BULLETIN OF MARINE SCIENCE, 47(1): 149-158, 1990

ECOLOGY OF PRJAPULUS CAUDATUS LAMARCK, 1816 () IN AN ALASKAN SUBARCTIC ECOSYSTEM

Thomas C. Shirley

ABSTRACT Priapulid () were a conspicuous component of the meiofauna «0.500 mm) and macrofauna (>0.500 mm) in Auke Bay, Alaska (58°N, I34°W) from 1985- 1988. The smallest priapulid collected had a lorica of 0.050 mm length, while total lengths of adults exceeded 150 mm. Priapulids were usually the third most abundant mei- ofaunal organisms, following and harpacticoid copepods, with densities to 58,000' m-2 at subtidal stations of 25-55 m depth. Priapulids were much less abundant in the

macrofauna, never exceeding 85, m-2• Large interannual variation in densities of larval pria- pulids occurred. The smallest larval stages were found during the winter months, the apparent spawning period, with greatest densities of larvae occurring in early spring. Growth rates suggested by length-frequency distributions support a 2-year larval period.

The Priapulida are a small of benthic, marine worms of obscure phy- logenetic affinity that have been allied with a variety of phyla (Candia Carnevali and Ferraguti, 1979; Conway Morris, 1977; Lang, 1963; Malakhov, 1980; Por and Bromley, 1974; Shapeero, 1961); most recently they have been aligned with the (Land and Norrevang, 1985). Approximately 15 extant and 6 species have been described (Land and Norrevang, 1985; Conway Morris, 1977), with Priapulus caudatus Lamarck undoubtedly the best studied (Carlisle, 1958; Lang, 1939, 1948a; 1948b; Nyholm and Borno, 1969; Shapeero, 1962a; Wesenberg-Lund, 1929). Priapulids are generally considered rare and biologically important only in marginal environments (Calloway, 1982; Land, 1970). However, in a long-term benthic study in Auke Bay, Alaska (Shirley et al., 1988), Priapulus caudatus were common and conspicuous members of the meiofauna as larvae and macrofauna as adults. Auke Bay is not a marginal environment, but has a diverse ichthyofauna (Haldorson et al., 1988), macrofauna and meiofauna (Shirley et al., 1987, 1988; Coyle and Shirley, in press). Despite its widespread northern circumpolar distri- bution (Land, 1970), few quantitative data exist regarding the ecology of P. cau- datus. New quantitative data available on long term densities and seasonal size- frequency distributions of priapulids were the impetus for the present paper.

MATERIALS AND METHODS

Sampling was conducted in Auke Bay, Alaska (58°22'N, 134°40'W), approximately 20 km north of Juneau (Fig. I). Auke Bay is a partially enclosed embayment with an area of approximately 16 km2 in a system of fjords connected with the Pacific Ocean. Approximately 50% of the bay is between 40- 60 m depth and has a muddy substrate. The bay has semidiumal tides with amplitudes of 3-7 m. Bottom water temperatures at 25 m depth vary between 3-7°C annually and salinities vary between 29 and 320/00annually (Coyle and Shirley, in press). A modified Kajak corer, the Tom-Tom corer, was used to collect meiofauna samples. The corer

149 ISO BULLETIN OF MARINE SCIENCE, VOL. 47, NO.1, 1990

STI:PHENS PASSAGE

o 2 • I KM

Figure I. Map of Auke Bay, Alaska with locations of sampling stations: ABM = Auke Bay Monitor, E = East, FC = Fritz Cove, I = Intertidal, N = North, NE = Northeast, UAF = Juneau Center for Fisheries and Ocean Sciences" University of Alaska Fairbanks. collects four replicate cores 2.5 cm in diameter each time it is deployed and has proved to be reliable and effective (Chandler et aI., 1988). All cores were collected between 0800 and 1100. Macrofauna were retained fi'om meiofauna core samples on all sampling dates in 1986 and examined for priapulids. Additional macrofauna samples were collected with four 0.1 m' Petersen grabs on the same dates. Meiofauna samples werc collected from four subtidal stations in Auke Bay and one station in adjacent Fritz Cove (Fig. I) from 1985-1988: East (E, 27 m water depth), Auke Bay Monitor (ABM, 55 m), North (N, 42 m), Northeast (NE, 42 m) and Fritz Cove (FC, 25 m). In 1985 eight cores were collected at 14-d intervals from E and FC from 3 April-IO June, and in 1986 at lO-d intervals from 8 May-7 August at E. The most intensive sampling was conducted in 1987 from ABM, E, Nand NE: four replicate cores were colle(:ted at 2-week intervals prior to the spring phytoplankton bloom, weekly during the bloom and for 6 weeks after its sedimentation. Cores were collected at 2-week intervals thereafter throughout the summer. Samples were collected monthly during the fall and winter from E and ABM, yielding a total of 20 sampling dates during 1987. During 1988, four cores were collected weekly from E and ABM from 16 March to 16 June, a total of 14 sampling dates. Priapulid data were therefore collected during 13 of 15 months from April 1987 to June 1988 on 34 sampling dates. Intertidal meiofauna was sampled at 2-week intervals from March 1987 to July 1988 from a single station (0 m) in Auke Bay with four cores collected with hand-held 50-cc syringes (McGregor and Shirley, 1988). The upper 5 cm of sediment from all cores and the overlying water was returned to the laboratory for processing where samples were sieved with seawater, except in 1986, when the entire core was processed. All material passing through a 0.500 mm mesh sieve but retained on a 0.063 mm mesh sieve was then extracted live with sucrose centrifugation (Fleeger, 1979). Sediment pellets resulting from centrifugation were examined weekly for sorting efficiency, which was estimated to exceed 95%. Macrofauna remaining on thc~0.500 mm sieve were narcotized with MgCl and preserved in 10% buffered formalin. All organisms were identified to major taxon under a stereo-dissection microscope. Nematodes were subsampled with sorting trays and a technique which employs a triply-balanced square design described by Sherman et al. (1984). Lorica lengths of all available priapulid larvae to a maximum of 50 were measured with an ocular micrometer each month. SHIRLEY: ECOLOGY OF AN ALASKAN PRIAPULID lSI

RESULTS Priapulid larvae and adults were present at all subtidal stations sampled in Auke Bay and Fritz Cove; none were present in intertidal samples (Fig. 1). Densities ofpriapulids were not significantly different by depth among the subtidal stations. Priapulid larvae were present only in meiofaunal samples, as they were not re- tained on macrofauna sieves (0.500 mm mesh). Small numbers of metamorphosed priapulids having adult morphology, e.g., lacking the lorica and possessing ru- dimentary caudal appendages, were also present in meiofaunal samples. Large interannual variations in mean density of larval priapulids, from < 1.0' 10 cm-2 in 1985 to >50.0· 10 cm-2 in 1988, were recorded from the single station (E) sampled all 4 years (Fig. 2), with densities generally increasing from 1985 through 1988. Intraannual variation also occurred, with larval densities usually being highest in the early spring and declining in the summer months, although large variation often existed between dates. Densities of the smallest size class of larvae increased in December, 1987, the only year that winter samples were collected (Fig. 3a). Total meiofauna also had interannual variations in mean density during the 4 years. Mean total meiofauna density varied by an order of magnitude at E from 1985-1988, from a minimum of 520·10 cm-2 in 1987 to a maximum of 5,900' 10 cm-2 in 1986 (Fig. 2). Total meiofauna were heavily predominated at all subtidal stations by nematodes, which comprised 97% of the total meiofauna across all dates and stations (Shirley et aI., 1989). Harpacticoid copepods were usually the second most numerically abundant taxon at subtidal stations, even though they rarely comprised more than 2% of the total meiofauna. The percent composition of harpacticoids in the total meiofauna was consistently higher from late May through August. Standard errors were unusually low for meiofauna, seldom exceeding 15% of the mean, despite the average sample size of only 4 cores per station per collection date in 1987 and 1988 (Shirley et aI., 1988; Shirley et aI., 1989; Fleeger and Shirley, 1990). The density relationships between the predominate meiofaunal components, excluding nematodes, did not vary greatly between years or subtidal stations (Walls et aI., 1987). At E, the numerically predominate meiofauna, in order of abundance, were harpacticoids, priapulid larvae, , and ostracods. After the har- pacticoids, only priapulids contributed significantly to total meiofaunal density and at times replaced harpacticoids as the second most abundant taxon. The same relationships were true at ABM, with the exception that kinorhynchs supplanted ostracods in abundance, and at Nand NE stations, where the maximal density of the less abundant taxa was <2.5·10 cm-2• Approximately 35 larval size classes, lorica total lengths ranging from 0.050 to 1.050 mm, were found in Auke Bay (Fig. 3). Many of the larval sizes undoubtedly represent variation around modes, and many were not present in all months. The smallest size class of priapulid larvae first appeared in December (Fig. 3a). A comparison oflength-frequency distributions ofpriapulids in December and March implies that the larvae of the predominate size classes had increased by 0.100 mm (Fig. 3a). The 0.400 mm shift in size-frequency distributions of the predom- inate size class between April and August similarly presents a tenable growth schemata representing four molts with approximately a 25% molt increment (Fig. 3b). Nearly identical size-frequency distributions ofpriapulid larvae, particularly with the predominate size class, were present in the same months in 1987 and 1988 (Fig. 3c). Bottom water temperatures and salinities varied between years in Auke Bay. 152 BULLETIN OF MARINE SCIENCE. VOL. 47. NO.1, 1990

60 6000 -E -E u u 50 5000 cr A cr (/) (/) 0 Meiofauna 0 40 T""" T""" ,,~1 4000 - I ->. .•...>. I ;:: , (/) (/) 30 I 3000 I c: c: Q) Q) , I 0 0 20 , I 2000 ns "0 ,I c: :3 :3 cu c. 10 1000 cu .Q .;:: -Q) a.. 0 0 ~ 1985 1986 1987 1988

60

-E 50 1988 (.) I Priapulid larvae C"' 40 en B 0 ~ 30 ->. ..- 20 1987 en \ c o Q) 1986 • 0 10 / •• •• •• 0 60 90 120 150180 210240 270300 330360 Julian Date Figure 2. Density of priapulid larvae and total meiofauna from 1985-1988 at East Station, Auke Bay, Alaska. A) Sampling dates are noted with open and closed circles for meiofauna and priapulid larvae, respectively; B) larval priapulid densities, 1985-1988. SHIRLEY: ECOLOGY OF AN ALASKAN PRIAPULID 153

Size Distribution of Priapulids 50 Dec 9 Mar A 40 ~ :: ~ :: " ->. 30 · , 0 ,'· ,. c: , Q) · , '· ~ ,. 0" 20 ,· . Q) ... ,: 0' LL · . 10 o \.. 0 0 200 400 600 800 Lorica Length (pm)

50 B 40 ~ -~ >. 30 --0-- Apr 0 ....•.... c: Aug Q) ~ 0" 20 ...Q) LL 10

0 0 200 400 600 800 1000 1200 Lorica Length (pm)

60 C 50 ~ -0 40 ->. --+- April1988 0 c: 30 ····0···· April1987 Q) ~ 0" ...Q) 20 LL 10 a o a 200 400 600 800 1000 1200 Lorica Length (pm) Figure 3. Length-frequency distributions of priapulid larvae in selected months from East Station, Auke Bay, Alaska, in 1987-1988. 154 BULLETIN OF MARINE SCIENCE, VOL. 47, NO.1, 1990

At 30 m depth, water temperatures had increased by only 0.5°C between mid- March and mid-May in both 1985 and 1986, while salinities of the bottom water varied by less than 0.5%0 (Ziemann, 1986; Laws et al., 1988). In 1987 and 1988, temperatures remained about 1°C higher through May (Ziemann et al., 1988).

DISCUSSION Land (1970) reported Priapulus caudatus as having a northern circumpolar distribution, being common along the coasts of Scandinavia, Scotland, Iceland, Spitsbergen, Greenland, and the far eastern coast ofthe USSR, and having a more scattered occurrence in more southern regions, including the southern North Sea, England, Ireland, eastern Canada, the New England and western coasts of the U.S. A few unusual southerly records also exist from the Mediterranean Sea, and southern Japan and California (Land, 1970). Only a single collection of P. caudatus has been reported from the Alaskan coast, near Pt. Barrow (MacGinitie, 1955; Shapeero, 1962b), although Land (1970) hypothesized its scarcity to be due to a paucity of collections and stated that it was probably common. My examination of 32 adult priapulids fi:om 17 different collections contained by the Museum of the University of Alaska Fairbanks confirmed Land's (1970) supposition. In ad- dition to my collections in Auke Bay and Glacier Bay in southeastern Alaska, P. caudatus has also been collected from Prudhoe Bay, numerous locations in the central and eastern Bering Sea, and in the Gulf of Alaska. Few quantitative ecological data exist on priapulid larvae or adults. Albeit some ecology papers have been published (reviewed by Land, 1970), most priapulid research has been on morphology, and systematics (reviewed by Land and Norrevang, 1985). The extant ecological literature specifically concerning Priapulus caudatus was reviewed by Land (1970). Priapulus caudatus has been collected from the intertidal to 7,500 m depth (Land, 1970); the optimum depth range reported by Wesenberg-Lund (1929) is approximately that of Auke Bay, with 50% of the bay being between 40-60 m. No priapulids were found in the intertidal zone of Auke Bay during biweekly sampling for a year (McGregor and Shirley, 1988), but were present at all other depths sampled. The other specimens from Alaskan waters reported above were collected in depths from 16 to 187m depth. The bottom sediments of Auke Bay are primarily soft, clayey muds with shell fragments, similar in composition to that reported to be preferred by Priapulus caudatus in Danish and adjacent waters (Wesenberg-Lund, 1929). Auke Bay does not have periods of oxygen depletion, nor are the sediments rank with , conditions deemed optimum for P. caudatus because of their potential for decreasing competition (Land, 1970). The benthic infaunal community containing P. caudatus in Auke Bay is diverse, but is characterized as a Macoma sp.- community. The epibenthic community includes large crabs (predominantly Paralithodes camtschaticus, Pa- gurus spp. and Chionocetes bairdi), snails (Fusitriton oregonensis and Neptunea spp.), and flatfish (Limanda aspera, Lepidopsetta bilineata, Platichthys stellatus, Hippoglossus stenolepis and Hippoglossoides elassodon). Salmon fry and adults (Oncorhynchus keta, O. nerka, O. gorbuscha, O. kitsutch and O. tshawytscha) are seasonally abundant in the bay. The meiofaunal community is predominated by nematodes, but also exhibits a diverse harpacticoid community of greater than 35 species (Shirley et al., 1989). Adult Priapulus caudatus are characterized as carnivorous, feeding on slow moving prey such as polychaetes, and ingesting mud only incidently along with SHIRLEY: ECOLOGY OF AN ALASKAN PRIAPUUD 155 prey or in cases of emergency (Lang, 1948a; Land, 1970). Larvae are reported to be detritus feeders (Lang, 1939; Land, 1970). Although adult priapulids undoubt- edly consume macrofaunal prey in some parts of their distribution, all adult priapulids from Auke Bay had only mud in their intestines, while larvae contained undefined detritus. Many benthic feeding marine fishes (particularly flatfishes and gadids) and mammals have been reported to feed incidentally on priapulids (reviewed by Land, 1970). Although several species of salmonids and postlarval flatfish in Auke Bay feed principally on meiofauna, particularly harpacticoid copepods, no priapulids were found in fish stomachs in Auke Bay (Landingham and Motherhead 1988; Sturdevant, 1987; McGregor and Shirley, 1988). Conflicting information exists concerning the reproductive biology of Priapulus caudatus. Sexually mature females of P. caudatus from the west coast of Greenland in the summer months had fully ripe ovaries (Wesenberg-Lund, 1929), however breeding was thought to occur in the spring (Wesenberg-Lund, 1939). In contrast, Lang (1948a) reported that spawning of P. caudatus near the Kristineberg Zoo- logical Station in the Baltic Sea occurred at the end of the year. Few had spawned by 5 December and all (13 individuals) had spawned by 8 January (Lang, 1948a). The first appearance of the smallest size class of priapulid larvae in December samples from Auke Bay supports Lang's (1948a) observations that spawning occurs late in the year. The presence of nearly identical size-frequency distributions of priapulid larvae from the same months in 1987 and 1988 (Fig. 3c) suggests that spawning occurs at approximately the same time annually in Auke Bay. Growth of priapulids has received little attention. Priapulid larvae and adults molt in order to grow, but the number of instars in P. caudatus has not been determined. Lang (1939) reported the smallest "shelled" larvae of P. caudatus to be 0.500 mm and the largest to be 1.3 mm. Lang's measurements were of total larval length, which varies depending on the state of relaxation of the larvae, whereas measurements in this paper represent only the totallorica length, which varies only slightly with degree of relaxation or state of preservation. The fully extended introvert contributes about 35% ofthe total larval length (Lang, 1948b). Approximately 35 larval size classes, lorica total lengths ranging from 0.050 to 1.050 mm, were found in Auke Bay. The total length of the largest larvae from Auke Bay would approximate the maximum size reported by Lang (1939), but the smaller instars have not been previously reported, perhaps due to collection or concentration techniques. Land (1970) cautioned against interpretation of a bimodal scattergram of larval P. caudatus lengths, based on measurements of approximately 50 specimens from Denmark, to imply only two larval stages existed, but made no statements about the total number of instars which might be present. Descriptions of molting by Wesenberg-Lund (1929) and Lang (1948a) were of adults, and no suggestion of molt increments or increases in size were given. A larva of 0.050 mm lorica length which had a 25% increase in size with each molt would molt 14 times before reaching the postlarval stage of approximately 1.1 mm lorica length. Assuming this molt increment, which is sustained by the length- frequency distributions, the 0.100 mm increase of the predominate size class of larvae in Auke Bay between December, 1987 and March, 1988 represents three molts with a 25% increase per molt (Fig. 3a). The 0.400 mm shift in size-frequency distribution of the predominate size class between April and August could sim- ilarly be interpreted as representing four molts (Fig. 3c). With only seven molts having occurred between December and August, the two year larval period pro- posed by Lang (1939) appears to be substantiated. Lang derived his long larval 156 BULLETIN OF MARINE SCIENCE, VOL. 47, NO. I, 1990

period primarily on observations of the molting infrequency of larger larvae he maintained in the lab and not on sustained culture until metamorphosis. Coull (1985) reported interannual variations in meiofauna density that changed by an order of magnitud{: during an 11 year study in South Carolina. Therefore, the variability in meiofaunal densities from year to year in Auke Bay may not be exceptional. The small interannual differences in temperature and salinity of bot- tom waters are probably not responsible for the pronounced interannual differ- ences present in the meiofauna community. Interannual variability in composition and density of meiofauna in Auke Bay was not related to the amount or timing of primary production and sedimentation; a relationship did exist between the timing of reproductive events ofharpacticoid copepods and phytoplankton sedi- mentation (Shirley et aI., 1989). The successive density maxima of priapulid larvae occurring intraannually may be artifacts of spatial heterogeneity. The long term trend of increasing density of priapulids is not definitive; it does not parallel that of the meiofaunal community and must be controlled by other variables. Priapulids are widespread, common components of the marine benthos of northern latitudes, particularly as larvae within the meiofauna, and should not be considered as rare and relict forms restricted only to marginal environments. Given the new information on their larval densities within the meiofauna and extremely long larval p{:riod, their role in the ecology of the marine benthos warrants further study.

ACKNOWLEDGMENTS

Thanks are due to L. McNutt and S. McGregor for assisting with data collection; J. Fleeger, J. McCall, G. T. Chandler and A. Decho also helped with sample collection and sorting. D. Erickson was the boat skipper. S. Shirley offered many helpfu] suggestions to the manuscript. This work was supported by contract NA-85-ABH-00022 from the U.S. Department of Commerce (NOAA) to the School of Fisheries and Ocean Sciences, University of A]aska Fairbanks, as part of APPRISE (As- sociation of Primary Production with Recruitment in a Subarctic Ecosystem).

LITERATURE CITED

Calloway, C. B. ]982. PriapUl]ida. Pages 94]-944 in S. P. Parker, ed. Synopsis and classification of living organisms, Vol. I. McGraw-Hill Book Co., New York. Candia Camevali, M. D. and M. Ferraguti. 1979. Structure and ultrastructure of muscles in the priapulid Halicryptus spinulosus: functional and phylogenetic remarks. J. Mar. BioI. Assoc. U.K. 59: 737-744. Carlisle, D. B. 1958. On the exuvia of Priapulus caudatus Lamarck. Arkiv for Zoo]ogi ]2(5): 79-81. Chandler, G. T., T. C. Shirley and J. W. Fleeger. 1988. The Tom-Tom corer: a new design of the Kajak corer for use in me:iofauna sampling. Hydrobiologia 169: 129-134. Conway Morris, S. 1977. Fossil priapulid worms. Special Papers in Paleontology No. 20. The Paleontological Assoc., London. (4): ]-95, 30 pis. Coull, B. C. ]985. Long-term variability of estuarine : an II year study. Mar. Ecol. Prog. Ser. 24: 205-2] 8. Coyle, K. O. and T. C. Shirley. In Press. A summary of the fisheries and oceanographic research in Auke Bay, Alaska and vicinity. Association of primary production with recruitment in a subarctic ecosystem. Fleeger, J. W. 1979. Population dynamics of three estuarine meiobenthic harpacticoids (Copepoda) in South Carolina. Mar. BioI.. 52: 147-156. --, and T. C. Shirley. ]990. Meiofaunal responses to sedimentation from an Alaskan spring bottom. II. Harpacticold population dynamics. Mar. Edol. Prog. Ser. 59: 239-247. Haldorson, L., J. D. Watts, D. A. Sterritt, M. S. Pritchett and L. R. McNutt. ] 988. Larval fish production in Auke Bay, Alaska- 1987. Pages 299-360 in Summary of 1987 APPRISE Research Program, Ann. Rpt. to NO.'\A. School of Fisheries and Ocean Sciences, Univ. of Alaska, Fairbanks. Vol. I. SHIRLEY: ECOLOGY OF AN ALASKAN PRIAPULID 157

Land, J. van der. 1970. Systematics, zoogeography, and ecology of the Priapulida. Zool. Yerhandl. Leiden 112: 1-118. --- and A. Norrevang. 1985. Affinities and intraphyletic relationships of the Priapulida. Pages 259-273 in S. Morris, J. D. George, R. Gibson and H. Platt, eds. The origins and relationships of lower . The Systematics Association Special Volume 28. Clarendon Press, Oxford. Landingham, J. H. and P. D. Motherhead. 1988. Feeding habits of pink salmon in nearshore and offshore areas of Auke Bay, Alaska. Pages 449-470 in Summary of APPRISE 1987 Research Program, Ann. Rpt. to NOAA, SFOS APP87-100. School of Fisheries and Ocean Sciences, Univ. of Alaska, Fairbanks. Lang, K. 1939. Uber die Entwicklung von Priapu/us caudatus Lam. Kung!. Fysiogr. Sallsk. Lund Forh. 9: 80-87. --. 1948a. Contribution to the ecology of Priapu/us caudatus Lam. Ark. Zool. 41A(5): 1-12. --. 1948b. On the morphology of the larva of Priapu/us caudatus Lam. Arkiv for Zoologi 41A(9): 1-8. ---. 1963. The relation between the Kinorhyncha and Priapulida and their connection with the Aschelminthes. Pages 256-262 in C. Dougherty, ed. The lower Metazoa, comparative biology and phylogeny. Univ. Calif. Press, Berkeley, Calif. Laws, E. A., P. K. Bienfang, D. A. Ziemann and L. D. Conquest. 1988. Phytoplankton population dynamics and the fate of production during the spring bloom in Auke Bay, Alaska. Limnol. Oceanogr. 33: 57-65. MacGinitie, G. E. 1955. Distribution and ecology of the of Point Barrow, Alaska. Smithsonian Misc. Collections 128(9): 1-201. McGregor, S. B. and T. C. Shirley. 1988. Ontogenetic dietary changes and selectivity of post meta- morphic flatfish in a subarctic ecosystem. American Zoologist 28(4): 169A. Malakhov, V. V. 1980. Cephalorhyncha, a new type of uniting Priapulida, Kinorhy- cha, Gordiacea, and a system of Aschelminthes worms. Zool. Zh. 59: 485-499. Nyholm, K. G. and C. Borno. 1969. Introductory studies ofthe oxygen studies of Priapu/us caudatus Lam. Zoologiska Bidrag fran Uppsala 38: 262-264. Por, F. D. and H. J. Bromley. 1974. Morphology and anatomy of Maccabeus tentacu/atus (Priapulida: Seticoronaria). J. Zool. Soc. London 173: 173-197. Shapeero, W. L. 1961. Phylogeny of the Priapulida. Science 133: 879-880. ---. 1962a. The epidermis and cuticle of Priapu/us caudatus Lamarck. Trans. Am. Microscop. Soc. 81(4): 352-355. ---. 1962b. The distribution of Priapu/us caudatus on the Pacific coast of North America. Am. MidI. Nat 68: 237-241. Sherman, K. M., D. A. Meeter and J. A. Reidenauer. 1984. A technique for subsampling an abundant taxon while completely sorting other taxa. Limnol. Oceanogr. 29(2): 433-439. ---, J. W. Fleeger, A. W. Decho and M. V. Sturdevant. 1987. Association of meiofauna with primary production and recruitment in a subarctic ecosystem-1986. Pages 361-412 in Summary of 1986 APPRISE Research Program, Annual Report to NOAA. School of Fisheries and Ocean Sciences, Univ. of Alaska, Fairbanks. Vol. I. ---, ---, S. W. McGregor and L. McNutt. 1988. Association of meiofauna with primary production and recruitment in a subarctic ecosystem-1987. Pages 381-429 in Summary of 1987 APPRISE Research Program, Ann. Rpt. to NOAA. School of Fisheries and Ocean Sciences, Univ. of Alaska, Fairbanks. Vol. I. ---, ---, --- and ---. 1989. Interannual variability in density and recruitment of meiofauna in Auke Bay, Alaska. Pages 255-285 in Summary ofl988 APPRISE Research Program, Ann. Rpt. to NOAA-1988. School of Fisheries and Ocean Sciences. Univ. of Alaska, Fairbanks. Yol. 1. Sturdevant, M. V. 1987. The role of meiofauna in the diets and feeding ecology of post metamorphic flatfish. M.Sc. Thesis, Univ. Alaska, Juneau. 195 pp. Walls, S., M. Sturdevant, L. McNutt and T. Shirley. 1987. Changes in meiofauna composition and density with depth and season in an Alaskan subarctic ecosystem. Amer. Zool. 27(4): 164A. Wesenberg-Lund, E. 1929. Some remarks on the biology and anatomy of the genus Priapu/us. Yidensk. Meddel. Dansk. naturhist. Foren. 88: 165-202. --. 1939. Polseorme (Gephyrea). Sipunculider, Priapulider, Echiurider. Danmarks Fauna 45: 1-60. Ziemann, D. A. 1986. Patterns of oceanographic conditions and particulate sedimentation in Auke Bay, Alaska, during the spring bloom, 1985. Pages 143-244 in Summary of 1985 APPRISE Research Program, Ann. Rpt. to NOAA, UAl SFS-8603. School of Fisheries and Sciences, Univ. of Alaska Juneau. Vol. 1. --, L D. Conquest, P. K. Bienfang and J. Kanda. 1988. Patterns of primary production and sedimentation during the 1987 spring bloom in Auke Bay, Alaska. Pages 29-144 in Summary of 158 BULLETINOFMARINESCIENCE,VOL.47, NO.I, 1990

APPRISE 1987 Research Program, Ann. Rpt. to NOAA SFOS APP87-100. School of Fisheries and Ocean Sciences, Univ. of Alaska, Fairbanks. Vol. I.

DATEACCEPTED: January 30, 1989.

ADDRESS: Juneau Center for Fisheries and Ocean Sciences, University of Alaska Fairbanks, 11120 Glacier Highway, Juneau, Alaska 99801. -