Q^SjyMmr-s afiC-iH NOAA TR NMFS CIRC-387 NOAA Technical Report NMFS CIRC-387 U.S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service Marine Flora and Fauna of the Northeastern United States. Crustacea: Stomatopoda RAYMOND B. MANNING SEATTLE, WA FEBRUARY 1974 NOAA TECHNICAL REPORTS National Marine Fisheries Service, Circulars The major responsibilities of the National Marine Fisheries Service (NMFS) are to monitor and assess the abundance and geographic distribution of fishery resources, to understand and predict fluctuations in the quan- tity and distribution of these resources, and to establish levels for optimum use of the resources. NMFS is also charged with the development and implementation of policies for managing national fishing grounds, develop- ment and enforcement of domestic fisheries regulations, surveillance of foreign fishing off United States coastal waters, and the development and enforcement of international fishery agreements and policies. NMFS also assists the fishing industry through marketing service and economic analysis programs, and mortgage insurance and vessel construction subsidies. It collects, analyses, and publishes statistics on various phases of the industry. The NOAA Technical Report NMFS CIRC series continues a series that has been in existence since 1941. The Circulars are technical publications of general interest intended to aid conservation and management. Publica- tions that review in considerable detail and at a high technical level certain broad areas of research appear in this series. Technical papers originating in economics studies and from management investigations appear in the Circular series. NOAA Technical Reports NMFS CIRC are available free in limited numbers to governmental agencies, both Federal and State. They are also available in exchange for other scientific and technical publications in the ma- rine sciences. Individual copies may be obtained (unless otherwise noted) from NOAA Publications Section, Rock- ville, Md. 20852. Recent Circulars are: 315. Synopsis of biological data on the chum salmon, Oncorhynclms keta (Walbaum) 1792. By Rich- ard G. Bakkala. March 1970, iii + 89 pp., 15 figs., 51 tables. 319. Bureau of Commercial Fisheries Great Lakes Fishery Laboratory, Ann Arbor, Michigan. By Bureau of Commercial Fisheries. March 1970, 8 pp., 7 figs. 330. EASTROPAC Atlas: Vols. 4, 2. Catalog No. 1 49.4:330/ (vol.) 11 vols. ($4.75 each). Avail- able from the Superintendent of Documents, Washington, D.C. 20402. 331. Guidelines for the processing of hot-smoked chub. By H. L. Seagran, J. T. Graikoski, and J. A. Emerson. January 1970, iv + 23 pp., 8 figs., 2 tables. 332. Pacific hake. ( 12 articles by 20 authors. ) 1970, iii + 152 pp., 72 figs., 47 tables. March 333. Recommended practices for vessel sanitation and fish handling. By Edgar W. Bowman and Alfred Larsen. March 1970, iv + 27 pp., 6 figs. 335. Progress report of the Bureau of Commercial Fisheries Center for Estuarine and Menhaden Research, Pesticide Field Station, Gulf Breeze, Fla., fiscal year 1969. By the Laboratory staff. August 1970, iii + 33 pp., 29 figs., 12 tables. 336. The northern fur seal. By Ralph C. Baker, Ford Wilke, and C. Howard Baltzo. April 1970, iii + 19 pp., 13 figs. 338. Bureau of Commercial Fisheries Biological Lab- oratory, Auke Bay, Alaska. By Bureau of Com- mercial Fisheries. June 1970, 8 pp., 6 figs. 339. Salmon research at Ice Harbor Dam. By Wesley J. Ebel. April 1970, 6 pp., 4 figs. 340. Bureau of Commercial Fisheries Technological Laboratory, Gloucester, Massachusetts. By Bu- reau of Commercial Fisheries. June 1970, 8 pp., 8 figs. 341. Report of the Bureau of Commercial Fisheries Biological Laboratory, Beaufort, N.C., for the fiscal year ending June 30, 1968. By the Lab- oratory staff. August 1970, iii + 24 pp., 11 figs., 16 tables. 342. Report of the Bureau of Commercial Fisheries Biological Laboratory, St. Petersburg Beach, Florida, fiscal year 1969. By the Laboratory staff. August 1970, iii + 22 pp., 20 figs., 8 tables. 343. Report of the Bureau of Commercial Fisheries Biological Laboratory, Galveston, Texas, fiscal year 1969. By the Laboratory staff. August 1970, iii + 39 pp., 28 figs., 9 tables. 344. Bureau of Commercial Fisheries Tropical Atlan- tic Biological Laboratory progress in research 1965-69, Miami, Florida. By Ann Weeks. Oc- tober 1970, iv + 65 pp., 53 figs. 346. Sportsman's guide to handling, smoking, and pre- serving Great Lakes coho salmon. By Shearon Dudley, J. T. Graikoski, H. L. Seagran, and Paul M. Earl. September 1970, iii + 28 pp., 15 figs. 347. 337. Program of Division of Economic Research, Bureau of Commerecial Fisheries, fiscal year 1969. By Division of Economic Research. April 1970, iii -(- 29 pp., 12 figs., 7 tables. Continued on inside back cover, Synopsis of biological data on Pacific ocean perch, Sebastodes alutus. By Richard L. Major and Herbert H. Shippen. December 1970, iii + 38 pp., 31 figs., 11 tables. ATMOSP/, '^lENJ Of U.S. DEPARTMENT OF COMMERCE Frederick B. Dent, Secretary NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION Robert M. White, Administrator NATIONAL MARINE FISHERIES SERVICE Robert W. Schoning, Director NOAA Technical Report NMFS CIRC-387 Marine Flora and Fauna of the Northeastern United States. Crustacea: Stomatopoda RAYMOND B. MANNING ^oUJT/O/v '6-1 SEATTLE, WA FEBRUARY 1974 For sale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C. 20402 FOREWORD This issue of the "Circulars" is part of a subseries entitled "Marine Flora and Fauna of the Northeastern United States." This subseries will consist of original, illustrated, modern manuals on the identification, classification, and general biology of the estuarine and coastal marine plants and animals of the Northeastern United States. Manuals will be published at irregular intervals on as many taxa of the region as there are specialists willing to collaborate in their preparation. The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates of the Woods Hole Region," edited by R. I. Smith, published in 1964, and produced under the auspices of the Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole, Mass. Instead of revising the "Woods Hole Keys," the staff of the Systematics- Ecology Program decided to expand the geographic coverage and bathymetric range and produce the keys in an entirely new set of expanded publications. The "Marine Flora and Fauna of the Northeastern United States" is being prepared in collaboration with systematic specialists in the United States and abroad. Each man- ual will be based primarily on recent and ongoing revisionary systematic research and a fresh examination of the plants and animals. Each major taxon, treated in a separate manual, will include an introduction, illustrated glossary, uniform originally illustrated keys, annotated check list with information when available on distribution, habitat, life history, and related biology, references to the major literature of the group, and a system- atic index. These manuals are intended for use by biology students, biologists, biological ocean- ographers, informed laymen, and others wishing to identify coastal organisms for this region. In many instances the manuals will serve as a guide to additional information about the species or the group. Geographic coverage of the "Marine Flora and Fauna of the Northeastern United States" is planned to include organisms from the headwaters of estuaries seaward to approximately the 200-m depth on the continental shelf from Maine to Virginia, but may vary somewhat with each major taxon and the interests of collaborators. When- ever possible representative specimens dealt with in the manuals will be deposited in reference collections of the Gray Museum, Marine Biological Laboratory, and other universities and research laboratories in the region. After a sufficient number of manuals of related taxonomic groups have been pub- lished, the manuals will be revised, grouped, and issued as special volumes. These vol- umes will thus consist of compilations of individual manuals within phyla such as the Coelenterata, Arthropoda, and Mollusca, or of groups of phyla. 11 CONTENTS Page Introduction \ Key to Stomatopoda of the Northeastern United States 2 Annotated list, order Stomatopoda 4 Selected references 4 Index to scientific names 4 Acknowledgments 6 Coordinator's comments 6 The National Marine Fisheries Service (NMFS) does not approve, rec- ommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales pro- motion which would indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. Ill Digitized by the Internet Archive in 2013 http://archive.org/details/marineflorafaunaOOmann MARINE FLORA AND FAUNA OF THE NORTHEASTERN UNITED STATES. Crustacea: Stomatopoda RAYMOND B. MANNING 1 ABSTRACT This manual includes an introduction on the general biology, an illustrated key, an annotated systematic list, a selected bibliography, and an index to the stomatopod Crustacea of the inner continental shelf of the Northeastern United States. Four species are treated. INTRODUCTION The crustacean order Stomatopoda is a small group of primitive crustaceans found almost exclu- sively in shallow tropical waters. More than 300 species in four families are known. Although 70 species have been recognized in the western Atlan- tic, only four are found in the Northeastern United States. Two of these, N annosquilla grayi (Chace) and Squilla empusa Say, live in shallow water, whereas the other two, Heterosquilla armata (Smith) and Platysquilla enodis (Manning), live in depths of 30 m or more. All four species are included in the key given below. The stomatopods or mantis shrimps can be recog- nized by the presence of two anterior movable so- mites bearing eyes and triramous antennules, a carapace which covers the anterior portion of the cephalothorax dorsally but leaves the four posterior thoracic somites (No. 5-8) free, five pairs of subche- late thoracic appendages (one of which is greatly enlarged as a raptorial claw), and abdominal gills (Fig. 1). The common name of these organisms, mantis shrimps, alludes to the resemblance of the raptorial claws to those of the praying mantis (Fig. 2). Adult stomatopods are primarily benthic or- ganisms that usually live in burrows. So far as is 1 National Museum of Natural History, Smithsonian Institu- tion. Washington, D.C. 20560. known, all species are predators, and some leave their burrows at night to feed. The female carries the oval egg mass between her maxillipeds until the young hatch. The planktonic larvae, known as alima or erichthus larvae, go through a number of distinct stages before settling on the botton; each species may have as many as 10 separate larval stages. The larvae of most species are unknown. Our knowledge of most aspects of the general biology of stomatopods and their interrelationships with other organisms at best can be characterized as scanty. Often our knowledge is restricted to the ex- ternal morphology of a few individuals of a species. For example, many of the specimens of the two species, Heterosquilla armata and Platysquilla enodis, known from deeper waters off the North- eastern United States were found in stomach con- tents of fishes, and this is practically all that we know about these species ; P. enodis is known from parts of three specimens. In contrast, specimens of the other two species, N annosquilla grayi and Squilla em- pusa, living off the Northeastern United States are comparatively common. Judging from collections made by M. B. Gray in the 1950's, N annosquilla grayi was abundant in the Bass River, Yarmouth, Cape Cod, and material might be collected readily for a variety of studies. Similarly, Squilla empusa, which seems to be abundant throughout its range, could serve as the subject for many investigations. Abdominal somites Thoracic somites Carapace Rostral plate •Eye •Antennule Telson. Antenna Antennai scale First' J maxilliped Mandibular palp Raptorial claw Third maxilliped Fifth maxilliped Epipod Male copulatory tubes Ventral spine Third pereiopod Figure 1. — Morphological structures and terms used in descriptive accounts of stomatopods (from Manning, 1969). Carpus Merus Dorsal ridge of carpus Proximal. Movable teeth Pectinations Teeth of doctylus Ischium Propodus Dactylus Figure 2. — Structure of raptorial claw (from Manning, 1969). KEY TO STOMATOPODA OF THE NORTHEASTERN UNITED STATES 1. Carapace with median carina terminating an- teriorly in a bifurcation (Fig. 3). Telson lacking movable marginal teeth. Family Squillidae: Squilla empusa Figure 3. — Squilla empusa. Anterior portion of body, dorsal view. 1. Carapace lacking median carina. Telson with one pair of movable marginal teeth. Family Lysiosquillidae 2 (1) Posterior armature of telson submar- ginal, not completely visible in dor- sal view. (Fig. 4, 5) N annosquilla grayi Figure 4. — Nannosquilla grayi. Telson in ventral view. Figure 5. — N annosquilla grayi. Sixth ab- dominal somite and telson in dorsal view. 2 (7) Marginal armature of telson entirely visible in dorsal view 3 3 (2) 3 (2) Telson with four pairs of fixed marginal projections (Fig. 6). Posterior margin of abdominal somites and dorsal surface of tel- son smooth Platysquilla enodis Figure 6. — Platysquilla enodis. Sixth ab dominal somite and telson in dorsal view. Telson with two pairs of fixed marginal projections (Fig. 7). Posterior margin of fourth, fifth, and sixth abdominal somites and dorsal surface of telson with spinules Heterosquilla armata Figure 7. — Heterosquilla armata. Sixth abdominal somite and telson in dorsal view. ANNOTATED LIST ORDER STOMATOPODA Family SQUILLIDAE Squilla empusa Say, 1818. (Fig. 3, 8). Mas- sachusetts to northern South America in depths to 154 m, usually in shallower water. This is the most common American species of Squilla. Size moderate to large, total length of adults to 185 mm. Family LYSIOSQUILLIDAE Heterosquilla armata (Smith, 1881). (Fig. 7, 9). Off New England in 96-218 m. Rare. Size moderate, total length of adults 20-70 mm. Nannosquilla grayi (Chace, 1958). (Fig. 4, 5, 10). Off Massachusetts and Georgia, sublittoral to 15 m. Common in Bass River, Yarmouth, Mass., where it was first found by M. B. Gray. Size small, total length of adults 20-42 mm. Platysquilla enodis (Manning, 1962). (Fig. 6). Off Vineyard Sound and North Carolina, in 31-49 m. This species is known only from speci- mens collected in the late 1800's. Size mod- erate, total length of adults about 60 mm. SELECTED BIBLIOGRAPHY CHACE, F. A.. JR. 1958. A new stomatopod crustacean of the genus Lysios- quilla from Cape Cod, Massachusetts. Biol. Bull. 114:141-145. MANNING, R. B. 1969. Stomatopod Crustacea of the western Atlantic. Stud. Trop. Oceanogr. (Miami) 8: viii + 1-380. SMITH, R. I. (chief editor). 1964. Keys to marine invertebrates of the Woods Hole re- gion. Systematics-Ecology Program, Mar. Biol. Lab., Woods Hole, Mass., Contrib. No. 11, x + 208 p. INDEX TO SCIENTIFIC NAMES Heterosquilla armata 1,3,4 Nannosquilla grayi 1,3,4 Platysquilla enodis 1,3,4 Squilla empusa 1,2,4 Figure 8. — Squilla empusa. Dorsal view (from Manning. 1969). Figure 9. — Heterosquillo armata. Dorsal view (from Manning, 1969). Specimen 3.8 cm long. Figure 10. — Nannosquilla grayi. Dorsal view (from Chace, 1958). Specimen 4 cm long. ACKNOWLEDGMENTS Preparation of the "Marine Flora and Fauna of the Northeast- ern United States" is being coordinated by the following Board: Coordinator: Melbourne R. Carriker, Marine Biological Laboratory, Woods Hole, Mass. (Address after September 1, 1973: College of Marine Studies, Field Station, University of Delaware, Lewes, Delaware 19958.) Wesley N . Tiffney, Department of Biology, Boston University, Boston, Mass. Ruth D. Turner, Museum of Comparative Zoology, Harvard University, Cambridge, Mass. Roland L. Wigley, National Marine Fisheries Ser- vice, Biological Laboratory, Woods Hole, Mass. Robert T. Wilce, Department of Botany, Univer- sity of Massachusetts, Amherst, Mass. Advisers: Marie B. Abbott, Marine Biological Laboratory, Woods Hole, Mass. Arthur G. Humes, Boston University Marine Pro- gram, Marine Biological Laboratory, Woods Hole, Mass. The Board established the format for the "Marine Flora and Fauna of the Northeastern United States," invites systematists to collaborate in the preparation of manuals, reviews manuscripts, and advises the Scientific Editor of the National Marine Fish- eries Service. COORDINATOR'S COMMENTS Publication of the "Marine Flora and Fauna of the North- eastern United States" is most timely in view of the growing universal emphasis on environmental work and the urgent need for more precise and complete identification of coastal organisms than has been available. It is mandatory, wherever possible, that organisms be identified accurately to species. Accurate scientific names unlock the great quantities of biolog- ical information stored in libraries, obviate duplication of re- search already done, and make possible prediction of attri- butes of organisms that have been inadequately studied. Raymond B. Manning began his study of the Stomatopoda of the Western North Atlantic in 1957 as a graduate student at the Institute of Marine Sciences, University of Miami. His studies on the American species were subsequently expanded to include investigations on all of the known species, the ma- jority of which live in the Indo-West Pacific region. His studies have resulted in more than 50 papers on the systema- tics of stomatopods, including three reviews of classification at the generic level, and a monographic revision of the western Atlantic species which was published in 1969. After complet- ing his graduate studies at the University of Miami, Manning joined the staff of the National Museum of Natural History, Smithsonian Institution, in 1963. Manuals are available for purchase from the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. The manuals so far published in the series are listed below. COOK, DAVID G., and RALPH O. BRINKHURST. Marine Flora and Fauna of the Northeastern United States. Annelida: Oligochaeta BORROR, ARTHUR C. Marine Flora and Fauna of the Northeastern United States. Protozoa: Cilio- phora MOUL, EDWIN T. Marine Flora and Fauna of the Northeastern United States. Higher Plants of the Marine Fringe McCLOSKEY. LAWRENCE R. Marine Flora and Fauna of the Northeastern United States. Pycnogonida MANNING, RAYMOND B. Marine Flora and Fauna of the Northeastern United States. Crustacea: Stomatopoda •6- U.S. GOVERNMENT PRINTING OFFICE: 1974-798-369/14 REGION 10 349. Use of abstracts and summaries as communica- tion devices in technical articles. By F. Bruce Sanford. February 1971, iii + 11 pp., 1 fig. 350. Research in fiscal year 1969 at the Bureau of Commercial Fisheries Biological Laboratory, Beaufort, N.C. By the Laboratory staff. No- vember 1970, ii + 49 pp., 21 figs., 17 tables. 351. Bureau of Commercial Fisheries Exploratory Fishing and Gear Research Base, Pascagoula, Mississippi, July 1, 1967 to June 30, 1969. By Harvey R. Bullis, Jr., and John R. Thompson. November 1970, iv + 29 pp., 29 figs., 1 table. 352. Upstream passage of anadromous fish through navigation locks and use of the stream for spawn- ing and nursery habitat, Cape Fear River, N.C 1962-66. By Paul R. Nichols and Darrell E. Louder. October 1970, iv + 12 pp., 9 figs., 4 tables. 356. Floating laboratory for study of aquatic organ- isms and their environment. By George R. Snyder, Theodore H. Blahm, and Robert J. Mc- Connell. May 1971, iii + 16 pp., 11 figs'. 361. Regional and other related aspects of shellfish consumption — some preliminary findings from the 1969 Consumer Panel Survey. By Morton M. Miller and Darrel A. Nash. June 1971, iv + 18 pp., 19 figs., 3 tables, 10 apps. FOURTH CLASS ■« UNITED STATES PHIlimTC DEPARTMENT OF COMMERCE ^nlT^SSSSf i^SS 1 ^ MATIONAl OCEANIC AND ATMOSPHERIC ADMINISTRATION NATIONAL MARINE FISHERIES SERVICE SCIENTIFIC PUBLICATIONS STAFF ROOM 450 1107 N.E. 45TH ST. ■ SEATTLE, WA 98105 \, OFFICIAL BUSINESS ADDDDTaDiabTT