RsRDir. Skr. Ser. r-lavUnders. 14(5):229-556 A Quantitative Study in Puget Sound, WASHINGTON, USA, IN 1963-1964 BY ULF LIE Department of Oceanography, University of Washington, SeattZe, Washington, USA With a section on polychaetes by KARL BANSE, KATHARINE D. HOBSON AND FREDERIC H. NICHOLS UNIVERSITETSFORLAGET 0 The Norv,egian Research Cou~lcilfor Scietlce and the Huinailities 1965 (Norges almenvitenskapelige forskningsrid) Section: D. 69. C0-3Y. Printed ill Norway by UNIVERSITETSFORLAGETS TRYKNINGSSENTRAI OSLO, NORWAY CONTENTS A . Purpose of the present investigation ...................... 237 B . Literature review ...................................... 237 1. Review of concepts and methods .................... 237 2 . Subtidal investigations from the west coast of the United States .................................. 239 C . Area and environment .......................... ........ 241 1. Topography and sedimentology ...................... 241 2 . Hydrography ...................................... 241 3. Phytoplankton production ........................... 244 MATERLAL AND METHODS ............................... 245 A . Choice of stations ...................................... 245 B . Description of stations .................................. 245 Station 1 (47°41'33"~. 122~24'18"~)................ 247 Station 2 ($7°42q16"~. 122~26'24"~) ................ 247 Station 3 (49°44'044"~. 12~~31'53s'~) ................ 247 Station 4 (47°44131q'~.122~32'41"~) ................ 247 Station 5 (47*10'48"~. ~22~50'00"~)................ 247 Station 6 (47°10'40"~. 122~48'48"~)................ 247 Station 9 (47°13'30"~. 122'49'36"~) ................ 248 Station 8 (41°15'10"~7 1220 50'06"W) ................ 248 C . Choice of sampler and sampling procedures ............... 248 D . Number of samples .................................... 249 E . Sieving ......................................... 253 8" . Laboratorymethods ................................... 253 f . Sediment particle size analysis .................... 253 2 . Sorting and identification of fauna .................. 253 3 . Size measurements and weighings .................. 254 4 . Electronic data processing ......................... 255 III . RESULTS AND DISCUSSION .............................. 256 2 A . Performance of the 0.1 ni van Veen grab ............... 256 1. Repeatability ..................................... 256 2 . Sampling efficiency ............................... 261 3 . Comparison of the van Veen and the Smith- McIntyre grabs .................................. 262 B . The substrate at the stations ........................... 263 C . Composition of the faunal assemblages .................. 271 Station 1 (Tables 5 and 6) .......................... Station 2 (Tables 7 and 8) .......................... Station 3 (Tables 9 and 10) ......................... Station 4 (Tables 11 and 12) ........................ Station 5 (Tables 13 and 14) ........................ Station 6 (Tables 15 and 16) ........................ Station 7 (Tables 17 and 18) ........................ Station 8 (Tables 19 and 20) ........................ D . Number of species and specimens ...................... 324 1. Seasonal variatioils it1 numbers .................... 324 2 . Distribution of taxoiloniic groups in relation to sediments ..................................... 335 3 . Patchiness of ~lumericallydominant species ......... 335 E . Standing crop .........................................339 1. Standing crop of benthic infauna .................... 339 2 . Seasonal variation in standing crop ................. 341 F . Similarity among the faunal assemblages ................. 345 6 . Relationships between the faunal assemblages in Puget Sound and known benthic communities .............. 350 H . Diversity .............................................. 351 1. Indices of diversity as measurable parameters of communities .................................... 351 2 . MARGALEFts index of diversity ..................... 352 3 . The SEANNON-WIENER function as an index of diversity ....................................... 354 4 . Redundancy ....................................... 358 5 . Comparisons of indices of diversity .................. 360 6 . Indices of diversity with polychaetes included ......... 363 IV . THE NUMERICALLY DOMINANT SPECIES ................. 367 A . Selection of dominant species ........................... 367 B . Biological and ecological information about the nunlerically dominant species ........................... 368 Harmothoe imbricata (LINNB) ...................... 368 Lepidasthenia berkeleyae PETTIBONE ............... 368 Malmgrenia lunulata (DELLE CEIAJE) .............. 370 Peisidice aspera JOHNSON ......................... 370 Pholoe minuta (FABRICIUS) ........................ 372 Sigambra tentaculata (TREADWELL) ................ 372 Pionosyllis uraga IMAJIMA ........................ 372 Platynereis bicanaliculata (BAIRD) .................. 374 Nephtys ferruginea !&ARTMAN ...................... 374 Glycera capitata OERSTED ........................ 376 Lumbrineris bicirrata TREADWELL ................ 376 Lumbrineris californiensis HARTMAN .............. 378 Lumbrineris cruzensis HARTMAN .................. 378 Lumbrineris luti BERKELEY and BERKELEY ....... 380 Haploscoloplos pugettensis PETTIBONE ............. 380 Laonice cirrata (SARS) ............................ 382 Laonice sp. I ..................................... 382 Prionospio cirrifera WIR~N........................ 382 19 . Prionospio malmgreni CEAPAR~DE ................ 385 20 . Prionospio pinnata ENLERS ....................... 385 21 . Caulleriella alata (SOUTHERN) ..................... 387 22 . Chaetozone setosa MALMGREN ..................... 387 23 . Chaetozone sp. I ................................... 387 24 . Armandia brevis (MOORE) ......................... 389 25 . Travisia pupa MOORE ............................. 389 26 . Euclymene zonalis (VERRILL) ...................... 389 27 . Praxillella affinis pacifica BERKELEY .............. 392 28 . Praxillella gracilis (SARS) ......................... 392 29 . Pectinaria californiensis HARTMAN ................ 392 30 . Pectinaria granulata (LINN~)....................... 394 31 . Golfingia pugettensis FISHER ....................... 394 32 . Euphilomedes carcharodonta (V . Z . SMITH) .......... 397 33 . Euphilomedes producta POULSEN ................... 401 34 . Paraphoxus variatus BARNARD ..................... 404 35 . Heterophoxus oculatus (HOLMES) ................... 408 36 . Byblis veleronis BARNARD ......................... 408 37 . Leptochelia dubia (KROYER) ....................... 417 38 . Eudorella pacifica HART .......................... 421 39 . Pinnixa schmitti RATHBUN ........................ 422 40 . Lophopanopeus bellus (STIMPSON) .................. 431 41 . Nucula bellotii ADAMS ........................,. .. 438 42 . Crenella columbiana DALL ......................... 442 43 . Psephidia lordi BAIRD ............................. 446 44 . Mysella tumida (CARPENTER) ...................... 451 45 . Axinopsida sericata CARPENTER ................... 456 46 . Macoma carlottensis (WHITEAVES) .................. 461 47 . Macoma alaskana DALL ........................... 466 48 . Macoma calcarea GMELIN ......................... 466 49 . Semele rubropicta DALL ........................... 469 50 . Myaarenaria LINN~ .............................. 475 51 . Amphiodia urtica (LYMAN) ......................... 479 52 . Leptosynapta clarki HEDING ........................ 489 53 . Brisaster townsendi PGASSIZ ....................... 490 V . SUMMARY ............................................... 498 VI . ACKNOWLEDGEMENTS ................................... 501 VII . REFERENCES ............................................ 503 VIII . APPENDIX I . "MISCELLANEOUS GROUPS" ................ 518 M . APPENDIX II . ANNOTATED LIST OF POLYCHAETES BY KARL BANSE. KATHARINE D . HOBSON AND FREDERIC H . NICHOLS ............................................. 521 X . TAXONOMIC LISTING OF NONPOLYCHAETES .............. 549 I. INTRODUCTION A. Purpose of the present investigation In 1962 the Department of Oceanography, University of Washington, planned to start benthic investigations in the Northeast Pacific Ocean, but as the Depart- ment was inexperienced in the field of benthos research it was felt that a pilot study in Puget Sound would be necessary. The pilot study would be particularly concerned with sampling problems, processing of material, and spatial and sea- sonal variability. Methods and concepts of modern synecology, such as statisti- cal methods for delimiting communities and the concept of species diversity, have received little attention from benthos researchers, and one of the objec- tives therefore has been to test the validity and applicability of some of these methods. It was realized that future studies of the dynamics of productivity and energetics of benthic communities would have to be limited to the dominant spe- cies population. One of the objectives of the present investigation therefore has been to determine which species are the most important and to compile ecologi- cal and biological information about those species. B. Literature review 1. Review of concepts and methods A truly quantitative study of the bottom fauna is a product of this century, and is mainly attributed to the invention of a quantitative sampler, the Petersen grab (PETERSEN and JENSEN 1911). The advantage of grabs over the dredges that had been used in the last century is that they enable the scientist