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Taxonomy and Distribution of the Marine Calanoid Copepod Family Euchaetidae

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Taxonomy and Distribution of the Marine Calanoid Copepod Family Euchaetidae BULLETIN OF THE SCRIPPS INSTITUTION OF OCEANOGRAPHY UNIVERSITY OF CALIFORNIA SAN DIEGO Volume 29 Taxonomy and Distribution of the Marine Calanoid Copepod Family Euchaetidae Taisoo Park University of California Press Taxonomy and Distribution of the Marine Calanoid Copepod Family Euchaetidae Taisoo Park UNIVERSITY OF CALIFORNIA PRESS Berkeley • Los Angeles • London CONTENTS v vi vii viii BULLETIN OF THE SCRIPPS INSTITUTION OF OCEANOGRAPHY OF THE UNIVERSITY OF CALIFORNIA, SAN DIEGO LA JOLLA, CALIFORNIA Advisory Editors: Charles S. Cox, Gerald L. Kooyman, Richard H. Rosenblatt (Chairman) Volume 29 Approved for publication July 1993 UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES, CALIFORNIA UNIVERSITY OF CALIFORNIA PRESS, LTD. LONDON, ENGLAND © 1995 BY THE REGENTS OF THE UNIVERSITY OF CALIFORNIA PRINTED IN THE UNITED STATES OF AMERICA Library of Congress Cataloging-in-Publication Data Park, Taisoo. Taxonomy and distribution of the marine calanoid copepod family Euchaetidae/Taisoo Park. p. cm. — (Bulletin of the Scripps Institution of Oceanography, University of California, San Diego; v. 29) Includes bibliographical references (p. ). ISBN 0-520-09802-1 (pbk.:alk. paper) 1. Euchaetidae—Classification. 2. Euchaetidae—Geographical distribution. I. Title. II. Series. QL444.C72P37 1994 595.304—dc20 94-28941 CIP The paper used in this publication meets the minimum requirements of American National Standard for Information Sciences—Permanence of Paper for Printed Library Materials, ANSI Z39.48-1984. ABSTRACT The marine calanoid copepod family Euchaetidae and its two genera, Euchaeta and Paraeuchaeta, are redefined. Fourteen species of Euchaeta and 61 species of Paraeuchaeta, including 13 new species, are described and their geographic ranges defined from specimens found in midwater trawl and plankton net samples collected throughout the Atlantic, Pacific, and Indian oceans. The species of Euchaeta are classified into three species groups - marina, concinna, and acuta groups - and an independent species, E spinosa; those of Paraeuchaeta were classified into six species groups malayensis, pavlovskii, norvegica, glacialis, hebes, and antarctica groups - and three independent species - P. biloba, P. bisinuata, and P. grandiremis. Each of the nine species groups is defined with detailed descriptions of its representative species and each of the four independent species that could not be grouped is also described in detail. Phylogenetic relationships among the species groups and independent species are discussed. Keys are presented for identification of the species groups in each genus and the species in each species group containing three or more species. The geographic distribution of the species groups and common species is discussed. Key words: Taxonomy, phylogeny, geographic distribution, worldwide, Copepoda, Calanoida, Euchaetidae Euchaeta, Paraeuchaeta, species groups, new species, keys. ix ACKNOWLEDGMENTS This study is based on plankton and midwater trawl collections at Scripps Institution of Oceanography, Woods Hole Oceanographic Institution, the Smithsonian Oceanographic Sorting Center, and the University of Tokyo Ocean Research Institute. Many people at these institutions helped make the collections available for my study and I am particularly indebted to the following persons: M. Mullin, E. Brinton, M. Ohman, G. Snyder, and A. Townsend of Scripps Institution of Oceanography; J. Craddock of Woods Hole Oceanographic Institution; F. D. Ferrari of the U. S. National Museum of Natural History; M. Terazaki and S. Nishida of the University of Tokyo. I am also indebted to H. Weikert of the University of Hamburg, Germany, and N. Copley of Woods Hole Oceanographic Institution who allowed me to study their sorted specimens. This study was supported by National Science Foundation Grant OCE-8915094 and most of the study was carried out at Scripps Institution of Oceanography. I wish to acknowledge the cooperation of M. Mullin, director of the Marine Life Research Group, and M. Ohman, curator of the Planktonic Invertebrates Collection. I wish also to thank my wife, Insuk, for assistance in preparing the illustrations. This paper benefited from discussions with and reviews by E. Brinton and R. H. Rosenblatt of Scripps Institution of Oceanography and F. D. Ferrari of the U. S. National Museum of Natural History. xi INTRODUCTION Giesbrecht (1892) erected the subfamily Euchaetina, which was subsequently raised to the rank of a family by Sars (1902). The family Euchaetidae comprised a single genus, Euchaeta Philippi 1843, until Scott (1909) proposed a division of the family into two genera, Euchaeta Philippi 1843 and Paraeuchaeta Scott 1909. He placed species of the Euchaeta marina type in Euchaeta and those of the Euchaeta norvegica type in Paraeuchaeta. The distinction between the two genera was based on the structure of the endopodal setae of the female maxilla and the exopods of the male 5th pair of legs. In the species assigned to Euchaeta, Scott found that 2 of the 6 endopodal setae of the female maxilla are armed with long spinules in addition to rows of very short spinules and the exopods in both the right and left 5th legs of the male taper into long spines. In the genus Paraeuchaeta, on the other hand, he found that none of the endopodal setae of the female maxilla has long spinules and none of the exopods of the male 5th pair of legs terminates in a long spine. Sars (1925) found another character by which the females of the two genera could be distinguished, viz., the appendicular caudal setae, which were found to be straight and much larger than the marginal caudal setae in Euchaeta but geniculated and much thinner than the marginal caudal setae in Paraeuchaeta. With (1915) was the first author to disagree with Scott’s proposal. He pointed out the problem of assigning Euchaeta hebes to either Euchaeta or Paraeuchaeta, because 1 of the 6 endopodal setae of its female maxilla is armed with long spinules as in Euchaeta, while its male 5th pair of legs is of the Euchaeta norvegica type. After examining a large number of euchaetid species, Vervoort (1957) also disagreed, mainly for the same reason as With (1915), to Scott’s division of the Euchaetidae. The species Vervoort cited as intermediate and thus problematical include Euchaeta pubera, E. spinosa, E. hebes, and E. russelli. Vervoort also mentioned that the appendicular caudal setae vary in size and form without a definable correlation with the proposed division of the family into Euchaeta and Paraeuchaeta. For the sake of convenience, however, he arranged the species into the following 5 groups based on the presence or absence of spinulose setae on the female maxillary endopods and the structure of the male 5th pair of legs: 1) Euchaeta marina group, 2) E. flava group, 3) E. hebes group, 4) E. pubera group, and 5) E. norvegica group. The last was further subdivided into 2 subgroups according to the length of the digitiform process on the 2nd exopodal segment of the male left 5th leg. Of Vervoort’s 5 species groups of Euchaeta, only the marina group has subsequently been found to be a natural group (Bradford 1974), which now comprises 4 species: E. marina (Prestandrea 1833), E. rimana Bradford 1974, E. marinella Bradford 1974, and E. indica Wolfenden 1905. More recently Fontaine (1988) proposed that the 5 southernmost species of Euchaeta (E. antarctica Giesbrecht 1902, E. austrina Giesbrecht 1902, E. similis Wolfenden 1908, Paraeuchaeta erebi Farran 1929, and E. tycodesma Park 1978) form another natural species group, the antarctica species group. In studies of the geographic distribution of bathypelagic calanoids based on plankton net and midwater trawl samples collected mostly from depths exceeding 1000 m throughout the Atlantic, Pacific, and Indian oceans, which are presently available at Scripps Institution of Oceanography, the Smithsonian Oceanographic Sorting Center, Woods Hole Oceanographic Institution, and the University of Tokyo Ocean Research Institute, I have found 75 species, including 13 new species, attributable to the family Euchaetidae. The identification of species was extremely difficult because descriptions in the literature are in most cases either inadequate or too general and thus applicable to more than one species. Furthermore, the proper diagnoses of new species and inadequately known species in many cases entailed redefining some of the previously known closely related species. When all of the species found in the studies were closely examined and compared with one another for their diagnoses, they were found to be classifiable into 9 species groups and 4 independent species that could not be grouped. Of the 9 species groups, 2 correspond to the marina and hebes groups of Vervoort (1957) and one to the antarctica group of Fontaine (1988). When these species groups and independent species were analyzed using the structure of the female genital somite, appendicular caudal setae, maxillule, maxilla, and maxilliped and the male 5th pair of legs, the distinction between 1 the 2 genera Euchaeta and Paraeuchaeta became clear. In this paper I attempt to rediagnose the family, redefine its 2 genera, characterize all species groups recognized in the family, describe in detail with figures each of the species found and provide information on their geographic distribution. 2 MATERIALS AND METHODS The material on which this study is based consists of Isaacs-Kidd midwater trawl (IKMT) and plankton net samples selected from the collections available at Scripps Institution of Oceanography,
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