Zootaxa 3371: 1–307 (2012) ISSN 1175-5326 (Print Edition) Monograph ZOOTAXA Copyright © 2012 · Magnolia Press ISSN 1175-5334 (Online Edition)

ZOOTAXA

3371

Decapod Crustacea of the Californian and Oregonian Zoogeographic Provinces

MARY K. WICKSTEN Department of Biology, Texas A&M University, College Station TX 77843-3258 U.S.A. E-mail: [email protected]

Magnolia Press Auckland, New Zealand

Accepted by P. Castro: 16 Mar. 2012; published: 4 Jul. 2012 MARY K. WICKSTEN Decapod Crustacea of the Californian and Oregonian Zoogeographic Provinces (Zootaxa 3371) 307 pp.; 30 cm. 4 Jul. 2012 ISBN 978-1-86977-935-1 (paperback) ISBN 978-1-86977-936-8 (Online edition)

FIRST PUBLISHED IN 2012 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

© 2012 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use.

ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition)

Abstract ...... 5 Introduction ...... 5 Methods and coverage ...... 6 Zoogeography ...... 10 Habitats ...... 13 Symbiotic species ...... 14 Introduced species ...... 15 Shrimps ...... 15 Key to the families of shrimps...... 16 Suborder Dendrobranchiata...... 17 Superfamily Penaeoidea ...... 17 Family Benthesicymidae ...... 17 Family Penaeidae ...... 24 Family Sicyonidae ...... 25 Family Solenoceridae...... 26 Superfamily Sergestoidea ...... 26 Family Sergestidae ...... 26 Suborder Pleocyemata ...... 32 Infraorder Stenopodidea ...... 32 Family Stenopodidae ...... 32 Infraorder Caridea ...... 34 Superfamily Pasiphaeoidea ...... 34 Family Pasiphaeidae...... 34 Superfamily Oplophoroidea ...... 41 Family Oplophoridae ...... 41 Superfamily Nematocarcinoidea ...... 48 Family Nematocarcinidae ...... 48 Superfamily Stylodactyloidea ...... 50 Family Stylodactylidae...... 50 Superfamily Atyoidea ...... 50 Family Atyidae...... 50 Superfamily Palaemonoidea...... 51 Family Palaemonidae ...... 51 Superfamily Alpheoidea...... 57 Family Alpheidae ...... 57 Family Hippolytidae...... 69 Family Lysmatidae ...... 71 Family Thoridae ...... 72 Family Ogyrididae ...... 95 Superfamily Processoidea ...... 96 Family Processidae...... 96 Superfamily Pandaloidea ...... 99 Family Pandalidae ...... 99 Superfamily Crangonoidea...... 106 Family Crangonidae ...... 106 Family Glyphocrangonidae ...... 121 Infraorder Astacidea ...... 123 Superfamily Astacoidea ...... 124 Family Astacidae ...... 124 Family Cambaridae...... 127 Infraorder Palinura ...... 128 Superfamily Palinuroidea ...... 128 Family Palinuridae ...... 128 Infraorder Polychelida...... 129 Superfamily Eryonoidea ...... 129 Family Polychelidae ...... 129 Infraorder Axiidea...... 130 Family Axiidae ...... 131 Family Callianassidae...... 133 Family Calocarididae ...... 135 Family Ctenochelidae...... 136 Family Eiconaxiidae...... 137 Infraorder Gebiidea ...... 137 Family Laomediidae...... 137

CALIFORNIAN AND OREGONIAN DECAPODA Zootaxa 3371 © 2012 Magnolia Press · 3 Family Upogebiidae ...... 139 Infraorder Anomura...... 141 Key to the families of the Anomura ...... 142 Superfamily Galatheioidea ...... 142 Family Chirostylidae ...... 143 Family Galatheidae...... 145 Family Munididae...... 146 Family Munidopsidae...... 149 Family Porcellanidae ...... 153 Superfamily Hippoidea...... 160 Family Albuneidae ...... 160 Family Blepharipodidae ...... 161 Family Hippidae ...... 163 Superfamily Lithoidea ...... 163 Family Hapalogasteridae ...... 163 Family Lithodidae...... 165 Superfamily Paguroidea ...... 171 Key to the species of the Paguroidea ...... 172 Artificial key to species of common intertidal and shallow subtidal hermit crabs ...... 173 Family Diogenidae ...... 174 Family Paguridae ...... 179 Family Parapaguridae...... 199 Infraorder Brachyura ...... 200 Key to the families of the Brachyura ...... 201 Superfamily Homoloidea ...... 202 Family Homolidae ...... 202 Superfamily Dromioidea ...... 202 Family Dromiidae ...... 202 Superfamily Cyclodorippoidea ...... 203 Family Cyclodorippidae...... 203 Superfamily Calappoidea ...... 205 Family Calappidae ...... 205 Superfamily Leucosioidea ...... 206 Family Leucosiidae ...... 206 Superfamily Majoidea ...... 206 Family Epialtidae ...... 207 Family Inachidae ...... 214 Family Inachoididae...... 216 Family Oregoniidae ...... 218 Family Pisidae ...... 219 Superfamily Parthenopoidea ...... 226 Family Parthenopidae...... 226 Superfamily Cancroidea...... 227 Family Cancridae ...... 227 Superfamily Portunoidea ...... 234 Family Portunidae ...... 234 Superfamily Xanthoidea...... 237 Family Panopeidae ...... 238 Family Xanthidae ...... 242 Superfamily Pilumnoidea...... 244 Family Pilumnidae ...... 244 Superfamily Pinnotheroidea...... 245 Family Pinnotheridae ...... 245 Superfamily Ocypodoidea ...... 260 Family Ocypodidae ...... 260 Superfamily Palicoidea ...... 261 Family Palicidae...... 261 Superfamily Grapsoidea...... 262 Family Grapsidae ...... 263 Family Varunidae ...... 264 Acknowledgments ...... 267 Reference...... 268 Index of Species...... 289 Plates ...... 293

Approximately 325 species of decapod crustaceans are reported from the Californian and Oregonian zoogeographic provinces, a figure that includes all freshwater, estuarine, and marine (intertidal zone to 4000 m) decapods from the area. At least six of these species have not been reported from California since 1921, three species cannot be recognized from their descriptions and lack type material and illustrations and another five species may have ben reported from mistaken localities or are the result of misidentification. The area is mostly inhabited by cold-temperate species. Genera endemic to the northeastern Pacific include the anomurans Janetogalathea (Galatheidae), Acantholithodes, Phyllolithodes, and Rhinolithodes (Lithodidae); and the brachyurans Mimulus (Epialtidae), Loxorhynchus, and Scyra (Pisidae). Families that are particularly diverse in species include carideans of the families Thoridae, Pandalidae, and Crangonidae; anomurans of the families Hapalogasteridae, Lithodidae, and Paguridae and brachyurans of the families Epialtidae, Cancridae, and Pinnotheridae. Crayfishes of the genus Pacifastacus (Astacidae) were endemic to the area prior to human introductions elsewhere. At least three estuarine and two freshwater decapods belonging to the families Palaemonidae, Cambaridae, Panopeidae and Varunidae have been introduced into the area and maintain reproducing populations. Keys are provided to all the families, genera, and species treated. A major synonymy, short description, and information on habitat, biogeography, type locality, and color in life are provided for each species. References are provided to the original descriptions of all taxa mentioned. Additional remarks on taxonomy, symbiotic associations, characteristic behavior, and other information that may help in identification also are given. New generic designations for Hippolyte affinis Owen, 1839 and H. layi Owen, 1839 (Thoridae) are supported. Expanded diagnoses are given for Heptacarpus franciscanus (Schmitt, 1921) (Thoridae), Isocheles pilosus (Holmes, 1900), and Paguristes parvus Holmes, 1900 (Diogenidae).

Key words: Decapoda, Dendrobranchiata, Penaeoidea, Sergestoidea, Pleocyemata, Stenopodidea, Caridea, Astacidea, Palinura, Polychelida, Axiidea, Gebiidea, Anomura, Brachyura, Californian zoogeographic province, Oregonian zoogeographic province.

Introduction

Approximately 325 species of decapod crustaceans live along the west coast of North America between Puget Sound, Washington, U.S.A. and Magdalena Bay, Baja California, Mexico, the Californian and Oregonian zoogeographic provinces. Garth & Wicksten (1993) gave an account of the history of crustacean studies in the area. Studies of northeastern Pacific decapods started in the early 1800's. The last complete guide to the decapods of California was Marine Decapod Crustacea of California, by W.L. Schmitt (1921). Schmitt's pioneering book, largely based on collections by the U.S. Fisheries steamer Albatross, is now badly out of date. Shallow-water decapods are also mentioned in more recent guidebooks to intertidal animals, such as those by Morris et al. (1980), Ricketts et al. (1985), and Carlton (2007). Jensen (1995) published good color photographs of shallow-water species, along with brief information on range and identifying features. These more recent works nevertheless provide limited information on the species. There are few recent works on the species of deeper benthic habitats (outside of the range of scuba diving, or 40 m). Wicksten (2002) listed pelagic species. Revisions of the nomenclature, new systematic interpretations of families and higher taxa, descriptions of new species, range extensions, and natural history information published since 1921 are scattered in the literature of at least seven nations and written in four languages. Many valuable works on decapods are old and unavailable except through major libraries. Even with the use of the Internet, information may be difficult to find because many works have not been scanned. Starting in 1991, J. Haig and J. Garth of the University of Southern California and I initiated plans to write an updated synthesis on the decapods of California. Changes in editors, loss of funding, constant changes in computer format and other problems delayed publication. Other potential co-authors declined to undertake the task of writing the sections of the manuscript after the deaths of Garth and Haig. I eventually enlarged and updated the original manuscript to include areas to the north and south of California. The Scripps Institution of Oceanography Library produced the manuscript as a website (http://repositories.cdlib.org/sio/lib/26) in 2008. The present work updates, corrects, and expands on the material in the website in a format suitable for publication.