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(Polychaeta) of the Gulf STUDIES ON THE FAUNA OF CURAÇAO AND OTHER CARIBBEAN ISLANDS: No. 129. Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea by Nancy Marie Foster (Dunbarton College, Washington, D.C.) page figures INTRODUCTION History 3 Acknowledgements 5 Genera] Taxonomic Characters of the Spionidae (PI. 1-2) . 5 Development 11 Ecology and Distribution 12 Methods and Materials 14 Family Diagnosis 14 Key to Spionid Genera of the Gulf of Mexico and Caribbean Sea 15 TAXONOMIC SECTION Polydora 18 1—12 — commensalis Andrews 20 1-12 — ligni Webster 22 13-21 — plena Berkeley & Berkeley 24 22-29 — websteri Hartman 26 30-36 Pygospio 28 — elegans Claparfede 29 37-47 Spio 32 — 35 pettiboneae sp. n 48-56 Scolecolepides 37 — viridis (Verrill) 37 57-65 Spiophanes 40 — bombyx (Claparede) 40 66-75 — wigleyi Pettibone 43 76-85 Malacoceros 47 — vanderhorsti (Augener) 48 86-92 — indicus (Fauvel) 50 93-99 — glutaeus (Ehlers) 53 100-111 — in 57 112-117 flatus sp. n 2 Scolelepis j 58 — squamata (Miilleri 59 118-131 — texana sp. n 63 132-142 Aonides 65 — mayaguezensis Foster 66 143-154 Laonice 69 — cirrata (Sars, 69 155-160 Dispio 72 — uncinata Hartman 73 161-174 Prionospio 79 — steenstrupi Malmgren 84 175-185 — cristata sp. n 87 186-199 — heterobranchia Moore 89 199-212 Apoprionospio 93 — pygmaea (Hartman; 94 213-225 — dayi Foster 97 226-236 Paraprionospio 100 — pinnata (Ehlers) 102 237-261 Aquilaspio 105 Minuspio 106 — cirri fera (Wir6n) 108 262-275 Streblospio 112 — benedicti Webster 112 276-283 ZOOGEOGRAPHY 116 284-285 DISCUSSION OF TAXONOMIC CHARACTERS 118 LITERATURE CITED 121 INTRODUCTION HISTORY Although there have been several collections of polychaetous annelids from the Gulf of Mexico and Caribbean few Sea, very spionids have been included in the published species lists. This is not because they are poorly represented in this area but probably a of their result small size and the fact that they are easily overlooked both in collecting and in sorting of samples. It is also probable that their small size renders them an unpopular groupwith which to work. Very few spionids were reported from collections of the early large scale collecting cruises. This can be, at least in part, attributed to that the fact they are more common in littoral habitats than in deeper waters. SCHMARDA (1861) reported three species as a result of his world cruise (1853—57) during which he visited several Caribbean Islands. TREADWELL (1901, 1931a + b, 1939a, 1939b) reported only three species and this included his treatment of the annelids collected during the Scientific Survey of Puerto Rico and the Virgin Islands. Other contributions include works by AUGENER (1906, 1927, 1933), HORST (1922), KAVANAGH (1940) and BEHRE (1950). HARTMAN (1951) reported on a collectionof littoral polychaetes from the Gulf of Mexico. The latter included what is probably the largest number of dealt with in that time. Twelve spionids a single study up to species were reported, most of which represented new records. Later, small papers were contributed by CARPENTER (1956) from the north- 4 ern Gulf, FRIEDRICH (1956) from Central and South America and WESENBERG-LUND (1958) from the Lesser Antilles. Finally in 1962, JONES reported on a collection of polychaetes from Jamaica which included two spionid species. Although few spionids have been reported in faunal studies from the Gulf of Mexico and have dealt with this Caribbean, many authors family systematically. MCINTOSH dealt with British His (1909, 1914, 1915b) specifically spionids. papers, and the discussed often inde- however, are occasionally quite confusing species are CHAMBERLIN workable the then known terminable. (1919) presented a key for spionid genera. The most useful monographic works are those by MESNIL (1896, 1925) and SSDERSTROM (1920). FAUVEL (1927) treats the Spionidae in his monographic work on the polychaetes of France. He it includes eleven presents a good generic key, although only genera. OKUDA (1937a) dealtwith spioniformpolychaetes (Spionidae,Magelonidae, Owe- niidae) from Japan. A later paper by HARTMAN (1941) was concerned with California and treated with spionids some fourteen species, observations on reproductive in few PETTIBONE characterized biology a species. (1963) revised the spionid genera by branchiae and pointed prostomia and branchiae and frontal horns. In addition the there have been innumerable with selected in to above, papers dealing genera this family. The majority of these can be found in the synonymies included in the present paper. HARTMAN (1966) summarized what was known about Antarctic spionids and DAY included the in reported twenty-three species. (1967) spionids a monograph of South African and polychaetes presented a generic as well as species key. The present study is primarily a faunal study of spionids of the Gulf of Mexico and Caribbean Sea. It is obvious from the literature thatthis is unstudied with to its survey area virtually regard spionid faunayet this family is usually one of the most abundantin benthic communities. With the increasing amount of ecological work being done in the the lack of this study area, knowledge concerning group The work will become more and more significant. present represents the that a starting point toward filling this gap. In spite of fact written with numerous papers have been dealing spionids, a great deal of confusion exists at the generic level. It has become in- that of is creasingly apparent a faunal study a group impractical unless the systematics of that group are well-defined. This study, therefore, has become somewhat more than a strict faunal survey and, of necessity, includes considerations of species outside the insure study area to more significance to later faunal studies. The present collections represent by no means the entire spionid 5 fauna in the Gulf and Caribbean. They are comprised of littoral samples collected by the author and those solicited from other workers. The latter often included samples of a single specimen. The species herein reported probably represent no more than one-third of those in in- actually present the study area. Twenty-six species, cluding sixteen genera, are describedand figured. This includesfour new species and the revision of the genus Prionospio to include three new genera. ACKNOWLEDGEMENTS This study was supported by a Smithsonian Pre-doctoral Fellowship from the Smithsonian Research Foundation. I wish the to express my appreciation to many who contributed collections from the Gulf of Mexico and Caribbean persons spionid Sea for use in this work, particularly Dr. MEREDITH L. JONES (Smithsonian Insti- tution) and Dr. P. WAGENAAR HUMMELINCK (Zo6logisch Laboratorium, Utrecht). I wish to thank Dr. MARIAN H. PETTIBONE for her (Smithsonian Institution) many helpful suggestions and willingness to advise and for the kind loan of library materi- als. I thank the members of consultative committee at The my George Washington University, Drs. I. B. HANSEN, and E. E. MORTENSEN, and I am especially gratefulto Dr. MEREDITH JONES for his guidance as director of this research and for access to his library and research facilities. In appreciation for the use of type material I also thank Drs. OLGA HARTMAN (Allan Hancock Foundation), JOHN DAY (University of Capetown), GESA HARTMANN-SCHRODER (Staatsinstitut and Zoologi- sches Museum, Hamburg), LUCIEN LAUBIER (University de Paris, Biologie Marine), G. HARTWICH (Berlin Museum), DAVID GEORGE (British Museum), and HERBERT LEVI (Museum of Comparative Zoology, Harvard). I wish to to the staff at the Smithsonian Institution express my gratitude library with special thanks to Mr. JACK MARQUARDT for his never-endingpatience. Finally, I to E. for the of the am especially grateful my sister, JUDITH FOSTER, typing manuscript. to This work was submitted the Faculty of the Graduate School of Arts and Sciences of The George Washington University, in partial satisfaction of the require- ments of the degreeof Doctor of Philosophy, September30, 1969. GENERAL TAXONOMIC CHARACTERS OF THE SPIONIDAE fall within that of Spionids group polycheates often designated the collectively as subclass Sedentaria; however, the classification of polychaetes in the subclasses Sedentariaand Errantia quickly breaks down when one considers individual species. There do not seem to be definite characters any by which the two "subclasses" can be con- The of sistently separated. family Spionidae represents a group poly- 6/1 PLATE 1 and . - A. Anterior end ofPrionospio, third subsequentparapodiaomitted. - B. Anterior end of Malacoceros (Rhynchospio), third and subsequent parapodia end Anterior end omitted. - C. Anterior of Spiophanes.— D. of Scolelepis, second and = = = omitted. - ntl noto- subsequentparapodia pr prostomium, pe peristomium, nrl = = fh = frontal br = podial lamella, neuropodial lamella, p palp, horn, branchia. 7 chaetes which does not exhibit most of the typical characters of the Sedentaria. The following is a typical diagnosis of Sedentaria as a subclass. The spionid deviations will become obvious later in this discussion and, because of these inconsistencies, Sedentariaand Er- rantia are not recognized as formal taxa, but merely as generic terms in continuum general language to indicate the two ends of a repre- from the forms the senting progression more primitive errant to more highly specialized sedentary forms. Sedentaria (USHAKOV, 1965, p. 3) into "Body segments as a rule dissimilar. Body divided distinct sections (thorax, abdomen and sometimes pygidium). Prostomium secondarily modifiedand
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