorded 6 to 183 m. northwest of Vai~couverIsland, Brit- Corticitciif sp. Sublittoral, 34 n~. ish Columbia (Ball, 1974). Geodinella robtcstn Lendenfeld, 1910. ~Vesionnrcticuiii Hyman, 1956. Upper Sublittoral, 274 111. communities; f, 1.0; c, 0.09; a, 2. Order Polycladida community; f. 1.0, c. 0.33, a. Noto/~/a~inntomntn (0. I?. h,liiIler, 2. Asiatic. 1776). Alarin--HedophyN~tm com- /Itrtolytus sp. Alaria-Hedopl~ylli~~tf munity; f, 1.0; c, 0.08; a, 1. community; f. 0.66, c. 0.17, a. 6, ?Leptopla?zn vesic~tlatnHyman, 1939. 1-11, Juvenile. Lai~titzaria community; f, Ex ogo 71 e ge,i~?itifercr Pagenstecher, 1.0; c, 0.33; a, 1. 1862. Identified by R. M. O'Clair. Lolver communities; f. 0.36; C. 0.33, Phylum Nemertea a. 127,46-889. Class Enopla Spl~nerosyllis sp. A. cf. S. pirifera Tetrastemtna sp. Identification to genus Clapar&de, 1868. Lailti?iaria cotn- by E.N. ICozloff of specimens col- munity;f. 0.8, c. 0.33, a. 5, 1-9. lected by J. F. Palmisano. Sphaerosyllis sp. B. Alarin-Hedo- Phylum Nelnatoda pltyllt~mcomnlunity; f. 0.66, c. 0.17, Unide~ltified species. In the intertidal a. 54, 1-64. area, nematodes were most llulllerous ?Syllis (Typosyllis) nltet-rlntn h,loore, among the boldfasts of Lnv~itmrin 1908; Banse and Hobson, 1974. loiigi/)es and rllnria crispn, reaching Upper con~munities;f. 0.5, c. 0.09, a. densities as high as 261 per 0.062 m2. 202, 24-380. Syllis (ryposyllis) articillaris (0.F. Phyl~rmAunelida RfK~ller,1776). Identified by R. h'l. Class Polychaeta Har,iiotl~oeiti~briccrta (Linnaeus, 1767). O'Clair. Lorver Communities; f. 0.5, c. 0.4, a. 30, 5-318. 'iYidely distrib- Collected and identified by K. utcd. ICimura. Lepidoilotus I~elotypzts(Grube, 1877). Syllis (Typosyllis) stezoarti Berkeley, Identified by R. h,I. O'Clair. Sublit- 1942. Verified by R.h'I. O'Clair. toral, 183 m. Hn1osaccio1~-Fttclrs conlmunity; f. Plzoloe mi7zzcta (Fabricius, 1780). Veri- 1.0,~.0.09, a. 2. fied by R. b1. O'Clair. Lnttiiiiarin com- Syllis (T)~posyllis)sp. Verified to sub- munity; f. 0.8, c. 0.33, a. 50, 22-123. genus by I<. Banse. Lal?zit~ariacom- Dyspo?tetvs p)rg?tlaeirs Levinsen, 1879; munity; f. 0.52, c. 1.0, a. 90, 9-668. Ushakov, 1955. Lnmi~mria com- Trypnt~os)~llisgetit?t~iparn Johnson, munity; f. 0.8, c. 0.33, a. 50, 11-88. 1901. Identified by R. hs1. O'Clair. Arctic-boreal. Sublittoral, 34 m. The specimen Eitpl~rositie Dorenlis Oersted, 1843. examined is 315 mm long and 5 mm Identified by R. M. O'Clair. Sublit- wide with 596 setigers. To my knowl- toral, 34 m. edze it is the largest recorded speci- Eteone flnva (Fabricius, 1780). Identi- men of this species. Deposited in ficd by R, h81. O'Clair. Lai~tinnrincom- Nh4NS. munity; f. 0.8, c. 0.33, a. 40, 23-56. Nereis pelagica Linnaeus, 1758. Identi- Ezrlnlin (Ezclnlia) viridis (Linnaeus, fied by R. &I. O'Clair. Lnnzitiarin com- 1767). Lao~illarincommunity. Widely munity; f. 0.42, c. 0.47, a. 4, 1-62, distributed. and sublittoral to 183 m. Ei~lnlin(Ezrnrirla) longicor~i~ctah,Ioore, 1906. (Banse and I-Iobson, 1974). Nereis vesillosa Grube, 1851. Identified fllnria-Herlophyllt~it~ comn~unity. by R. hc1. O'Clair. Collected by K. Ki- North American. mura. Intertidal. Arctic-boreal. ~\~oto/)l~yll~t~~~sp. Identified to genus by Nepl~tyssp. Identification to genus by R. M. O'Clair. Sublittoral, 64 111. R. h'l. O'Clnir. Sublittoral, 183 m. Phyllodocidae sp. La~~ti~znriacommu- S/~l~crerodoro~sissp., near S. ttiitiuta nity; f. 0.57, c. 0.33, a. 1. (Webster and Benedict, 1887). Lower A u tolytits pristt~aticus (Fabricius, communities; f. 0.35, c. 0.67, a. 62, 1780). Identified by R. h'I. O'Clair. 2-381. Hn1osaccioi~-firczts comm~rnity. Eunice kobiei~sish,IcIntosh, 1885. Iden- A u tolytirs beritigiail~ts Annenkova, tified by R. M. O'Clair. Sublittoral, 6 1934; Ushakov, 1955. I.an~btarin to 270 m (see Gustus, 1972). d.lnrirte Iitvertebrates in Rocky Intertidal Contttttr~tities 441 Lzr 17tbrbteris inflatu Moore, 1911. twelfth notopodia; body as in C. Identified by R. &,I. O'Clair. Lanti- Type I (0.Hartman, personal corn- ttnria community. See Tablc 2 for muni cation). Alarin-Hedoplt)~Ilz~t~~ population statistics. Widely distrib- community; f. 0.38, c. 0.42, a. 54, uted. 1-762. A'aineris qtmdricttspida (Fabricius, Cirrattrltts Type IV. Acicular uncini 1780). Identified by R. ht1. O'Clair. present from first notopodia and Latni~taria community; f. 0.44, c. from sixth to eighth neuropodia; 0.33, a. 37, 25-454. Arctic-boreal. tentacular cirri sinlilar to C. c. spec- Polydoru sp. (see Hobson and Banse, in tabilis. Included with C. Type 111 in preparation). Lnttti~zuria community; Figs. 6 and 12. Lami~tnria commu- f. 0.66, c. 0.33, a. 38, 23-117. nity; f. 0.5, c. 0.33, a. 1, 1-23. Bocca~dia pro boscidea Hartman, Cirmtttltrs sp. Alaria-Hedo/?lt)'lItt??t 1940. Identified by R. M. O'Clair. community; f. 1.0, c. 0.08, a. 4. Alaria-Hedophyllttn community; f. Cirratulidae sp. Upper communities; f. 1.0, c. 0.08, a. 1. North American. 0.5,~.0.09, a. 96, 64-127. Pygospio eleguits Clapar;de, 1863. Brada graittrlata hlalmgren, 1867. Iden- Identified by R. AcI. O'Clair. Alavia- tified by R. hl. O'Clair. Sublittoral; Hedopltylltetn community; f. 0.6, c. 183 m. 0.25, a. 212, 54-508. Widely distrib- Flabelligera i~tjitnclibttluris Johnson, uted. 1901. Identified by R. Ac1. O'Clair. Spio filicorttis (0. F. Miiller, 1776). Lanri~mria community. North Ameri- Identified by R. AcI. O'Clair. Upper can. communities; f. 0.5, c. 0.09, a. 20, Sculibveg~~zuittfluttt~~t Rathke, 1843. 1-38. Widely distributed. Identified by R. hl. O'Clair. La~iti- Chaetopterns varioperlutus (Renier, ~tariucommunity. 1804). Identified by R. O'Clair. Capitella cu/)ituta (Fabricius, 1780). Sublittoral; 2 to 183 m. Identified by R. M. O'Clair. Aluriu- Caulleriella alata ntaczclutu (Annenkova, Hedophyllto~tcommunity; f. 0.38, c. 1934). Verified by R. Acl. O'Clair. Sce 0.42, a. 127, 1-444. Table 2 for population statistics. Abure~ticolu cluparedii oceu~ticnHealy Chaetozotte sp., near C. bevkeleyonolt and Wells, 1959. Verified by R. hc1. Banse and Hobson, 1968. Upper O'Clair. Intertidal in moat (see Fig. 3 communities; f. 0.5, c. 0.09, a. 212, in Lebednik and Palmisano, Chap. 17, 169-254. this volume). North Pacific. ?Cir.ntulzts cirrattrs spectubilis (I(i11- Praxillella praeteri,tissn (hlalmgrcn, berg, 1866) (Hartman, 1969). Lunti- 1866). Identified by R. AsI. O'Clair. ttaria community; f. 1.0, c. 0.06, a. Sublittoral, 34 m. 20. North American. Idur~tltgvstts arlnatus Kinberg, 1867. Cii.ratzclus Type I. Determination by 0. Identified by R. h1. O'Clair. Sublit- Hartman. Near C, c, spectubilis. Acic- toral, 55 ~n. ular uncini present from the first /I sa bellides sibirica (\Yirkn, 1883) notopodia and neuropodia; body [= Pseudosubellides littoralis Berkeley short with about 40 setigers (0. Hart- and Berkeley, 19431 (see Hartman, man, personal communication). 1959, p. 492). Laini~tariacommunity; Aluria-Hedophylltm comnlunity. f. 0.57, c. 0.33, a. 38, 38-276. Civratultts Type 11. Determillation by Amp/zigletta pucifica Annenkova, 1934. 0. Wartmail. Near C. c. spectabilis. Identified by R. hil. O'Clair. Upper Acicular uncini present from first comm~unitics;f. 0.5, c. 0.56, a. 127, notopodia and neuropodia; body long 27-1562. Endemic. The type locality t\rith 50 setigers (0.Hartman, per- of /I. pacifica is Bering Island, not sonal communication). ITalosacciott- Bering Strait as listed in Hartman Fttctcs community. See Table 4 for (1959). population statistics. Cltotte cittctn Zachs, 1933. Identified Cirrattcltts Type 111. Determination by by R. h3. O'Clair. Alaria-Hedop1t)rl- 0. Hartman. Acicular uncini present lunz community; f. 0.46, c. 0.83, a. from first neuropodia and from 190, 4-1544. Asiatic. 442 O'Clair C. itrftt!tdibtcliforuris ICrGyer, 1856. Hnlosnccioir-Fzicus community. See Identified by R. h'I. O'Clair. Sublit- Table 4 for population statistics. toral, 34 In. North American. Dci~to~tns?sp. Lnminarin co~nmunity. Collisella strigntella (Carpentel; 1864). See Table 2 for populatiotl statistics. Verified by J. N. McLean. Hnlosnc- fibricin snbefln (Ehrenberg, 1836). cion-Fzcctis community; f. 0.59, c. Identified by R. h'l. O'Clair. illnrin- 0.36, a. 4, 1-36. North American. Hedo/~h)dl~oncommunity; f. 0.41, c. Notoncnren scuttrnt (Rathke, 1833). 0.58, a. 3594, 2329,274. Lantinaria community. See Table 2 11f)~xicolninfirndibztlt~nt (Renier, 1804). for population statistics. North Identified by R. h'l. O'Clair. Sublit- Pacific. toral, 34 m. Rltodopetnla rosea (Dall, 1872) Potnrnilla rcniforntis (Leuckart, 1849). (= ~Vaccllarosea Dall, 1872). Verified Identified by R. h.1. O'Clair. Sublit- by J. H. McLean. This species was toral, 34 In. usually found on the holdfasts and Snbella crnssicornis Sars, 1851. Identi- fronds of Lantinaria yeroensis. One fied by R. h'l. O'Clair. Sublittoral, specimen collected in May 1974 was 34 m. brooding young (404 individuals) in Desiospira scmidentntn (Bush, 1904). its ~lilchalcavity. Other adults in the Identified by R. hf1. O'Clair. Lnnti- vicinity of this specimen at tlte time rtarin community; f. 0.66, c. 0.33, a. of collection were surrou~lded by 1, 1-24. North Pacific. many tiny, apparently newly re- Dcsiospira sp. Laininarin cotnmunity; f. leased, limpets. Asiatic. 0.62, c. 0.67, a. 32, 11-6632. Diadora sp. Sublittoral, 6 111. Class Oligochaeta (All Oligochaeta identified by Pzcnctzrrella (Puncturelln) noachinn D. G. Cook.) (Lintlaeus, 1771). [= P. longifissn Dall, Pnranais littornlis (hli~ller,1784). Halo- 1914.1 Verified by J. H. McLean. sacciorr-Fucics con~munity. See Lnntinnrin community. Arctic- Table 4 for populatio~lstatistics. boreal. Tubificidae sp. Immatore specimens. i\fargarites beringcnsis (Smith, 1899). Laminaria community; f. 0.57, c. [= 1t.1. helicinus ercavnttcs Dall, 1919. 0.67, a. 38, 1-124. J. H. McLean (personal communica- Lto~tbricillas sp. ~llaria-Hcdoplr)1I1tr~~t tion).] Lanrinaria community; f. community. See Table 3 for popula- 0.46, c. 0.6, a. 4, 1-468. Arctic- tion statistics. boreal. Phylum h'lollusca (Oldroyd, 1924-1927). 1 vorticifcrn (Dall, 1873). Idetltificd Class Gastropoda [The nomenclature of species by J. H. AtcLean. Lniirinaria com- of the superfa~~liliesPatellacea munity. and Fissurellacea generally fol- Aioelleria qundrae Dall, 1897. Veri- lows h'1cLezm (1966; 1969)l. fied by J. H. A4cLean. Laininnrin Problacnraea s)rbaritica (Dall, 1871) community. North American. [= Ac~naca(Tccttrrn) syba~Onchidella borealis Dall, 1871. Com- 1848). Sublittoral, 6 to 30 m. pared with USNA4 specimens. Tro/~laono/~sis(= Boreotroplzon) sp. Alaria-lledop/~ylltr~)t community. Sublittoral, 18 to 38 m. North American. Tlrais lima (hlartyn, 1784). Alaria- Class Amphineura Herlo/)hyllr~t~tcommunity; f. 0.48, c. Cry/)toclzito~t stelleri (Middendorff, 0.52, a. 4, 1-16. North Pacific. 1846). Shell plates found by L. Sotvl. Bttcci~zzon baeri )norchia~tunz Fisclier, Lepiflocltitona aleutic~ts Dall, 1878. 1858. Identified by A. M. Iceen. Compared with USNh,1 specimens. Lower coinmunities; f. 0.41, c. 1.0, a. most abundant in the upper com- 45, 1-296. Asiatic. munities. It appears to be ecologically Katltari~ta tlo~icata(\'\'ood, 1815). similar to Lasaea cistrcla Iceen (see AIaria-Hedopl~)~llnt~~.community. Glyin~, 1965). Ttcrtol~ia nri~ltcta is See Table 3 for population statistics. similar in size and morpl~ologyto L. North Pacific. cisttcla. Both spccies are ncstlers, Atnic~tlasp. Subtidal, 37 m. although T. vri1ttcta appears to prefer ~Wopaliasp. Sublittoral, 3 in. areas of high siltation. Both species Naci/~lrorellasp. Subtidal, 5 to 24 In. brood their young. Scl6i~0p1a.x bra~~dtii(Middendorff, Serripes groet~la~~dicits(BruguiPre, 1846). Alaria-lIedophyllrtt~t com- 1789). Sublittoral. Shell only. munity; f. 0.45, c. 0.36, a. 3, 1-42. Spisula poly~tyt~~a(Stimpson, 1860). North Pacific. Sublittoral. Arctic-boreal. Ha~lleya? sp. Latni~~ariacommunity. ?Silirllta media Sowerby, 1839 [= S. Class Bivalvia alto? (Brodcrip and So~ver 11.lodiohts modioltcs (Linnaeus, 1758) by, 1829) (Abbott, 1974)l. Shell (Soot-Ryen, 1955). Sublittoral. only. Sublittoral. 1l.ltcsc1~1tcsdiscors var. laeuiynt~ts(Gray, Macottta calcaren (Gmelin, 1791). Iden- 1824). Idciitified by A. bl. ICeen. tified by A I I

Psetrdopnlleite circularis (Goodsir, Order Tanaidacea 1842). Subtidal, 24 to 30 m. Taimis alascei~sis Richardson, 1899. Pltosichiliditti~t feittowttr~it (Rathke, Identified by G. J. Tutmark. Sublit- 1799). Snbtidal, 6 to 183 m. toral. Endemic. Achelia 1atifi.011~(Cole, 1904). Lanti- Anatnitnis ? sp. Lower communities. i~nriacommunity. Tanaidae sp. kllaria-Hedof)hy/ltr??~ A. pribilofeiuis (Cole, 1904). Lo\ver community;f. 1.0, c. 0.08, a. 1. intertidal con~munities. Order Isopoda (Richardson, 1905) (Schultz, A. alaskeitsis (Cole, 1904). Lai?tiimria 1969). community. Li?it?toria ligitortri~t Rathke, 1799. See A. sp. 1. Lai~tir~nrinconnnunity. Richards and Belniore (1976) for A. sp. 2. Subtidal, 24 in. collection data. A. sp. 4. Subtidal, 12 tn. Exos/~ltaeromn a~~tplicnitrln(Stimpson, Colosseildeis orientalis Losina-Losin- 1857). Alaria-Hedo[>hyllui?t com- sky, 1958. Subtidal, 274 m. munity; f. 0.39, c. 0.82, a. 4, 1-96. Rl~of~alorl~y itclttts c/~itiitostcin(Nilton, North American. 1943). Subtidal, 183 m. G~~orii~tospltaeromaoregotieftsis (Dana, Pycitogo~tzct~tcrassirostre Sars, 1888. 1855). Halosaccioi~-Fuczcs commu- Lalitillaria coinlnunity to 24 m. nity; f. 0.83, c. 0.09, a. 1, 1-5. North P)~ci~ogo~tuiitcf. P. ungellattam Loman, Pacific. 1911. Subtidal, 183 m. Dy?~amettellasheareri (Hatch, 1947). Class Crustacea Alaria-Nedopl~yll~cin community; Subclass Copepoda f. 0.43, c. 0.5, a. 310, 32-1026. North Order Harpacticoida American. Unidentified species. Nearly all indi- ?Arcttcrtcs loitgis/)ittus (Benedict, 1898). viduals passed through the 1-mm Sublittoral, 183 m. sieve. Idotea (Idotea) ochotensis Brandt, Order Notodelphyoida 1851. Verified by E. Rafi. Lnntiitaria Blnkeaints cor,~(qerWilson, 1921. Idell- community. North Pacific. tified by P. L. Illg. I11 the branchial Ido tea (Peittido ten) zuosneseitskii basket of Styelidae sp. Lower coni- (Brandt, 1851). Verified by F. Rafi. munities. Alaria-fido/)hyllz~ti~ contmunity; f. 0.36, c. 0.84, a.6, 1-74, North Subclass Cirripedia American. Bala7ttts cariosus (Pallas, 1788) (Pilsbry, Iai~iropsiskittcaidi derjtrgijti Gurjanova, 1916). Alaria-Hedo/>/tyll~c~t~com- 1933 (Rlenzies, 1952). Lniitiitaria munity. See Table 3 for populatioil community. North Pacific. Females statistics. North Pacific. collected in October 1971 were carry- B. gla~tdtcla Dar\\rin, 1854; Pilsbry, ing eggs and young. 1916. Halosnccio~t-Fttcus commu- Ligia pallasii Brandt, 1833. Supralit- nity; f. 0.64, c. 0.17, a. 3, 1-1760. North American. toral fringe, North American. Lepas attatifera (Linnaeus, 1758). On lVltr~rna(~\'eoi~tuilita) ste/)lteitseili Gurja- debris washed ashore. nova, 1933. L(cmr'ttnrin community; f. 0.8, c. 0.33, a. 2, 1-2. North Pacific. Subclass hIalacostraca ?11'11cituasttbite,~lectn - Gurjanova, 1936. Order Nebaliacea Ln~itiitariacommunity; f. 0.8, c. 0.33, Nebalia sp. Lalitinaria community. a. 44, 1-88. Asiatic. Order Mysidacea ~Vlt~nttnsp. c. f. I\{. (~\'eoin~ot~ta)ar~~Itol(Ii Gurjano\~a, 1933. Lariliitnrict corn- Gitathop/tattsia gkas \Willemoes-Sul~m, munity; f. 1.0, c. 0.33, a. 1. 1875. Identified by G. J. Tutmark. Ordcr Amphipoda Pelagic. Odirts sp. La?tii~?arincommutlity; f. Q8, Parucnntltomysis ktoileitsis Ii, 1936. c. 0.67, ;I. 1. Ideiltification by C. I-Iolmquist of Ainplrithoe sp. fllnrirr~-Hedophylltt~it specimells collected by P. N. Slattery. conimunity; f. 0.38, c. 0.75, a. 42, Neritic. 1-500. Callio/~iella sp.A. Collected by P. N. See Table 3 for population statistics. Slattery. An undesc~ibed species North American. (Slattery, unpublished manuscript). Allorcltestes sp. Near A. carinata I~vasa, CalliopieNa sp.B. Collected by P. N. 1939. Mature males unavailable for Slattery. An undescribed species study. Lantinaria community; f. 0.6, (Slattery, unpublished manuscript). c. 0.06, a. 1, 1-2. Calliopiella sp.C. Collected by P. N. Parallorcl~estes ocl~ote~uis(Brandt, Slattery. An undescribed species 1851) (Barnard, 1962). Lamittaria (Slattery, unpublished manuscript). commutiity; f. 0.49, c. 0.67, a. 8, Cnllio/)ieNa ? sp. Ha1osaccio1~-Fuctts 1-137. North Pacific. community. See Table 4 for popula- Ischgroce~xsType I. Verified to geilits tion statistics. by U. Lie. Lower commui~ities;f. 0.5, Calliopiidae Type I. IIalosaccioti- c. 0.2, a. 2, 1-10. Fttcus community. Ischyrocertcs Type 11. Lanti~taria com- Calliopiidae Type 11. Halosacciolt - munity; f. 0.66, c. 0.33, a. 2, 1.3. Flcctts community; f. 1.0, c. 0.09, Jassn sp. Verified to genus by U. Lie. a. 1. Lanti~taria coinintui~ity; f. 0.66, c. Calliopiidae Typc 111. Alaria-Hedo- 0.33, a. 16, 1-32. pltyllton community; f. 1.0, c. 0.08, Orclzo~ilene ? sp. Lat~tittaria commu- a. 1. nity; f. 0.5, c. 0.67, a. 100, 1-318. Coropltitc~~~brevis Shoemaker, 1949. Parapltoxtts spi~tosus Holtnes, 1903 Identified by U. Lie. Alaria-Hedo- (Bamard, 1960). Alaria-Hedoplt.)tl- phyllron comn~unity.See Table 3 for lirtn community; f. 0.46, c. 0.42, a. population statistics. North Anieri- 106, 13-381. Amphiboreal. can. Paraple~fstespugettettsis (Dana, 1853). Poitto.geiteia nndrijasclievi Gurjanova, Identified by U. Lie. Lantinaria com- 1951. Tentative ideiltification by munity; f. 0.57, c. 0.67, a.2, 1-4. E. L. Bousfield of specime~is col- North American. lected by P. N. Slattery (Slattery, Pletlstes sp. Lai~ti~la~iacommtui~ity. unpublished manuscript). Asiatic. S)~nt/>leustessp. IJerified to genus by P. ivnltovi Gurjanova, 1951. Tentative U. Lie. La~~tinariacomn~unity; f. identificatioil by E. L. Bousfield of 0.66, c. 0.33, a. 2, 1-85. specimens collected by P. N. Slattery ~l.Jesonteto/>a? sp. Verified to family by E. L. Bousfield. Lantittaria commu- (Slattery, unpublished manuscript). nity; f. 1.0, c. 0.33, a. 1. Asiatic. Parantetopella stelleri Gurjanova, 1948. ?P. i~tter~itecliaGurjauova, 1938 (Gurja- Tentatisely identified by E. L. Bous- nova, 1951). In pool on uplifted field. La~rri~tariacommunity; f. 0.8, c. rock bench, probably stranded in a 0.33, a. 1. Asiatic. storm. North Pacific. ?A'aji~a co~isiliovzrt~tDerzhasin, 1937 P. t~iakarouiGurjanova, 1951. Identifi- (Bulycheva, 1957) (Banlard, 1962). cation by E. L. Bousfield of speci- Verified to genus by U. Lie. Lai~ti- mens collected by P. N. Slattery. itctria community; f. 0.88, c. 0.33, La~itinaria community; f. 0.47, c. a. 2, 1-3. North Pacific. 0.27, a. 4, 1-69. Asiatic. Orchestia sp. Supralittoral fringe. P. rostrata Gurjanova, 1938. Identified Ca/)re/la lziizcaidi Cole, 1910. Lantiitaria by P. N. Slattery and confirmed by community; f. 0.66, c. 0.67, a. 1, \\I. S. Gray. North Pacific. 1-19. Asiatic. C. laevi~ascula h'layer, 1903 (Laubitz, Poittogeiteia sp. Confirmed to gellus by 1970). Ln~~tittariacomni~u~~ity. North P. N. Slattery. La~ititiariacommunity; Pacific. f.0.65,c.0.2,a.2,1-8. Cerco/)s compactus Laubitz, 1970. Eusiridae sp. I3alosaccio~t-Fttctrs com- Verificd by D. Lanbitz. Lai~ti~tnria munity; f. 1.0, c. 0.02, a. 1. community. See Table 2 for popula- Attisogaintitartcs coitfe~vicolt~s(Stimp- tion statistics. North American. son, 1857). Identified by U. Lie. Caprelln sp. Lat~tirtnria community; f. A laria -Hedopltyll~t~it commtuiity. 1.0, c. 0.33, a. 2, 1-2. llhritte Inuertebrates in Rocky Intertidal Coinn~tinities 447 Order Eupl~ausiacea(All eupl~ausidsidenti- P. setosus (Benedict, 1892). Identified fied by G. J. Tutmark.) by G. J. Tutmark. Sublittoral. T/i)~sa~toessalottgif~es Brandt, 1851. P. stevensae Hart, 1971. Identified by Pelagic. C. Nyblade. Sublittoral. T. itter~nis(lCr$yer, 1846). Pelagic. P. tr&o~~ocheirus(Stimpson, 1858). Eupltausia pncifica Hansen, 1911. Identified by C. Nyblade. Sublittoral, Pelagic. 55 In. Tessarobrachiou oculatznii Hansen, Cryptolitltodes typicus Brandt, 1849. 1911. Pelagic. See Barr (1973). North American. Order Decapoda Dertitatzcn~s iuartdtii Brandt, 1850 Sergestes sitnilis Hansen, 1903. Identi- (Makarov, 1938). Intertidal to 5 m. fied by G. J. Tutmark. Pelagic. Hapalogaster grebnitzkii Schalfce\v, Hymenodora frontalis Rathbun, 1902. 1892 (A,Iakarov, 1938). Sublittoral, Identified by G. J. Tutmark. Pelagic. 5 m. i\rotostomtts ja/)otticus Bate, 1888. Lithodes aeqzrispinn Benedict, 1894 Identified by G. J. Tutmark. Pelagic. (Makarov, 1938). Sublittoral, 91 to Pandaltts montagrti tridei~s Rathbun, 302 m. 1902. Sublittoral, 55 to 183 m. Oedignatltnts inennis (Stimpson, 1860). Leb beus groenlandica (Fabricius, Identified by G. J. Tutmark. Sublit- 1775). Sublittoral, 55 to 183 m. toral. North Pacific. Leb berrs gmndi~na~ttrs(Brashnikov, Paralitltodes camtschatica (Tilesius, 1907). Identified by G. J. Tutmark. 1815) (R,Iakarov, 1938). Sublittoral, Sublittoral, 5 m. 18 to 192 m. Ezcalzts fabricii (Ies (Sars, 1865) Callopora sp.A. May be a geographical (Fisher, 1911). Sublittoral, 192 m. form of C. Iiorrida (k1. Dick, personal Pteraster sp. Sublittoral, 6 to 12 m. communication). Sublittoral, 55 m. Aletctiaster scltefferi (Clark, 1939). Callo/)ora sp.B. Sublittoral, 55 n1. Identified by R3. E. Downey. La~iti- Bidenlza/>iaspitsbergensis var. alaskeiwis riaria community; f. 0.72, c. 0.4, a. 1, Osbum, 1953. Sublittoral, 55 m. 1-9. Speci~nens deposited in the Tegella sp. Sublittoral, 55 m. USNAfl and the British A,Iuseum fiicellnria erecta (Robertson, 1900). (Natural Ilistory). Lnntiitco.ia cornmunit)'. He~triciatzoiiidn Verrill, 1909 (D'yako- Uer~drobenrtia mrcrra)~n~ta (Johnson, not7, 1950). Laniiimria community; f. 1847). La~iliilariacommunit)~. 0.64,~.0.27,a.1,1-4. D. 111. var. fittiticosa (Packard, 1863). 11. levh~scltla(Stimpson, 1857) (Fisher, Sublittoral, 55 m. 1911). Lairtinaria community; f. 1.0, De~tdrobeartia sp. Sublittoral, 55 m. c. 0.06, a. 1. Iitvertebrates in Rocky Intertidal Cotittitttnities 449 Order Forcipulata endtooth of globifcrous pedicellariae Leptasterias alaske~tsis (Verrill, 1909) and the shape of the valves of the (Fisher, 1930). La11tiizaria commu- ophicephalus pedicellariae (see Mor- nity; f. 0.75, c. 0.06, a. 2, 1-2. tensen, 1943). Specimens collected L. aleutica (Fisher, 1930). Laminaria subtidally at Amchitka by NhrlFS community. See Table 2 for popula- biologists from Auke Bay, Alaska, tion statistics. I could not distinguish have been tentatively identified as S. small L. alaske~tsis from I,. aleutica polyacatlthus by D. L. Pawson (L. without exatnilling the pedicellariae. Barr, perso~lal con~n~unication).h,I. These two species were not distin- Jellsell verified several intertidal spec- guished in field observations; hoxv- imens. Lai~ziitarici community. See ever, Leptasterias in random samples Table 2 for population statistics. Also from the intertidal area were mostly sublittoral to 35 m. L. aletttica. De~zdraster? sp. Sublittoral. Class Opltiuroidea Class Holothuroidea Gorgonocephaltis calyi (Lyman, 1860) Cucttiitaria uegae (Th6e1, 1886). Veri- (Clark, 1911). Sublittoral, 128 m. fied by D. L. Palc'son. Alaria- ?Ai~t/~lti/~holissqttantata (Dellc Chiaje, He~lopltyllttn~community; f. 0.41, c. 1828) (Clark, 1911; D'yakonov, 0.92, a. 212, 1-4128. 1954.) Sublittoral, 34 m. Psol~issp. Sublittoral, 183 in. /l?nphipho/is si. Sublittoral, 18 1n. Ainpltiura sp. Sublittoral, 12 to 18 In. Phylum Chordata O/>hiopltolis actcleata (Linnaeus, 1767) Class Ascidiacea (Clark, 1911) (D'yakonov, 1954). Styela clavata (Pallas, 1774). Identified Lantiitaria community; f. 1.0, c. 0.06, by D. P. Abbott. Lai1zi1zaria commu- a. 2. Also sublittoral to 274 m. nity; f. 1.0, c. 0.13, a. 2, 1-2. Also Ophittra titaculata (Ludwig, 1886) sublittoral to 12 m. (Clark, 1911) (D'yakonov, 1954). ?S. coriacea (Alder and Nancock, Inf~alittoralfringe to 37 m. 1848) (Van Name, 1945). Lainiitaria 0. sarsii (Liitken, 1854) (Clark, 1911). conlnlunity; f. 0.88, c. 0.33, a. 8, Sublittoral, 91 111. 6-82. Opltittra sp. Sublittoral, 18 nl. Styela ? sp. (Van Name, 1945). Lanti- ?Opltiacatttha eniteactis Clark, 1911. izaria community; f. 0.57, c. 0.27, Some specimens have characters a. 23, 1-218. nearer 0. ~tutrixBaranova, 1954. The Aplidittnl Type I (Van Name, 1945). number of arms is variable. Several Lai~ti~tnriacommunity; f. 0.82, c. 0.2, speciillcns were brooding young, a. 1, 1-7 colonies. which to my knowledge is the first Aplidilri~t Type 11. Near A. trans- record of brooding in this species. hrcidttm Ritter, 1901 (Van Name, Sublittoral, 183 In. 1945). Lniitiitaria community; f. 1.0, Class Echinoidea c. 0.06, a. 1 colony. Stro~tg)~ioce?ttrotuspolyacaittltus Suborder Aplousobranchia Agassiz and Clark, 1907. Specinlens Unidentified species (Van Name, 1945). examined were identified prinlarily Lantitzaria community; f. 0.67, c. 0.13, on the basis of the size of the a. 1, 1-4 colonies. This page intentionally left blank

Marine fish Charles A. Sirnenstad Communities * John S. Isaksont Roy" E. Nakatani Fisheries Rc:search h~stitute,Utliversity of \Vashington, Seattle, \Vashington

Intensiue mtrltipla-gear saarpling of the fish fnrrnn occupy- n n~njorrole in tlre transfer of nlgne-bused detritus to the ing the marine zunters ndjacetrt to Anlcllitkn island sho~ued inshore fislr communities; eaphausiids contribute similnrly 92 fish species distributed over seven basic commu?rities to offshore fish commr~nities. The Pacific sand lance according to their characteristic hnbitnls-littoral, inslrore/ appears to be n significant importer of offshore-deriued rock-algae, ir~skorelsa~~d-gravel,offshore demersal/rock- +ooplanktoa bion~nssinto the inshore fislr contmu,~ities. sponge, offsl~oor demersal/saf~d-grauel, epipelngic, mzd Among the three underground nuclear tests at Amchitka, mesopelngic. Reclrrrext group m~olysisdisclosed 13 statisti- Canniki~zproduced the most sigtrificant impact by kills of cnlly distinct species nssocintiotrs sitl~i~ifive of these several fish species and changes b~ ltabitat through uplifting communities ns iuell as several distinct predator-prey of some littoralrock benches. Althouglr the recoueq of the 1elatioasl~ips.North Asrericnn species predominated in the affected fish communities appeared to be complete within a composition of the fish fauna, which reflects the principal yenr, the habitat and prey resources formerly provided by dispersal role of the Alnskn Streom carrent, zuhich flows the bench that is sow ttplifted will remain u,~uuailablefor westward nbost Amchitkn. Benthipelngic ninplzipods and many years. mysids are the predominant prey orgnnisms, ar~dboth serue

The Aleutian arcl~ipelago occupies a distinctive limits east\vard expansion of Asiatic species but zoogeographic position as a shallo~v-waterbridge also limits the \vestward cxpa~lsiollof North Pacific joining temperate Asia and North America. As a species. This filter bridge is further acce~ltuatcdby result the assemblages of ~narinefishes inhabiting Amchitka Pass [500 to 1000 In (275 to 550 the waters adjacent to Amcllitka Island are distinct fathoms) deep], \vhich separates Amchitka Island within ichthyofauna of the North Pacific and from the Aleutian Islands to the east and \shich is a within that of the Alcutian Isla~lds of Alaska. nlajor point of exchange betwee11 the North Pacific Although deepwater species common to Ocean and Bcring Sea water masses (kLcAIistcr, the Aleutians are distributed througl~outthe North 1971; hkAlister and Favorite, Chap. 16, this Pacific basin, the Aleutian fish fauna of nearshore volume). and intertidal \\raters is characterized predottli- The composition of Amchitka's fish faiuna also nantly by North America11 forms. This predomi- reflects the island's distinctive littoral and near- nance appears to be due to the Alaska Stream, the shore topography. The extensive wide littoral nearshore mest\vard-flowing surface current that benches and the broad sublittoral shelf on the originates in the Gulf of Alaska and eventually North Pacific Ocean side of the island provide loses its distinctness as it is diverted north and east suitable habitat for large populations of littoral and through the tllajor passes in the \vestern portion of inner sublittoral fishes. the chain (Fig. 1, Chap. 16). The tllaska Stream, rather than the east\\rard-flo\ving subarctic HISTORICAL REVIEW I),Jordan and Gilbert (1883; pelts, many Russian scientific expeditions carried 1899), Nelson (1887), Gilbert (1895), Tanner naturalists \\rho collected and described much of (1892-1896), Jordan and Starks (1904), aild the newly encountered flora and fauna. Sarychcf Evermanit and Goldsborough (1907). of the 1790-1792 Billi~tgsexpedition, Tilesius and U. S. Fish and Wildlife Service expeditions to Langsdorff of tlie 1803-1806 ICrusenstem ex- the Aleutians front 1936 to 1938 assessed birds pedition, Chamisso and Esclrscholtz of the and lrtanttnals and also studied marine fishes and 1815-1818 Kotzbuc cspedition, Volt Icitlitz, invertebrates (Schcffer, 1959). Tltese investigations Postels, and i\,Ierteits of the 1826-1829 Lutke documented 48 species of fishes indigertous to the expedition, and Vosnesensky, on an expeditio~l archipelago and included descriptions by Schultz sponsored by the Leningrad h'luscum, all contrib- (1939) of a ne\v genus and two new species. uted significantly to the natural history of the The most coinprchc~isi~~cdescription of Aleutians and Beri~tgSea. Ichthgofaunal specime~ts Aleutian ichthyofauna esisti~igat the start of our collectcd by Steller and succeeding Russian natu- xvork was that by Wili~novsky(1964); he described ralists were for~ually described in the scientific the distribution of 135 littoral fishcs throughout literature by Tilesius, Pallas, Cuvier, Shmidt, and the length of the tlleutian chain. Earlier, others (Jordan and Gilbert, 1899), the most Wilimovsky (1954) had listed 222 marine fishes comprehensive \\rork being that of Pallas [Zoo- from "all waters south of the Beri~tgStrait and grnplzin Kosso-Asiriticn (1811-1842)j and Sltmidt north of the Alaska Peninsula, including the [Pisces ~IilnritintO?ie~~talizoit (1904)l . Aleutian chain, Pribilof Islands, St. h,Iatthele Scientific iit\~cstigatioitsof Alaska and adjacent Island, and St. Lawrence Island," and had pro- waters were instituted by tlie U. S. go\~ernmeitt duced a pro\risional key to 241 species occurring soon aftcr the purchase of Alaska front Russia in \sititin that range or Inore generally in the North 1867 (see Fuller, Chap. 7, this volume, for a more Pacific Ocean (!\'ilimovsky, 1958). A sitrvey of co~ttpletediscussion of scientific investigatioits in Antcbitka's lvildlife was also co~tductcdby Fish the western Aleutians after 1867). William H. Dall and \,Vildlife Service biologists during the summer was in the first U. S. Coast and Geodetic Sulvey of 1962, in which they recorded 12 a~tadromous hydrogfi~plticand geographic reconnaissance of the and miwine species and several ~~~iidelttified Aleutia~tsfront 1871 to 1874, during ~vliichtime blennies, sculpins, liparids, a~tdflounders (h,lcCann, he collected faunal specimens for deposit in tlte 1963). U. S. National h~Iuscum(Jocltelson, 1925); he Tvas Seventeen marine and anadroinous fish species the first American naturalist to study Amchitka's \$,ere observed in 1965 during the bioenviron- nlariile fisltcs (Dall, 1874 aitd 1877). From 1880 ntental safety program associated with the Long through the mid-1890s, the U. S. Coast Survey Shot nuclear test 011 Anichitka (Seymour and schooner I'rckon, tlte U. S. Treasury re\reilue Nakatani, 1967). stcaittcrs Corzuitz and ~\.loliicn?~,a~td the U. S. I.'isli Contemporary descriptive syntheses of North Co~trmissionsteamer Albatross cruised the North Pacific ichthyofauna, including soine of the species Pacific Ocean and Bering Sea waters making encou~ttered in these studies (Quast and Hall, h y drograpliic and ~tteteorologic observations, 1972; &at, 1973), have been used in the discus- patrolling for illicit fur seal hunters, a~tdcon- sioit in this chapter of tlie zoogeograpltical distribu- ducting biological collections. A. B. Alesander, ti011 of inarine fishes that occur around Amcltitka Tarleton I-I. Bean, William 13. Dall, Henry \V. Island. Elliott, Barton \\I. Evermann, Charles H. Gilbert, N. B. Miller, Charles H. To\snsend, aitd L. hfl. RECENT INVESTIGATIONS Turner were among thc notable collectors, natu- ralists, a~tdichthyologists accompanying these sur- The Fisheries Research Institute (FRI) of the veys; they ;$Is0 collected for the U. S, National Uttiversity of Wasliiiigton studied and evaluated the iVluscum. Included in the records from this period impact of two underground nuclear explosions ~lfarirteFish Coi,!t~!ntlities 453 (Milro\v on Oct. 2, 1969, and Cannikin on Nov. 6, \\?th a 10-cm (4-in.)* mesh cod end of 96-thread 1971) on the nlarilie fishes off Amchitka Island. nylon. Chafing xiiats protected the bottom and cod The FRI fish investigations \irere conducted from end, and rollers of various sizes were used on tlie the sumliier of 1967 to the fall of 1973 for the footrope, clepcuding on suspected bottom rougli- U. S. Atoniic Energy Co~nmissio~i(AEC) by sub- ness. The tiet was fished \vitIi Formosan-type otter contract \\'it11 tlie Colunibus Laboratories of boards that \veiglied 430 kg (950 lb) in 1967 and Battelle R'leniorial Institute (BCL). Reports of 340 kg (750 lb) in 1969 and later years. investigatio~ts\\.ere subniitted annually to BCI, aiid ~l,lidzuaterTrzul The niid\vatcr trawl \\,as a are summaxized in Nakatani and Burgner (1974). 27-m(90-ft)-long nlodified herring traxvl \\(it11 a Although previous information on ~nariucfishes 6-m(20-ft)-square opening \sliich \\,as fishetl with off Amchitka Islalid nras largely liniited to compila- 227-kg (500-lb) otter bo2irds at :tpproximately 4 tion of species check lists, FRI fish sampling also knots. \\'ebbing raliged fro111 7.5-cm (3-in.) \\rings deterniiued the distribution, relative abiindance, to 1.25-c~n (0.5-in.) cod end. A standard 3-111 and life-history aspects of fish species found in tlie (10-St) Isaacs-I

MATERIALS AND METHODS Salnzon Gill A'ets. The surface salmon gill nets were composed of six to ten 91-m(300-St)-long by 5.5-m(18-ft)-deep shackles of multifila~nentnylon Fishing Vessels that ranged in mesh size from 6.4 to 11.4 cn1 (2.5 In 1967 and 1969 tlie Ajl. 1'. Paragon, a 27-~n to 4.5 in.). (90-ft) tran~lcr,rvas used in the deeper ~ffshore Bottoin Gill i\rets. R,lonofilament gill nets were waters [depths greater than 38 m (125 ft)], and a 2.7 m (9 ft) deep with six 12-~n(40-ft, hung variety of s~nalloutboard boats \\,ere used in tlie nleasure) panels of 2.5-, 3.8-, 5.0-, 6.4-, 7.6-, and nearshore \\raters. Fro~li1970 to 1972 the M. V. 10-cm (I-, 15,2-, 2.5-, 3-, and 4-in.) web and nritli Con~ii~ai~der,a 26-111 (85-ft) purse seiner \vith 21 lo\\,-buoyancy float line, \vhich allo~vcdtlie net multiple-gear citpabilities, \\'as used principally to sink to tlie bottom. duriug fall months. Larger inboard-outboard boats were available during the latter period, ~vhich Swrface Gill ~\'ets. R~lonofilament surface gill permitted nearshore sampling at greater distances nets were identical to the sinkitig gill net except fro111boat-launching sites. that they contained niorc floats. The offshore sampling from the research \~cssels Tra~mirel~\rets. The tratnniel nets used lvere included bottoni and mid\sater tra~vling, pnrse nlade of green-dyed nylon outer \rails of 50-cm seining, saln~o~lgill-netting and long-lining, bottom (20-in.) web and inner \valls of 10-cm (4-in.) web dredging, and the fishing of crab traps. Nearshore aud were 91 m (300 ft) long and 1.8 111 (6 ft) deep. sa~iiplitigi~tcludcd the use of bottom and surface They \\,ere \veigl~ted to siuk but \\,ere buoged experimental gill nets aud tra~nnlelnets, bottom sufficiently to hold them vertically above the and surfacc longlincs, shrimp, crab, minno\v, and bottom. antphipod traps, plankton nets, rotenone, aud misccllaueous hook-and-line and hitlid collections. Snl~~~o~~Lo~~gI:li~~es.s.Japanese surface salnloli longlincs \\,ere used to catch salnion offshore during the 1967 through 1969 sampling period. Gear Types This gear generally consisted of 10 skates, each Bottoin Trazul. The bottoni tranrl, the priniary tia 22.6-kg test vinylon niain line that \\.as offshore san~pliliggear, \\'as a 400-mesh t~vo-seam eastern bottom (otter) trawl with a 22-111 (71-ft) headrope and a 29-m (95-ft) footrope constructed *All mesh sizes are stretched measure. BERING SEA

Sea Otter Point BERING SEA

PACIFIC OCEAN

Sand Beach Cove Crown Reefer Point

0 12345 PACIFIC OCEAN KILOMETERS GRAPHIC SCALE

Rifle Range Point AMCHITKA ISLAND

Fig. I-Location of coastal features of Amchitka Island, Alaska. ~VlarilzeFish Co??znrt~nities 455

approximately 138 111 (450 ft) long with 49 eve~lly nets were of standard 0.5-nl (2-ft) No. 3 mesh spaced 12.3 nl (7.5 ft)] size-8 hooks on (0.324-mm apcrture) with a flowmeter attached. I-m(3-ft)-long 10-kg (22-lb) test monofilament Both vertical and horizontal tows were made. leaders. Ten cvc~llyspaced \\rooden floats buoyed Rotenoite. This poison was applied in 500-g each skatc. Thc bait used \ires 7- to 10-cm(3- to warmwater-paste batches to isolated tide pools 4-in.)-long dried salted anchovy (E~zgractlis \vlierc suitable concentrations could be maintained ja/>o~cica).'The total length of the longline set was and confined; larger tide pools required two approximately 1.4 km (0.75 nautical mile). batches. Tide pools selected rangecl from less than Surface Longli~tes. Surfitce longlines fished 2.8 m3 (100 ft3) to over 5.7 1u3 (200 ft3) in nearshore were made from standard 49-hook Japa- volumc, differed in algal cover and substrate, and nese salmo~ilongline gear with floats spaced five yielded a \vide diversity of species. hooks apart. Octopus was most often used as bait. ~Vliscella~teotts:Iloolz-a~zd-Line and Hand Col- Bottonz Lo~zglbzes. Longlines consisting of the lections. These sampling methods were prodoctivc standard 49-hook Japanese salmon lo~igli~tewith- but were colisidered incidental and more selective out floats or a he;~v~~-duty8- to 10-hook anchored than other sampling procedures. IIancl collections halibut longline were fished on the bottom ill included fishes capturcd during scitba surveys with nearshore waters. Squid or octopus was usually a "slurp gun" or by hand during i~ivcrtebrateand used as bait. algae collections in the littoral zone. /Inchor Dredge. An anchor dredge with an 89- by 38-cm (35- by 15-in.) opening from the inclitlcd Sampling Schedules and Areas plate into the 10-cm (4-in.) nylon 96-mesh col- Sampli~igschedules varied \\fit11 the type of gcar lecting bag was cast from the offshore vessel alld used. Offshore bottom tra\vlitig and purse seining rctricvcd after digging into the bottom or briefly were usually done duriltg the day and midwater to~vedalong the bottom surface. Se\feral fish \\'ere trawling at night, An echo sou~tderwas used during caught with the be~lthos brought up with this bottom trawling to detect the prescnce of fishes, to dredge. select the fishing depth ;old sititable bottom types Slrrin~/?Trafls. Experimental shrimp traps [45 for nearshore set net fishing, and to locate fishes by 45 by 75 cm (18 by 18 by 30 in.)] were uscd in duri~lgmid~vatcr tra\vli~tgand purse seining. Sur- water 3 to 60 In (10 to 200 ft) deep. Fishes caught face sal~no~tgill nets were fished at night, and nearshore \\rere uscd as bait. Several species of fish, surface salmo~ilo~tgli~les were usually fished for a ~vliichwere probably attracted to the amphipods 1- to 3-lir period before daybreak. that were feeding on the bait, were captured in the Night sampli~lg in small boats was not con- traps. sidered safe. Therefore most ~learshorcsampling took place during tlic day ~vhenweatlier allo\ved. Crab Traf,s. Comniercial king crab pots [1.8 Nets a~idlonglines were commo~tlyfished over- m (or 6 ft) in diameter] were used in 1967 but in night except during winter and spring sampling later years were replaced by 86-cm(34-in.)-di;tme- \vlien the unpredictable weather conditions limited ter Taylor crab traps.,Nearshore fishcs were uscd as sets to 2 to 5 hr during daylight. Thus fishing time bait. Fishes and varlous invertebrates as \veil as per set varied from 2 to 24 hr, depending 011 safety crab were captured in these pots. considerations, the gcar type, and season. O\ving to ~VJi~z~zozoTraps. Comn~ercially built polypro- the destructive predation of captttred fishes by pylene traps, 43 cm (17 in.) long by 23 cm (9 in.) amphipods, bottom-set nets could not be fished for in diameter with 2.5-cm (1-in.) openi~igsin each more than a couple of hours when stomachs or end, were baited with nearshore fish alld usually other critical salliples were to be retained. anchored in littoral pools. Co~lstraintson the sampling program included thc selectivity of the fishing gear; inability to fImphi/?od Traps. Prior to the initiation of sample at all seasons, particularly offshore where plankton pump sampling, amphipod traps made an expensive chartered vessel was necessary; a11d from 3.7-liter (1-gal) wide-mouth nalgene bottles selection of sit~nplingsites near undergrou~~dnu- with an inverted cone opening were used to collect clear test sites, which concentrated nearly all scavenging crustaceans. Bait included octopus and marine fish sampling effort around tlic southeast ~tcarshorefishes. A few small fishes were captured part of Amchitka Island. Thcse limits should be along \vith the invertcbrates. co~tsidered in tlic follo\ving discussion of Pla?zkton l\'ets. Plankton nets were used to Amchitka's marine fishes and in the interpretation collect fish eggs and larvae in nearshore areas. The of results. %~mpleProcessing. All catches \\,ere countcd tagged with green "spaghetti" tags ~neasi~ring3.8 (or total 5vcight estimated for largc tra\\,l hauls) cni (1.5 in.) long, and then released in the general and recordcd by species along with such inf~)rma- area of capture. Posters rcq~restingthe return of tion as date, depth, location, time of sct, and time tagged rock greenling from sport fisher~nenat of recovery. Tow and \\rind directions \irere also Amclnitka Island were displayed. recordcd for all tralvl catches. During the first fe~vyears of the program, all RESULTS AND DISCUSSION fishes captured in nearshore waters lvere retained for lifc-history, meristic, and food-\vcb studies. In Fish Occi~rrence later yczxs some species \\,ere retained for con- tinuing food-\veb studies whercas the rest were Table 1 lists the 92 identifiable species docu- comntcd and releascd alive. mented from Amchitka's nearshore and adjacent All the fishcs returned to the field laboratory offshorc marine \\raters. In addition, three species at Amchitka nrcrc ~neasuredfor total length to the of cottids xvcre identified from ichtlnyoplankton nearest nlillinleter and \\reighed to the nearest but do not necessarily represent established popu- gram. Sotnc specics of particular interest \Irere lations at Amchitka. preserved \vInole in 10% buffered Formalin or Contctnporary accounts of North Pacific fish frozen and returned to the Seattle laboratory as fauna [Quast and Hall (1972) and Hart (1973)l muscum specinicns or for further examination. were used to group the specics caught at Amchitka Ho\vever, most fishcs were processed in the according to thcir geographical distributions Amchitka 1;iboratory for such infor~natiotnas sex, (Table 2). According to these sources, nine of the gonad weight, age (otolith or scale), and stomaclt species were kno\crn to range throughout the sample. All fishes and accolnpanying infornlation ternperatc and torrid zones of the North Pacific- were identified by code uunlbcrs and sanlpling set nlostly epipelagic, mesopelagic, and bathypclagic niunlbers. Data \Irere trausfcrred to computer cards fishes ranging from near the sitrface to 3000 111 for easc of analysis. Several thousand fishes were (1650 fin) and deeper. Of the Amchitka species, 32 processed each year. have been recorded as occurring tlnrougltout the Three basic tasono~nickeys to fishes of the temperate* and subarctic waters of the North North Pitcific-A~ndriya~l~c\~(1954), Clemens and Pacific basin (amphi-Pacific) and 21 are confined \\'ilby (1961), and \\Tilin~o\rsky(1958)-were used to the subarctic water nnasses of the North Pacific to identify the fishcs collected at Amchitka. Occan and Bering Sea. T~venty-fourspecics have Further verifications \sere made from more specific previously been reported in the eastern portion of tasonomic keys in the ichtl~yologicalliterature. the temperate and snbarctic North Pacific only; of these, Raja trbncltz~rbn, Cor)~pltae~~oirlesfiliferbn, and Food-Web Studies. The description of the Rad~tli~t~lsas(~rellus had not bccn reported in the Amchitka lnarinc food \veb resulted from the Aleutians before the FRI Amchitka studies, and qiiantitati\,e atlalysis of the stomach contents from Cl~irolophis ttzcgbntor, Sebastes bbnbcocki, and ahnost 2000 fish spccimc11s obtained during the Glyptoceplzahts racl~ir~ishad not bccn recorded tlmchitka collections. The !vet weight, number, of Unalaska Island in the eastcrnnlost region and tasa of prey organislns and rank evaluations of of the archipelago. Three species arc considered to sto~~lacltfiillncss and digestion \\'ere deternlincd for occur universally in arctic and subarctic norther11 each stomach sample (Burgner ct al., 1969; hemisphere waters, one in north tcmpcrate waters. Simenstad, 1971a). Tlic ranr data were analyzed The longnose lancctfish (Alepisa~~~z~sferox) is specifically for the percentage of total weight of known throughout the tropic to subarctic regions stomach contcnts by food orgallism with fre- of both the Atlantic and Pacific Oceans and the quency of occurreuce and numerical abundance h'leditcrranean Sea. The toothed cod (Arctogadus providing supporting indexes of feediug habits. The bolisovi), previously knolvn from the arctic seas of ensemble of stomach sannplc data was analyzed North Anncrica only, was also captured durilng our \vith digital-computer programs dcsigncd specifi- bottom trawling at Amchitka, a significant south- cally to address the Amchitka data set. ern extension of its documented range. Three additional cottid species, Furciltn Tagging Studies. Prompted by the great abun- ishiknzuae, 0c)lltectes ~~tbnscl~a~ts,and Scor- dance of rock greenling (Hescryra~~a~~toslago- p(~e~ticlttl~ysntbn7?norat1rs (the cabezon), \\.ere iden- ce/)hnlus) in nearshore \\raters, in 1968 through 1971 we undertook to study their gro\vth and movcmcnts by tagging. Rock greenling captured in Temperate waters are here defined as extending to thc sampling uets were measured, weighed, sexed, 45'i\' latitude. ~VIari~reFislr Co~~z~~rri~zities457 Table 1-Marine Fishes Collected at Amchitka, 1967 Through 1972*

Scientific name Con~n~onname

Class Cyclostomi Order Petromyzontiformes Pamily Petromyzonidae Lampreys Entosphe~tsstridentnlus (Gairdner in Richardson, 1836) Pacific larnprey Class Cl1rondrichth)res Order Squaliformes Fanlily Lamnidae h,Iackerel sharks I,nmnn ditropis (Hubhs and Follctt, 1947) Salmon shark Order Kajiformes Fateily Rajidae Skates Rajo binocslnta (Girard, 1854) Big skate Rnjn parmifern (Bean, 1881) Alaska skate Raja tracl~urn(Gilbert, 1891) Iloughtail skate Class Osteichthyes Order Salmonifortnes Family Saln~onidae Saln~onand trout Oncorhynchas gorbuscho (!\'\'albaum, 1792) Pink salmon Oncorlryncl~usketn (\\'albaum, 1792) Cham salmon Oncorlrynchtzs kisutch (\\'albautn, 1792) Coho salmon 0acorhy1,ckus rterka (IYalbaum, 1792) Sockeye salmon 0,tcorhynchus tshawytscha (\\'albaum, 1792) Chinook salmon Snluelinas n~nlma(\Yalhaum, 1792) Dolly \larden I2amily Bathylagidae Deepsea smelts llnthylagus millerit uordan and Gilbert, 1898) Stout blacksmelt Hntlzylogus stilbius (Gilbert, 1890) California smoothtongue Pamily Gonostomatidae 1,ightfishes C~~clotl~oseinic~odortt (Gunther, 1878) \leiled anglemouth Fanlily Chauliodontidae Viperfishes Clmztliodus mncouni (Bean, 1890) Pacific viperfish Order Myctophiformes Family Alcpisauridae Lancetfishes /Ilepisnarus ferox (Lowe, 1833) Longnose lancetfish Family hlyctophidae 1,anternfishes Dinphus lhetn (Iiigenn~annand Eigenmann, 1890) California headlightfish Electronn arcticot (l,utken, 1892) Bigeye lanternfish Lnmpanyctt~sregalis (Gilbert, 1892) Pinpoint lampfish Stenobmchius leucopsnrrcs (Eigenmann and Eigenmann, Northern lampfish 1890) Tnrletonbennin crentrlnris Uordan and Gilbert, 1880) Blue lanternfish Order Gadiformes Family Gadidae Codfishes Arctogodus borisoui (Drjagin, 1932) Toothed cod Gndus ,nncrocephn/us (Tilesius, 1810) Pacific cod Tlterngra chnlcogrammo (Pallas, 1811) Walleye pollock Family lllacrouridae Grenadiers Coryphnenoides ncrole/>is~(Bean, 1883) Roughscale rattail Coryphnenoides filiferat (Gilbert, 1895) Filamented rattail Order Gasterosteiformes Family Gasterosteidae Sticklcbacks Gnsterosteus nculentus (Linnaeos, 1758) Threespine stickleback Order I'erciformes Family Trichodontidae Sandfishes li-ichodon tricltodon (Tilesius, 1811) Pacific sandfish Family Uathymasteridae Ronquils Hnthyrnnsler caeruleofnscintas (Gilbert and Burke, 1912) Alaskan ronquil Hnthp~nnstersignnntus (Cope, 1873) Scarcltcr Ro,zr/uilss jordani (Gilbert, 1888) Northern ronquil Family Sticllaeidac Pricklebacks iileclrins nlectrolophus (I'allas, 1811) Stone cockscomb A~~oplorclzt~spurprrrescens(Gill, 1861) High cockscomb Cltirolo/~hisnugotor (Jordan and \\'illiams, 1895) I\losshead warbonnet Gpiirnocliizus crisfulatus (Gilbert and Burke, 1912) 'Trident pricklcback

(Table continuer on page 458.) 458 Si?~ienstnd,Isakso~~, and ~Vakatn~ti Table 1 (Contiaued)

Scientific nalne Conunon name

Plrytichthys chirtrs (Jordan and Gilbert, 1880) Ribbon prickleback Porocli~tusrothrocki (Bean, 1890) Whitebarred prickleback Family Pholidae Gunnels Pholis dolichogaster (Pallas, 1811) Stippled gunnel Pltolis laeta (Cope, 1873) Crescent gunnel Family Ptilichthyidae Quillfisl~es Ptilichtlrysgoodei (Bean, 1881) Quillfish Family Zaproridac Prowfisl~es Zaprora silenus (Jordan, 1896) Prowfish Panlily Amrnadytidae Sand lances Artnnodytes hexapterus (Pallas, 1811) Pacific sand Lance Family Scorpaenidae Scorpiot~fislles Sebastes aleutianus (Jordan and I<\-ermann, 1898) Rougheye rockfish Sebastes alutus (Gilbert, 1890) Pacific ocean perch Sebastes babcocki (Thompson, 1915) Redbanded rockfish Sebastes ciliatus (Tilesius, 1810) Dusky rockfish Sebastes polyspirtis (Taranetz and hloisee\r, 1933) Northern rockfish Sebnstes proriger (Jordau and Gilbert, 1880) Redstripe rockfish Sebnstolo611salascanas (Bean, 1890) Shortspine thornyhead Family Anoplopomatidae Sablefishes A~~oplopomnfirnbri (Pallas, 1811) Sablefish Family Hexagrammidae Greenling Hesagramnros decagrannaus (Pallas, 1810) Kelp greenling Hexugrar~~moslagocephalus (Pallas, 1810) Rock grccoling Pleurogrammus monoptarygius (Pallas, 1810) Atka mackerel Familv Cottidae Sculpins ~ledsiascirrhosus (Pallas, 1811) Silverspotted sculpin Clinocottus acuticeps (Gilbert, 1895) Sharpnose sculpin Cli~rocottusentbryrrrn (Jordan and Starks, 1895) Calico sculpin E~ophrysdiccraus(Pallas, 1787) Antlered sculpin Gvrnnocnntkur paleatus (Bean. 18811 Armorhead sctrlpin ~;~~~~r~oca~ttlras~istilli~e~§(pillas, 1811) Threaded scolpin Herrtilebidott~shemile~idotus (Tilesius. 18101 Red Irish lord Yellow Iris11 lord Bigmouth sculpin Icelur cnnoliculatus$ '(dilbert, 1895) ~\lnlacocotteskincaidi (Gilbert and ?'hompson, 1905) Hlackfin sculpin ~\fyosocephnlr~spolyacn~~fhocepltal,rs (Pallas, 1811) Great sculpin ~\'aatichtltysichtltys~ribilouitrs(Jordan and Gilbert, 1899) Eyeshade scolpin Rnduli~r~rsnspreNtrs(Gilbert, 1890) Slim sculpin ,~.lrrglopsforficata (Gilbert, 1895?) Scissortail sculpin Triglops r~refopins§(Gilbe~t and Burke, 1912) ..lrrglops . pingeli (Reinhardt, 1832) llibbed sculpin Triglops scrpficus (Gilbert, 1895) Spectacled sculpin Family Agonidae Poachers Aeontrs acil~e~~seri~rrrs(Tilesios, 1811) Sturgeon poacher ~~~,idoplro;oidesbnrtdni (Gilbert, 1895) Aleutian alligatorfish Ilypsago~~asquadricornis (Cuvier, 1829) I'ourhorn poacher Pnllosi>tnbnrbnta (Steindachaer, 1876) 'l'ubenose poacher Sarritor leDtorltvrrchus (Gilbert. 1895) Longnose poacher Fanlily Cycloptcridae I.u~npfishesand soailfishes llptocycl~rsuortricosus (Pallas, 1770) Smooth lumpsucker Careproclss gilbertit (Burke, 1912) Smalldisk snailfish Careprocfusphasrtta§ (Gilbert, 1895) CrvstallicAth~~scvc/oshi/us (Gilbert and Burke. 19121 Blotched snailfish ~u.microtrent't~shis icunthcr, 1861) I'acific spiny lumps~tuker Letholremus nltlticus8-. (Gilbert. 1893) Lipuris callyodorr (I'allas, 1811). Spotted snailfish Libaris nreenceDltales6 (Burke. 1912) Tadpole snailfish 1.obefin snailfisll ~Vlarit~eliiS12 Cottl?~ttolities 450

Table 1 (Continued)

Scientific name Colrmon ,lame

Order Pletlro~lectifor~lles Family Pleuronectidae Righteye flounders /Itheresthes stomins (Jordan and Gilbert, 1880) Arrowtooth flounder Glyptoce/>Aolusrachirus (Lockington, 1879) Rex sole Hippoglossoides elnssodon (Jordan and Gilbert, 1880) Flathead sole Hil~~oeiossussle~toleDis (Schmidt. 1904) Pacific halibut .. lchthyoplankton specimens; not captured as adults: Order Perciformes Family Cottidae F~lrcinaishikazuae (Jordan and Starks, 1904)q Ocynectes maschn~rs(Jordan a~tdStarks, 1904)n Scorpnenichtltys mnrmomtus (Aycrs, 1854)

*Nomenclature and species arrangement from the American Fisheries Society (1970) and Green!%-oodet al. (1966) except +Hart (1973). $Renamed from P. thontpsoni (Chapl~~an,1944), Hart (1973). SQuast and Hall (1972). IJordan and Starks (1904). tified* from Amchitka ichthyoplankton. Ft~rci~m influencing the composition of Amchitka's fish ishilzawae and Ocynectes ?naschaus represent fauna. The occurrence of tmo exclusively Asiatic species hitherto bclie\red to be restricted to the cottids in our ichthyoplankton samples indicates, western North Pacific basin and unreported in however, limited immigration from faunal sources North America, including the Aleutians. Scor- to the west. Why these species have not becomc puetticlrthys Fttarmoratus is a tcnlpcratc eastern established remains a matter for speculation. Possi- North Pacific species not previously reported \vest bilities inclitde competition from already cstab, of the Gulf of Alaska. Although the identification lishcd species, en\~ironmentalconditions preventing of the larvae of the two Japanese species is maturation of larvae or juveniles, and predation. reasoi~ablypositive, the larval cabezon could be The distribution of Amchitka's marine fishes confused with some species of the genus A4yo.uo- according to the depth from which \vc collected cepltal~ts.In view of the lack of adult forms in the them is shown in Fig. 2. Complete dcpth rangcs arc collections, the existence of viable reproducing indicated by thin lines, depths of relatively high populations of these particular cottids in the abundance by the heavy portion of these lines, and Amchitka vicinity is doubtful. It1 a true ecological the location of infrequent captures by circles. sense, then, they are not confirnled as components Sampling methods produce biases in these patterns; of the Amchitka fish fauna. so they must be considered as only approximate. \Ve note that, although 24 spccics of fishes at (Fishes may occur at depths iunsampled, fishes may Amchitka are represcnted as being near the western change to other depths at other seasons, and some limit of their Alaskan distribution (including sis gears sample incompletely or sainple during ascent westward extensions of known range), none of the from or descent to the dcsircd depth.) species collected as adults at Ainchitka are cxclu- sively Asiatic species near the eastern limit of their Alllcllitka Fisll Co,lll,lilllities distribution. The distincti\le ~ositionof Amchitka Island anlidst the North Pacific cltrrent systems, Follo\ving \\'hittaker (1970), \\re define a biotic pointed out by \Vilimousky (1964) and discussed community as a i~atitralasselnblage of populations e.at .I' I~I . herein, appears to be an important factor that live together in a distinguishable habitat and interact <\pith each other, "forming togethcr a distinctive living-. system with its own composition, structure, environmental relations, development, *Taxonomic determinatio~~of ichthyoplankton speci- mens was made W,E, Barmclo,lgh, Fisl,crics Research and function." In earlier reports \ire discussed Board of Canada, Nanai~no Uiological Station, Wanairno, Amchitka fish communities subjccti\~ely on the B.C., Canada. basis of the species distribution according to four 460 Siuzenstad, Isakso11, and ~\'aknta~ti

Table 2-Zoogeographic Ilistributions of hlarine Fishes Collected at Anlchitka Island, Alaska, 1967 Througll 1973 * Distribution Species Ilistl.ibutioa Spccies . Tempcrate and sub- Oncorhy~tchusgo,.buschn Careproctus phnsmn arctic North 0.keta Crystallichtlt ys cyclospilus Pacific Ocean; 0.kisutch Eastern portion Rnjn trnchurat Japan to western 0. nerka of the telnperate Cor),phaenoidesfiIifera$ North America 0. tsltnwptschn and subarctic Ctt iroloplzis nugator$ (amphi-Pacific) Saluelinus mnlmn North Pacific, Ronquilrcs jordarti (32 species) Cltnuliodus ntncou~ti including the Raja binocrllntn Cndtts mncrocephal16s Aleutian Islands Anoplnrchus purptrrescens Tlterngro chalcopetnra (24 species) Gpntnoclinus cristtrlatus Trichodon trichodon Phytichthys chiras Rathy~nastersig~rntas Poroclines rotlrrocki Ptilick thys goodei Pholis laetn Znproro silenzts Sebnstes nleutianus Sebastes nltttus S. babcockis Sebastolobrrs alascanus S. ciliatus Anoplopoma fierbria S. proriger Hezngrommos Ingocephnlus Ilexngrar~tmos decaqammas Pleurograrnntus mottopterygitrs Cli?~ocottusacuticeps Blepsins cirrhosus C, embrytmt Jlalacocottus kincaidi Radulinss nsprelhcs$ ~\lyosoceptraluspolyacanthocephalss Triglops metopins Ilypsagonus qundricornis Letltotrentus maticas Pnllasinn bnrbotn Liparis cnllyodo~z Aptocyclus uentricosus L. megflcepltnlus Cnreproctas gilberti Polypera qeeni Eztmicrotremss orbis Glyptocepl~alssznchirtrss i\'ecto/iparis pelngic~ts 'Torrid and Bnthylagus milleri Atheresthes stotains temperate North B. stilbius Hippoglossoides elnssodon Pacific (9 species) Cyclotltone rnicrodo~l Hippoglossus stenolepis Diaphas ttretn I.nntnn ditropis Electronn arctica Lepidopsetta bilii~eatn Laetpa~zgctusregnlis Si~harcticNorth Entosphe>nrstride~ttatus Stenobrachius leucopsar~s Pacific Ocean Raja pnrntifern Tarletoabeani crenulnris and I%el.ing Bnthyntaster caeruleofnsciatus CorypItne,roides ncrolepis Sea (21 species) Alectrias alectrolophus Universal in the tlmmodytes hesapteres Pholis doliclrogaster North Pacific Triglops pingeli Sebnstes polyspinis I3asi0, Bering Reinhardtius ltippoglossoides Ertophrys diceraus Sea, and Gymnocanfhusgnlentus Arctic Ocean G. pistilliger Universal in the Gnsterosteus ncttleatus Henrilepidotus hemilepidotss 11. jordani north temperate oceans Heezitriptenrs bolini Icelur cannlicelntus Arctic seas of ilrctogcdas borisouis iVaeticltthys pribilouius North America Triglogs forficntn .Iropical ~ to sub- dlepisnurus feros T. scepticus arctic regions Ago,rrcs acipetrseri~tus of tltc Atlantic Aspidophoroides bnrloiri and Pacific Scwitor leptorlr),nchtcs Oceans

Wl'ltis list does not includc three unrecorded ichthyoplankton species as discussed in the test. tEstensioo of known range: previously unrecorded west of the Gulf of Alaska. ]:Extension of knolvn range: previously unrecorded west of Unalaska Island. §Extension of knorvn range: ne~vlyrecorded in the subarctic North Pacific Ocean. ~Vlnri~teFish Cov~t~~lolities461 distiilct habitats (Isakson, Sitncnstad, and Burgner, Amchitka \\,ere preponderantly 3-year-old fish 1971). Quantitati\~e association analyses of the (having spent 2 years in freshwater and 1 year in Amchitka collection data with the use of recurrent the ocean) with a scattering of 4 year olds (2 years group a~valysis (Pager, 1957) has identified 13 in fresh\vater and 2 years in the ocean). Both distinct groups alid associated species with an sockeye and chum are immature fishes moving in a affinity index of at least 0.40 (Fig. 3). Thesc general ~vesterly drift exploiting the feeding groupings dcfine the structure of four offshore grounds of the Bering Sea and the North Pacific coininunities aud one inshore community, a littoral Ocean immediately south of the west-central assemblage. L,lany species, especially those cap- Aleutians. The only hexagrammid truly pelagic as tured in the inner sublittoral and neritic waters an adult, the Atka mackerel, was also an abundant around Amchitka, did not fall illto significant lneinber of the cpipelagic community. The Atka assenlblages ~vhcnan affinity index of 0.40 was used mackerel traveled it1 dense schools, especially \vhen as a test of significance. \\'hen these results are entering the inshore \craters of the islands during iucorporated rvith a detailed examination of June and July for spawning. The rock greenling, species distributions, the existeilce of seveu identi- the adult dominant of the inshore/rock-algac fiable fish connnit~nities, identified here by their co~nmutlity,was also found in epipelagic schools of characteristic habitats, is suggested: Epipelagic, 1- to 3-yea~oldjuveniles. Pacific sand lance were mesopelagic, offshore demersal/rock-sponge, off- frequently observed in extensive rapidly slvimining shore demersal/sand-gravel, iilshore/sand-gravel,* schools near the surfacc at night. inshore/rock-algae,* and littoral (Fig. 4 and hlesopelagic Community. A distinct com- Table 3), with some variations in species associa- munity of fishes apparently esists between 200 tious detectable within several of these. The coin~nullity descriptions that follo~vare basically and 1000 111 in the oceailic inesopelagic zonc. \Ve assembled \\,it11 tlte use of these qnalitativc charac- sainpled only the upper region of this zonc (200 to terizations; more subtle relationships, such as 300 m; 324 111 lnasi~numdepth) and only during heterogeneity ~vithin a habitat, mechanisms of nighttime midwater tra\vling, concentrating in the species associations, and microhabitats, cannot be productive np~\rellingwaters of the Bering Sea off identified from the limited scope of our data. the easteril end of the island. Only slightly more diverse (18 species versus 11) than the Epipelagic Community. Fishes comprising the epipelagic comtnunity, the inesopelagic community epipelagic community occi~rin the oceanic waters is dominated by lailter~lfish(family Ivlyctophidac), off Atnchitka bet\\~eenthe surface and 200 111. This deepsea smelt (family Bathylagidae), lightfish community was sampled by salmon longline, (family Gonostomatidac), and viperfish (family surface-gill-net, and purse-seine fishing in the upper Chattliodontidae). All arc typically zooplankton- 10 111 of thesc waters; the mid\\.ater traxvl salnpled feediilg fishes and rvith their predators, grenadiers the deeper portion of the habitat. (family h,lacrouridae) aud longllose lancetfish This coinniunity was characterized by a rela- (family tllepisauridae), apparently perform exten- tively lo~vdiversity of fishes, only 11 species sive nightly vertical migrations for feeding in the lla\~ing been found in it. The fishes are highly upper mesopelagic layers. mobile, often representing migratory species, such We opti~nizedour midwater trawl catches in as five of the six species of Pacific salmon salnpling this community by usiilg echo-sounding (Oncorlzy~~cl~vsspp.). Only one t\\ro-species eqilipment to find thc depth of the deep-water group, which was coinposed of O~~corhy~~cltzis plankton and the associated fishes (deep-scattering lteta, the chuln salmon, and 0. ,~erlza,the sockeye layer). In the rich np\velling area, lnasimuln salmon, was defined for this habitat by the catches exceeded 300 fish per hour compared to recurrent group analysis. The majority of the 10 fish per hour in other areas sampled. Peak salmonids fouud in the epipelagic community \\.ere abundance aud species diversity in thc salnples chuin aud sockeyc salmon. Other FRI studies have occitrrcd bct~veen2400 and 0400 (BDT). In tlte indicated that most of the chum salinoll in the inost productive tra~vl sets, northern lampfish, Amchitka area are of Asian origin and that the California lleadlightfish, bigeye lanternfish, and majority of the sockeyc salmon are fro111 Bristol Pacific viperfish were the most abundant fishes. Bay, Alaska, \\ratersheds (Hartt, 1966; Nakatani Catches of fishes corresponded closely with and Burgner, 1974). The sockeyc caught at catches of euphausiids, mysids, and carid shrimp, coinlnon prey organisins of thcse fishes. *No significant recurrent groups were found in thcse The recurrent group analysis suggested a micro- commitnities at the affinity index selected as a test of structure to this community and defiuite associa- significance. tions bctwccn some midwater fishes and associated LITTORAL DEPTH DISTRIBUTION, m EPIPELAGIC $

Raja binowlata Raja parmifera Raja rrachura Oncornynchus gorbuscha Oncorhynchus Xera Oncorhynchus kisurch Oncorhynchus nerka Oncorhynchus tshawyrscha Salvelinus malma

Bathylagus milleri Barhylagus stilbius

Cyclorhone microdon

Chauliodus macouni

Alepisaurus ferox Oiaphus theta Electrons arcrica Lampanyctus regalis Srenobrachius leocopsarus Tarletonbesnia crenularis Arctogadus borisovi Gadus macrocephalus Theragra chaicogramma

Coryphaenoides acrolepis Coryphaenoides filifera

Gasterosteus aculearus Trichodon rrichodon

Ptilichrhys goodei Zaproia silenus

Ammodyrer hexapterus Marine Fish Comntt~nities 463 I OFFSHORE OFFSHORE DEMERSALI DEMERSALI EPIPELAGIC MESOPELAGIC SAND-GRAVEL ROCK-SPONGE LITTORAL

Pl?olir laeta Anaplarchchos purpr,rercens Phyrichthys chirur

polyacanllzo-

liuvenilerl

acuticepr Liparis

1 Lithodid crab 2 King crab 3 Euphauriidr 4 Shrimps 5 Squids 6 Octo~ur

Pig. 3-Species associations of marine fishes and some associated invertebrates (dashed lines), Amchitka Island, 1967 to 1973, according to Fager's (1957) recurrent group analysis (affinity index, 0.40; sample size, 468).

ROCK-ALGAE MESOPELAGIC SAND-GRAVEL

OFFSHORE DEMERSALI OFFSHORE OEMERSALI SAND-GRAVEL ROCK-SPONGE

Fig. 4-1)iagrammatic sketch showing marine habitats associated with Amchitka fish conlmunities. ('file 200-m depth division in the pelagic zone does not apply to the demersal habitats or commttnities.) i\rlarine Fish Con~71lu1zities 465

Table 3-Relative Distribution of Amcllitka Marine Pisbes According to Seven Identifiable Co~n~n~tnitiesIdentified by Their Characteristic Habitats

Offshore Offshore de~nersall demersal/ Inshore/ Insllorel rock- sand- sand- rock- Epipelagic hlesopelagic sponge gravel gravel algae Intertidal

Lamnn ditropis Oncorliynchrles gorbuscha 0. keta 0. kisutch 0. nerka 0, tshnwytsclra Alepisarcrus feros Antmodytes hexapterus Plercrog/ntnmus monopterygitcs Entosphenus tride~ttatus Bnthylagas milleri B. stilbirrs Cyclotltotte microdon Chauliodus macouni Diaphrrs theta Electroita arctica Lonipanyctus regalis Stenobrachius leucopsarus Tnrletonbeani crenularis Arctogndus borisoui Coryphaeaoides ncrolepis C. filifero Ptilichthys goodei Znprora silenus ~Vectoliparispelagicus Eunzicrotrem~rsorbis Rnjn binoculntn R. pnrmifera R. trachurn Gndus macrocephalus Tlzeragra chnlcogronzma Ronquilus jordani Alectrias alectrolopk~rs Gymtioclitius cristulatus Porocli,rns rothrocki Sebnstes n/eutianus S. nlutus S. babcocki S. polyspiiiis S. proriger Sebastolobas nlascnnus A~~oplopomafimbrin Enoplrrys dicerous Hemilepidotus jordani Ilenzitripterus bolini Icelas cnnnliculotus ilIalococotttrs kincaidi ilfyosocephnhcspolyaca~rtlroceplralus Triglops forficata 1: nretopias

(Table continues on page 466.) Table 3-(Continued)

Offshore Offshore de~nersall demersall Insl~ore/ Inshore/ rock- sand- sand- rock- Epipelagic Mesopelagic sponge gravel gravel algae Intertidal

T, scepticus Cnreproctusgilberti C,phasma Lethotremus muticus Liparis megacepkalus Gymtfocanthtrsgaleatus Hypsagonus quadricornis Sarritor leptorlrynchus Atlterestltes stomias Glyptocephalus zaclrirus

~e'inhdrdtiushippoglossoides Saluelinus malnta Trichodon trichodon Gy mt~ocanthuspistilliger Radulinus nsprellus Pallosina barbata Bathymaster cneruleofasciattrs B. signatus Chiroloyhis nugator Sebastes ciliattcs Hexagrammos decagmmmus H, lagocephal~cs Blepsias cirrltosas Henfilepidotus henrilepidotus Nautichtltys pribilouiris Triglops pingeli Agontcs acipenserinus Aspidophoroides bartoni Aptocycltis ventricosus Crystallichthys cyclospilus Polypera greeni Casterostefcsaculeatvs Anoplarchtcs purpurescetts Plry tichtltys chirus Pholis dolichogaster P. laeta Clinocottus acuticeps C. earbry utn Liparis callyodon

*Uncommon in A~ncllitkawaters. tlnfrequentlv encountered; seasonal or suspected (but undocumented) occurrence. E: l:requently eorountered. Ghbundant in iuvcnilc life states. j~bundant;representative of this community. Marine Fish Communities 467 invertebrates, many of which appear to be their were distinctly biased because only the least-rugged major prey organisms. Several groups exist in the terrain was fishable by trawl nets. Echo-sounding lolver (150 to 330 m) strata of our sampling depth tracings of the sideivalls of submarine canyons and regime. The California headlightfish and the north- pinnacles showed large aggregations of fishes that ern lampfish and their principal prey species, could not be san~pledwith our nets. Their behavior Tltysattoessa lottgipes, T. ittermis, Elrphnzisia pa- indicated that these fishes were rockfish (family cifica (cuphausiids), Sergestes sitnilis, Hyntenodora Scorpaenidae, probably Pacific ocean perch) or frontalis (shrimps), and Gonatus magister and walleye pollock. The most abundant schooling Galitetrtlcis arinata (squids), occur with another fishes of the offshore demersal/rock-sponge com- fish group that is co~nposedof the Pacific viperfish munity were Pacific ocean perch, redstripe rock- and the bigeye lanternfish and another invertebrate fish, Pacific cod, and walleye pollock. The most group of Gotlatzrs fabricii (squid) and Bolitaetta abundant bottom fishes were yellow Irish lord, lteatlci (octopus). A fish-shrimp group of the blackfin sculpin, ribbed sculpin, spectacled sculpin, pinpoint lainpfish and Notostontnrs japotticus,an northern ronquil, roughtail skate, and arrowtootb oplophorid shrimp, is defined for the deeper (220 flounder. to 330 m) strata, and an intermediate-depth (110 Bottom trawl catches from this coinmunity to 146 m) group of Pacific sand lance and tadpole ranged between 200 and 1000 fish per hour, the snailfisl~is also designated. higher catch rate occurring when schools of Pacific Demersal fishes, such as the \valIeye pollock, ocean perch, Pacific cod, or walleye pollock were appeared to enter the mesopelagic co~nnlunity encountered. Catch composition typically ranged when the plailktonic scattering layers composed of from 40 to 900 Pacific ocean perch, 30 to 55 euphausiids, shrimps, mysids, and squids migrated walleye pollock, 20 to 45 Pacific cod, 20 to 30 within their feeding range. Some of the species sculpins (family Cottidae), 6 to 12 arrowtooth noted as being uncommon in the mesopelagic flounders, 6 to 7 rock sole, 2 to 6 Pacific halibut, community lnay actually have been members of and 0 to 1 poacher (family Agonidae) per trawling the epipelagic comm~ulity \vhich were captured hour. Lithodid crabs, predominantly Litltodes during the movement of the midwater trawl aeqncispina, were taken in many trawls, their catch through epipelagic waters. The occurrence of the averaging 4 to 9 crabs per hour with maximum Pacific lamprey, smooth lun~psucker, and rock catches of 40 to 50 crabs per hour. gieeilling in this community may well be explained Three groups of fishes were designated by the in this way. The Pacific lamprey, however, has recurrent group analysis as existing within this been noted at depths of 300 to 400 m by community, primarily as found on the Pacific Abakumov (1964) and may range through both Ocean side of Amchitka. A cottid group, which communities. includes the blackfin sculpin, the yellow Irish lord, Offsl~ore Demersal/Rock-Sponge Com- and several Triglops species and associated with the munity. \Yl>at we originally described as one northern ronquil and the lithodid crab (Litl~odes entity, the offshore/bottom conlnlunity (Isakson aeqzrispina), occurs doininautly in the 150- to et al., 1971), has since been redefined as two 200-nl depth region. The Pacific ocean perch and coiuinunitics on the basis of expanded data fro111 walleye pollock occur in association with the king catches over two physically different deepwater crab (Paralitltodes canrtschatica) in deeper water bottom habitats. The habitat of the offshore (from 250 to 400 m) and a distinct rockfish den~ersal/rock-sponge community iilcludes those group, which includes the rougheye rockfish and areas adjacent to Amchitka which are rocky and sl~ortspine thornyhead, was found in the 220-m rough bottomed, often with precipitous cliffs aud depth region. pinnacles, and which form much of the bottom The offshore comtnunity is vulnerable to dis- area between 55 and 180 m (180 and 590 ft) in turbance, as indicated by replacement of rockfish depth on the southeastern side of Amchitka. The by walleye pollock after fishing by foreign vessels bottom material retrieved incidentally ~viththe in fall 1972 and spring 1973 (Nakatani and fishes during tra~vling included an abundance of Burgner, 1974). If \vc assume that the Pacific sponge (family Axinellidae), coral (family ocean perch populatioi~s were cropped to low Stylasteridae ?), and tunicates. Observations made population levels by the fishing fleets [and had not while scuba diving over the extensive shoals indi- just moved out of the area (Chikuni, 1975)], it cate that the bottom (at least to 30 nl deep) is also appears that populations of walleye pollock, appar- extensively blanketed by coralline algae (Clathro- ently a successf~~lcompetitor, had succeeded thc ntorplt~cnt spp.) (P. A. Lebednik, University of rockfish. After an extensive period for recruitment \\lashington, personal coinmunication). The catches to build up its population again, the longer lived Pacific ocean perch may eventually suppla~ltthe sand-gravel substrate that possesses little stabilized wallcyc pollock. benthic gro\vth. This type of habitat is prevalent in embayments, such as Constalltine IIarbor, South Offsliore Demersal/Satid-Gravel Com- Bight, Cyril Cove, and h,Iakarius Bay (Fig. I), and mutiity. The offshore demersal/satld-gravel com- occasioi~ally occurs as narrotv areas along the lnu~iityis the assemblage of fishes common to the exposed coastliue bet\veen kelp beds or reef offshore low-relief expwscs of silt, sand, and rock pro~nontories along the bottom of submarine cobbles found between approximately 55- and canyons. In areas where rock pilinacles \\,it11 95-m depths in the Bering Sea north of the associated abunda~it algal growth stand in the hlilrow aud Canniki11 test sites 011 At~ichitka nliddle of a sand-gravel expanse, fishcs of the two (Fig. 1). There appears to be very little ellcrusting inshore subtidal communities often intermingle, benthic gro\vth, such as sponges, corals, and especially at night. This may indicate the direct llydroids, although sea pens (family Virg~lariidae) relationship bet~veenspatial heterogeneity of this occur on tlie outer (deeper) inargiils of this habitat. habitat and the diversity of its residcnt fish Cobbles brought up in the trawling operation were community. often entirely covered by the coralline alga ~VIeso- Pacific cod, walleye pollock, and Pacific hali- plzyllz~nz sp., \vhicIi suggests that the habitat's but appear to be seasonal transients of the inshore/ cobble substrate is relatively loose and perhaps not sand-gravel community; ho\vever, litnitcd tvinter stable (P. A. Lebednik, University of \Vashington, sampling precluded the determination of whether personal comniunication). The diversity of the fislt these are actual migratory movements. Cod and faltua comprisi~lg this commuuity is lower thall pollock are abundaut fro111 spring until mid- that in the other offshore bottom habitat. Catch October. Pacific halibnt occur com~no~llythrough- compositioil in the offshore demersal/swd-gravel out the year except dnring January, February, and community ranged to a maximum of 320 rock solc, March when they apparently move offshore for 94 Pacific halibut, 50 armorhead sculpins, 5 search- spa\vning. Juvenile Pacific halibut and Pacific sand ers, and 10 sturgeon poachers per hour ot tralvling lance \\.ere observed regularly in nearsbore waters effort. The stomach contents of top carnivores, year round. No sigiiificant recurrent groups were such as the Pacific I~alibut(see follo\ving food-web section), from this communit)~ indicated that defined tvitliin this habitat. pelagic schools of Pacific sand lance also were I~ishore/Rock-Algae Co~n~l~i~~~ity.Tlie colnmon members of this demersal community. inshore/rock-algae colnnlunity is characterized by Unlike fishes of the other offshore demersal fish a diverse abundant assemblage of fishes iutilnately community, two fishes, Pacific halibut and rock associated wit11 the exte~~sivealgal gro~vthdominat- sole, in this commitnity are relatively mobile, their ing the rocky nearshore coast. Approximately 60 concentrations shifting about locally, probably for to 70% of Amchitka's shoreline is composed of feeding. They also change depth seasonally; this exposed rock outcrops, cliffs, or flat-rock benches movement is perhaps associated tvitli spaurning extending iu~derwateras subtidal rock terraces out activity. In the Bering Sea liortheast of Amchitka, to depths of as much as 100 tn (Gard, Chap. 2, this Pacific halibut maintain denser schools during tile volume). Abutida~ltsubmarine algal growth, such winter, dispersiug and uiigrating into the shallower as encrustiiig corallincs, species of fragile Rho- \\raters in late spring (Novikov, 1964), and rock dophyta, and various kelps cover these terraces; sole concentrations dispcrse during a late summer- inost co~lspicuousare the dense kelp beds of Alaria fall feeding migration (Sliitbnikov and I.isovenko, fistulosa, which sometimes exteud to tlie 20-m 1964). depth contour (Lebednik and Palmisano, Chap. 17, Tl~erecurrent group analysis indicated that a this volume). Alaria coiltributes to the bottom basic group of three species, the rock sole, Pacific habitat by its massive holdfasts and sporophylls halibut, and Pacific cod, existed in distinct associa- and by its long fronds, which form a protective ti011 wit11 two other species, the arrowtooth floun- forest canopy reaching to the surface where the der and sturgeon poacher. The Pacific cod associa- large floating blades form a microhabitat in them- tioil ~vith the Pacific halibut and rock sole, selves (Fig. 5). The kelp forest habitat may however, was Inore pro~lounced on the Pacific contribute the multilayered quality of a terrestrial Ocean side of Amcliitka than in satnples from the forest that has been shown to illfluence bird Berixig Sea habitat. species diversity through the vertical partitioning of niches (MacArthur and i\IacArtIlur, 1961; Cody, Inshore/Sand-Gravel Commiuiity. Like its 1968; Orians, 1969). In addition to the luxuriant offshore counterpart, the habitat occttpied by the macropllytic growth, sessile benthic animals- inshore/sand-grax~el connnu~lity consists of a sponges, bryozoaus, anthozoans, and ascidians- dusky rockfish, siherspotted sculpin, and some of the less abundant liparid species, occupy the pelagic kelp bed canopy and near-surface micro- habitats. Although the latter fishes move frccly about the ~Ilariablades and surfacc canopy either singly (silverspotted sculpiil and liparids) or in schools (dusky rockfish), the bottom-associated fishes appcarcd restricted to a particular site. Scuba observations indicatc that during daylight these fishes seldolll lnove beyond the protectiou of the dense bottom algal cover, but at night thcy arc seen among the benthipelagic swarms of mysids and amphipods. The tra~nlllel net was the most productive sampling gear used it1 the inshore/rock-algae habi- tat. Sets of short duration, \vhich avoided net saturation, yielded up to 36 fish of lnixed species per hour, which averaged 75 to 80% rock greenli~lg with the relnaiuder red Irish lord, sturgeon poacher, Pacific cod, northern ronquils, and great Fig. 5-Alnria fistulosa kelp bed (underwater pho. sculpins. Dusky rockfish move in scl~oolsand \\rere tograph). Note school of pelagic fish. not collsistently encountered; but, \vhen they isere encoiuntered, they made up a significant propor- tion of the catch. Sil\rerspotted sculpins and encrust ally firm rock substrate, which creates a liparids were captured \vith surface plankton net bottom habitat of incredible diversity in forin, tolvs through the il1al.ia fronds and were obser\red texture, and color. duriug scuba divcs. During the xvinter, \\,hen the Associated \\'it11 the algac for food aud protec- illaria lose many of their surface fronds from wave tion are ab~uldantcrustaceans, echinoderms, and molluscs. The benthipelagic* crustaceans [princi- pally mysids (Acal~thon~ysissp.) and amphipods] were often found in extremely dense underwater swarms (Fig. 6). Thcsc and other abundant invertebrate popula- tioils comprise a diverse array of food orgallisms a~~ailableto the fisbes frequenting the habitat (see later food-web discussion). The spatial hetero- geneity of the entire habitat aid the diversity of its algal groxvth and associated food resources arc jointly responsible for a11 abundant diverse fish asscinblagc. Representative species in the ioshorc/rock- algac community include the rock greenling, red Irish lord, ilorthcrn ronquil, silverspotted sculpin, great sculpin, and dusky rockfish. Pacific cod \irere also captured in the inshore/rock-algae com- munity habitat of the island's exposed coast, \\.hich they were assumed to lta\rc entered from the adjacent inshore/ or offshore/sand-gra\.el com- nlunity. No recurrent groups \\,ere found in the inshore/rock-algae habitat salnplcs \\rith the use of the affiility index of 0.40. This assemblage for the most part consists of sed- entary botto~nfishes; l~on~e\,er,a felt,, notably the

Fig. 6-hfysid, Acnntho~nysissp., swarms in Ainria *i\ssociated with bottom but rising pelagically off it. kelp bed (underwater photograph). action, tlie kelp forest is greatly thinned, and the Icagalaska Island, 290 km (180 miles) east of pelagic fishes descend into the sublittoral fringe Amchitka, often included significant numbers of zone and its lush Lamijln,'in gro\vth. Other species, juvenile rock greenling. After 2 to 3 years, they such as the dusky rockfish, also move from their apparently migrate back into the inshore areas and siunmer habitat into deeper \\rater during the becollie solitary (Gorbunova, 1962; Simenstad, winter, perhaps to avoid wave action or to follolv 1971a). This unquantified obselvation suggests their food resources. randoln recruitment from the rock greenling stocks Thc bottom-associated fishes are cryptically of the \vliole Aleutian archipelago. Rock greenling colored to rcsemble the reds and browns of the food habits and feeding ecology are discussed in bottom habitat's flora and fauna. Even the distinc- more detail in tlie following section on food webs. tive silverspotted sculpin, \vhich swims freely amidst thc kelp fronds, is difficult to tell from a Littoral Community. Tlie littoral fish com- shred of algae whcn obscrvcd within its habitat. munity is that assemblage of fishes cli;~racteristicof The male rock greenling, tunlike the female, is thc surge channels and tide pools of the rocky marked with bright red bars that possibly are littoral bench that typifies much of Amchitka's associated lvith reproductive or otber social bc- shore (Fig. 7). Although this assemblage can logi- havior. Altliough not so well camouflaged, it cally be considered a sliallow-water extension of nevertheless blends well with the rcd algae of the the inshore/rock-algae community, it can be understory. The rock greenling, red Irish lord, and distinguished from that comm~unityby tlie pres- sturgeon poacher are further concealed by their ence of distinctive species. This habitat also pro- remaining motionless on tlie bottom supported by vides a nursery pound for juvenile fishes. their broad pectoral fins. The suctio~idisk on the This community was most often sampled at ventral surfacc of liparids enables these small fishes lotide; so tlie entries of Table 3 do not com- to adhere to kelp fronds in the constant \vave pletely represent the community at high tide when motion of the exposed kelp beds. species undetected in our sampling could have been present. Littoral-occurring fishes were collected Tlie rock greenling is by far the most abundant with the use of rotenone in the tide pools (see and widely distributed number of this community. method discussed by \\'ilimovsky, 1964), by hook- 'l'l~isobservation stimulated a detailed study of its and-line fishing at higll tide, atid by setting gill nets life history, especially its feeding ecology on tlie littoral bench through a tidal cycle. As (Simenstad, 19712). Scuba observatiolis indicate noted by \Vilimovsky (1964) and generally con- that adult rock greenling are territorial all months firmed in our sampling, tlie fishes of the littoral of the year and especially so during the spa\vning commimity responded to the rotenone sequentially and incubating periods of late spring and early according to their oxygen demands. \\Je found that, summer. Thc eggs are laid in purple masses that aftcr rotenone had been applied to a tide pool, fish adhere to algae (usually I.(o,ii~znrin sp.) in the succumbed in tlie follo\ving order: (1) silverspotted sublittoral fringe zone and are guarded by both the and geat sculpins (the smaller specimens re- male and female. Even during ilicubitti~ig ailcl sponded later), (2) sharpnose sculpins, (3) spotted nonspa\\~ningpcriods, individuals appear to main- tain their established homesites, to which they return end rest aftcr brief feeding forays. Out of about 910 rock greenling in tagging studies, none of those recaptured moved along the shoreline more tlian 75 in (Burgner et al., 197 la). Simcnstad (1971a) suggests that at high tide, especially at night, rock greenling leave their territories to forage on the invertebrate community of the littoral bench. The results from tagging imply that such depmt~~resfor feeding arc merely temporary excursions. After the ju\~enilerock greelili~ighatch in the siunmcr, they form pelagic scliools and apparently lnove into offshore oceanic waters and into the epipelagic comtnunity. Although juvciiile rock greenling were seldonl captured in Amchitka's Fig.7-Littoral bench at Rifle Range Point, Amchitka Island, Alaska. Note tidepools connected inshore \\raters, the purse seine catches of FKl's to sea by surge chaniicls. Visible bench ltcre averages research vessels 65 to 95 km (40 to 60 miles) off 150 m in width. ~MnrbzeFish Coitriilili~ities 471 snailfisli and lobefill snailfish, (4) smooth Irtmp- serve thc primary objective of our investigations, suckers, (5) ribbon pricklebacks and high cocks- i.e., prediction of tlie pote~itialconsequences of a 1 combs, end (6) northern ronquils. The response nuclear test or tests on Amchitka. Test-induced bcha\,ior also varied betwcc~ispecies, as with the changes in tlic organizational structitre of the liparids, \vhich habitually s\tram to the tide-pool m.nrinc food web, direct or secondaxy, cotuld surface, in contrast to the rock greenling, n~liich conceivably result from loss of habitat or from loss s\sam excitedly among the algnc at the bottom of or change in ~iumbers of i~nporta~itpre)' or tlie pool and e\>entually died there. Such varied predator species. Also, tlic food web is a potential bcliavior and response rate may have biascd the route for transfer of radionuclides through the collectio~~;lion,ever, an effort was made to obtain ecosystem and ultimately to man in the event of a and record every fish in a treated tide pool. releasc of radioactivity to the environment. Food- The recurrent group analysis i~ldicateda single \veb studies were tlicrcfore u~idertakc~ito identify group of littoral fishes, \vhicli included tlie crescent prey organisms of Amchitka marine fishes that (1) g~~nnel,high cockscomb, ribbori prickleback, juvc- are most uni\~ersallyexploited througl~uutthe food nile great sculpin, sharpnose sculpin, and spottcd web, (2) have a major role in energy transfer or s~iailfish.During high-tide periods \\~Iicntlic littoral conversion, and (3) import significant amounts of bench is flooded, adult rock greenling and anadro- energy into the food web from tlie high seas 111ous Dolly Varden also cruise over the bench, around the islalid. feeding on the mysids, amphipods, and other Results of preliminar)~food-\seb analyses werc ~iiacroi~i\,ertebratcsexposed or brought in with tlie prcsented in Burgtier et al. (19G9; 1971a), Burgner flooding tide. and Nakatani (1972), Nakatani et al. (1973), Species richness was greatest during the Nakata~iiand Burgner (1974), and scheniatically in summer season, at which time the comiiion- lsakson et al. (1971), ~vliereinthc tropliic interac- resident littoral fishes werc tlie high cocksco~nb, tions among fish commuu~~iticsand other exploiters crescent gicnnel, sharpnose and silverspotted scul- (marine nia~nmals,littoral-fccding birds, and man) pins, and spottcd snailfish. In tlie few large tide Ivcrc sIio\vn. 111 a practical sense, a relatively simple pools or surge channels, adult rock greenling \verc scliematic cannot adequately display such co~llplex also found in abundance, especially dttri~igthe relatiotiships. Ho\vever, it is possible to diagram the June to August spa\vning period ~vlienthey laid major tropliic path\va)ls and significant prey re- their eggs amolig the Lnitlitiarin in the sublittoral sources asailable to thc upper-level compotients of fringe zone. In the spring juvenile great sculpi~ls tlie ecosystem. These aspects of the niarinc fisli begen appearing in profusion in the littoral area comnii~~litiesof Amcliitka are sIio\vn in Figs. 8a and remained until fall before migrating off the and 8b. Although these figures are constructed bench. These small juveniles constituted up to 55 to from actual data, each link in tlie web represents 85% of all fish fo~111din the mids~u~iimertide-pool an average derived from all sa~iiples,often collccted population. Similarly, j~uveiiile smooth lump- over several seasons, including daytime and night- suckers, Dolly Varden, coho salmon, and rock titile collections, and a range of pred;~torsizes. This grce~ilitig\\,ere also abundant periodically during schetiiatic modcl, ;I si~iiplificd condensed reprc- the summer accordi~~gto the size of tlie tide pool, scntatiort of the upper level of a very co~nplex extent of algal cover, proximity to frcshnnter tropllic strttcturc, indicates the dominant ~ne~nbers strca~ns, substrate type, and distance from tlie of the Amchitka fish cotnn~uniticsrelative to their sea\vard mixgin of the bench. Fish densities in the more significant connections to other fishes and to tide pools averaged 98 fisli (range, 20 to 250) per the prey components ol thc system. 3- to 6-ni3 tide pool. Sollie of this variability in Altliougli we detected so~iiesei~sonal and year- these catchcs undoubtedly was due to varying to-year variations in the food habits of a few temperature, \vliicli is kno\\.n to greatly influcncc species, the limited extent of winter sampling and tlie effectiveness of the rotenone. Reduction of tlie the small number of years of sampli~igpreclude tlie littoral algal cover and of available prey resources incorporation of sucli variations into our model. as well as the reduced effecti\~encssof rotenone in Where appropriate, we have connected prey organ- colder \\raters were factors co~itributingto the Ion, isms to several predators. i\ltl~ouglithcse predators ~iumbersatid diversity of fislies encot~ntercdduring liiay use colnmo1i prey resources, this does not the winter ~nontlis. necessarily imply competition among these fishes for the same populations of prey or ~~~o\~c~~icntinto A~~~cl~itkaMarine Food Web adjaccnt communities for their esploitation. For Some kno\vledge of the food-\veb structure in ~iian)~prey tasa (;~niphipods,shrimp, ceplialopods, the marine ecosystem was deemed necessary to molluscs, and copcpods), these gc~ieralcategories Fig. 8a-Offshore epipelagic, mesopelagic, demersallsand-gravcl, and demcrsai/rock-sponge fish communities illustrating semiquantitativc food-web relationships.

474 Sitnettstad, Isakson, and ~\'akatani incltcdc Inany diverse species representing different "r~p~ling-po~~~~ing" . sounds (Simenstad, 1971a). populations, niches, microhabitats, and/or bc- They \\,ere s\\~i~nmingacross the bcnch picking up havior; it was impossible to determine, much less benthic macroiit\'ertebrates--gastropod and bi- to indicate, where or hob\, a fish captured any valve molluscs, ampllipods, isopods, and poly- specific prey since we kno\s little of each prey chaete annelids-nrhcn they were cncot~ntered. organism's distributio~tand behavior. Instead of confining their feeding to periodic Frequency of occurrence of prey items intensive activity during the subinersioll of the (Tablc4) provides a11 index of a fish's actual intertidal bench, the grecnling appear to foragc to exploitatioil of the prey complementary to the some degree continuously throughout the day and biomass-pcrccntagc of total stomach contents night. Altltough the grecnling tcnd to feed slightly indicated in Figs. 8a and 8b. The two kinds of more during the day, at night they call safely leave information together show the trophic relation- the algal protectioil at thc bottom and forage in ships among the components of Amchitka's marine the s\crarms of henthipclagic amphipods and mysids fish communities. of the illaria forest. Simenstad (1971a) suggested i\rlarinc fishes in the littoral region rely on thc that, apparently because of the metabolic or diverse invertebrate commttnities associated with survi\'al cost of thc uncertainty of encounter xvith the extensive algal growth of that habitat. Espe- the patchy swarms of crustaceans, the rock green- cially in this habitat, the atuphipod populations ling maintain a geueralistic feeding behavior uiltil provide the major source of trophic energy to the they encounter dense concentrations of amphipods littoral fishes and high-tidc immigrants from adja- or mysids. Then the fish feed on these crustacean cent communities. These amphipod populations s\varlns cxclusi\'ely until satiation or until thc reach Itigh densities in the littoral area during the crustaceans arc too sparse to make it \vortli the summer \vhcn the \\rindrows of detached kelp and effort. Simenstad considered predation pressures othcr algae pilcd on the beaches during the winter by t11c sea otter and othet. carnivores ('Table 4) and storm period contribute a tremendous source of ~nainteiranceof reproductivc territories important detritus used by these crustaceans. The usitally factors in maintaining this feeding behavior. This sedentary territorial character of many of the pattcrn of feeding was often sho\v11 by rock littoral fishes \vould restrict them to the food greenling stomach contents, \\,here homogeneous resources within their nticrohabitat or those \vhich concentrations of mysids or amphipods wcre fo~tud drift into it. Wit11 this restriction it is possible that preceded by or follo\ved by a heterogeneous they prcfcrcntially exploit the abundant "rcnew- :tccumulation of ingcsted gastropods, isopods, a~td able" prey resources, the shifting ainpllipod, annelids. Similar feeding strategies have been dis- harpacticoid copcpod, and drift insect populations, cussed by Schoener (1969;1971), Rapport (1971), which pass through their territory. Most of the aud Pi~lliam (1974) in their discussion of opti- fishes that feed as specialists on thcse dense but mality theories in dict strategies. patchy swarms of orgauis~nsmake very limited usc Although none of the other dominant fishcs of of the cpibenthic in\rertcbratcs of this habitat. the iilshore/rock-algae commiuitity are such gen- Thus the sedentary invertebrates are not heavily eralistic feeders as the rock greenling, most also exploited except by the high cockscomb, lobefin rely on the abundant bcntbic macroin\~ertebratcs snailfish, and that mobile forager from the adjacent or benthipelagic crustaceans com~nonto that habi- inshorc/rock-algae community, the rock greenling. tat. Of these, the great sc~ilpi11aud Alaskan roilqoil The diet and feeding ecology of the rock seem to be the most opportunistic, thc former greenling, the most abundant fish of Amchitka's being the only fish that is at all pisci\,orons; red inshore ~narincenvironntent, were the focus of ;in Irish lords, sturge011poachers, and searchers tcnd to intensi\re study (Simenstad, 1971a). As an opportu- be more specialistic in their feeding preferences. nistic feeder the rock greenling csploits the diverse The only fishes with a narrow ntnge of choice in community of macroin\,ertebrates found in abun- their dict are the pclagic fishes of thc contntunity, dance in the inshore/rock-algae and intertidal the dusky rockfish and sil\~crspottedsculpin, \vhich habitats. The fact that its stomach contains organ- exploit solely the bcnthipelagic mysid and amphi- isms from both littoral and inner sublittoral habi- pod populations \vithin the /llaricr forest. tats suggests that the grecnling move over the The sinooth lu~npst~ckcrsccms to exploit food littoral bcnch during high tide for feeding. resources throughout the water colu111n from the Although the adult rock greeuling is not typically a shrimp and polychactc ai~itclidsassociated \vith the free-s\viinming schooling species, schools of this bottom to the benthipelagic amphipod aud mysid fish (often only males) were observcd foraging over swarms bcncatlt the Alaria canopy. This fish may the bench, often rising to the surface and making once have been a domin;lnt component of this ncarshorc community, but apparently it was Lithodes aeqltns/>itla and king crab Paralitltodes sc\~erelyreduced iri abundance by sea ottcr prcda- canttschatica. Armorhead sculpi~~srely predomi- tion since it is an extremely sluggish species and nantly on benthic amphipods and isopods and not efficient at escaping a predator like the sea sccorrdarily on polycliaetc annelids. These sci~lpi~ls ottcr (l:'itlzy(l~a lj~tris).Fish collections made in the are somc\vltat opportunistic, as indicated by tl~e 1950s Goduced abundant catches of the smooth dotuinance of Oil~o/)leura in their diet during an lumpsucker (ICeuyon, 1969, and perso~ml com- ahnormally high occurrence of these larvaceans in munication); our recent intensive sampling \\,it11 tlie inshore watcrs in August 1969. siniilar methods seldom produced more than t\vo The diets of the demersal fishcs in the offshorc of tliese fishes a year, one-tx\~entictliof a day's portions of tlie sand-gravel habitat sIio\\~some catch of 20 years ago. That the lumpsucker still interesting diffcrctices from their inshore diets in persists in the Amcliitka nearshore environment, those species cornmoll to both ('Table 5). Offshore- despite tlie intcnse sea otter foraging pressure, is a caught rock sole consunled more opliiuroids rcsult of its rcproducti\,e strategy of producing than polycliaete annelids and amphipods but there are hundreds of thousands of pelagic eggs per fish. Eggs comprised 41% of the total ~vciglttof a female fewer opliiuroids in ncarshore arcas. Skalkin captured in late April 1972! As a result juve~iile (1963) and Shubnikos and Lisovenko (1964) smooth lumpsuckers are fouitd iu tremendous indicated tliat rock sole in the southeastern Bcring numbers in the littoral zoue every spring. Sea dcpetld on polychaetes and ~nolluscsas their Guryanosa (1935) also ~iotcdthe abimdance of major food items, with crustaccaos, Pacific sa~id tliis fisl~in tlie Co~nnla~lderIslallds during the lance, and ecliinodcrms of secondary importance. This may reflect, I effect, diffcrcnces in the spring and summer. benthic macrofauna essociatcd with this habitat in In thc inshore/saod-ga\,el community is a the two regions. Offshore Pacific halibut prey on t\vo-compartment food web, one oriented to the \vallcye pollock, armorhead and other unidentified neritic fishcs, the salmo~iids,Atka mackerel, and sculpins, and sturgeon poachers rather than on Pacific sand lance, and tlie other oriented to the Pacific sand lance and rock sole, even though the bottom-i~ssociatedfishes, tlic gadids, plcuronectids, latter occur abiuidantly in this irabitat. This appar- armorhead sculpin, ;uid Pacific sandfish. The prey ent selectivity also occurred in this species' preda- or~a~iisrnsin the neritic compartment arc plank- tion on decapod crustaceans, primarily horse crabs tonic crustaceans, hypcrid amphipods (mostly and sltrinip, althougli king crabs appear to occur in Paratlten~isto abyssorton), lnysids and copcpods cqi~dabundance. The reason that rock sole and (C(11autls c~istatns,Psett~localn~rtts elolzgat~~s), a~id king crab were not preyed on is not apparent. Tlie adult a~idlarval Pacific sa~idlance. Oikopleura sp. dict composition of offshore halibut was similar to (a larvacean) are heavily preyed on during infre- that found by Gordecva (1954) and Novikov quent periods of high abundance in the plankton. (1964) in tlie western, nortli\\,estcnl, a~iclcentral In addition, Dolly Vardcri and pink salmou in Bering Sea, where fishes (pollock, smelts, eelpouts, inshorc waters occasionally take drift insects that Pacific Iierri~lg,and Pacific sand lance) and crusta- have bee11 blown out from the adjacent land mass. ceans [spidcr crabs (Cltio~~oecetes?), fidcllcr crabs, Bottom-associated fishes are almost all mac- hermit critbs, and shrimp] are tlic primary prey rophagous. Prime examples of tliese are tlie Pacific and ~iiolluscs(octopuses) are secondary. A trcrtd of halibut and Pacific cod. All but the rock sole are to cliangi~igdiet composition by size, as notcd by some extent piscivoroirs in their dict. Partitioning Noviko\l (1964), \\'as also indicated at of food resources among fishcs of similar habitats is Amcltitka-increasil~g dependcnce on fishcs and evident \vhen the gadids and pleuronectid members molluscs anti decreasing emphasis on crustaceans of this commu~lityare compared. The Pacific cod with increasing fish size (30 to 60 cm to 790 cm). is dependent on armorhead sculpins and mobilc If one ct~nsidersthe Pacilic sand lance to be the be~ithiccrustaceans, shrimp, and the horse crab westem Aleutian ecological cquivalcnt of smelt and (Erit~tncrus ise~tbeckii), \\,liereas the walleye herring, the diets of halibut throoghout thc Bering pollock concentrates on niysids, with Pacific sand Sea are very simikir. Even \\,it11 a small sample size, lance arid rock solc providi~lgincidental cotltribu- the indications arc that ncarshorc tlic halibut mttch tions to tlie diet. Tlie rock solc is a bcnthopl~agc ~wcfcrs to eat a~iiphipods(primarily bottom spe- that preys on polycl~aete anclelids and benthic cies) tvltereas offshore it is more pisci\~orous.Thc amphipods. The Pacific halibut is a higher order arro~vtoothflounder, not found nearshore, appears colisumer that feeds principally on rock solc and to divide its diet bet\\,ccn shrimp and an uniden- Pacific sand la~lceand secondarily on squid, octo- tified cottid, possibly the armorheat1 sculpin; again pus, wd the horse crab and t\vo lithodid crabs, this is an inference from a small sample.

Table 5-Percentage Coniposition by 18iomass and Percentage Occurrence (in l'arentl~eses) for General Prey Taxa of Important Species Occurring in Both Offshore and Inshore Conlni~~nities - - Prey Predators No. Fish Decapods Molluscs Amphipods hlysids Copcpods hliscellaneo~ts

Pacific cod Offshore 4

Inshore 112

\$'alleye pollock Offshore 22

Inshore 48

Great sculpin Offshore 4

Inshore 37

Red Irish lord Offshore 6

Inshore 110

Pacific halibut Offshore 5 7

Inshore 69

Rock sole Offshore 4

Inshore 16

"Less than 0.05%.

The offsliore deniersal/rock-sponge cont- fish, California smoothtongue, and juvenile rock- munity includes four fishes that are also found in fish, Sebnstes). illsltore comii~unities:(1) Pacific cod, (2) \\,alleye Offshorc cottids also differ from their inshore pollock, (3) great sculpin, and (4) red Irish lord counterparts in their food habits (Table 5). Red (Table 5). Cod iiisliorc and offshore arc equally Irish lord feed selectively on horse crabs with pisci\~orous, but offsliore they feed esclusively on supplemental contribntions by shrimp, anipliipods, red Irish lord and Pacific sand lance \vliereas and polychaete annelids. Large adult great sculpins inshore they prey on armorliead sculpi~lsmore feed exclusively on tlie txvo large decapods, the than on aiy other fish. I~isltore, slirimp \\.ere horse crab (E~iri~acrus)and tlie tanncr crab proniinent in their diets, and squid and octopus (Cltioi~occetessp.). The blackfin sculpin, an abun- had replaced Eritilncrus. Pacific cod in both inshore dant cottid esclusi\~e to this habitat, is a and nearshore habitats seen1 not to exploit the bcntliiphagc that preys on polychaete annclids, abinlda~ltbenthipelagic amphipods. t\lthouglt \\'all- ophiuroids, bi\,alvcs, gastropods, squid, octopus, eye pollock in the offshore co~nmtuiityappear to deepwater isopods (Arctririrs set~rsrrs?),horse and be characteristically associated \\,it11 the Pacific cod tanner crabs, amphipods, shrimp, and unidentified aid sculpi~lsalong the bottom, benthic organisms cottids, tlie morc important prey being polycliactc are not do~tiinantin their diet. Instead, tlie walleye annelids, anipltipods, and the fishcs. pollock apparently move up off the bottom diur- 'I'he dominant fish of the deepwater rockfisli trally to exploit mesopelagic euphausiids (contribu- niche, the Pacific ocean perch, is plankti\,orous. ting 87% of the stomach contents biomass), coilcentrating on the euphausiid (86% composition shrimp, squid, octopus, and fislies (northern lamp- by weight) and liypcriid ampliipod (6% composi- tion by \\'eight) components of the coastal lUcutians arc euphausiids, hyperiid anipliipods, and zooplankton tiincidental exploitation of calanoid copepods \\it11 fish and squids forming mesopelagic fishes-Pacific ~ipcrfisli and only secondary food resources, indicating their lanternfishes-ancl demersal cott~tls. A large seasonal patchy distributions aroiuntl the North ktthypelagic calanoid copepod (Calai~~tscristntus) Pacific. Allen (1956b) further indicates that, where luid the chaetognatli [Sagittrr plr~~ctoi~is(?)I also food is most abundant, both species become niore appear in the Pacific ocean perch diet. This diet selective in the feeding habits. composition generally agrees \\,it11 the findings of Longne.;e lancetfish are ocemiic alcpisaurs and Lyubimo\~a (1963), Paraketsov (1963), Skalkin were represented in the Amchitka collectiori by (1964), and Chikmii (1975), \\rho found crusta- two fish that were blown ashore during stormy ceans to be the niajor prey items ~vitliplanktonic ~vcatlicr. T'.eir stomach contents indicated that [calanoids (C. cristat~rs), eupliausiids, a~idhy- these fish :c omnivorous predators that range periids] and nektobentliic (mysids, amphipods, through the epi- and mesopelagic water colutnn ant1 shrimps) organisms predominating. These in- and deeper, feeding on nckton resembling in vestigators also found sonle squids and small fish in composition that captured in our niid\vater tratvl- tlie rockfish stomachs. ing. Prey specifically includes Pacific spiny liunp- suckers, octopus (Bolitaetla lientlti), and squid h~Icsopclagic fishes include plankti\rorous ban- (Go?~ntirssp.). Other nlinor co~iil~o~ientsincli~de ternfishes and bathylagids and their predators, tlie polycliaetc annelids, Pecific herring (Clurpen Pacific viperfish and rot~ghscderattail. Calanoid I~are~tguspallasi), and juvenile cottids, possibly red copepods and hypcriid amphipods constitute the Irish lords. This diet composition indicates that the major food sources of tlie lanternfishcs with longnose lancetfish feeds opportunistically on the supplemental input fro111 euphausiids slo\v-movitng coniponents of the nekton, neglecting (Tl~ysnitoessa lo~tgipes,T. iiterirtis, and E~r/~hnusin tlie abundant but niore niobile lanternfishes and /~acifica), chaetognaths (Sagitta sp.), squid and other forage fishes (Pacific sand lance and juvenile pteropods, and carid shritiip (Sergestes sirrtilis). The Iiexegrammids) as \\,ell as tlie dense zooplankton viperfish and rattail prey sclccti\,ely on the north- populations, all apparently available within the ern lampfish, tlie most abundant lanternfish of that fish's habitat. This conclusion conforiiis wit11 fish community. The roughscale rattail also preys Haedrich (1964), Haedrich and Nielson (1966), and on its snialler competitor, tlie Pacific viperfish, and Parin, Nesis, and \Gnogi.adov (1968). Parin et al. niesopelagic nlysids (Gi~atophausiagigas), isopods described these fish as feeding at depths of 100 to (Arctttri~s spp.), euphausiids, and squid (Gal- 300 In, at the boundary of the epi- and mesope- ite~~tlfitrsarluata). California s~noothtongueprey lagic regions. This depth in the Indian Ocean is in on chaetognaths and calanoid copepods 'ivith a the lower levels of tlie upper isothermal layer and secondary contribution fro111 euphausiids. the upper levels of the principal thermocline. The maturing niigrating sockeye and chuni Without a niore thorough kno\vledge of the salmon of the epipelagic community are primarily distributions and abundances of tlie prey organisms piscivorous, preying on both tlie forage fish resi- or quantitative c\~alaationof predator exploitation, dent in that co~n~nunity(the Pacific sand lance) a cornplete sunimary of the dynamics of the and tlie planktivorous northern lanipfish of the trophic relationships betxireen tlie colnponcnts of adjacent niesopelagic community; tlie sockeye Amchitka's ~iiarinefood web is not possible. Given salmon probably captures these lanternfish during the first-order semiquantitative results of these that prey's diurnal ~nigrationinto the upper regions invcstig;itions, ho\vever, some important generaliza- of the deeper habitat. Hyperiid amphipods and tions can be made. calanoid copepods contribute to the diets of both The nearshore environment harbors a rich species of salmon, calanoid copepods being an diverse community of marine organis~iisprr)viding important secondary coniponent (25% composi- abnndant food resources to resident and immigrant tion) of tlie chum sal~nondiet. The pisci\~oroos fishes. This is, in part, a result of the diverse feeding habits of Amchitka's sockeye and chum macropliytic algal flora of the Amchitka littoral salmon may well reflect the patchy seasonal and inner s~tblittoralzones. The intricacy of this z~bunclance of fish [especially myctophids (Ian- colnmunity appears to be in response to the ternfislies)] occurring in the ul)welling area of the predation pressure on herbivores fro111 tlie systcm's offshore waters. Previous investigatiotis top carnivore, the sea otter (Palmisano and Estes, (Andrievskaia, 1957; Allen, 1956a; 1956b; Chap. 22, this volume). This comples algal com- Foerster, 1968; Le Brasscar, 1959; and Synkova, niunity not only provides a diversity of habitats for 1951) have sho\vn that the most common compo- benthic and benthipelagic macroin\~ertebratesbut nents in the diets of tliese salmon in the western also provides considerable autotmphically accumu- lated energy to the inshore marine food \veb constitutes an unpredictable rapid-s\\~immingre- through linkages to the abundant herbivorous source, it contributes to the diets of 8 of the 10 molluscs, echinoderms, and crustaceans. Direct species studied from the inshore/sand-gravel com- herbivory, however, is not as significant a meclta- munity. Its larval and prejitvenilc forms are also in nis~nof energy transfer in this system as indirect the diets of three fishes of the inshore/rock-algae herbivory, as was indicated by the lo\\. trophic community. This forage fish is the system's maill contributiotls of hcrbi\~orousorganisms to inshore means of energy transfer between zooplallktoll and fish. Even the opportunistic benthophagous rock the piscivorous fishes of the community. No greenling fail to prey significantly on herbivores in predator can afford to prey selecti\~elyon sand areas where other prey are abundant. Specifically, lance since its occurrence is apparently not certain the ntajor herbivore of the system, the sea urchin enough to guarantee a reliable food resource. (Stro?1g3docefztrotrts polyaca?~tl~zts)occurs only Holvever, \vlten it does occur, it appears to be the incidentally in fish diets. Again, the effect of sea selected preferred prey. Sto~nachsof predators otter predation 011 the inshore sea urchin popula- were found to be ~u~~usuallydistended with sand tions is to severely liltlit the availability of urchins lance \\'henever their scl~oolswere in the inshore to othcr inshore predators (Palmisano and Estes, waters. Even the rock greenling was obser\red to Chap. 22, this volunte). capture sand lance to distention during a large On the other hand, indirect energy transfer inshore influx of schools of sand lance in August through the production of detritus does appear to 1973. Studies of the avian predators of Amchitka be significant. Seasonal die-off and attrition, espe- inshore fish have also noted the occurrence of sand cially during the winter storm period, create large lance in the diets of many species (IVilliamson, masses of detached macroalgae in the upper littoral Emison, and \\'ltite, 1972). Offshore the sand lance region. Their decay releases great quantities of appears to suffer little predation pressure from detritus and nutrients into the inshore waters and eplpelagic piscivores. prontotes prodigious amphipod and mysid popula- The relative ilnportance of various components tions in the inshore habitats. The trophic intpor- of tlte A~ttchitka inshore and offshore marine tance of these benthipelagic lnacroiltvertebrates ill ecosysteltls can be fi~rtlterseen by cnulnerating and tlte food web is evident. Antphipods contributed a rveightitlg the food-web links identified in Fig. 8. mean 39% by ~veightto the diets of the 25 inshore Table 6 sho\vs the number of trophic contributions fish, and their mean frequency of occurrence Ivas (arrows in Figs. 8a and 8b) it1 each of four 72%; mysids, not as uridely distributed as the categories of percentage relative biomass; the four diverse amphipod taxa, contributed 13% by weight percentage categories are weighted and added to and 20% in frequency of occurrence. Energy provide a total weigl~tedcontribution of this prey transfer through brcakdo\vn of lnacroalgae is not to the ecosystem's fishes. Organisms in the 0 to limited to insltore amphipods and mysids. 24% column were weighted 1, those in the 25 to Palmisano and Estes (Chap. 22, this volume) indi- 49% column were weighted 2, etc. Theoretically, in cate that kelp fr,lgments collect around Hedo- this simplistic approach tlte higher the total 11holdfasts and thus provide detrital food weighted contribution, the greater the role these supplies to the special communities of inver- prey resources play in the structure and energy- tcbratcs associated \vithin and under the lloldfasts, flo\v dynamics of the ecosystem. As indicated in cspccially the isopods Idotea iuos~tese~~skiiand Table 6, the ltlost significant contributors are Esos/~l~ae~o~~~aa~n/~/ica~rda, \vlticlt also contribute amphipods, mysids, polychaete annelids, and to the diets of rock greenling and great sculpin. Pacific sand lance in tlte inshore ecosystem and Simenstad (1971a) suggested that inshore- amphipods, copepods, euphausiids, and northern offshore circulation along the bottom carries dc- lampfish in the offshore ecosystem. tached intertidal algac from the littoral bench into Spatial heterogeneity of the habitats studied inner sublittoral habitats xvherc fishes prey both on influences not only the structure of the associated littoral i~lvertebratescarried with this algac and on ani~ttalcom~nu~lities but also the colnplexity of the those sublittoral forms foraging on the algae. It xvas rood web involved. The homogeneous epipelagic also denlonstrated that the decaying masses of portions of both offshore and insltore habitats have algac in the upper littoral zones support Ilea\,)' much simpler patterns of trophic links between the populations of beach fly (Dq~terasp.) larvae \vhich fishes and their food resources than the spatially are then available at high tides to foraging rock heterogeneous inshorc/rock-algae or offshore greenling, spotted snailfish, and Dolly Varden. demersal/rock-sponge communities. This is as- The pelagic trophic equivalent of the amphipod siuned to be a result both of the greater diversity is the Pacific sand lance. Although this fish of food resource available in the heterogeneous ~VlariueFish Cowti1tt~nities 481

Table 6-Tropllic Contributions by Percentages of Relative Bio~llassand Total Weighted Co~~tributiorlsof Prey Organisms to InsLore and Offshore Amcllitka hlarin~Fi&cs

Contribution and xveigl~tingfactors* Weighted Prey organisms Total contributio~~s Inshore r\mphipods hIysids Shrimps Copepods Isopods Brachyuran crabs Lithodid crabs Cumaceans Gastropods Pelecypods Amphineorans Cephalopods Ophiuroids Polychacte annelids Insects Pacific sand lance Armorhead sculpin Rock sole Red Irish lord Offshore Amphipods klysids Shrimps Copepods Isopods Brachytcran crabs Litbodid crabs Euphausiids Gastropods Pelecypods Cephalopods Pteropods and heteropods Ophiuroids Polychaete annelids Chaetognatl~s Pacific sand lance Armorhead sculpio Red Irish lord Northern lampfish Pacific viperfish \Valleye pollock Pacific ocean perch California smoothtongue Bigeye lanternfish Pacific herring Spiny li~mpsucker Sturgeon poacher Tadpole snailfislt Unidentified cottids Unidentified fish

*The factors used in weighting are given in parentheses. 482 Sin~e~~starl,Isnkson, n11d ~Vakatn~~i conimunities and the conconlitant more gen- avian and mammalian top carnivores is sitnilarly eralistic feeding behavior of the fishes there. The limited. Williams (1938), Murie (1940), Lensink pelagic specialists (salmon, Pacific sand lance, and (1962), Burgner and Nakatani (1972), and Atka mackerel) exclusively exploit the patchy Palniisano and Estcs (Chap. 22, this volume) ill1 abundance of planktonic prey, \vIiich enables them report predation on marine fishes by the sea otter to lnaxinlize energy ingested per encounter with at Amchitka. ICenyon (1969) specifically identified prey (copepods, pelagic amphipods, and mysids) the smooth lcuiipsucker, red Irish lord, Atka concentrations. mackerel, and rock greenling as important con- The large specialist consulners of the dclnersal stituents of sea otter dict at Amchitka in tlie fish communities (great sculpins, arro!vtooth flotm- 1960s, \\.it11 the smooth hunpsucker occurring ders, and Pacific halibut) fccd on large high-energy- most frequently in the stomachs examined. As per-item prcy (fish and large crnstaceans), \vhicIi noted earlier, predation pressure may have sharply enables then1 to extract more energy per unit of reduced the abundance of this spccies in recent feeding time than if they esploited tlie more times. common snialler prey available in their habitats \Villiamson, Emison, and \\We (1972) have (Scliocncr, 1969). 'file sniall specialists of the reported fish predation by littoral and pelagic littoral commnnity (pricklebacks and gunnels) feed feeding birds, among them the Red-throated Loon on tlie abundant crustaceans. Their diets and (Gnuia stcllata), Pelagic Cormorant (P/talncrocorccx opportunistic feeding behavior reflect the necessity f~eln~qicus),Red-feccd Cormorant (P. lrrile), Ked- for tllcsc species (small, territorial, and situated in breasted A'lerganser (~l.Jergtissevrator), Bald Eaglc a constantly fluctuating environment) to monitor (Hnliaeet~cs/et~cocepltnlzts), Glaucous-winged Gull prey, predators, colnpetitors, and tnates within a (Lccrtcs gla~rcesce~~s),Arctic Tern (Ster~tapnra- liniiting habitat. Most of the generalists (rock disnen), Aleutiw Tern (S. nlelrticn), Cotnmon greenling, red Irisli lord, Pacific cod, ~\zalleye &lurre (Uria aalge), Thick-billed Aclurre (U. lootvia), pollock, and blackfin sculpin) arc species that Pigeon Guillenlot (Cepf~hzts cobo~zbn), Holned occi111y habitats ~vithconsistent heterogeneous Puffin (Frntercztlcc cornictclata), and Tufted Poffin concentrations of prey. These fishes are thus able (Lzotcla ciwltata). The common prcy of these birds to maximize energy consuliled by feeding continu- \\,as the Pacific sand lance, but the rock greenling, ously on a diverse diet of prey organisms. Atka mackerel, arlnorhead sculpin, red Irish lord, The benthipelagic atnpllipod and niysid popul;~. and spotted and lobefin snailfish were also repre- tions appear to be the n~ostnni\,ersal prey and the sented in their diets. Pacific cod and rockfish principal converters of plant detritus energy into species tverc also reported prey organisms, but it is energy sources avi~ilableto the secondary consumer suspected that tliese were scavenged from waste level, and the offshore eupliausiid populations froni a foreign fishing vessel offshore. play, to a lesser degree, a siniilar role; the principal energy inlporter apparently is the Pacific sand Impact of Amchitka Nuclear Tests lance, the mobile populations of wl~icl~continu- on Marine Fishes ously introduce energy derived fro111 the zooplank- ton resources of the surrounding oceanic region Principal interests of tlie U. S. Atomic Energy into the timchitka inshore ecosystem. Commission in FRI's study of Amchitka's marine Other investigators (see Palmisano and Estes, fishes \\'ere establishing pretest base lines of spccies Chap. 22, this volume) have documented the effect occurrence and abundance and of ecological rela- of the sea otter as a "keystone species" (Paine, tionships so that posttest colnparisons could be 1969) and its herbivorous prey, tlie sea urchin, an niade to evaluate the inipact of proposed nnder- influential "fou~idationspecies" (Dayton, 1972) in gromnd nuclear tests at Amchitka. This information structuring the inshore biotic community. The role and information fro111 the literature concerning of this interaction, thong11 indirect, in dctcrmining shock effects \sere cotnbined to provide inputs for energy flo\v in the system's food \veb should not the predictions and assessments by Battclle Memo- be ignored because it is the lower level conversion rial Institute and the AEC of the environmental of energy that eventually determines the coniposi- impacts of the i\,Iilro\v and Cannikin tests tion of the upper lc~~clconsumers, si~cllas the fish (ICirk\vood, 1970; Merritt, 1970; Kirk\vood and communities. Fuller, 1971; 1972). In the establisl~mentof thesc The lack of data on food sources of the prcy base lines, a great deal of nenr infc)rma t'1011 tvas organisms illustrated in Fig. 8 prevented us from obtainecl about the marine fishes of Atnchitka extending corresponding linkages to lower trophic Island. This section summarizes the observed levels. Docnmentatioll of predation by Amchitka's impact of the two tests (h,Iilrow, Oct. 2, 1969; Cannikin, Nov. 6, 1971) on Amchitka's marine fish com~nunities. A rcvic\v of the literature from pre\ions under- \\rater explosions and related experiments or phe- nomena (Simenstad, 1971b; 1974) indicated that the shock \vavc from an undergronnd nuclear explosion could inflict damage on marine organ- isms throi~gl~several IIICC~~~I~~SIIIS.Directly, it could damage or kill individual organisms through the mechanical impact or pressure cl~angcproduced when the shock wave passes through the water. It could also indirectly alter species composition, distribution, and/or abundance in the affected fish commnnitics through the alteration of the marine habitat, wltich \vould affect changes in food resources, living space, and reproductive substrate. The hydrostatic changes i~nposed by the shock Fig. 9-Aerial photograph of marine waters sur- xva\le \vere considered, ho~vever,to be potentially rounding test site during hlilrolv test, Oct. 2, 1969, the most danlaging to indi\idual fish. Two specific approximately 1.5 sec after the detonation. Note nlcchanislns are applicable it1 these circumstances: white rvater areas xvhich are regions of cavitation (1) Pressures over and under the anlbient pressure caused by stlock wave produced by the underground and (2) bulk cavitation. The literatnre docunlcnts nuclear detonation. (Photograph courtesy of Edgerton, Germesl~ausenand Grier, Inc.) that, for lot\?-frequency shock \\raves, such as those produced by the nnderground tests (10 to 15 1-12), the negative presstrre pltases of the hydrostatic fishes ctbser\,ed were t'ivo Pacific cod in the \\raters pressure history \vould be the most damaging to off Cyril Cove. On the basis of the live-box fishes, nlore so to physoclistous* than to phy- cxperi~ncntsand the observed fish kill, the damage sosto~noustand non-gas-bladdered species. Except to fish life caused by Long Shot \\'as judged for fast-rising shock waves incident on gas- insignificant (Seymour and Nakatani, 1967). bladdered fishes (as from dynanlite explosions), \\'it11 the background of the Long Shot cxperi- colnpression or overpressure was not shown to be cnce, in the sunliner of 1967 FRI began stitdies at detrimental. A phenomenon that limits the maxi- Amchitka, including base-line surveys of existing mum underpressure attainable, as a fnnction of fish populations for pretest and posttest compari- water depth, is cavitation, i.e., the reduction of sons of species composition and abundance of pressure to the point of producing gas bubbles fishes, and live-box experiments were conducted at within the water. Cavitation of an organism's body h4ilro\\~test time. Snrveys continued after the 1969 fluids was thought to be a1 important mechanism h~lilrow test through the period of preparation for of fish kill possible from the underground nuclear the Cannikin test in 1971 and continued thereafter tests (Fig. 9). The observed effects of the h,filro\v until the spring of 1973. and Cannikin events were evaluated with this - - Nakatani and Burgner (1974) and Isakson background of direct and habitat-associated ef- (1974) have reported on the impact of the h,lilro\\. fects. and Cannikin tests on the overall marine en\riron- An nnderground nllclear test (Long Shot, ment. Burgner, Isakson, and Lebednik (1971b) Oct. 29, 1965) of 80 kt at Amchitka Island su~nmarized the impact of the b,Iilrow test on occurred prior to our investigations. A group of Amchitka's lnarine environment, including fishes. biologists from several organizations conducted We repeat the major findings belo~v. su~veysand experinlents to evaluate the impact of this test on the farina of Amchitka. Seymour and hlilrow. Fish surveys were conducted botll Nakatani (1967) reported no fish killed in live- inshore and offshore before and after the h,Iilro\v boxes in the ocean as close as 1.6 km (Cyril Cove) test. Studies \\,ere concentrated on the Rifle Range to Long Shot surface zero (SZ).$ The only dead Point-Duck Cove area (Fig. 1). Analysis of the catch-per-unit effort (fish per hour) before and *Possessing no pneumatic duct bettveen the gas bladder after the test for all gear types used revealed no and the alimentary canal. significant or nlarked changes in fish abundance tpossessing an open pneumatic dact bet>veerr the gas bladder and the alimentary canal. offshore or inshore (Burgner et al., 1971a). $That place at the island's surface \vhere the emplace- Immediately prior to the test, three pairs (six ment hole was started. total) of live-hoses containing rock greenling and Fig. lOa-Predicted overpressure [in megapascals (pounds per square inch)] at bottom in Bering Sea adjacent to Cannikin SZ.

red Irish lord Itrere placed in Duck Cove at capture for the Milrolv holding experiments. The distances from 2.6 to 4.0 km (8,500 to 13,000 ft) three injured fishes seen after the test may have from A~lilro~vSZ. These fishes were exposed to been Pacific cod or rockfish, \vhich possess gas overpreswres of 0.59 to 1.17 h'lPa (85 to 170 psi), bladders. imderpressures of about -0.57 h(1Pa (-83 psi), and No accurate estimate of fish kill by Milrow is acceleratio~lsof 2.5 to 5g (hkrritt, 1970; 1973). possible from such insufficient information. \Ve Other than two rock greenling, which apparently believe there may have been a small kill of marine escaped, all fishcs in the live-boxes survived. After fishes, but it could have had little impact because it the test these fishes were held for 4 xveeks in \\,as not detectable in pretest and posttcst com- seawater tanks and then released, no further or parisons. Indirect effects related to disruptioll of delayed illjuries having been noted. marine fish habitat \\,ere not detected. Three hours after the test, from a helicopter three unidentified and apparently injured fishes Cannikin. After h,lilrow, sampling effort was wcre seen thrashing on the oceau surface, two off shifted to the Bering Sea adjacent to Caullikill SZ. Rifle Range Point, about 1.9 km (1.2 miles) Locations ranged from Petrel Point to Crown southwest of ivlilrow SZ, and one off Crown Reefer Reefer Point and also illcluded a seco~ldarea Point, about 5.6 km (3.5 miles) north of Milro\v adjacent to ICirilof Point aud ICirilof Bay. SZ. These fishes could not be collected because the The salnplil~gdesign iitlcludkd both test-time hclicoptcr could not safely land. No other dead or live-box experiments and pretest and posttest injured fishes were seen during the posttest beach surveys in the areas where fishes \\'ere thoaght surveys. most vuherable to pressure changcs. Sampling and Only rock greenling and red Irish lord \\'ere live-box positions were established with the use of pl;tced in the test-time live-boxes because they maps of peak water-pressure changes (Figs. 10a and were the only species that survived the rigors of lob) predicted to occur during the test. The theory Fig. lob-Prcdicted underpressure [in megapascals (pounds per square inch)] at the bottom in Bering Sea adjacent to Cannikin SZ. and calculations behind these predictions resulted of handling, possibly duriug cage retrieval, rather from the kIilro\\, pressure-time data (A,Ierritt, than a test effect. All experimental fishes and 1970; 1973; 1974; personal comn~unication, control fishcs xvcrc hcld for 8 days after the test, 1971). and no delayed mortalities or injuries were noted A severe storm, with winds of 110 kin/l~s,gust- in the experinlental fishes. ing to over 150 km/lir, throughout the day and night Immediately aftcr the test, no dead or injured before the Cannikin test resulted in heavy seas, fishes were seen from a helicopter or fro111 a ship. \vhich prevented the placement of live-boxes as Ho\vever, viexving was poor because of heasy scas. planned (Nakatani, Isakson, and Burgner, 1973, Beginning about 4 hr aftcr Cannikin and during Appendix B; Isakson, 1974). Instead, 7.5 hr before daylight hours for 3 days after the tcst, biologists the test a single live-box string \\'as placed in from several organizations surveyed the Bering Sea 18-m(60-ft)-deep water in Constantine Harbor and Pacific Ocean coasts adjacent to the Cannikin about 15 km (9 miles) from CanElikin SZ. A SZ area. The areas surveyed and the fishcs se- live-box at 16-~n(52-ft) depth contained sis rock covcrcd are shoxvn in Fig. 11. Almost all the dead grceuling, four Pacific cod, three Pacific halibut, fishes found were stranded on tidal benches or two red Irish lord, one rock sole, one great sculpin, beachcs uplifted as much as a meter. Otherwise, and one dusky rockfish. Another liuc-box at 2-111 owing to the north\vesterly winds prevailing during (7-ft) depth contained 10 rock greenling and 6 the tcst, fishes washed up on the shore were found Pacific cod. Three hours after the tcst, only one ~nostlyon the Pacific Ocean side of Amchitka. fish, a 67-cm (26 in.) Pacific cod, shelved any Of the 295 fishes recovered during beach \valks, abnormality, a "bubblc" in the right eye and the majority (277) were rock greenling that \crere difficulty in equilibrium maintenance. Because no stranded on uplifted intertidal benches and washed simiiar problems \\'ere seen in the other fishcs, it into co~~es011 sand and cobble beaches. Autopsies was assumed that thcse symptolns were the result by Dr. Robert Rausch (U. S. Public I-Iealth Service, Alaska) of 23 rock greenling revealed that about cxpectcdt posttest data were directly compared to half had hemorrhaged or sustained other da~nagein plxtest data For the salnc areas. The geometric the brain cavity and/or the viscera as the rcsult of mean catch of rock sole (average catch depth) water-pressitre changes, acceleration \vhile over the significantly (t-test statistic, 2.52) declined from littoral bench, or in struggling once the beach had about 100 fish per hour pretest to a mean catch of lifted.* The other autopsied rock greenling ex- about 31 fislt per hour posttrst. [See Nakatani et hibited no noticeable i~lj~u)'. al. (1973) for further explanation of statistical 'The Pacific sandfish \\,ere found stranded in catch comparisons.] DiKcrcnces bet\veen Pacific uplifted sandy beach areas and in drift zones of halibut ;nld total bottom tra~vl catches were Sand Beach Cove (Fig. 11). This spccies commonly statistically insignificant. Dcclines were obscr\~ed burroxvs upright in the sand of the littoral zone, for sahnon species, Pacific cod, rockfishes and the fish were probably strandcd there by the (Sebastes), and inidwater fishes, but occitrrcnce beach uplift during the test. Since longnose and abundance of these species in thc catches were lancetfish are often ~vashedashore after storms, the too lo\\< to pcrmit statistically valid comparisons of one specimcn fo~n~dafter Cannikin is assu~ncdto prctest and posttest catches. No decrease in catch be a result of the pretest storm and not the test. of the nlore vulnerable physoclistous species (Pacific cod and rockfishes) \\,as detected by this The remains of Pacific cod and rockfish found method, although the facts of dead cod having on the Pacific shore had been stripped nearly to been found on the Pacific Ocean shore and grcater skeletons by amphipods. Because both of these fish pressure changes being cxpccted on the Bering Sea species arc physoclistous, they \\,ere probably side imply that some decrease might have been killed by \\rater-pressure changes, and a portion of expected. This species is not a donlinant of the fish those ~vhichsurfaced were blown onto the shore. community in the sa~nplingarea, ho\vever. Cannikin being nearer the Bering Sea, grcater We estimate the size of the marine fish kill pressure cltanges (both in magnitude and in area) from Cannikin to be in the thousands (I

DATE Fig. 12-Trammel net catchcs of rock peenling per hour, for short (<5hr) sets in inshore/rock-algae fish community adjacent to intertidal sites IA-2 and IA-3, ,\ugust 1971 through October 1972. A range is indicated where niore than one set was made on a given day. le\rels by fall 1972 wes lien catches averaged about impact on Bering Sea and Pacific Ocean marille 8.5 rock greenling per hour (Fig. 12). As with tlie fislies \vithin a 6-kni (4-mile) radius of the test site. offshore rock solc, rock greenling repopulatio~iwas Complete detectable recovery of these populations, most likely by thc i~ii~iligratio~iof rock greenling probably by immigratio~lfrom unaffected areas, from nearby ~maffcctednearsbore waters. occurred after 1 year. Because of adverse ~veathes Finally, two a~iomalies \irere noted in the co~lditio~isand tlie tendency for dead fishes to spccies composition in tra~ninelnet catches in the sink, few dead fish (Pacific cod and rockfish inshore fish commiu~iities before and after species) were recovered after the test, but the Cannikin. In the inshore/rock-algae conimunity, i~nplications of our other data are that marine four species (searcher, great sculpin, Pacific hali- fishes ~lu~nberi~lgin tlie order of thousands were but, and Alaskan ronquil) werc not captured after killed in the vicinity of Can~iiki~iSZ. Although this Cannikin; a fc\v indi\.iduals of each of these species may seem a major impact, its sig~lifica~iceto tlie had been caught before. I11 the inshore/sand-gravel marine ecosystem of Amchitka Island must be community, two species (Atka mackerel and dusky considered as minor, especially \vIien there was a rockfish) were caught before but not after documeuted recovery witl~ina year or less. Cannikin. In both cases, considering seasonal dif- The ~upliftiligof a section of the littoral bench ferences, differences in tlie duration of sets, and and co~lsequentloss of habitat had only a slight tile infrequent occurrelice of these species in the illdirect inlpact on inshore ~iiarine fishes. The area, the species co~npositionand abundance are uplifted area no longer available to these fishes is coiilparable (Nakatani arid Burgner, 1974). but a small percentage of the total littoral habitat of Amchitka. Again, as 114th fish kills, this impact Snmmary of 111ipacts on Marioe Fishes \\,as co~lfi~ledto tlie Cannikin SZ vicinity.

I11 revie\\,, the data available from Cannikin esperimeuts aud obser\~atiotisdo not per~iiitprc- cise quantification of tlie impact of that tcst on marine fish communities. Camliki11had both direct Research on tlie marine fishes in the seas and indirect impacts on marine fishes inhabiting aro~und Anicbitka Island was co~lducted over a the Bering Sea littoral bench, which \\?as uplifted 7-year period, the primary objective being to during tlie tcst. These areas have been denied for evaluate the prompt and long-term impacts of some time, perhaps forever, as habitat for marine xlirclear testing on marine fish populations. Despite fishes, both those tvhich are resident members of the mission-oriented nature of this research, nrhich the comm~mityand those of adjacent colnmunities imposed some restraints on the scope of the \vliich use the bench for feeding at high tide, e.g., in\~estigations,the studies hase led to iie~vunder- the rock greenling. A precise quantitative assess- standings of the role of the Amchitka niarine fish meut of fishes actually killed duri~igthe ilplifti~ig fau~ia in the structure and fu~ictio~iiligof the cannot be with the data available; however, niarinc ecosystem surrounding the island. Informa- hundreds of dead fish were collected, which tion collected on the 92 fish species taken from iliiplies a significant short-ten11impact. marine xvaters around Amchitka has significantly The direct i~iipactsobserved after Ca~i~iiki~l espauded kno\vledge of Aleutian ichthyofauna. \\.ere not fully espected (Fig. 11). The majority of Collect ions demo~istrated the presence at the dead fishes recovered \\,ere found oil or near Amchitka of several fish species not previously the uplifted Bering Sea littoral be~~chor along tile reported beyond the eastern part of the North Pacific Ocean coastline. One trawl in the Bering Pacific. The nelv i~lfortiiationreinforces the con- Sea, 6 km (4 miles) from Ca~miki~iSZ, captured a cept of an Aleutian "filter bridge," which was dead rockfish, a significant datum since we have suggested earlier by l\rilimo\~sky(1964) and others. never before, in over 4 years of trawling about The Alaska Stream al)parently serves both as the Amchitka, e~icou~ltcrcdan obviously dead fish in pri~liary dispersal mechanism for pelagic eggs, our catches. Co~iiparison bet\\reen pretest and larvae, and juvc~iilesof indigenous fishes and as a posttest catches in the Bering Sea off Cannikin SZ b,drller .' to east\vard espansiou of western Pacific sholt'ed marked dcclines in rock sole in offshore species. However, this barrier is not an absolute otter tralvl hauls and rock greenling in inshore one because we have tentatively identified larvae of trammel net catches. These two species of fish typically Asiatic species in collections made off reco\~ercdto pretest lc\,els within a year after the Amchitka. We can only speculate about possible test. exclusion nlechatiisms respo~isiblefor the occur- From these observations \\re conclude that lence of otily larval for~iisof these species in the Catl~iiki~ihad a significant short-tcrlli prompt vicinity of Amchitka. The structure of se\len fish communities asso- shock wave of 10 to 15 HL coupled 114th bulk ciated \\,it11 specific epipelagic, nlid\\rater, aud ca\ritation of the marine waters in a relatively slnall bottom habitats has been characterized in rela- area adjacent to the Cal~nikilltest site resulted in tively broad and subjective terms. Recurrent group the death of large numbers of several species of fish analysis of the data identified 13 species associa- in offshore waters. This conclusion xvas based on tions in five of the colnlnullities and provided an observed posttest decline in catches of rock additional understanding of the internal structure sole, a fish that theoretically, by its lack of an air of the species groupings, especially ~vithrespect to bladder, should be little affected by such pressure some significant relationships between inid~vater changes. Populations of more susceptible species, predator species and their invertebrate prey. such as Pacific cod aild several rockfish species, Our sainplillg methods were not sufficiently \\'ere assumed to have been affected also, although precise to detect ally statistically sigilificant direct e\ridence to support this assumption is changes in community compositioll during the meager. A gear after the test, catches of rock sole 7-year collection period. Ho\vever, catch data equaled pretest catches; rccstablishmetlt of the rc\.ealed what appeared to be one population shift population was probably by immigration from of collsiderable interest, which occurred after an nearby cullaffected stocks. intensive harvesting of Pacific ocean perch near In the littoral en\~ironment,catches of rock Amchitka by foreign fisheries in the late 1960s. greenling declined sharply in Bering Sea waters Following this intensive fishing effort, there ap- adjacent to the Cannikill site iminediatcly posttest. peared to have becn a reversal of dominance in the The decline was attributed to a substantial kill of demersal Pacific oceail perch-~valleye pollock asso- rock greenling it1 the area resulting from a com- ciation, with \\.alleye pollock becolning the billation of forces generated by the Cannikin dominant member of the assemblage. detonation. Catches posttest indicated that the Our studies indicate that the algae-based detri- affected population had begull to recover within 2 tus component of the insl~oremarine ecosystein is weeks after the detonation and had fully recovered largely responsible for the structure and energy by fall of the next year. Recovery was judged to flow in the food web of that ecosystem. Results of have been by inlnligratioll of greellliilg from stocks stomach analyses of fishes suggest that the es- outside the affected area. tremely high densities of epipelagic crustaceans Althougll both offshore and inshore fish popu- that use this food base (amphipods and mysids) lations in waters adjacent to the Amchitka test support a large standing crop of benthipelagic- sites were adversely affected by the iluclear deto- feeding fishes, most uotably rock greenling and nations, especially by the Cannikin test, the Atka mackerel. The ~vholequestion of the stability changes were short-term. \\'ithitl a year after the of this particular food-web structure takes 011 ile\v Cailnikin detonation, no effects on fish popula- dimensions when the rolc of the top carllivore of tions were detectable. Since the total area of the system, the sea otter, in regulating llcrbivore marine l~abitataffected by the tests xvas such a populations is realized (Palmisano and Estes, Chap. small fraction of the total habitat around the 22, this \.olunlc). island, the o\rerall effect of the tests on the Amchitka marine eeosysteln should be considered As a major prey organism of nearshore fishes, transitory and insignificant. the Pacific sand lance \\.as second only to amphi- pods in the number of idelitified food-\vcb links to inshore predators. Although \\re were unable to REFERENCES evaluate fully the abundance of the Pacific sand lance, we consider that the role of this plank- Abakumov, V. A., 1964, On the Oceanic Period in the Life Cycle of the Lamprey Estosphenus tl.ide~~tntus tivorous fish in importing biomass from offshore (Richadson), in Soviet 1:isheries Investigations in the waters into the inshore ecosystelll is an itnportallt Northeast I'acific, I'art 11, translated from Tr. I/ses. feature of the inshore portion of the lnarillc food Nnuch.-Issled. Inst, rlforsk. Kvb. Khor. Okeaaow,.-. xveb. \'ol. 49, by Israel Program for Scientific Translations, TT67-51204. VD. 268-270. The effects of the undergromnd nuclear testing i\llen, G. H., 1956a, Food of Salmonid Fishes of the Nortll prograin \\,ere very difficult to e\,ali~ate,but our Pacific Ocean. a. Food of Sockeyc Salmon (0. nerkn), efforts in documenting the impact of pressure University of \\'ashington, Department of Oceanogra- pnlses and shock wa\res 011 Amchitka lnariue fishes phy, Collection and Analysis of Oceanographic Data have coiltributed to the Iinlited knorvledge of the from the North Pacific Ocean, Fishery Report No. 1. -, 1956b, Food of Salmonid Fishes of the Norti, I'acific prompt effects of such perturbatious on aquatic Ocean. b. Food of Chum Salmon (0. keta) (with Notes organisms. Bottoin pressure cllanges of 300 psi on the Food of Sockeye and Pink Salmon), University over and 200 psi under aillbient pressures in a of \l'ashington, Department of Oceanography, Collec- tion and Analysis of Oceanographic Data from the Gilbert, C. H., 1895, The Ichthyolo$cal Collections of the North Pacific Ocean, Fishery Report No. 2. Steamer Albntross During the Years 1890 and 1891, Americati Fisheries Society, 1970, A List of Commof~njld U. S. Comniission on Fish and Fisheries, Report of the Scientific Arnt,ies of Pishes frotn the United States and Comniissioner for the Year Ending June 30, 1893, Part Ca~tnda,3rd ed., Spec. Publ. No. 6. 19, pp. 393-476. hndrievskaia, L. D., 1957, [The Food of Pacific Salmon in Gorhunova, V. N., 1962, Spa\rrning and Development of the North\vestern Pacific Ocean.] hlaterialy po Biologii Greenlings (Family tleragallmlidae), in Greenlings- hlorskogo Perioda Zhizni Dal'ne~~ostochnykhLososei, Taxonomy, Biology, Interoceanic Transplantation, T. S. pp. 64-75, translated from Tr. I'ses. i\'ouch.-Issled. Rass (Ed.), pp. 121-183, translated from Acnd. Sci. Inst. i\lorslr. Ryb. Khoz. Okensogr., Fisheries Research USSR, Trnss, h~st.Ocennob, Vol. 59, by Israel Program Board of Canada Translation No. 182. for Scientific Translations, 1970. i\ndriyashev, A. P., 1954, Fishes of the Northern Seas of Gordeeva, K. T., 1954, Pitanie Paltusov v Beringovon~hiore the USSR, translated from Zool. Inst. USSR Acnd. Sci., (Food of llalibut in the Bering Sea), Izu. TII\'RO, No. 53. bv Israel Prom'am for Scientific Translations. Voi. 39. ~~63-11160,1964. " Grccnnfood, P. H., D. E. Rosen, S. 11. Weitzman, and G. S. Bean. T. 11.. 1882a. Descrintions of Nerv Fishes from hlyers, 1966, Phyletic Studies of Teleostean Fishes, hlaska atid ~ibert the Ensterl~Pncific munity Resilience and the Potential Effects of Enrich- Ocean, U. S. Departtnent of the Interior, Bureau of ments to the Benthos at hlcMurdo Sound, Antarctica, Sport Fisheries and \\'ildlifc, North American Fauna, in Proceedhrgs of the Colloqtlium on Co~~seruotioa No. 68. Problettts in ,I~ttnrcficn, B. C. Parker (Ed.), pp. 81-95, IIchthyology, pp. 418-427. Gilbert) in tllc Gulf of i\laska, in Soviet Fisheries Pulliam, 11. R., 1974, On tlre Theory of Optimal Diets, Investigations in the Northeast Pacific, Part I, P. A. Anrer. ~Vntur., lOX(959): 59-74. h'loiseev (Ed.), pp. 308-318, Tr. Vses. ~\'nrzch.---Issled. Quast, J. C., and E. L. Hall, 1972, List of Fishes of Alaska Inst. dforsk. Ryb. Khor. Okea~togr.,Vol. 48. and Adjacent \\'atcrs with a Guide to some of Their hlacArthor, R. H., and J. \\I. hlacArthur, 1961, On Bird Literature, NOAA Technical Report NbIFS SSRF-658. Species Diversity, Ecolog)~,42: 594-598. Rapport, D. J., 1971, An Optimization hlodcl of Food hlakarova, R. I., 1975, Rtrssinns on the Pncijic, Selection,Arner. Nntur., 105: 575-587. 1743-1799 (translated and edited from the 1968 Scheffer, V. B., 1959, Invertebrates and Pishes Collected in Russian original by R. h. Pierce and A. S. Donnelly), the Aleutians, 1936-1938, in Fnuan of the Aleutinn Lin~estonePress, Kingston, Ontario (hlaterials for the Ixlnt~ds n,rd Alaskn Peizbtsuln, OlausJ. hlurie, Study of Alaska History, No. 6). pp. 365-406, with Notes on invertebrates and Fishes hlcAlister, \\I. B., 1971, Oceanography in the Vicinity of Collected in the Aleutians, 1936-1938, U. S. Depart- t\rnchitka Island, Alaska, BioScience, 21(12): 646-651. ment of the Interior, Bureau of Sport Fisheries and blcCaon, J. A,, 1963, A Summary of Biological Field IVork IVildlife, North i\merican Fauna, No. 61. done on ilrnchitka Island, Aleutian Islands National Schoener, T. \V,, 1971, Theory of Feeding Strategies, Aans. \\'ildlife Refuge, Cold Bay, Alaska, unpublished. Rev. Ecol. Syst., 2: 369-404. blerritt, M. L., 1974, Characteristics of Shock \Saves in -, 1969, Optimal Size and Specialization in Constant and \Vater Near Underground Explosions, in Proceedings of Fluctuating Environments: l\n Energy-Time Ap- the First Conference on the Environmental Effects of proach, in Symposium on Diversity and Stability in Esplosives and Explosions, \Vhite Oak, hid., klay 30- Ecological Systems, Upton, N. Y., hlay 26-28, 1969, 31, 1973, George A. Young (Comp.), Report NOLTR- USAEC Report BNL-50175, pp. 103.144, Brookhaven 73-223, pp. 78-85, Naval Ordnance 1,ahoratory. National Lahoratoly. -, 1973, Pressures in \\later on and Near r\mchitka Schultz, 1.eonard P., 1939, A New Genus and Two Nerv Island, hlilrolv and Cannikin, USAEC Report Species of Cottoid Fishes from the Aleutian Islands, SC-RR-720547, Sandia Laboratories. Proc. U. S. A'atl. illus., 85(3038): 187.191. -, 1970, Physical and Biological Effects: i\lilron~Event, Sevmour. A. 1.1.. and R. E. Nakatani. 1967, Lone Shot USAEC Report NVO-79, Neeada Operations Office. Bioenvironmental Safety Program, Final Report, h,lurie, 0. J., 1940, Notes on the Sea Otter, J. ~\Ian~rnnlogy, USAEC Report TID-24291, University of \\'ashinaton.- 21(2): 119-131. Shmidt, P. U., 1904, Pisces zllnritctn Orientnlirtnr, St. Nakatani, R. E., and R. I.. Burgner, 1974, Amcl~itka Petersburg. Bioeneironmenta! Program. Research Program on Shubnikov, D. A,, and L. A. I.isovenko, 1964, Data on the Marine Ecology, t\mchitka Island, Alaska, Annual Biology of Rock Sole of the Southeastern Bering Sea, in Progress Report, July 1972 'l'hrough September 1973, Soviet Fisheries Investigations in the Northeast Pacific, and Summary Report, 1967 'l'hrot~gl~1973, USAEC Part 11, P. A, hloiseev (Ed.), pp. 220-226, Tr. Irses. Report BhIl-171-156, 13attelle hlemorial Institute. Arnach.-Issled. Inst. Morsk. Hyb. Kltor. Okcnttogr., -,J. S. Isakson, and R. I,. Burgner, 1973, A~nchitka Vol. 49. Bioenvironn~cntal Program. Research Program on Simenstad, C. A,, 1974, Biological Effects of Underground h'larine Ecology, Amchitka Island, Alaska, Annual Nuclear Testing on hlarine Organisms. 1. Review of Progress Report, July 1, 1971-June 30, 1972, USAfiC Documented Effects. Discussion of hlcchanisms of Report BM1:171-150, Rattelle hlemorial Institute. Damage, and l'rcdictions of Amchitka Test Effects, in Nelson, E. \Y., 1887, 1;ield Notes on Alaska Fishes, rvitll Proceedings- of the First Confcrencc on the Environ- Additional Notes by Tarleton H. Bean, in Report Upolt mental Effects of lixplosives and Lixplosions, \!'ltite ~Vaturnlllistory Collections i11ode i~rAlaskn Betrueen Oak, hld., hlay 30-31, 1973, George A. Yot~ng the I'ears 1877 nnd 1881, Henry I\'.IIenshaw (Ed.), (Comp.), Report NOLTR-73-223, p1). 86-97, Naval pp. 295-322, Arctic Scries Pi~hlicationsNo. 3 Issucd in Ordnance Laboratory. Connection with the Signal Service, U. S, Army. -, 1971a, The Feeding Ecolo~rof the Rock Greenling, Novikov, N. P., 1964, Basic Elements of the Bio1o.w of the Ilesagranrnros Ingoceplrnlrrs, in thc Inshore Waters of 1,.,ICI~I< :. Il.tlil>ttt (//i/,po,~lo~cr~rhi/,/,oglos,rrr rlc~,rolr/,i~ Amchitka Island, r\laska, b1.S. Thesis, University of S<.l~nliFlatfishes in the Southeastern Bering with the Signal Service, U. S. Army. Sea, in Soviet Fisheries Investigations in the Northeast U. S. Bureau of Fisheries, 1907, Dredging and Hydro- Pacific, Part 1, P. I\. hloiseer, (Ed.), pp. 235-250, Tr. graphic Records of the U. 8. Fisheries Steamer e.1Vuuch.-Issled. Inst. 11Iorsk. Ryb. Khot. Albntross for 1906, Bureau of Fishcries Document 621. Okeamtogr., Val. 48. \Yhittaker, R. H., 1970, Co~n,nunitiesn~~dEcosystcms,The Stcjneger, L., 1936, Georg Il'ilhelnz Steller: The Pioneer of h~lach~lillanCompany, London. lllasknn i\'ataml History, Harvard University Press, \lrilimovsky, N. J., 1964, Inshore Fish Fauna of the Cambridge, Mass. Aleutian Archipelago, in Proceedi~rgsof the 14th Alaska Synkova, I\. N., 1951. [On the Food of Pacific Salmon in Scimrce Corrference, pp. 172-190. Ka~ncllatka\\'aters], Izu. TIArRO, 39: 105-121 (Fish- -, 1958, Provisional Keys to tile Fishcs of Alaska, U. S. eries Research Board Translation No. 415). Tanner, Z. L., 1892-1896, Report Upon the h~vestigations Fish and \lrildlife Service, unpublished. of the U. S. Fish Commission Steamer Albatross for -. 1954. List of the Fishes of l\laska. Stmrford Ictrth. the Year Ending June 30 (1889-1894), U. S. Commis- iull., 4(5): 279-294. sion on Fish and Fisheries, Reports to the Commis- Willian~s.C. S.. 1938. Notes on Food of the Sea Otter..