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FRONTISPIECE.—A, Malmgreniella variegata with Ophionereis reticulata; B, M. puntotorensis with Ophiophragmus cubanus; C,D, M. puntotorensis with Ophiophragmus pulcher. (Photographs by John E. Miller.)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 538

Scaled Polychaetes (Polynoidae) Associated with Ophiuroids and Other Invertebrates and Review of Species Referred to Malmgrenia McIntosh and Replaced by Malmgreniella Hartman, with Descriptions of New Taxa

Marian H. Pettibone

SMITHSONIAN INSTITUTION PRESS

Washington, D.C.

1993

ABSTRACT

Pettibone, Marian H. Scaled Polychaetes (Polynoidae) Associated with Ophiuroids and Other Invertebrates and Review of Species Referred to Malmgrenia McIntosh and Replaced by Malmgreniella Hartman, with Descriptions of New Taxa. Smithsonian Contributions to Zoology, number 538, 92 pages, frontispiece, 55 figures, 1993. Included in this report are new distribution records and descriptions of 44 species of Polynoidae distributed between two subfamilies: Lepidonotinae, with Hermenia verruculosa Grube and Lepidonopsis humilis (Augener); and Harmothoinae, with Subadyte pellucida (Ehlers), and three species and one new combination of Hololepidella Willey. The study includes a replacement of Malmgrenia McIntosh, 1874 (= indeterminable) with Malmgreniella Hartman, 1967, emended, with a redescription of M. dicirra Hartman, 1967, the type species, and a review and revision of the species that have been referred to Malmgrenia, resulting in 13 new combinations (with 2 new synonyms) and 19 new species of Malmgreniella. In addition, three new genera are added, including Paragattyana, for P. micropoides (Augener), new combination, and P. intesi, new species; Rullieriella, for R. monoechinata (Rullier), new combination, with synonym Parahalosydna chryostictus Hutchings and Rainer; and Wilsoniella for W. furcosetosa (Loshamn), new combination. Most of the species show a more or less close association with other invertebrates, including echinoderms (ophiuroids, spatangoid echinoids, synaptid holothurians), coelenterates (hydrocorals), and other polychaetes (arenicolids, terebellids, and chaetopterids).

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging-in-Publication Data
Pettibone, Marian H.
Scaled polychaetes (Polynoidae) associated with ophiuroids and other invertebrates and review of species referred to Malmgrenia McIntosh and replaced by Malmgreniella Hartman, with descriptions of new taxa / Marian H. Pettibone.
p.  cm. — (Smithsonian contributions to zoology ; no. 538
Includes bibliographical references (p.    ).
1. Polynoidae—Classification. 2. Malmgreniella—Classification. 3. Ophiuroidae—Ecology. 4. Commensalism. I. Title. II. Series.
QL1.S54 no. 538 [QL391.A6] 591 s–dc20 [595.1′47] 92-35329

art The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48—1984.

Scaled Polychaetes (Polynoidae) Associated with Ophiuroids and Other Invertebrates and Review of Species Referred to Malmgrenia McIntosh and Replaced by Malmgreniella Hartman, with Descriptions of New Taxa

Marian H. Pettibone

____________

Marian H. Pettibone, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Introduction

Many polynoid polychaetes have been reported living in close associations with other animals, including ophiuroid echinoderms. Reports in the literature of the association of ophiuroids and commensal polynoids include publications by Millott (1953), Spooner et al. (1957), Devaney (1967, 1974a), Gibbs (1969, 1971, 1972), Boesch (1977), and Weston (1984).

In connection with his studies on the Ophiuroidea, Gordon Hendler collected commensal polynoids on different species of burrowing ophiuroids from the following localities: Galeta Point and Punto Toro, Panama; Dry Tortugas, Looe Key, and Virginia Key, Florida; and Belize. These polynoids were given to me for study, along with the identifications of the ophiuroids. During his study of the bioluminescent ophiuroid Ophiopsila californica on Santa Catalina Island, Lawrence Basch collected commensal polynoids and sent them to me for identification. Additional polynoids were received from Liliana Fomeris from the Bay of Flamengo, Säo Paulo, Brazil; from John Taylor and Stuart Santos from Tampa Bay, Florida; from Marvin Wass from the York River and Chesapeake Bay, Virginia; from Anne McCrary from Wrightsville Sound, North Carolina; from Susan Oldfield from the Perlas Islands. Gulf of Panama; from Nancy Maciolek from Corpus Christi Bay, Texas; and from K. Banse from the Puget Sound, Washington area. In addition, numerous polynoids from the Plymouth area, England, were sent by Peter Gibbs, including some collections by G.M. Spooner.

ACKNOWLEDGMENTS

I am indebted to the above-mentioned people for their cooperation and help and for allowing me to examine the material upon which this study is based. The numerous collectors mentioned in the text, who have deposited their collections in the Smithsonian Institution, National Museum of Natural History, and furnished collecting data, also have contributed greatly to the study. Special thanks go to Gordon Hendler for the numerous polynoids, along with the identifications of their ophiuroid hosts, and to Stephen Caims for the polynoids and the identifications of their stylasterine hydrocoral hosts.

In addition to the collections in the Smithsonian Institution (USNM = the former United States National Museum, collections now in the National Museum of Natural History), material was obtained on loan or in exchange from the following sources: Allan Hancock Foundation, Los Angeles (LACM-AHF), through J. Pilger; American Museum of Natural History, New York (AMNH), through E. Kirsteur and H. Feinberg; Australian Museum, Sydney (AMS), through P. Hutchings and P. Coleman; The Natural History Museum, London, formerly the British Museum (Natural History) (BMNH), through J.D. George and A.I. Muir; Los Angeles County Museum of Natural History (LACM), through G. Hendler; The Laboratory, Citadel Hill, Plymouth, England (LCHP), through P. Gibbs; Natural History Museum of Gothenburg (NMG), through A. Eliason; National Museum of Victoria (NMV), through B.J. Smith; Museum National d'Historie Naturelle, Paris (MNHN), through J. Renaud-Momant; Naturhistoriska Riksmuseet, Stockholm (NRS), through R. Oleröd; University Zoological Museum, Copenhagen (UZMC), through J.B. Kirkegaard and M. Petersen; Zoological Institute of Science, Leningrad (ZIASL), through V.C. Averincev; Zoologisches Museum, Universiteit van Amsterdam (ZMA), through S. van der Spoel; Zoologisches Museum, Berlin (ZMB), through G. Hartwick; Zoologisches Staats-museum, Hamburg (ZMH), through G. Hartmann-Schröder; Zoological Museum, Trondheim (ZMT), through A.-A. Loshamn; Zoologisk Museum Universitet, Bergen (ZMUB), through J. Kjennerud. The manuscript benefited from the careful reviews of Thomas H. Perkins, Gordon Kendler, and an anonymous reviewer.

Subfamily LEPIDONOTINAE Willey, 1902

Genus Hermenia Grube, 1856

Hermenia verrruculosa Grube, 1856

Hermenia verruculosa Grube, 1856:44.—Devaney, 1974a:122, 140.—Pettibone, 1975:235, figs. 1,2 [synonymy].—San Martin et al., 1986:7.

MATERIAL EXAMINED.—WESTERN ATLANTIC OCEAN: Virgin Islands: St. John, Brown Bay, West Point, 3 Jul 1984, L. Leward, collector, found crawling on pavement type substrate, 1 specimen. Belize: Carrie Bow Cay, 16°48′N, 88°05′W, 23 Apr 1974, K. Ruetzler, collector, 1 specimen (USNM 80540). Panama: S. of Holandes Cay, 30–42 m, Anton Bruun cruise 10, 18 Sep 1965, S.A. Earle, collector, 1 specimen (USNM 71540). Florida: Dry Tortugas, west side of Loggerhead Key, spur, groove, and seagrass habitats, 1–5 m, 29 Sep 1982, G. Hendler, collector, 1 specimen (USNM 80060).

REMARKS.—Devaney (1974a:122, 140) reported H. verruculosa as a possible commensal on the ophiuroid Ophiocoma pumila Lütken from Belize. The ophiuroid had transversely banded brown and white arms and the polynoid was similarly banded on its dorsum. The dorsal side of the polynoid also was covered with papillae, similar in size and color to the disc granules of the ophiuroid, giving the polynoid a similar appearance.

DISTRIBUTION.—Western Atlantic Ocean, West Indies and Caribbean Sea, Bahamas, Central and South America, Gulf of Mexico, Atlantic Ocean off New Jersey. Intertidal to 223 meters (Pettibone, 1975).

Genus Lepidonopsis Pettibone, 1977

Lepidonopsis humilis (Augener, 1922)

Lepidonotus humilis Augener, 1922:40.—Fauchald and Reimer, 1975:80 [key].—Fauchald, 1977:6.

?Malmgrenia lunulata variety.—Gardiner, 1976:90 [part; not Delle Chiaje, 1830].

Lepidonopsis humilis.—Pettibone, 1977:51, figs. 5. 6 [synonymy].

MATERIAL EXAMINED.—WESTERN ATLANTIC OCEAN: North Carolina: Bird Shoal, western tip, intertidal, on oral surface of keyhole sand dollar Mellita quinquiesperforata, 28 Apr 1975, C. Jenner, collector, 1 specimen (USNM 52850, as ?Malmgrenia lunulata var. by Gardiner, 1976). Florida: Looe Key, 24°32′N, 81°24′W, sta LK-22, 8 May 1985, 6–8 m, on Ophiothrix lineata, with sponges and large rubble slabs, G. Hendler, collector, 1 specimen (USNM 133565). Panama: Galeta Reef, coralline algae and Laurencia zones, 1970–1973, A.A. Reimer, collector, 4 specimens (USNM 54376, 54377, 66008, K. Fauchald, ident.). Turks and Caicos Islands: Rockawash Cay, calcareous alga wash, 13 Apr 1989, R.W. Heard, collector, 5 specimens (USNM 133566).

EASTERN PACIFIC OCEAN: Panama: Whorehouse Reef, Panama Survey sta 133-1, 5 Apr 1973, low tide, M.L. Jones, collector, 1 specimen (USNM 71710). Mexico: Ixtapa Island, Playa Quieto, Guerrero, about 100 miles WNW of Acapulco, intertidal, among sabellarid tubes, 26 Feb 1982, C. Kerby, collector, 1 specimen (USNM 72819).

BIOLOGY.—Free-living or commensal with irregular echinoids and ophiuroids.

DISTRIBUTION.—Western Atlantic Ocean, off North Carolina to Florida, West Indies, and Caribbean Sea, Gulf of Mexico, Central America. Eastern Pacific Ocean from Southern Califomia to Peru. Intertidal to 40 meters (Pettibone, 1977).

Subfamily HARMOTHOINAE Willey, 1902

Genus Subadyte Pettibone, 1969

Subadyte pellucida (Ehlers, 1864)

FIGURE 1

Polynoe pellucida Ehlers, 1864:105, pl. 2: fig. 10; pl. 3: figs. 5, 7–13; pl. 4: figs. 1–3.

Hermadion fragile Claparède, 1868:383, pl. 5: fig. 2.

Scalisetosus communis.—Cuénot, 1912:40.

Scalisetosus pellucidus.—Fauvel, 1923:74, fig. 27a–f [synonymy].—Spooner et al., 1957:112.—Bellan, 1960:6; 1964:29.—Guille, 1965:286.—Hamond, 1966:388.—Hartmann-Schröder, 1971:69, fig. 21a–e.

Harmothoe crosetensis.—Monro, 1930:57 [part; not McIntosh, 1885].

Scalisetosus sp.—Hartman, 1939:6, fig. 1f–k.

Subadyte pellucida.—Pettibone, 1969a:8, fig. 4a–e [synonymy].—Intes and Le Loeuff, 1975:274.—Hartmann-Schröder, 1977:68.—Barel and Kramers, 1977:54.—Campoy, 1982:52.

Adyte pellucida.—Tebble and Chambers, 1982:63, figs. 5a, 20c,d, 56b.

MATERIAL EXAMINED.—SOUTH ATLANTIC OCEAN: South Africa: Simon's Town, False Bay, Basin of H.M. Dockyard, 0–2 m, Discovery sta 90, 10 Jul 1926, 1 specimen (BMNH; mixed with Harmothoe crosetensis by Monro, 1930).

EASTERN PACIFIC OCEAN: Galápagos Islands: James Island, Sullivan Bay, shore and tide-pool collecting, W.L. Schmitt, collector, Presidential cruise sta 13, 24 Jul 1938, 1 specimen (USNM 20517, as Scalisetosus sp. by Hartman, 1939). Isla Isabella, Tagus Cove, in Amphiroa mat, Feb 1978, W.D. Hope, collector, 1 specimen (USNM 75375).

FIGURE 1.—Subadyte pellucida, specimen from Galápagos (USNM 20517): A, dorsal view of anterior end, pharynx partially extended; styles of median antenna, right dorsal and left dorsal and ventral tentacular cirri missing; B, right elytron showing veination, with detail of border and surface papillae; C, left elytrigerous parapodium from segment 2, anterior view, acicula dotted; D, left elytrigerous parapodium from segment 7, anterior view, acicula dotted; E, left cirrigerous parapodium from segment 8, posterior view; F, long and short notosetae, with detail of tip; G, upper, middle, and lower neurosetae. (Scales = 1.0 mm for A; 0.5 mm for B–E; 0.1 mm for F,G.)

DESCRIPTION OF SPECIMEN FROM GALÁPAGOS ISLANDS (USNM 20517, as Scalisetosus sp. by Hartman, 1939).—Body flattened, elongate-oval, with 40 segments, last one minute, 14 mm long, 3 mm wide including setae. Elytra 15 pairs, on segments 2, 4, 5, 7, then on alternate segments to 23, 26, 29, and 32. Elytra oval, delicate, transparent, dotted with oval sensory papillae and some larger papillae, with “veins” eminating from place of attachment to elytrophores (Figure 1B; Hartman, 1939, fig. 1g,h).

Prostomium bilobed, with minute cephalic peaks and 2 pairs of small eyes in posterior half of prostomium; ceratophore of median antenna in anterior notch, style absent; ceratophores of lateral antennae inserted ventrally, with short papillate styles; palps long, tapering; tentaculophores of first segment lateral to prostomium, without setae, each with pair of dorsal and ventral papillate tentacular cirri (Figure 1A; Hartman, 1939, fig. 1f). Second or buccal segment with small nuchal lobe, first pair of elytrophores, biramous parapodia, and papillate ventral buccal cirri longer than following ventral cirri (Figure 1A,C).

Parapodia biramous, notopodium short, conical, with projecting acicular lobe on lower side; neuropodium with long conical presetal acicular lobe from which tip of aciculum projects, and shorter rounded postsetal lobe (Figure 1C–E). Notosetae subequal in width to neurosetae, with spinose pouches and blunt tips, shorter ones with entire tips, longer ones with minute bifid tips (Figure 1F; Hartman, 1939, fig. 1j). Neurosetae each with basal semilunar pocket and distal spinose region, upper ones longer than lower ones, all with bifid hooked tip (Figure 1G; Hartman, 1939, fig. 1i,k). Dorsal cirri with cylindrical cirrophores on posterior sides of notopodia and long papillate styles; ventral cirri short, tapered (Figure 1E).

BIOLOGY.—Subadyte pellucida has been reported on the discs or arms of two species of Ophiothrix: (1) O. fragilis (Abildgaard) from Plymouth, England (Spooner et al., 1957), including postlarvae in the plankton (7–12 setigers); Norfolk, England (Hamond, 1966); NE Atlantic (Barel and Kramers, 1977); English Channel (Bellan, 1964); Arcachon, France (Cuénot, 1912); Portugal (Bellan, 1960); Iberian Peninsula (Campoy, 1982); and (2) O. quinquemaculata (Delle Chiaje) from the Mediterranean by (Bellan, 1964); Banyuls, France (Guille, 1965), including many young stages (15–20 setigers).

Some records of S. pellucida were indicated as free-living or their associations were not noted. The species also is commensal with asteroids.

DISTRIBUTION.—Eastern Atlantic Ocean, North Sea, English Channel, Adriatic and Mediterranean Seas, West and South Africa; Indian Ocean, Bay of Bengal; Eastern Pacific Ocean, Galápagos Islands. Intertidal to 100 meters (Pettibone, 1969a).

Genus Hololepidella Willey, 1905

Hololepidella nigropunctata (Horst, 1915)

FIGURES 2, 3

Polynoe nigropunctata Horst, 1915:20; 1917:104, pl. 21: figs. 15–17.

Allmaniella marquesensis Monro, 1928b:469, figs. 1–4. [New synonymy.]

Malmgrenia marquesensis.—Day, 1962:628; 1967:50, fig. 1.4.p–s.—Intes and Le Loeuff, 1975:273.—Kudenov, 1975b:79

Hololepidella nigropunctata.—Devaney, 1967:287, figs. 1–5; 1974b:154.—Pettibone, 1969a:50, fig. 2a–g.—Gibbs, 1971:117, 120; 1972:203.—Uchida, 1975:19, figs. 1–3.—Amoureux et al., 1978:67, 156.—Sloan et al., 1979:94.—Hendler and Meyer, 1982:742.—Hartmann-Schröder, 1984:66 [key].—Bailey-Brock and Hartman, 1987:234, fig. 3.II.3a–g.—Devaney and Bailey-Brock, 1987:97, 98.—Hanley, 1987:156.

Hololepidella minuta.—Gibbs, 1969:448, 449, fig. 131 [not Polynoe minuta Potts, 1910].

?Harmothoe minuta.—Amoureux et al., 1978:63 [not Polynoe minuta Potts, 1910].

Harmothoe lunulata spp. synaptae.—Amoureux et al., 1978:63 [not Saint-Joseph, 1888].

MATERIAL EXAMINED.—CENTRAL PACIFIC OCEAN: Hawaii: North coast of Oahu, underside of ledge in cave, 7 m, on asteroid Mithrodia sp., 15 Aug 1975, A. Birtles, collector, 1 small specimen (USNM 52793). North shore of Oahu, Pupukea Bay, Beach Park at Kahuku, 2–13 m, sandy pothole, 27 Jul 1976, C. Roper and M. Sweeney, collectors, 1 small specimen (USNM 54147). Marquesas Islands: Tai O Hae Pool, C. Crossland, collector, 6 syntypes of Allmaniella marquesensis (BMNH 1928.1.11.1–3; USNM 45567). Solomon Islands: Graham Point, Marau Sound, on ophiuroids Ophiarthrum elegans Peters and Ophiocoma brevipes Peters, 2 Oct 1965, P.E. Gibbs, collector and identifier, 5 specimens (BMNH 1970.115,118,121,127; USNM 48053).

WESTERN PACIFIC OCEAN: Okinawa: 26°30′N, 127°51′E, 54–57 m, diving, from Ophiuroidea, 14 Mar, 8, 10 Apr 1981, R.F. Bollard, collector, 6 specimens (USNM 100443).

INDIAN OCEAN: Red Sea: Gulf of Elat, from toxic sponge Latrunculia magnifica Keller, 1 specimen (TAV 6928, as H. minuta by Rullier); on solitary coral Fungia scutaria Lamarck, 1 specimen (TAV 9565, as H. lunulata synaptae by Rullier). Mozambique: Inhaca Island, west shore Delagoa Bay, commensal with asteroid Linckia multifora (Lamarck), from J.H. Day, 2 specimens (USNM 45568, as Malmgrenia marquesensis). Australia: Western Australia, Rosemary Island, Dampier, 2 m, on echinoid, 24 Sep 1972, N. Coleman, collector, 1 specimen (AMS W5475).

DESCRIPTION OF SYNTYPES AND SPECIMENS FROM INHACA ISLAND.—Largest complete syntype 9 mm long with 41 segments and 19 pairs of elytra, elytrophores on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, 31, 34, 36, 38, 40. Smallest complete syntype 7 mm long, 37 segments, and 17 pairs of elytra, in similar arrangement. Complete specimen from Inhaca Island with 36 segments and 17 pairs of elytrophores (15 pairs indicated by Day). Elytra oval, darkly pigmented, smooth except for group of microtubercles on anterior region (Figure 2B; Monro, 1928b, fig. 2).

Prostomium bilobed with lobes subtriangular, without distinct cephalic peaks, darkly pigmented except for light areas around 2 pairs of small eyes; ceratophore of median antenna in anterior notch, with long style; ceratophores of lateral antennae inserted terminoventrally, with short styles; palps stout, long, tapered; tentaculophores of first segment lateral to prostomium, each with single seta and pair of dorsal and ventral tentacular cirri (Figure 2A; Day, 1967, fig. 1.4.r).

FIGURE 2.—Hololepidella nigropunctata, syntype of Allmaniella marquesensis (USNM 45567): A, dorsal view of anterior end; right palp and styles of antennae, tentacular cirri, and dorsal cirri of segment 2 missing; B, right elytron, with detail of microtubercle; C, right elytrigerous parapodium, anterior view, acicula dotted; D, right cirrigerous parapodium, posterior view, style of dorsal cirrus missing; E, notoseta; F, upper, middle, and lower neurosetae. (Scales = 0.5 mm for A; 0.5 mm for B; 0.2 mm for C,D; 0.1 mm for E,F.)

Parapodia biramous, smaller notopodium with projecting acicular lobe on lower side, larger neuropodium with subtriangular presetal acicular lobe and shorter rounded postsetal lobe (Figure 2C,D; Monro, 1928b, fig. 2; Day, 1967, fig. 1.4.s). Notosetae slightly stouter than neurosetae, with single row of 10–16 widely spaced spines on slightly convex side and abruptly pointed tip (Figure 2E; Monro, 1928b, fig. 3; Day, 1967, fig. 1.4.p). Neurosetae with faint spinose rows, upper ones with longer spinose regions, all with bifid, hooked tip (Figure 2F; Monro, 1928b, fig. 4; Day, 1967, fig. 1.4.q,q′). Dorsal cirri with cylindrical cirrophores on posterior sides of notopodia, with long styles extending to tips of neurosetae or beyond; dorsal tubercles nodular; ventral cirri short (Figure 2D; Monro, 1928b, fig. 2; Day, 1967, fig. 1.4.s).

DESCRIPTION OF YOUNG SPECIMEN FROM CAVE ON OAHU, HAWAII (USNM 52793).—Body in life pinkish orange with pale elytra (noted by A. Birtles, in litt., who collected it on the asteroid Mithrodia sp.). Body flattened, tapered posteriorly, 5.5 mm long, 2 mm wide including setae, with 31 segments. Elytra 15 pairs, on segments 2, 4, 5, 7, then on alternate segments to 23, 26, 29, 31 (small on last small segment). Elytra oval, delicate, transparent, each with group of conical microtubercles on anterior part (Figure 3B).

Bilobed prostomium with subtriangular anterior lobes, without distinct cephalic peaks; ceratophore of median antenna in anterior notch, with style wider basally and with long slender tip; ceratophores of lateral antennae inserted terminoventrally, with short subulate styles; palps long, stout, with filamentous tips; eyes small, 2 pairs; tentaculophores of first segment lateral to prostomium, each with few setae, with pair of dorsal and ventral tentacular cirri similar to median antenna (Figure 3A). Second segment without nuchal fold, with first pair of elytrophores, with biramous parapodia and long ventral buccal cirri similar to tentacular cirri (Figure 3A). Notopodium smaller than neuropodium, rounded basally with digitiform acicular lobe on lower side, aciculum projecting; larger neuropodium with conical presetal acicular upper part with projecting aciculum, rounded lower part, and shorter rounded postsetal lobe (Figure 3C,D). Notosetae, about 15, forming radiating bundle, each with single row of 7–10 widely spaced spines along curved side (Figure 3E). Neurosetae relatively few, about 11, subacicular, with bifid tips and faint spinose rows, upper ones with longer spinose regions than lower ones (Figure 3F). Dorsal cirri with cylindrical cirrophores posterior to notopodia, with long tapered styles extending beyond neurosetae; dorsal tubercles small, nodular; ventral cirri rather large, extending to tip of neuropodium (Figure 3C).

FIGURE 3.—Hololepidella nigropunctata, young specimen from Hawaii (USNM 52793): A, dorsal view of anterior end, style of right tentacular cirrus missing; B, left elytron, with detail of microtubercles; C, right cirrigerous parapodium, posterior view; D, right elytrigerous parapodium, anterior view, acicula dotted; E, long and short notosetae; F, upper and lower neurosetae. (Scales = 0.5 mm for A; 0.2 mm for B; 0.2 mm for C,D; 0.1 mm for E,F.)

REMARKS.—The young specimen differs from the adults in the shape of the neuropodia, the subacicular distribution of the neurosetae, and paler pigmentation. Perhaps the dark pigmentation is accumulated as the polychaete grows larger.

BIOLOGY.—Hololepidella nigropunctata is commensal with ophiuroids, asteroids, echinoids, and other invertebrates. On the Hawaiian Islands, Devaney (1967:296) reported numerous specimens of H. nigropunctata on brittle stars of the family Ophiocomidae. They were especially common on Ophiocoma dentata Müller and Troschel, occasionally on O. brevipes Peters, and rarely on O. erinaceus Müller and Troschel. Single specimens were found on the oral surfaces of the brittle stars in contact with the oral arm plates near or below the disc, often with the head toward the oral cavity or in the interradius and on the aboral surface of the disc (Devaney, 1967:298, fig. 5a,b). According to Devaney, it appeared to be an adaptation protecting the commensal from predation and affording a place to obtain food. Multiple polynoids (2–5) were found on single specimens of the multirayed starfish Acanthaster planci (Linnaeus) on Oahu by Devaney (1967:298). Uchida (1975:19) also described H. nigropunctata on A. planci from Japan.

On Eniwetok Atoll, Marshall Islands, a single specimen of H. nigropunctata was reported by Devaney (1967:297) on Ophiocoma anaglyptica Ely. In the Marquesas Islands, Devaney (1974b:154) reported the polynoid associated with O. doederleini Loriol. On the Solomon Islands, Gibbs (1969:450; 1971:120) reported specimens of H. nigropunctata living on the oral surface of the discs of Ophiocoma brevipes, O. dentata, and Ophiarthrum elegans Peters, under coral boulders, and in crevices on the reef platform. From the Cook Islands, Gibbs (1972:203) reported H. nigropunctata commensal with Ophiarthrum elegans and Ophiocoma dentata. On Aldabra, Sloan et al. (1979:94) reported H. nigropunctata on Ophiocoma brevipes.

DISTRIBUTION.—Indo-west and central Pacific Ocean. Intertidal to 57 meters.

Hololepidella sp.

Hololepidella nigropunctata.—Day, 1957:65, fig. 1a–f.—Balinsky, 1957:18.—Macnae and Kalk, 1962:101. [Not Horst, 1915.]

Polyeunoa nigropunctata.—Day, 1967:54, fig. 15r–u.—Amoureux, 1977:1101. [Not Horst, 1915.]

REMARKS.—The specimens from Inhaca Island, Mozambique, identified by Day (1957; 1967) as H. and P. nigropunctata, were found on the common brittle star Macrophiothrix hirsuta cheneyi (Lyman) by Balinsky (1957:18; polynoids identified by Day) and Macnae and Kalk (1962:101). As indicated by Devaney (1967:292) and Uchida (1975:29), the polynoids should be separated from H. nigropunctata based on the striking color pattern of dark stripes on the dorsum and the description by Day of the neurosetae. These polynoids need to be reexamined and more completely described.

Hololepidella ophiuricola Gibbs, 1969

FIGURE 4

Hololepidella commensalis.—Gibbs, 1969:451, fig. 132 [not Willey, 1905].

Hololepidella ophiuricola Gibbs, 1971:121, fig. 3A–H.—Hartmann-Schröder, 1984:66 [key].

MATERIAL EXAMINED.—CENTRAL PACIFIC OCEAN: Solomon Islands: Graham Point, Marau Sound, on oral surface of ophiuroid Macrophiothrix koehleri A.M. Clark, art Oct 1965, P.E. Gibbs, collector, holotype (BMNH 1970.20) and 3 paratypes (BMNH 1970.21–23). Kalota Island, Auki, Malaita, on oral surface of opiuroid Ophiarthrum pictum Müller and Troschel, 21 Nov 1965, P.E. Gibbs, collector, paratype (BMNH 1970.24).

DESCRIPTION.—Holotype 15 mm long, 3 mm wide including setae, with 53 segments and 25 pairs of elytra; 4 paratypes with 41–50 segments and 19–23 pairs of elytra, with arrangement as in H. nigropunctata. Pigmentation appearing on dorsal cirrophores and styles of dorsal cirri, on dorsum as broken or nearly continuous transverse bands, and as dark spots on ventral bases of parapodia and elytrophores (Figure 4A–C,E,F; Gibbs, 1969, fig. 132). Elytra oval, delicate, transparent, without microtubercles or papillae, with some scattered pigment spots (Gibbs, 1971, fig. 3B,C).

Bilobed prostomium with triangular lobes, pigmented medial part, and 2 pairs of small eyes; median antenna with dark ceratophore in anterior notch and long dark style; lateral antennae with small ceratophores inserted ventrally and short styles; palps long, tapered, pigmented on lateral sides; tentaculophores lateral to prostomium, without setae, each with pair of dorsal and ventral tentacular cirri similar to median antenna, minutely papillate (Figure 4A,B; Gibbs, 1971, fig. 3A). Second or buccal segment without nuchal lobe, with first pair of elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri; notosetae similar to those of following segments; neurosetae similar to upper ones of following segments (Figure 4A–D).

Notopodium of parapodium with projecting acicular lobe on lower side; larger neuropodium with tapered subtriangular presetal acicular lobe and shorter rounded postsetal lobe (Figure 4C,E,F; Gibbs, 1971, fig. 3D). Notosetae about same width as neurosetae, minutely serrated along one border (Figure 4G; Gibbs, 1971, fig. 3E). Few supraacicular neurosetae with prominent spinose rows on basal part and minute spinose row on distal part extending to near bare tips (Figure 4H; Gibbs, 1971, fig. 3F); more numerous subacicular neurosetae with faint spinose rows and slightly hooked, bare tips (Figure 4H,I; Gibbs, 1971, fig. 3G,H); all neurosetae with entire tips. Dorsal cirri with long cylindrical cirrophores on posterior sides of notopodia, styles extending close to tips of neurosetae; dorsal tubercles inconspicuous; ventral cirri extending near to tip of neuropodial lobe (Figure 4F).

BIOLOGY.—Hololepidella ophiuricola is commensal with ophiuroids. Individual specimens have been found on the oral surfaces of the discs of two species of ophiuroids: Macrophiothrix koehleri A.M. Clark and Ophiarthrum pictum Müller and Troschel, living under coral boulders and in crevices on the reef platform of the Solomon Islands. The pigment pattern of the polynoids differed somewhat: on M. koehleri the black bands were essentially longitudinal and on O. pictum, transverse, presumably imparting a cryptic coloration (Gibbs, 1969, fig. 132).

DISTRIBUTION.—Central Pacific Ocean, Solomon Islands. Low water.

Hololepidella arabica (Monro, 1937), new combination

FIGURE 5

Hamothoe arabica Monro, 1937:257, fig. 5a–f.—Fauvel, 1953:46, fig. 20c–g.

MATERIAL EXAMINED.—INDIAN OCEAN: Maldive Islands: East side of Kolumandulu Atoll, 02°13′S, 73°09′E, coral sand, John Murray Expedition sta 139, 25 Mar 1934, holotype (BMNH 1937.9.2.27).

DESCRIPTION.—Holotype, female with eggs (in 2 pieces), 8 mm long, 3 mm wide including setae, with 38 segments and 18 pairs of elytra, on segments 2, 4, 5, 7, then on alternate segments to 23, 26, 29, 31, 34, 36, 38, last pair of elytra very small. Body flattened, tapering slightly anteriorly and more so posteriorly, with transverse ciliated bands between elytrophores and dorsal tubercles, 2 per segment. Elytra round, delicate, transparent, without tubercles or papillae, with light brown mottled pigmentation on medial and posterior sides (Figure 5B).

Prostomium deeply bilobed, with triangular lobes, 2 pairs of small eyes, and light pigmentation on posterior half; ceratophore of median antenna in anterior notch, with very small style (regenerating); small ceratophores of lateral antennae inserted ventrally, with short subulate styles; palps stout, tapered; tentaculophores lateral to prostomium, each with single seta on inner side and pair of long dorsal and ventral tentacular cirri; antennae and cirri with delicate papillae (Figure 5A; Monro, 1937, fig. 5a). Second or buccal segment without nuchal fold, with elytrophores, with biramous parapodia and long ventral buccal cirri similar to tentacular cirri (Figure 5A).

FIGURE 4.—Hololepidella ophiuricola, paratype (BMNH 1970.22): A, anterior end, dorsal view; B, same, ventral view; C, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; D, neuroseta from same; E, right elytrigerous parapodium, anterior view, acicula dotted; F, right cirrigerous parapodium, posterior view; G, long and short notosetae; H, supraacicular neuroseta; I, subacicular neurosetae. (Scales = 0.5 mm for A,B; 0.2 mm for C,E,F; 0.1 mm for D,G–I.)

Parapodia with notopodium short, rounded, with projecting acicular lobe on lower side; longer neuropodium with projecting, conical presetal acicular lobe and shorter, rounded postsetal lobe; notosetae numerous, forming radiating bundle, extending to about tips of neuropodia; neurosetae numerous, forming fan-shape bundle (Figure 5C,D; Monro, 1937, fig. 5b). Notosetae about as stout as neurosetae, short to long, slightly curved, appearing smooth or very finely serrated (Figure 5E; Monro, 1937, fig. 5c). Upper few neurosetae smooth, tapering to slender slightly hooked tips; remaining supraacicular neurosetae coarsely spinose on lower half and finely spinose on distal half, with faintly bidentate hooked tips (Figure 5F; Monro, 1937, fig. 5d,e); subacicular neurosetae finely spinose, with faintly bidentate, hooked tips; lower ones with tips entire or faintly notched (Figure 5G; Monro, 1937, fig. 5f). Dorsal cirri with long cylindrical cirrophores and long papillate styles extending to tips of neurosetae or beyond, with brown pigmentation on basal half; dorsal tubercles small, nodular; ventral cirri short, tapering, extending to about tips of neuropodia, with brown pigmentation on basal half (Figure 5D). Pair of long anal cirri similar to dorsal cirri.

FIGURE 5.—Hololepidella arabica, holotype of Harmothoe arabica (BMNH 1937.9.2.27): A, dorsal view of anterior end, style of median antenna small, regenerating, right palp and left dorsal tentacular cirrus missing; B, right posterior elytron showing pigment pattern; C, right elytrigerous parapodium, anterior view, acicula dotted; D, right cirrigerous parapodium, posterior view; E, long and short notosetae; F, supraacicular neurosetae, with detail of tips; G, subacicular neurosetae, with detail of tip. (Scales = 0.5 mm for A, 0.2 mm for B–D; 0.1 mm for E–G.)

DISTRIBUTION.—Indian Ocean, Maldive Islands. Low water.

Genus Malmgreniella Hartman, 1967, emended

Malmgreniella Hartman, 1967:37. [Type-species: Malmgreniella dicirra Hartman, 1967, by monotypy. Gender: feminine.]

REMARKS.—Malmgreniella, as used herein and followed by Weston (1984), includes some of the species that have been referred to Malmgrenia McIntosh, 1874, and reviewed by Kudenov (1975b:79, key; 1977:95, key) and Hanley (1987:150, tables 1, 2). Malmgrenia whiteavesii McIntosh, 1874, from eastern Canada, the type species of the genus by monotypy, was poorly described by McIntosh (1874a:263, pl. 9: figs. 5–7) and the type specimen in the BMNH consists only of fragments. The genus Malmgrenia is considered to be indeterminable (also see Hanley, 1987:148). Malmgreniella (including Malmgrenia, as currently used) is close to Harmothoe Kinberg, 1856. As used herein, Malmgreniella includes harmothoid species having a close commensal relationship with other invertebrates. The elytra are delicate, with microtubercles, if present, confined to small anterior groups, sometimes with a few scattered on the surface; papillae are absent or confined to micropapillae, rarely with some longer lateral papillae. In Harmothoe, the elytra are tough, with numerous microtubercles and sometimes macrotubercles on the surface, and usually with long papillae. In Malmgreniella, the bilobed prostomium usually lacks cephalic peaks, with the lateral antennae inserted terminoventrally, compared with Harmothoe with prostomial cephalic peaks and ventral insertion of the lateral antennae.

DIAGNOSIS.—Segments up to 46. Elytra and elytrophores 14–15 pairs in adults, on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, and 32. Elytra delicate, with or without micropapillae, with or without anterior group of microtubercles, without macrotubercles. Prostomium bilobed, anterior lobes truncate, round or subtriangular, usually without distinct cephalic peaks, with 3 antennae, 2 palps, and 2 pairs of eyes; ceratophore of median antenna in anterior notch; lateral antennae with distinct ceratophores inserted terminoventrally, subterminally, or ventrally (converging midventrally). Tentaculophores of first segment lateral to prostomium, each with or without few setae on inner side, and pair of dorsal and ventral tentacular cirri. Second or buccal segment with first pair of elytrophores, subbiramous or biramous parapodia, and long ventral buccal cirri lateral to mouth. Parapodia subbiramous or biramous, with smaller notopodia and larger neuropodia, both rami with projecting acicular lobes; presetal acicular lobe of neuropodium longer, usually with supraacicular process; postsetal lobe shorter, rounded. Notosetae short, rodlike, similar in width to neurosetae, with tapering entire or notched blunt tips, with or without spinose rows. Neurosetae with spinose rows, with tips either all bifid, bifid and unidentate, or all unidentate. Dorsal cirri with cylindrical cirrophores and long styles; dorsal tubercles slightly developed or nodular. Ventral cirri short. Pygidium with pair of anal cirri. Pharynx with 9 pairs of border papillae and 2 pairs of hooked jaws.

Key to the Species of Malmgreniella from the Southern Ocean, South Atlantic Ocean (South Africa), Indian Ocean (South Arabian Coast), and South Pacific Ocean (New Guinea, SE Australia)

Malmgreniella dicirra Hartman, 1967

FIGURES 6–9

Malmgreniella dicirra Hartman, 1967:37, pl. 11A–D.—Cairns, 1983:130, 140, 145, 151.

Polynoe leioseta Averincev, 1978:65, pl. 7:55–59. [New synonymy.]

MATERIAL EXAMINED.—SOUTHERN OCEAN: Pacific Antarctic Ridge: 54°49′S, 129°48′W, 549 m, Eltanin sta 1346, 7 Nov 1964, 13 syntypes of M. dicirra (USNM 55493-4; BMNH 1986.57; ZMH); 1 specimen with Stylaster densicaulis Moseley (USNM 65951). 51°00′S, 162°01′E, 333–371 m, with Allopora eguchii, Boschma, Eltanin sta 1411, 8 Feb 1965, 1 specimen (USNM 65952). 56°21′S, 158°28′E, 1574–1693 m, with Calyptopora reticulata Boschma, Eltanin sta 1423, 12 Feb 1965, 5 specimens (USNM 65953). 54°09′S, 52°08′W, 419–483 m, with S. densicaulis, Eltanin sta 1521, 30 Jan 1966, 1 specimen (USNM 65954). 54°29′S, 39°22′W, 659–686 m, with S. densicaulis, Eltanin sta 1536, 8 Feb 1966, 1 specimen (USNM 65955). 54°43′S, 55°30′W, 1647–2044 m, with S. densicaulis and Conopora pauciseptata Brock, Eltanin sta 1592, 14 Mar 1966, 4 specimens (USNM 65956). 54°39′S, 170°22′E, 1862–2103 m, with Caltyptopora reticulata and Conopora pauciseptata, Eltanin sta 1991, 2 Jan 1968, 6 specimens (USNM 65957). Weddell Sea: Off Cape Norvegia, 71°50′S, 15°50′W, with Conopora pauciseptata and Lepidopora sp., Deep Freeze IV, USS Edisto TD-2, sta 14, 18 Jan 1959, 2 specimens (USNM 65950). Australia: Off Macquarie Island, 54°56′S, 158°49′E, 320 m, stony bottom, R/V D. Mendeleev sta 1294, 25 Jan 1976, paratype of P. leioseta (ZIASL 2/43284).

DESCRIPTION.—Syntypes of M. dicirra up to 23 mm long, 6 mm wide including setae, with 46 segments. Paratype of P. leioseta 12 mm long, 4 mm wide, with 40 segments. Additional specimens, commensal with stylasterine hydrocorals, 8–18 mm long, 3–5 mm wide, with 34–43 segments. Body colorless except for black eyes. Elytra 15 pairs, small, leaving middorsum uncovered, delicate, oval, without tubercles or papillae (Figure 8B,C; Hartman, 1967, pl. 11G; Averincev, 1978, fig. 7:56).

Prostomium bilobed, with truncate anterior border and 2 pairs of eyes on posterior half; median antenna with ceratophore in anterior notch, and long style with filamentous tip; lateral antennae with distinct ceratophores inserted terminoventrally and short styles; palps stout, tapered; tentaculophores each with pair of dorsal and ventral tentacular cirri similar to median antenna, without setae (Figures 6A, 8A; Hartman, 1967, pl. 11A; Averincev, 1978, fig. 7:55). Second segment with first pair of elytrophores, subbiramous parapodia, and long ventral buccal cirri similar to tentacular cirri; notosetae stouter than neurosetae, with 3–5 spinous rows; neurosetae tapering to slender, blunt, entire tips, with spinous rows (Figure 6A–D).

Parapodium subbiramous, with notopodium shorter and smaller than neuropodium, rounded with projecting acicular lobe on lower side and slightly projecting notoaciculum; longer neuropodium subconical with tapering presetal acicular lobe and short, thick, digitiform supraacicular process above projecting neuroaciculum and shorter, rounded postsetal lobe (Figures 6B, 7A,B,E, 8D,E, 9A,B; Hartman, 1967, pl. 11D). Notosetae short, relatively few in number, slightly stouter than neurosetae, with 2–6 spines along one side (none on posterior segments) (Figures 6C, 7C,F, 8F, 9C; Hartman, 1967, pl. 11E; Averincev, 1978, pl. 7:58). Neurosetae long, more numerous (25–30), with slightly enlarged spinose regions with close-set spines along cutting edges, with bifid tips (Figures 7D, 8G, 9D; Hartman, 1967, pl. 11F; Averincev, 1978, pl. 7:59). Dorsal cirri with short cylindrical cirrophores; styles either short, bulbous basally or longer, cirriform, former type occurring on segments 3, 6, 8, then more or less alternating regularly with latter type; dorsal tubercles slightly developed on anterior parapodia, bulbous on posterior ones; ventral cirri inflated basally on anterior parapodia, gradually tapering on posterior ones (Figures 7A,E, 8A,D, 9A; Hartman, 1967, pl. 11B,C). Pygidium with pair of long anal cirri.

FIGURE 6.—Malmgreniella dicirra, syntype (USNM 55494): A, dorsal view of anterior end, pharynx partially extended, right palp, left dorsal and ventral tentacular cirri and dorsal cirri of segment 3 missing; B, right elytrigerous parapodium from segment 2, posterior view; C, notoseta from same; D, upper and lower neurosetae from same. (Scales = 0.5 mm for A; 0.5 mm for B; 0.1 mm for C,D.)

FIGURE 7.—Malmgreniella dicirra, syntype (USNM 55494): A, right cirrigerous parapodium from segment 6, posterior view; B, right elytrigerous parapodium from segment 7, anterior view, acicula and newly formed internal notoseta dotted; C, notosetae from same; D, upper, middle, and lower neurosetae from same; E, left cirrigerous parapodium from segment 26, posterior view; F, notosetae from same. (Scales = 0.5 mm for A,B,E; 0.1 mm for C,D,F.)

FIGURE 8.—Malmgreniella dicirra, paratype of Polynoe leioseta (ZIASL 2/43284): A, dorsal view of anterior end, right palp and styles of median antenna and right dorsal tentacular cirrus missing; B, left elytron from segment 9; C, right elytron from segment 26; D, right cirrigerous parapodium from segment 10, posterior view; E, right elytigerous parapodium from segment 11, anterior view, acicula dotted; F, notosetae from same; G, upper and middle neurosetae from same. (Scales = 0.5 mm for A; 0.5 mm for B,C; 0.5 mm for D,E; 0.1 mm for F,G.)

FIGURE 9.—Malmgreniella dicirra, paratype of Polynoe leioseta (ZIASL 2/43284): A, right cirrigerous parapodium from segment 25; B, right elytrigerous parapodium from segment 26; C, notosetae from same; D, upper, middle, and lower neurosetae from same. (Scales = 0.5 mm for A,B; 0.1 mm for C,D.)

BIOLOGY.—Malmgraniella dicirra was found by Cairns (1983) living commensally with four stylasterine hydrocorals: Conopora pauciseptata Broch, Stylaster densicaulis Moseley, Allopora eguchii Boschma, and Calyptopora reticulata Boschma. According to Caims (1983:130), the polynoid induced Conopora pauciseptata to form a flattened tube about 3.8 × 2.2 mm in diameter (often considerably larger than the branch to which it was attached); the tubes were found on either the anterior or posterior sides and sometimes on both sides of the branch. On Stylaster densicaulis, the polynoid induced the coral to form a broad, flat tube on its anterior side. The tubes were quite long and up to 3 × 5.5 mm in diameter and even bifurcate at various points. The polynoid appeared to induce an anterior rectangular ridge, a preliminary step in tube formation (Caims, 1983:140, fig. 41A). On Calyptopora reticulata, the polynoids induced the coral to form flattened tubes, about 3 × 2.2 mm in diameter, on either the anterior or posterior sides or both sides of the colony (Cairns, 1983:151).

DISTRIBUTION.—Southern Ocean, Pacific Antarctic Ridge, off Macquarie Island, Weddell Sea. In 320–2103 meters.

Malmgreniella agulhana (Day, 1960), new combination

FIGURE 10

Harmothoe agulhana Day, 1960:277, fig. 1g–1; 1967:74, fig. 1.11.q–u.

MATERIAL EXAMINED.—SOUTH ATLANTIC OCEAN: South Africa: False Bay, 34°12′S, 18°37′E, University of Cape Town Ecological Survey, sta FAL 419, 15 May 1961, 48 m, sand and shell, identified by J.H. Day, 1 specimen (BMNH 1961.9.834).

DESCRIPTION.—Type material not available. According to Day, holotype 12 mm long, 2 mm wide without setae, and 36 segments. Anterior fragment from False Bay 3 mm long, 2 mm wide including setae, and 18 segments. Elytra large, oval to subreniform, smooth except for group of microtubercles near anteromedial margin (with faint network of brown pigmentation on exposed part, according to Day) (Figure 10B; Day, 1960, fig. 1k).

FIGURE 10.—Malmgreniella agulhana, specimen from False Bay, South Africa (BMNH 1961.9.834): A, dorsal view of anterior end, styles of median antenna, left dorsal and ventral, and right dorsal tentacular cirri missing; B, left elytron with detail of microtubercles; C, right elytrigerous parapodium, posterior view, ventral cirrus missing; D, short and long notosetae; E, upper, middle, and lower neurosetae. (Scales = 0.5 mm for A; 0.1 mm for B; 0.1 mm for C–E.)

Bilobed prostomium with lobes rounded to subtriangular, without distinct peaks; eyes small, anterior pair in region of greatest width, slightly larger than posterior pair; ceratophore of median antenna in anterior notch, with style about as long as prostomium; ceratophores of lateral antennae inserted terminoventrally, with short subulate styles; palps stout, tapered; tentaculophores lateral to prostomium, without setae, with dorsal and ventral tentacular cirri similar to median antenna (Figure 10A; Day, 1960, fig. 1j). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 10A; Day, 1960, fig. 1j).

Biramous parapodia with notopodia shorter than neuropodia, rounded, with projecting acicular lobes on lower sides; larger neuropodia with presetal conical acicular lobes with large digitiform supraacicular processes and shorter rounded postsetal lobes (Figure 10C; Day, 1960, fig. 1g). Notosetae about as stout as neurosetae, short, curved to longer, nearly straight, with spinose rows and tapered bare tips (Figure 10D; Day, 1960, fig. 1h). Upper neurosetae with longer spinose regions and bifid tips; middle ones with shorter spinose regions and bifid tips; lower neurosetae with entire tips (Figure 10E; Day, 1960, fig. 1i). Dorsal cirri with cylindrical cirrophores and papillate styles, extending beyond tips of neuropodia; dorsal tubercles nodular; ventral cirri short, subulate (Day, 1960, fig. 1g,j).

DISTRIBUTION.—Atlantic Ocean, South Africa. In 26–48 meters (see Day, 1960).

Malmgreniella dayi, new species

FIGURE 11

Harmothoe waahli.—Day, 1934:21, fig. 2a–d [not Kinberg, 1856].

MATERIAL EXAMINED.—SOUTH ATLANTIC OCEAN: South Africa: False Bay, intertidal, J.H. Day, collector, 1933, holotype (BMNH 1934.1.19.97, as Harmothoe waahli by Day, 1934).

DESCRIPTION.—Holotype in 2 pieces, with segments 15 and 16 missing, 13 mm long, 4 mm wide including setae, and 36 segments. Body flattened, tapering slightly anteriorly and posteriorly. Elytra 15 pairs, on usual segments, oval to subreniform, smooth, opaque, with group of microtubercles on anterior part, and some scattered micropapillae on surface; no color pattern visible (Figure 11B,C; Day, 1934, fig. 2b,c).

FIGURE 11.—Malmgreniella dayi, new species, holotype (BMNH 1934.1.19.97): A, dorsal view of anterior end, palps missing (drawn as dotted lines, taken from fig. 2a by Day, 1934); B, right 2nd elytron from segment 4, with detail of microtubercles; C, right posterior elytron; D, right parapodium from segment 2, anterior view, acicula dotted; E, upper, middle, and lower notosetae from same; F, right elytrigerous parapodium, anterior view, acicula dotted; G, right cirrigerous parapodium, posterior view; H, short and long notosetae; I, upper, middle, and lower neurosetae, with detail of tips. (Scales = 0.5 mm for A; 0.5 mm for B,C; 0.2 mm for D,F,G; 0.1 mm for E,H,I.)

Bilobed prostomium with anterior lobes truncate, without peaks; anterior pair of eyes anterolateral, larger than posterodorsal pair; ceratophore of median antenna large, in anterior notch, with papillate style longer than prostomium; ceratophores of lateral antennae inserted terminoventrally, with short, papillate, subulate styles; palps long, stout, tapering (now missing); tentaculophores lateral to prostomium, with single seta on inner side; dorsal and ventral tentacular cirri similar to median antenna (Figure 11A; Day, 1934, fig. 2a). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 11A,D). Notosetae numerous, similar to those of following segments; neurosetae more slender than those following, lower and upper ones with bulbous entire tips, middle ones each with secondary tooth (Figure 11E).

Biramous parapodia with smaller notopodia rounded, with projecting acicular lobes on lower sides; larger neuropodia with subconical presetal acicular lobes with thick supraacicular processes; postsetal lobes shorter, rounded (Figure 11D,F,G). Notosetae moderate in number, of several lengths, slightly stouter than neurosetae, each with faint spinose rows and short tapered bare tips (Figure 11H). Neurosetae moderate in number, upper ones with long spinose regions, upper few with tapered entire tips, rest with secondary tooth; lower neurosetae with shorter spinose regions, with small secondary tooth or entire (Figure 11I; Day, 1934, fig. 2d). Dorsal cirri with cylindrical cirrophores and styles, extending to tips of neurosetae or slightly beyond, with scattered short papillae; dorsal tubercles nodular; ventral cirri short, subulate, with short papillae (Figure 11G; Day, 1934, fig. 2a). Pygidium with dorsal anus medial to parapodia of posterior small segment, with pair of long anal cirri.

ETYMOLOGY.—The species is named for John H. Day, in recognition of his many contributions to the study of the Polychaeta.

DISTRIBUTION.—Atlantic Ocean, South Africa. Intertidal.

Malmgreniella capensis (McIntosh, 1885), new combination

FIGURE 12

Polynoe capensis McIntosh, 1885:114, pl. 4: fig. 4; pl. 15: fig. 1; pl. 19: fig. 4; pl. 9A: figs. 4, 5.

?Polynoe capensis.—Day, 1960:283 [examination of types].

Malmgrenia purpurea Day, 1960:281, fig. 2f–k.—1967:50, fig. 1.4.1–o.—Intes and Le Loeuff, 1975:273.—Kudenov, 1975b:79.—Hanley, 1987:160 [under incertae sedis]. [New synonymy.]

Parahalosydna capensis.—Day, 1967:86, fig. 1.15.a–d.

MATERIAL EXAMINED.—SOUTH ATLANTIC OCEAN: South Africa: Near Cape of Good Hope, 34°41′S, 18°36′E, 179 m, green sand, Challenger sta 141, 17 Dec 1873, 2 syntypes of Polynoe capensis (BMNH 1885.12.1.94). False Bay, 34°16′S, 18°40–42′E, 60–62 m, sand and shell, with Spatangus capensis Döderlein, University of Cape Town Ecological Survey, sta FAL 359/375, 24/25 Feb 1959, 2 syntypes of Malmgrenia purpurea (USNM 53540).

TYPE MATERIAL.—The two syntypes of P. capensis are fragmented, somewhat hardened, and in rather poor condition, measuring 19 mm in length, 7 mm in width including the setae, with 38–39 segments. The two syntypes of M. purpurea also are fragmented but in fair condition, the larger and smaller syntypes measuring 16 and 10 mm in length, 5 and 3 mm in width, with 38 and 37 segments, respectively.

DESCRIPTION.—Body elongated, flattened, tapering slightly anteriorly and posteriorly; 15 pairs of elytra, dull blackish grey where exposed (Mcintosh, 1885, pl. 19: fig. 1; figure incorrectly shown with 16 pairs of elytra); color of body dark purple, according to Day (1960); posterior most 6 or so segments of dorsum and ventrum with dark bands and dark pigment spots on anterior sides and bases of parapodia. Elytra large, oval to subreniform, with posterior two-thirds uniformly darkly colored and with group of microtubercles near anterior notch (Figure 12B; McIntosh, 1885, pl. 19: fig. 4; Day, 1960, fig. 2j).

Bilobed prostomium with lobes truncate, without peaks; eyes moderate in size, anterior pair anterolateral, larger than posterodorsal pair; ceratophore of median antenna large, in anterior notch, with large, club-shape style; ceratophores of lateral antennae distinct, inserted terminoventrally, with short, tapered styles; palps stout, tapered; tentaculophores lateral to prostomium with 0–1 seta on inner side, with dorsal and ventral tentacular cirri (Figure 12A; McIntosh, 1885, pl. 4: fig. 4; Day, 1960, fig. 2g). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 12A,C; Day, 1960, fig. 2g).

Biramous parapodium with smaller subconical notopodium with projecting acicular lobe on lower side; larger neuropodium with conical presetal acicular lobe with supraacicular triangular lip and shorter, rounded postsetal lobe (Figure 12C–E; Day, 1960, fig. 2k). Notosetae moderate in number, long to short, about as stout as neurosetae, with minute serrations and abruptly tapered tips, forming radiating bundle (Figure 12C,F: McIntosh, 1885, pl. 9A: fig. 4; Day, 1960, fig. 2h,h′). Neurosetae moderate in number, forming fan-shape bundle; upper ones with longer spinose regions and slight indication of secondary tooth; middle and lower neurosetae with shorter, faintly marked spinose rows, more developed hooked tip, and small secondary tooth (Figure 12D,G; McIntosh, 1885, pl. 9A: fig. 5; Day, 1960, fig. 2f,f). Dorsal cirri with cylindrical cirrophores and styles inflated subdistally, smooth, shorter than neurosetae; dorsal tubercles nodular; ventral cirri extending almost to tips of neuropodia (Figure 12D,E; Day, 1960, fig. 2g,k). Pygidium with pair of dark sausage-like anal cirri.

FIGURE 12 anterior end, styles of median and left lateral antennae and left dorsal and right dorsal and ventral tentacular cirri missing; B, right elytron, with detail of microtubercles; C, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; D, right middle elytrigerous parapodium, anterior view, acicula dotted; E, right cirrigerous parapodium, posterior view; F, short and long notosetae; G, upper, middle, and lower neurosetae, with detail of tip. (Scales = 0.5 mm for A; 1.0 mm for B; 0.5 mm for C–E; 0.1 mm for F,G.)

BIOLOGY.—The dark coloration of Malmgreniella capensis strongly suggests a commensal relationship. The types of the synonym M, purpurea Day were collected with the echinoid spatangid Spatangus capensis Döderlein, and this allowed Day (1960) to suggest it as the probable host for the polychaete.

DISTRIBUTION.—South Atlantic Ocean, South Africa. In 60–179 meters.

Malmgreniella inhacaensis, new species

FIGURE 13

Harmothoe lunulata.—Day, 1962:629 [not Delle Chiaje, 1830].

MATERIAL EXAMINED.—INDIAN OCEAN: Mozambique: Inhaca Island, Delagoa Bay, 25°50′S, 32°50′E, University of Cape Town Ecological Survey (IN 145A), commensal in burrows of Balanoglossus sp. and Ptychodera sp., holotype (BMNH 1963.1.4, as Harmothoe lunulata by Day, 1962).

DESCRIPTION.—Length of holotype 17 mm, width 5 mm including setae, 37 segments, last segment minute. Body flattened, tapering slightly anteriorly and posteriorly, with 2 transverse ciliated bands per segment. Ventral surface of posterior region with brown transverse bands. Eltyra 15 pairs, on usual segments, oval to subreniform, with rather large group of microtubercles on anterior part beginning on 2nd elytra, fewer in number on posterior elytra, and with some scattered micropapillae on surface; brown pigmentation pattern consisting of transverse bands and bands along medial and posterior borders (Figure 13B–E).

Bilobed prostomium with lobes truncate, without peaks; anterior pair of eyes anterolateral, nearly hidden from view dorsally, smaller posterior pair posterodorsal; ceratophore of median antenna large, in anterior notch of prostomium, style missing; lateral antennae with ceratophores inserted terminoventrally, with styles short, subulate, with short papillae; palps stout, tapered; tentaculophores lateral to prostomium, each with single seta on inner side and long tapered papillate dorsal and ventral tentacular cirri (Figure 13A). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri; notosetae similar to those of following parapodia; neurosetae more slender than following, with entire and bifid bulbous tips (Figure 13F,G).

Biramous parapodia with notopodia much shorter than neuropodia, rounded with projecting acicular lobes on lower sides; larger neuropodia with subconical presetal acicular lobes with thick supraacicular processes and shorter, rounded postsetal lobes (Figure 13H,I). Notosetae very numerous, of several lengths, forming spreading bundle, about as stout as neurosetae, with numerous spinose rows and tapered to rounded tip (Figure 13J). Neurosetae numerous, forming fan-shape bundle, upper ones with longer spinose regions and long slender secondary tooth; middle and lower ones with shorter spinose regions and shorter secondary tooth (Figure 13K). Dorsal cirri with cylindrical cirrophores; styles with short papillae and filamentous tips extending to about tips of neurosetae; dorsal tubercles bulbous; ventral cirri short, tapered, with short papillae (Figure 13I). Pygidium with dorsal anus medial to parapodia of posterior small segment, with pair of long anal cirri.

BIOLOGY.—Malmgreniella inhacaensis is found living commensally in the burrows of the enteropneusts Ptychodera and Balanoglossus.

ETYMOLOGY.—The species is named for the collection site of Inhaca Island.

DISTRIBUTION.—Indian Ocean, Mozambique, Inhaca Island.

Malmgreniella murrayensis, new species

FIGURE 14

Harmothoe lunulata— Monro, 1937:258 [not Delle Chiaje, 1830],

MATERIAL EXAMINED.—INDIAN OCEAN: South Arabian Coast: 18°03′N, 57°02′E, 38 m, Lithothamnion bottom, in cavity of sponge, John Murray Expedition, 29 Jul 1933, holotype (BMNH 1937.9.2.28, as H. lunulata by Monro, 1937).

DESCRIPTION.—Length 8 mm, width 3 mm including setae, 36 segments, last 2 segments very small. Body flattened, tapered slightly anteriorly and posteriorly, ventral side of posterior end with brownish streaks. Elytra 15 pairs, on usual segments, oval to subreniform, with group of microtubercles on anterior side of more anterior elytra, and scattered micropapillae on surface; brownish pigmentation forming almost complete circle, wider in some areas (Figure 14B–D).

Bilobed prostomium with anterior lobes truncate, without peaks; eyes small, anterior pair anterolateral, anterior to greatest width of prostomium, slightly smaller posterior pair posterodorsal; ceratophore of median antenna in anterior notch, style missing; ceratophores of lateral antennae inserted terminoventrally, with styles short, subulate, papillate; palps stout, tapered, appearing wrinkled; tentaculophores lateral to prostomium with single seta on inner side, tentacular cirri missing (Figure 14A). Segment 2 with first pair of large elytrophores, biramous parapodia, and long papillate ventral buccal cirri (Figure 14A). Pharynx not extended and not examined.

Biramous parapodium with notopodium shorter and smaller than neuropodium, rounded with projecting acicular lobe on lower side; neuropodium with subconical presetal acicular lobe with small supraacicular process and shorter, rounded postsetal lobe (Figure 14E,F). Notosetae moderate in number, of several lengths, with faint close-set spinose rows; shorter notosetae with longer tapered bare tips, longer notosetae with wider and shorter blunt bare tips, tips thinner on one side forming sort of flange (Figure 14G). Neurosetae moderate in number, with rather faint spinose rows, all with bifid tips, upper ones with longer spinose regions (Figure 14H). Dorsal cirri with cylindrical cirrophores, with slightly raised glandular areas on posterior sides; styles extending to tips of neurosetae, with filamentous tips and short clavate papillae; dorsal tubercles nodular, ventral cirri short, subulate, with short papillae (Figure 14F). Pygidium with dorsal anus medial to small posterior parapodia; anal cirri missing.

FIGURE 13.—Malmgreniella inhacaensis, new species, holotype (BMNH 1963.1.4): A, dorsal view of anterior end, styles of median and left lateral antennae and left tentacular cirri missing; B, right 1st elytron from segment 2; C, right 2nd elytron from segment 4, with detail of microtubercles; D, right middle elytron; E, right 13th elytron from segment 26; F, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; G, upper, middle, and lower neurosetae from same; H, right middle elytrigerous parapodium, anterior view, acicula dotted; I, right cirrigerous parapodium, posterior view; J, notoseta; K, upper, middle, and lower neurosetae. (Scales = 0.5 mm for A; 1.0 mm for B–E; 0.2 mm for F,H,I; 0.1 mm for G,J,K.)

FIGURE 14.—Malmgreniella murrayensis, new species, holotype (BMNH 1937.9.2.28): A, dorsal view of anterior end, styles of median antenna and tentacular cirri missing; B, right 2nd elytron from segment 4, with detail of microtubercles; C, left 5th elytron from segment 9; D, left posterior elytron; E, left elytrigerous parapodium, anterior view, acicula dotted; F, left cirrigerous parapodium, posterior view; G, short and long notosetae, with detail of tip; H, upper, middle, and lower neurosetae. (Scales = 0.5 mm for A; 0.5 mm for B–D; 0.2 mm for E,F; 0.1 mm for G,H.)

BIOLOGY.—Malmgreniella murrayensis was found living deeply embedded in the cavity of a sponge.

ETYMOLOGY.—The species is named for the John Murray Expedition.

DISTRIBUTION.—Indian Ocean, South Arabian Coast. In 38 meters.

Malmgreniella pettiti, new species

FIGURE 15

MATERIAL EXAMINED.—SOUTH PACIFIC OCEAN: New Guinea: Papua, Bootless Inlet, about 4 mi NE of Raine Point, 28 May 1981, removed from Alcyonacea, G.R. Pettit, collector, holotype (USNM 80508).

DESCRIPTION.—Holoptype 5 mm long, 1.5 mm wide including setae, and 34 segments. Body flattened, tapered posteriorly, colorless. Elytra 15 pairs, on usual segments, large, oval, covering dorsum; mottled brownish pigmentation on medial two-thirds and scattered microtubercles on anterior part (Figure 15B).

Prostomium oval, bilobed, with anterior lobes truncate, without peaks; anterior pair of eyes anteroventral, larger than posterodorsal pair, ceratophore of median antenna bulbous, in anterior notch, style missing; ceratophores of lateral antennae inserted terminoventrally, with styles short, subulate, minutely papillate; palps stout, tapered; tentaculophores lateral to prostomium, each with single seta on inner side; dorsal and ventral tentacular cirri about length of palps, with short papillae (Figure 15A). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 15A).

FIGURE 15.—Malmgreniella pettiti, new species, holotype (USNM 80508): A, dorsal view of anterior end style of median antenna, right dorsal and ventral tentacular cirri, and right palp missing; B, left elytron, with detail of microtubercles; C, right elytrigerous parapodium, anterior view, acicula dotted; D, right cirrigerous parapodium posterior view style of dorsal cirrus missing; E, short and long notosetae, with detail of tips; F, middle neuroseta, with detail of tip. (Scales = 0.5 mm for A; 0.2 mm for B; 0.1 mm for C–F.)

Biramous parapodium with notopodium short, rounded, with projecting acicular lobe on lower side; longer neuropodium with subconical presetal acicular lobe with digitiform supraacicular process; postsetal lobe shorter, rounded (Figure 15C,D). Notosetae numerous, of several lengths, forming radiating bundle. Notosetae about equal in width to neurosetae, with numerous spinose rows, shorter ones with tips entire and pointed or notched, longer ones mostly with notched tips (Figure 15E). Neurosetae numerous, forming fan-shape bundle, all with rather long, bare, bifid hooked tips, upper ones with longer spinose regions (Figure 15F). Dorsal cirri with cylindrical cirrophores and papillate styles extending to tips of neurosetae; dorsal tubercles nodular; ventral cirri short, subulate (Figure 15D).

BIOLOGY.—Associated with Alcyonacea.

ETYMOLOGY.—The species is named for the collector, G.R. Pettit.

DISTRIBUTION.—South Pacific Ocean, New Guinea.

Malmgreniella phillipensis (Knox and Cameron, 1971), new combination

FIGURE 16

Malmgrenia phillipensis Knox and Cameron, 1971:22, figs. 1–6.—Kudenov, 1977:85, pl. 1a–n.

Harmothoe phillipensis.—Hanley, 1987:153, fig. 31.

MATERIAL EXAMINED.—PACIFIC OCEAN: Australia: Victoria, Port Phillip Bay Survey, 1957–63, area 14 (175), 2 paratypes (NMV G1737).

DESCRIPTION.—Two paratypes examined, consisting of anterior fragments, 9 and 11 mm long, 6 and 7 mm wide including setae, with 18 and 22 segments (up to 20 mm long, 6 mm wide, with 38 segments in original description). Middorsum reddish brown to cream, ventrum pale cream; according to Kudenov (1977), posterior ventral half of each posterior segment with transverse brown bands. Elytra 15 pairs, on usual segments, large, overlapping, covering dorsum. Elytra oval to subreniform, stiff, slick and shiny, sometimes showing 1–3 indistinct parallel longitudinal ridges (due to preservation, according to Kudenov, 1977:88), with wide band of numerous microtubercles on anterior region and some scattered on surface; brownish pigmentation in form of large anteromedial patch, continuing as narrow band along medioposterior border (Figure 16C,D; Knox and Cameron, 1971, figs. 2, 3; Kudenov, 1977, pl. 1d–g).

Bilobed prostomium with anterior lobes truncate, without peaks; anterior pair of eyes anterolateral, anterior to widest part of prostomium, larger than posterodorsal pair; ceratophore of median antenna large, in anterior notch, style about length of prostomium; ceratophores of lateral antennae distinct, inserted terminoventrally, with styles short, subulate; palps large, stout, tapering; tentaculophores lateral to prostomium, with 0–1 seta on inner sides and pair of dorsal and ventral tentacular cirri similar to median antenna; antennae and tentacular cirri with scattered micropapillae; subconical facial tubercle between ventral bases of palps; antennae, tentacular cirri, facial tubercle, and anterior lip of ventral mouth showing brownish pigmentation (Figure 16A,B; Knox and Cameron, 1971, fig. 1; Kudenov, 1977, pl. la; Hanley, 1987, fig. 3I). Segment 2 with small, slightly bilobed nuchal fold, first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri; notosetae similar to following notosetae; neurosetae more slender than those following, upper ones with bifid tips, lower ones ending in rounded blunt tips (Figure 16A,E,F).

Notopodia of biramous parapodia rounded, with projecting acicular processes on lower side; larger neuropodia with subconical presetal lobes with projecting supraacicular processes and shorter, rounded postsetal lobes (Figure 16E,G,H; Knox and Cameron, 1971, fig. 4; Kudenov, 1977, pl. 1b,c). Notosetae very numerous, of 3 lengths, slightly curved, moderately stout, slightly stouter than neurosetae, forming radiating bundles. Notosetae with numerous rows of fine serrations, with rounded to tapered pointed tips, showing verying degrees of wear (Figure 16I; Knox and Cameron, 1971, fig. 5; Kudenov, 1977, pl. 1h–j). Neurosetae numerous, forming fan-shape bundle, upper ones with longer spinose regions, with tips entire or with small secondary tooth; middle ones with shorter spinose regions and bifid tips; lower ones with entire tips or slight indication of secondary tooth (Figure 16J; Knox and Cameron, 1971, fig. 6; Kudenov, 1977, pl. 1k–n). Dorsal cirri with cylindrical cirrophores, with long, tapered styles extending to about tips of neurosetae, with scattered micropapillae; dorsal tubercles slightly inflated to nodular; ventral cirri short, tapered (Figure 16H; Knox and Cameron, 1971, fig. 4; Kudenov, 1977, pl. 1c).

DISTRIBUTION.—South Pacific Ocean, SE Australia. Low water to 24 meters.

FIGURE 16.—Malmgreniella phillipensis, paratype of Malmgrenia phillipensis (NMV G1737): A, dorsal view of anterior end, turned slightly to right side and showing lateral position of anterior eye; styles of median antenna, right and left dorsal tentacular cirri missing; B, midventral view of anterior end, showing position of ceratophores of lateral antennae between palps and facial tubercle; C, right anterior elytron, with detail of microtubercles; D, right middle elytron; E, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; F, upper and lower neurosetae from same, with detail of tips; G, right elytrigerous parapodium, anterior view, acicula dotted; H, right cirrigerous parapodium, posterior view; I, short and longer notosetae; J, upper, middle, and lower neurosetae, with detail of tips. (Scales = 0.5 mm for A,B; 0.5 mm for C,D; 0.2 mm for E,G,H; 0.1 mm for F,I,J.)

Key to the Species of Malmgreniella from the Northeastern Atlantic Ocean (Spitzbergen, Norway, Sweden, England, France, Mediterranean Sea)

Malmgreniella andreapolis (McIntosh, 1874), new combination

FIGURE 17

Laenilla glabra Malmgren, 1865:73 [part, not same as lectotype].

Malmgrenia andreapolis McIntosh, 1874b:195; 1876a:377, pl. 67: figs. 20–23; 1900:382, pl. 28: fig. 8; pl. 31: fig. 3; pl. 33: fig. 11; pl. 40: figs. 27–30.

Harmothoe synaptae Saint-Joseph, 1906:147, pl. 1: figs. 1–6. [New synonymy.]

Harmothoe lunulata.—Cuénot, 1912:102, figs. 23–26.—Southern, 1914:52.—Orton, 1923:861, fig. 1 [part].—Cazaux, 1968:506, figs. 7, 8. [Not Delle Chiaje, 1830.]

Harmothoe lunulata var. andreapolis.—Fauvel, 1923:72, fig. 26k–o.

Harmothoe lunulata var. synaptae.—Fauvel, 1923:73.

Malmgrenia lunulata.—Pettibone, 1953:25 [part; not Delle Chiaje, 1830].

Harmothoe lunulata, var. E.—Spooner in Plymouth Marine Fauna, 1957:xxxviii, 112.

Malmgrenia castanea.—Amanieu and Cazaux, 1963:168 [not McIntosh, 1876a].

Harmothoe andreapolis.—Tebble and Chambers, 1982:49, figs. 16a–d, 46, 47.—Hanley, 1987:151, fig. 3D.

MATERIAL EXAMINED.—NORTHEASTERN ATLANTIC OCEAN: Sweden: Gullmaren, S. Lovén, collector, 1 specimen mixed with lectotype of Laenilla glabra Malmgren (NRS 5752). Scotland: St. Andrews, washed ashore after storms and in stomachs of cod and haddock, 17 syntypes of Malmgrenia andreapolis (BMNH 1921.5.1.5.0–511; USNM 54161). England: Plymouth area, var. E, with ophiuroid Acrocnida brachiata (Montagu): Salcombe, east side, G.M. Spooner, collector, 7 small specimens (LCHP–S-7); Salcombe, west side, 16 Mar 1966. P.E. Gibbs, collector, 4 small specimens (LCHP–G-5, USNM 55049). Plymouth area, var. E. synaptae, with Leptosynapta bergenensis (as L. inhaerens): Salcombe, west side, 26 Mar 1952, G.M. Spooner, collector, 1 specimen (LCHP–S-12); Salcombe, west side, 16 Mar 1968, P.E. Gibbs, collector, 2 specimens (LCHP–G-6; USNM 55050). Plymouth area var. E, with synaptid Labidoplax digitata (Montagu): Salcombe, east side, 25 Mar 1955, G.M. Spooner, collector, 1 specimen (USNM 55048, from LCHP–S-9); Salcombe, west side, 28 Mar 1967, P.E. Gibbs, collector, 1 specimen (LCHP–G-9). France: Terrenez, off Roscoff, commensal with synaptid, 15 Sep 1913, 1 specimen (MNHN).

TYPE MATERIAL.—Syntypes of M. andreapolis all fragmented, consisting of 17 anterior, 9 middle, and 2 posterior fragments. Anterior and posterior fragments of 12 and 23 segments, totaling 37, 25 mm long and 7 mm wide including setae. Pharynx extended on one of syntypes.

NONTYPE MATERIAL.—Larger specimens from Plymouth area, with synaptids, 18–29 mm long, 6–9 mm wide including setae, with 37–38 segments. Smaller specimens, with ophiuroids, 11–13 mm long, 4.5 mm wide, with 35–37 segments; smallest specimen 6 mm long, 3 mm wide, with 27 segments. Small specimen from Sweden 12 mm long, 5 mm wide, with 36 segments.

DESCRIPTION.—Body flattened, tapering slightly anteriorly and more so posteriorly. Dorsum of posterior segments with 2 dark bands per segment; posterior 10–15 segments ventrally with characteristic color pattern (Figure 17M; Cuénot, 1912, fig. 25). Fifteen pairs of elytra, large, soft, completely covering dorsum; first pair oval, with narrow dark band near anterior border, complete circle and dark spot on place of attachment to elytrophore, and some scattered micropapillae (Figure 17B); following elytra subreniform to oval, with complete or nearly complete circular band on posterior half and band of microtubercles near anterior border (Figure 17C,D; McIntosh, 1900, pl. 33: fig. 11); more posterior elytra without microtubercles and with pigmentation scattered, less concentrated (Figure 17E).

Bilobed prostomium with anterior lobes wide, truncate; eyes rather small, anterior pair anterolateral, about twice as large as posterodorsal pair; ceratophore of median antenna large, in anterior notch, with style about as long as prostomium; ceratophores of lateral antennae inserted terminoventrally, with styles short, subulate; antennae darkly pigmented, with scattered minute papillae; palps stout, tapered, smooth, variable in length; tentaculophores with 0–1 seta on inner side and pair of dorsal and ventral tentacular cirri, similar to median antenna (Figure 17A; McIntosh, 1900, pl. 28: fig. 8; Cuénot, 1912, fig. 26; Hanley, 1987, fig. 3D). Biramous parapodia of segment 2 with long ventral buccal cirri similar to tentacular cirri; notosetae similar to those following; neurosetae more slender than those following, upper and lower ones with entire bulbous tips, middle ones with long bare regions and bifid tips (Figure 17A,F,G). Extended pharynx with 9 pairs of border papillae and 2 pairs of reddish amber-colored jaws.

Notopodium short, rounded, with projecting acicular lobe on lower side; neuropodium with subconical presetal acicular lobe with thick digitiform supraacicular process and shorter rounded postsetal lobe (Figure 17H,I). Notosetae moderate in number, short to long, extending to about tip of neuropodia; slender, translucent, similar in size to neurosetae, with faint spinose rows and rounded bulbous tips (Figure 17J; McIntosh, 1900, pl. 40: fig. 27). Neurosetae moderate in number, forming fan-shape bundle, long, translucent; upper supraacicular neurosetae with longer spinose regions and entire bulbous tips and lower supraacicular neurosetae with secondary tooth (Figure 17K); upper subacicular neurosetae stouter, with shorter spinose regions, slightly hooked tips with secondary tooth, and lower ones with single knobbed tips (Figure 17L; McIntosh, 1900, pl. 40: figs. 28–30). Cirrophores of dorsal cirri long, cylindrical; styles tapered, extending to about tips of neurosetae, with scattered clavate papillae; dorsal tubercles nodular; ventral cirri short, subulate, with few clavate papillae (Figure 17I). Pygidium dorsal, medial to bases of small posterior parapodia, with pair of long anal cirri.

BIOLOGY.—In the Plymouth area, Spooner in Plymouth Marine Fauna (1957:112) and Orton (1923:861, fig. 1) observed Malmgreniella andreapolis (as Harmothoe lunulata var. E, with a strong circular pattern) commensal in the burrows of echinoderms. Smaller specimens were found with the ophiuroid Acrocnida brachiata (Montagu) whereas larger specimens were associated with the synaptids Leptosynapta galliennii (Herapath) (as L. inhaerens and L. bergenensis) and Labidoplax digitata (Montagu), hence M. andreapolis apparently changes hosts.

In Arcachon, France, Cuénot (1912:102, figs. 23–26) found the polynoids (as Harmothoe lunulata) living commensally with synaptids. The polynoids were found in the galleries excavated in the sand by the synaptids, crawling either on the body of its host or on the wall. When touched or when the water was agitated, the polynoids showed very brilliant white greenish luminescence.

Cazaux (1968:506–510, figs. 7, 8) studied the early development of the species (as Harmothoe lunulata) from the eggs to trochophores, metatrochophores I and II, nectochaetes I and II until when, after 3 weeks, they reached 10 segments and a length of 1.7 mm. Cazaux also noted that adults were very abundant on beaches of Arcachon with synaptids and were found in the galleries and along the walls of the hosts.

DISTRIBUTION.—Northeastern Atlantic Ocean from Sweden to France. Intertidal.

Malmgreniella arenicolae (Saint-Joseph, 1888), new combination

FIGURES 18, 19

Harmothoe arenicolae Saint-Joseph, 1888:174, pl. 6: figs. 22, 23 [with Arenicola marina].

Harmothoe lunulata var. arenicolae.—Fauvel, 1923:72, 73.

Harmothoe lunulata (var. nigra).—Orton, 1923:861 [part; with Golfingia vulgaris].

FIGURE 17.—Malmgreniella andreapolis, specimen from Plymouth area, with synaptid (USNM 55048): A, dorsal view of anterior end; B, right 1st elytron from segment 2; C, right 2nd elytron from segment 4, with detail of microtubercles; D, right 8th elytron from segment 15; E, right 15th elytron from segment 32; F, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; G, upper and middle neurosetae from same; H, right middle elytrigerous parapodium, anterior view, acicula dotted; I, right cirrigerous parapodium, posterior view; J, short and long notosetae; K, upper and lower supraacicular neurosetae; L, upper and lower subacicular neurosetae; M, ventral view of left side of segments 28 and 29, showing pigmentation pattern. (Scales = 0.5 mm for A; 1.0 mm for B–E; 0.2 mm for F,H,I; 0.1 mm for G, J–L; 1.0 mm for M.)

Malmgrenia alba.—Augener, 1928:694 [not Laenilla alba Malmgren, 1865].

Harmothoe lunulata var. and form A.—Spooner, in Plymouth Marine Fauna, 1957:xxxvi, 112 [with Arenicola marina.]

Harmothoe lunulata, form I.—Spooner in Plymouth Marine Fauna, 1957:112 [with Amphitrite johnstoni].

Harmothoe lunulata, var. G.—Spooner in Plymouth Marine Fauna, 1957:xxxvi, 112 [with Golfingia elongata and G. vulgaris.]

Harmothoe lunulata.—Plymouth Marine Fauna, 1957:xxxvi [with Notomastus latericeus].—Hamond, 1966:386 [with Lanice conchylega]. [Not Delle Chiaje, 1830.]

MATERIAL EXAMINED.—NORTHEASTERN ATLANTIC OCEAN: Spitzbergen: Grube, collector, 2 specimens (ZMB 1121, as Malmgrenia alba by Augener, 1928). Norway: Kreknag, with Arenicola, C. Dons, collector, 10 Aug 1938, 1 specimen (ZMT). England: Plymouth area: Salcombe Estuary, with Arenicola marina (Linnaeus): Salcombe, east side, 18 Mar 1947; Millbay, 12 May 1953; west side, 13 Apr 1960, G.M. Spooner, collector, 9 specimens (LCHP–S-11/13/16; USNM 55052); Salcombe, west side, 15 Oct 1966, G.E. Gibbs, collector, 1 specimen (LCHP–G-2). Salcombe Estuary, with Notomastus latericeus Sars: 20 Mar 1954, G. Spooner, collector, 1 specimen (LCHP–S-4). Salcombe Estuary, with Golfingia elongata (Keferstein): Ox Point, 5 Oct 1956, 2 specimens (LCHP–S-3/5, as var. J). Salcombe Estuary, with Amphitrite johnstoni Malmgren: below Marine Hotel, 29 Jul 1957, G. Spooner, collector, 1 specimen (LCHP–S-15, as var. J). Salcombe Estuary, with Loimia medusa (Savigny): east side, 18 Mar 1957, 26 Mar 1959, G. Spooner, collector, 3 specimens (LCHP–S-1/6, as var. J); Millbay, 25 Mar 1970, P. Gibbs, collector, 1 specimen (LCHP–G-4). Yealm Estuary, with Amphitrite johnstoni: River Yealm, 15 Apr 1953, 17 Mar 1961, G. Spooner, collector, 6 specimens (LCHP–S-14/17; USNM 55051, as var. J). Yealm Estuary, with Golfingia elongata: Yealm Mouth, 26 Mar 1963, G. Spooner, collector, 2 specimens (LCHP–S-2/8, as var. P); River Yealm, 16 Sep 1974, P. Gibbs, collector, 1 specimen (LCHP–G-8). Plymouth, 1 specimen (BMNH 1932.7.11.13, as Harmothoe lunulata by Monro). St. Peters Port, Guernsey, Channel Islands, 1 specimen (BMNH 1921.5.1.444, as H. lunulata by McIntosh). Southport, Lancashire, 4 specimens (BMNH 1921.5.1.445, as H. lunulata by McIntosh). Norfolk Coast, with Lanice conchylega (Pallas), R. Hamond, collector, 2 specimens (BMNH 1969:99, as H. lunulata by Hamond, 1966). Ireland: Blacksod Bay, Irish Fisheries, Eire sta W, shore, Sep 1905, 1 specimen (BMNH 1914.12.12.17, as H. lunulata). France: North coast, Roscoff, Grube, collector, 3 specimens (ZMB 1124, as Malmgrenia alba by Augener, 1928). St. Malo, Grube, collector, 2 specimens (ZMB 1127, as M. alba by Augener, 1928). Netherlands: East Friesland, Grube, collector, 1 specimen (ZMB 1123, as M. alba by Augener, 1928).

DESCRIPTION.—No type material available. Size given by Saint–Joseph (1888) for specimen with Arenicola marina, 25 mm long, 4 mm wide, with 36 segments. Additional specimens from Plymouth area: with A. marina, 16–29 mm long, 5–8 mm wide including setae, with 37–38 segments; with Amphitrite johnstoni, 18–42 mm long, 6–10 mm wide, with 32–38 segments; with Loimia medusa, 22–27 mm long, 9 mm wide, with 38 segments; with Golfingia elongata, 12–19 mm long, 4–5 mm wide, with 37 segments; with Notomastus latericeus, 14 mm long, 4 mm wide, with 37 segments. Specimen from Norfolk Coast, 12 mm long, 4.5 mm wide, with 37 segments.

Body elongate, flattened, tapered slightly anteriorly and posteriorly. Dorsum colorless, ventrad sometimes lightly pigmented in posterior region with one band per segment and spots medial to ventral cirri. Elytra 15 pairs, on usual segments, large, covering dorsum, round, subreniform to elongate-oval, without papillae, with transverse group of microtubercles near anterior border (absent on first pair and few or absent on posterior elytra), usually with transverse band of dark pigment in line with elytrophore and lighter mottled areas more posteriorly (Figure 18B–E). Elytra on specimens with Arenicola light to dark brownish on exposed part, with indications of darker band on anterior part.

Bilobed prostomium truncate anteriorly, without distinct peaks; eyes rather small, anterior pair anterolateral, larger than posterodorsal pair; ceratophore of median antenna large, in anterior notch, with style longer than prostomium; ceratophores of lateral antennae inserted terminoventrally, with styles short, subulate; antennae with minute papillae; palps stout, tapered, variable in length, minutely papillate; tentaculophores without setae, with dorsal and ventral tentacular cirri similar to median antenna (Figure 18A; Saint-Joseph, 1888, pl. 6: fig. 22). Biramous parapodia of segment 2 with long ventral buccal cirri, similar to tentacular cirri; notosetae forming thick bundle, similar to following parapodia; neurosetae more slender than following, upper and lower ones with entire bulbous tips, middle ones with bulbous bifid tips (Figure 18A,F,G). Extended pharynx with 9 pairs of border papillae and 2 pairs of reddish amber-color jaws.

Notopodium short, rounded, with projecting acicular lobe on lower side; neuropodium with longer presetal acicular lobe, diagonally truncate on lower side and thick conical supraacicular process; postsetal lobe shorter, rounded (Figure 19A,B). Notosetae numerous, forming very thick bundle of several lengths, not extending beyond tips of neuropodia, about as stout as neurosetae, with spinose rows, tapering to short bare blunt tips (Figure 19C). Neurosetae numerous, forming fan-shape bundle; upper ones with longer spinose regions and slender bifid tips; middle ones with shorter spinose regions and larger bifid tips; lower ones with minute secondary tooth or sometimes entire (Figure 19D; Saint-Joseph, 1888, pl. 6: fig. 23). Cirrophores of dorsal cirri long, cylindrical, wider basally, with style extending slightly beyond tips of neurosetae, with scattered short clavate papillae and filamentous tip; dorsal tubercles nodular; ventral cirri short, subulate, with minute papillae (Figure 19B). Pygidium medial to last pair of parapodia, with pair of long anal cirri.

BIOLOGY.—Malmgreniella arenicolae is commensal with sipunculids and polychaetes: arenicolids, capitellids, and terebellids. It was found in the burrows of Arenicola marina (Linnaeus) in Norway by C. Dons (ZMT), in Dinard, France, by Saint-Joseph (1888), and by Spooner (in Plymouth Marine Fauna, 1957) in the Plymouth area (as Harmothoe lunulata var. A). Also in the Plymouth area the species was collected by Spooner (1957) in the burrows of the capitellid Notomastus latericeus Sars and the sipunculids Golfingia elongata (Keferstein) and G. vulgaris (Blainville) (as var. G), and in the tubes of the terebellids Amphitrite johnstoni Malmgren and Loimia medusa (Savigny) (as var. J). On the Norfolk Coast, the species was found in the tubes of Lanice conchylega (Pallas) by Hamond (1966) (as H. lunulata).

FIGURE 18.—Malmgreniella arenicolae, specimen from Yealm Estuary, with Amphitrite johnstoni, as Harmothoe lunulata var. J (USNM 55051): A, dorsal view of anterior end, pharynx extended (only basal part shown); B, right 1st elytron from segment 2; C, right 2nd elytron from segment 4; D, right 10th elytron from segment 19, with detail of microtubercles; E, right 15th elytron from segment 32; F, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; G, upper, middle, and lower neurosetae from same, with detail of tips. (Scales = 0.5 mm for A; 1.0 mm for B–E; 0.5 mm for F; 0.1 mm for G.)

FIGURE 19.—Malmgreniella arenicolae, specimen from Yealm Estuary, with Amphitrite johnstoni, as Harmothoe lunulata var. J. (USNM 55051): A, right middle elytrigerous parapodium, anterior view, acicula dotted; B, right middle cirrigerous parapodium, posterior view; C, short and long notosetae; D, upper, middle, and lower neurosetae. (Scales = 0.5 mm for A,B; 0.1 mm for C,D.)

DISTRIBUTION.—Northeastern Atlantic Ocean, Spitzbergen, Norwegian Sea to France. Intertidal.

Malmgreniella castanea (McIntosh, 1876), new combination

FIGURE 20

Eunoa n. sp.—McIntosh, 1868:337, 338.

Malmgrenia castanea McIntosh, 1876a:376, pl. 67: figs. 15–19; pl. 68: fig. 15; 1900:379, pl. 28: fig. 15; pl. 30: fig. 5; pl. 33: fig. 10; pl. 40: figs. 23–26.—Allen, 1915:612.—Fauvel, 1923:48, fig. 17h–m.—Ditlevsen, 1929:5.—Støp–Bowitz, 1948:7, fig. 3.—Plymouth Marine Fauna, 1957:110.—Picard, 1965:53 [footnote].—Intes and Le Loeuff, 1975:273.—Kudenov, 1975b:79.

Laenilla castanea.—Giard, 1886:3, figs.

Polynoe (Malmgrenia) castanea.—Hornell, 1891:235.

Harmothoe castanea.—Saint-Joseph, 1898:236.—Ditlevsen, 1917:27, figs. 9, 10.—Tebble and Chambers, 1982:47, figs. 15, 44, 45.—Hanley, 1987:151, fig. 3E.

Malmgrenia lunulata.—Pettibone, 1953:25 [part; not Delle Chiaje, 1830].

Harmothoe lunulata.—Pérès, 1954:92.—Bellan, 1964:27 [part].—Picard, 1965:53 [part]. [Not Delle Chiaje, 1830.]

MATERIAL EXAMINED.—NORTHEASTERN ATLANTIC OCEAN: Norwegian Sea: 50°57′N, 10°46′W, 184 and 200 m, Michael Sars sta 96, 27 Jul 1910, 2 specimens (ZMUB 29582, 41504, identified by Støp-Bowitz, 1948). Shetland Islands: 25 mi NNE of North Unst, 155 m, shell and sand, 1868, on Spatangus purpureus O.F. Müller, G. Jeffreys, collector, 6 syntypes of Malmgrenia castanea (BMNH 1921.5.1.507; ZMB 1162; USNM 54714). SW Ireland: 50 mi west of Valentia, 146–201 m, G. Jeffreys, collector, 2 syntypes of Malmgrenia castanea (BMNH 1921.5.1.509). Channel Islands: Off St. Peter Port, Guernsey, 9–13 m, 1868, syntype of Malmgrenia castanea (BMNH 1921.5.1.508). English Channel: 50°21′N, 03°01′W, 53 m, Manihine sta 69, 30 Aug 1949, 1 specimen (BMNH 1950.6.6.1). England: Plymouth area, Eddystone, on Spatangus purpureus, shell and gravel, 26 Mar 1953, 24 Mar 1960, G.M. Spooner, collector, 2 specimens (LCHP–S-21). Plymouth area, P. Gibbs, collector, 2 specimens (LCHP–G-3; USNM 55047).

MEDITERRANEAN SEA: France: Marseille, 8 m, on oral surface of Spatangus purpureus, in Amphioxus sand, G. Bellan, collector, 1 specimen (USNM 59929).

DESCRIPTION.—Nine syntypes mostly fragmented. Length of complete syntype 15.5 mm, width including setae 6 mm, with 38 segments. Length of 2 specimens from Norwegian Sea 20–21 mm, width 7–8 mm, with 38 segments. Length of 4 specimens from Plymouth area 8–15 mm, width 2.5–8 mm, with 34–39 segments. Body elongated, flattened, tapering slightly anteriorly and posteriorly. Dorsum, exposed part of elytra, and dorsal cirri with light to dark brownish pigmentation. Elytra 15 pairs, on usual segments, large, covering dorsum, round to subreniform, without papillae, with group of microtubercles on anterior part but absent on first pair and some posterior elytra (Figure 20B–E; McIntosh, 1900, pl. 33: fig. 10).

Bilobed prostomium truncate anteriorly, without distinct peaks; eyes moderate in size, anterior pair slightly anterior to greatest width, slightly larger than posterodorsal pair; ceratophore of median antenna in anterior notch, with style longer than prostomium; ceratophores of lateral antennae inserted terminoventrally, with styles short, subulate; palps stout, tapered; tentaculophores without setae, dorsal and ventral tentacular cirri similar to median antenna (Figure 20A; McIntosh, 1900, pl. 28: fig. 15; Støp-Bowitz, 1948, fig. 3). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri; notosetae similar to following notosetae; neurosetae more slender than those following, with entire bulbous tips (Figure 20A,F,G). Extended pharynx with 9 pairs of border papillae and 2 pairs of jaws.

Biramous parapodia with translucent, pale setae; notopodium shorter than neuropodium, subconical, with projecting acicular lobe on lower side; neuropodium with subconical presetal acicular lobe with thick supraacicular process and shorter rounded postsetal lobe (Figure 20H,I; McIntosh, 1900, pl. 30: fig. 5). Notosetae moderate in number, short to long, slightly stouter than neurosetae, slightly curved, tapering toward tips, with faint spinose rows (Figure 20J; McIntosh, 1876a, pl. 67: fig. 15; 1900, pl. 40: fig. 23). Neurosetae moderate in number, with faint spinose regions, slightly hooked tips, some with very small secondary tooth (Figure 20K; McIntosh, 1876a, pl. 67: figs. 16–19, 1900; pl. 40: figs. 24–26). Cirrophores of dorsal cirri cylindrical, wider basally, with styles smooth, tapered distally, extending to tips of neurosetae or beyond; dorsal tubercles bulbous; ventral cirri short, subulate (Figure 20H; McIntosh, 1900, pl. 30: fig. 5). Pygidium medial to last pair of small parapodia, with pair of long tapered anal cirri.

BIOLOGY.—Malmgreniella castanea is commensal with spatangid echinoids. It has been found clinging to the test near the mouth of the burrowing spatangid Spatangus purpureus O.F. Müller.

DISTRIBUTION.—Northeast Atlantic Ocean, Norwegian Sea, Shetland Islands to Mediterranean Sea. In 8–878 meters (see McIntosh, 1876a).

Malmgreniella marphysae (McIntosh, 1876), new combination

FIGURE 21

Harmothoe marphysae McIntosh, 1876a:384, pl. 69: figs. 11–15; pl. 70: fig. 18; 1900:339, pl. 27: fig. 11; pl. 29: fig. 15; pl. 32: fig. 13; pl. 39: figs. 7–11.—Malaquin, 1890:283.—Orton, 1923:861.—Plymouth Marine Fauna, 1957:112.—Tebble and Chambers, 1982:43, figs. 13d,e, 40, 41a,b.

Harmothoe marphysae var. watsoni McIntosh, 1919:163; 1923:521, pl. 137: figs. 1, 1a, 1b.—Fauvel, 1927:406. [New synonymy.]

Harmothoe lunulata var. marphysae.—Fauvel, 1923:72, fig. 26i.—Bellan, 1962:89.

Harmothoe lunulata.—Pettibone, 1953:25 [part; not Delle Chiaje, 1830].

MATERIAL EXAMINED.—NORTHEASTERN ATLANTIC OCEAN: Channel Islands: English Channel, Guernsey, Jul, in galleries of M. sanguinea, lectotype (BMNH 1921.5.1.443). England: Plymouth Sound, in galleries of M. sanguinea (Montagu), rock crevices: off Mount Edgcume, 14 Apr 1961, G.M. Spooner, collector, 4 specimens (LCHP–S-10; USNM 55064); West Lose, 20 Mar 1969, P.E. Gibbs, collector, 1 specimen (LCHP–G-7). North Wales: In tube of Lagis koreni Maslmgren, Arnold Watson, collector, holotype of Harmothoe marphysae var. watsoni (BMNH 1921.5.1.485).

REMARKS.—The type material of Harmothoe marphysae in The Natural History Museum, London (BMNH 1921.5.1.443), consists of two syntypes. The larger specimen, a female filled with eggs, has 31 segments of which 22 are normal-size segments and 9 smaller, regenerating segments. It is 11 mm long, 4.5 mm wide including setae, with two elytra free in the vial. It agrees with the original description and is selected as the lectotype. The smaller syntype is an anterior fragment of 18 segments and belongs to another species. The holotype of H. marphysae var. watsoni in The Natural History Museum, London (BMNH 1921.5.1.485), is a small specimen, incomplete posteriorly, with 22 segments, 4 mm long, and 2.5 mm wide; elytra are now missing (described and figured by Mcintosh). It is considered to be a young specimen of M. marphysae.

FIGURE 20.—Malmgreniella castanea, syntype from Channel Islands (BMNH 1921.5.1.508): A, dorsal view of anterior end, left dorsal and ventral tentacular cirri missing; B, right 1st elytron from segment 2; C, right 2nd elytron from segment 4; D, right 5th elytron from segment 11, with detail of microtubercles; E, right middle elytron; F, right elytrigerous parapodium from segment 2, anterior view, acicula dotted, upper neurosetae mostly broken; G, upper and lower neurosetae from same, with detail of tips; H, right cirrigerous parapodium from segment 10, posterior view; I, right elytrigerous parapodium from segment 11, anterior view, acicula dotted; J, short and long notosetae; K, upper, middle, and lower neurosetae, with detail of tips. (Scales = 0.5 mm for A; 0.5 mm for B–E; 0.2 mm for F,H,I; 0.1 mm for G,J,K.)

DESCRIPTION.—Length of 2 complete specimens from Plymouth area 14–19 mm, width 5–6 mm, with 38 segments. Body flattened, tapering slightly anteriorly and more so posteriorly. Elytra 15 pairs, on usual segments, oval, subreniform to quadrate, smooth, pellucid, some with small group of microtubercles on anterior part and scattered micropapillae on surface; most of elytra with 1–2 longitudinal thickened ridges, longer one near inner border and smaller one near place of attachment to elytrophore; some elytra showing spot over elytrophore and transverse band of pigmentation (Figure 21B–E; McIntosh, 1900, pl. 32: fig. 13).

Bilobed prostomium with anterior lobes wide, subtriangular, without distinct peaks; eyes small, anterior pair anterolateral, anterior to greatest width of prostomium, posterior pair posterodorsal; ceratophore of median antenna in anterior notch, with style papillate, longer than prostomium; ceratophores of lateral antennae inserted terminoventrally, with short, papillate, subulate styles; palps stout, tapered; tentaculophores with 0–2 setae, dorsal and ventral tentacular cirri similar to median antenna (Figure 21A; McIntosh, 1900, pl. 27: fig. 11; Tebble and Chambers, 1982, fig. 40). Segment 2 with first pair of elytrophores, biramous parapodia, and long ventral cirri similar to tentacular cirri. Extended pharynx with 9 pairs of border papillae and 2 pairs of jaws.

Biramous parapodia with notopodia much shorter than neuropodia; notopodium small, subconical, with projecting acicular lobe on lower side; neuropodium large, with presetal acicular lobe diagonally truncate, with deep acicular notch, forming rounded longer upper and shorter lower processes; postsetal lobe shorter, rounded (Figure 21F,G). Notosetae relatively few, extending only slightly beyond tips of notopodia, with spinose rows, short ones with blunt tips, longer ones tapering to fine tips (Figure 21F,H; McIntosh, 1876a, pl. 69: fig. 11, pl. 70: fig. 18; 1900, pl. 39: fig. 7; Tebble and Chambers, 1982, fig. 13d,e). Neurosetae numerous, forming fan-shape bundle; upper ones with longer spinose regions, tapering to slender entire tips, some lower upper ones with bifid tips; middle neurosetae with shorter spinose regions, and small secondary tooth; lower ones with entire tips (Figure 21I; McIntosh, 1876a, pl. 69: figs. 12–15; 1900, pl. 39: figs. 8–11; Tebble and Chambers, 1982, fig. 41b). Cirrophores of dorsal cirri cylindrical, wider basally, with glandular area; styles extending to tips of neurosetae or beyond, with scattered short papillae; dorsal tubercles small, nodular; ventral cirri short, subulate, with few short papillae (Figure 21G; McIntosh, 1900, pl. 29: fig. 16).

REMARKS ON YOUNG HOLOTYPE OF Harmothoe marphysae var. watsoni, REFERRED TO Malmgreniella marphysae (Figure 21J–N; McIntosh, 1923, pl. 137: fig. 1).—The young specimen agrees in most respects with the adults. The elytra lack the longitudinal thickened ridges found on some elytra (McIntosh, 1923, pl. 137: fig. la). The anterior end is similar (Figure 21J). The parapodia are similar in general outline but lack the deep acicular notch on the neuropodial presetal acicular lobes (Figure 21K,L; McIntosh, 1923, pl. 137: fig. 1). The notosetae are of the same two types: short ones with a blunt tip and longer ones tapering to a fine tip (Figure 21M); the neurosetae have mostly a bifid tip (Figure 21N; McIntosh, 1923, pl. 137: fig. 1b), instead of the lower ones having an entire tip.

BIOLOGY.—Malmgreniella marphysae has been found in the galleries of the eunicid polychaete Marphysa sanguinea (Montagu) and in the tube of the pectinariid polychaete Lagis koreni Malmgren.

DISTRIBUTION.—Northeastern Atlantic Ocean, Irish Sea, English Channel to Northwest France.

Malmgreniella mcintoshi (Tebble and Chambers, 1982), new combination

FIGURE 22

Lagisca Jeffreysii McIntosh, 1900:305, pl. 27: fig. 10; pl. 32: fig. 7; pl. 38: figs. 7–9; pl. 42: fig. 30 [primary homonym; not Lagisca jeffreysi McIntosh, 1876b].

Harmothoe haliaeti.—Fauvel, 1914:57, pl. 3: fig. 17; 1923:57, fig. 19f–h. [Not McIntosh, 1876b.]

Harmothoe jeffreysi.—Ditlevsen, 1917:9 [not McIntosh, 1876b].

Malmgrenia jeffreysi.—Augener, 1925:108, figs. 1, la [not McIntosh, 1876b].

Harmothoe mcintoshi Tebble and Chambers, 1982:53, figs. 17a,b, 50.

MATERIAL EXAMINED.—NORTHEASTERN ATLANTIC OCEAN: Shetland Islands: 9 mi off Balta, 110 m, G. Jeffreys, collector, 1868, holotype of Lagisca Jeffreysii McIntosh, 1900 (BMNH 1921.5.1.310). Sweden: Gunnarskar, Kristineberg Zoological Station, 20 m, shelly sand, A. Eliason, collector, 13 May 1965, 1 specimen (USNM 43230, as Harmothoe haliaeti by Eliason). Ireland: Galway, 1 specimen (BMNH 1921.5.1.288, mixed with Lagisca floccosa by McIntosh).

REMARKS.—McIntosh (1876b:397) described Lagisca jeffreysi from specimens collected off the west coast of Iceland in 298 meters and in the Channel slope in 1262 meters. Later, McIntosh (1900:307) referred the above species to L. extenuata (Grube) and proceeded to describe a new species from the Shetland Islands in 110 meters as Lagisca jeffreysii. The latter species, being a homonym, was given a new name by Tebble and Chambers (1982:53), Harmothoe mcintoshi.

FIGURE 21.—Malmgreniella marphysae, A, F–I, lectotype of Harmothoe marphysae (BMNH 1921.5.1.443); B–E, specimen from Plymouth area (USNM 55064); J–N, holotype of Harmothoe marphysae var. watsoni (BMNH 1921.5.1.485): A, dorsal view of anterior end; styles of median antenna, left dorsal and ventral and right dorsal tentacular cirri, palps, and dorsal cirri from segment 3 missing; right parapodia of segment 2 and 3 had been cut off. B, right 1st elytron from segment 2, with detail of micropapillae, C, right 2nd elytron from segment 4, with thickened ridge on inner side and detail of microtubercles. D, right middle elytron, with 2 thickened ridges. E, right 15th elytron from segment 32. F, right elytrigerous parapodium, anterior view, acicula dotted; most of lower neurosetae broken. G, right cirrigerous parapodium, posterior view. H, long and short notosetae. I, upper, middle, and lower neurosetae. J, dorsal view of anterior end; styles of median antenna, tentacular cirri, and dorsal cirri of segment 3 missing; right parapodia of segments 2 and 3 had been cut off. K, right elytrigerous parapodium, anterior view, acicula dotted. L, right cirrigerous parapodium, posterior view, style of dorsal cirrus missing. M, long and short notosetae; N, upper, middle, and lower neurosetae. (Scales = 0.5 mm for A,J; 0.5 mm for B–E; 0.2 mm for F,G; 0.1 mm for H,I,M,N; 0.2 mm for K,L.)

McIntosh (1876a:384) described a posterior fragment without elytra from the Minch, Scotland, under the name Harmothoe haliaeti. Only the setae were described and figured (pl. 69: figs. 7–10). As pointed out by Tebble and Chambers (1982:70), who examined the fragmented holotype in The Natural History Museum, London, the species must be considered to be indeterminable. Harmothoe haliaeti has been used widely by Fauvel (1914, 1923) and others. At least some of these records, including those of Fauvel, agree with Malmgreniella mcintoshi as indicated in the above synonymy.

DESCRIPTION.—Holotype with 36 segments, posterior 4 segments small, regenerating, 18 mm long, 6.5 mm wide including setae. Incomplete specimen from Ireland with 28 segments, 13 mm long, and 5 mm wide. Specimen from Sweden with 36 segments, 9 mm long, and 3 mm wide. Specimens reported by Tebble and Chambers (1982) from Isle of Man up to 21 mm long, 6 mm wide, and 38 segments.

Elytra 15 pairs, on usual segments, large, oval to subreniform, some with anterior fold connected to place of attachment to elytrophore, with wide band of microtubercles on anterior part, lateral border with long papillae, and some scattered micropapillae on surface (Figure 22C,D; McIntosh, 1900, pl. 32: fig. 7; Tebble and Chambers, 1982, fig. 17a).

Bilobed prostomium with anterior lobes truncate, without peaks; anterior pair of eyes anterior to greatest width, posterior pair posterodorsal; median antenna with large ceratophore in anterior notch, with papillate style longer than prostomium; ceratophore of lateral antennae inserted terminoventrally, with short subulate papillate styles; palps long, tapered; tentaculophores with 0–2 setae on inner side, with dorsal and ventral tentacular cirri similar to median antenna (Figure 22A,B; McIntosh, 1900, pl. 27: fig. 10; Tebble and Chambers, 1982, fig. 50). Segment 2 with first pair of large elytrophores, biramous parapodia, and long papillate ventral buccal cirri similar to tentacular cirri (Figure 22A,B); notosetae similar to following notosetae; neurosetae more slender than those following, upper ones with long secondary tooth, lower ones ending in entire, slightly bulbous tip.

Biramous parapodium with notopodium shorter and smaller than neuropodium, subconical with projecting acicular lobe on lower side; neuropodium with subconical presetal acicular lobe, without supraacicular process, and shorter, rounded postsetal lobe (Figure 22E,F; McIntosh, 1900, pl. 42: fig. 30). Notosetae numerous, forming radiating bundle, short to long, with distinct spinose rows and short bare tapered tips (Figure 22G; McIntosh, 1900, pl. 38: fig. 7). Upper neurosetae with longer spinose regions, with tips entire or with small secondary tooth; middle neurosetae with long secondary tooth; lower neurosetae with shorter spinose regions and entire tips (Figure 22M; McIntosh, 1900, pl. 38: figs. 8, 9). Dorsal cirri with cylindrical cirriphores, wider basally, with glandular areas; styles papillate, extending beyond neurosetae; dorsal tubercles nodular; ventral cirri short, papillate (Figure 22F). Pygidium with pair of long anal cirri.

DISTRIBUTION.—Northeastern Atlantic Ocean, Sweden, Faeroe Islands, Shetland Islands to Ireland. In 21–128 meters (see Ditlevsen, 1917).

Malmgreniella lunulata (Delle Chiaje, 1830), new combination

FIGURE 23

Polynoe lunulata Delle Chiaje, 1830 (“1822”), pl. 79: figs. 5, 6; 1841:57, 106, pl. 144: figs. 5, 6.—Claparède, 1868:373, pl. 2: fig. 1, 1A–F.

Harmothoe lunulata.—Alaejos y Sanz, 1905:46, pl. 6: figs. 7–9; pl. 7: figs. 1–9.—Saint-Joseph, 1906:194, pl. 3: figs. 71, 72.—Fauvel, 1913:15; 1923:70, fig. 26a–h [part]. [Not Day, 1962.—Monro, 1937.—Katzmann. 1983.]

Malmgrenia lunulata.—Pettibone, 1953:25 [part].

REMARKS.—No type material is available for Polynoe lunulata Delle Chiaje from the Mediterranean, which was incompletely described and figured by Delle Chiaje and Claparède. The species has been widely reported as Harmothoe lunulata (Delle Chiaje) by McIntosh (1900), Fauvel (1923), and many others; and as Malmgrenia lunulata (Delle Chiaje) by Pettibone (1953) and others. As a northern species it has been reported from the eastern Atlantic and eastern Pacific, living commensally with a variety of hosts, including echinoderms, polychaetes, and balanoglossids. Far too many species have been lumped under M. lunulata.

As a part of this study, an available specimen in the Paris Museum from off Monaco in the Mediterranean is selected for study and referred to Malmgreniella lunulata. It is supplemented by the description and figures of Saint-Joseph (1906) on two specimens from the Mediterranean. The group will need further study to tie down the various records. Some records are dealt with herein. Others will need further study.

FIGURE 22.—Malmgreniella mcintoshi, specimen from Sweden (USNM 43230): A, dorsal view of anterior end; B, ventral view of anterior end; C, left 1 st elytron from segment 2, with detail of microtubercles and micropapillae; D, left middle elytron, with detail of microtubercles, lateral papillae and micropapillae on surface; E, right elytrigerous parapodium, anterior view, acicula dotted; F, right cirrigerous parapodium, posterior view; G, short and long notosetae; H, upper, middle, and lower neurosetae. (Scales = 0.5 mm for A,B; 0.5 mm for C,D; 0.2 mm for E,F; 0.1 mm for G,H.)

MATERIAL EXAMINED.—EASTERN ATLANTIC OCEAN: Mediterranean Sea: Off Monaco, 17–18 Mar 1903, 16–30 m, collection of S.A.S. le Prince de Monaco, sta Cap d′Ail, 1 specimen (MNHN, as Harmothoe lunulata by Fauvel, 1913).

DESCRIPTION.—Specimen from off Monaco, female with large yolky eggs in body cavity, 10 mm long, 4 mm wide including setae, and 37 segments, last one very small. Female specimen from Mediterranean, described by Saint-Joseph (1906), 25 mm long, 4 mm wide, with 37 segments.

FIGURE 23.—Malmgreniella lunulata, specimen from off Monaco (MNHNP): A, dorsal view of anterior end; styles of median antenna, right dorsal and ventral and left ventral tentacular cirri, and right ventral buccal cirrus of segment 2 missing. B, left 3rd elytron from segment 5, with detail of microtubercles. C, right 6th elytron from segment 11, with detail of microtubercles. D, right elytrigerous parapodium, anterior view, acicula dotted. E, right cirrigerous parapodium, posterior view, style of dorsal cirrus missing. F, long and short notosetae. G, upper, middle, and lower neurosetae. (Scales = 0.5 mm for A; 0.5 mm for B,C; 0.2 mm for D,E; 0.1 mm for F,G.)

FIGURE 24.—Malmgreniella darbouxi, new species, holotype (USNM 74370): A, dorsal view of anterior end, style of median antenna small, regenerating, styles of left dorsal and ventral tentacular cirri missing; B, right elytron, with detail of microtubercles; C, right elytrigerous parapodium, anterior view, acicula dotted; D, right cirrigerous parapodium, posterior view; E, long and short notosetae; F, upper, middle, and lower neurosetae, with detail of tips. (Scales = 0.5 mm for A; 0.2 mm for B; 0.2 mm for C,D; 0.1 mm for E,F.)

Elytra 15 pairs, on usual segments, large, oval to subreniform, last pair elongate-oval, nearly covering small posterior segments; posterior two-thirds of elytra nearly covered with mottled areolate brownish pigmentation; numerous microtubercles on anterior part and scattered globular micropapillae on surface (Figure 23B,C. Saint-Joseph, 1906, pl. 3: fig. 71).

Bilobed prostomium with anterior lobes truncate, without peaks; anterior pair of eyes anterolateral, larger than posterodorsal pair, ceratophore of median antenna in anterior notch, with papillate style about twice as long as prostomium; ceratophores of lateral antennae inserted terminoventrally, with very short papillate styles; palps long, stout, tapered; tentaculophores lateral to prostomium, with single seta on inner side; dorsal and ventral tentacular cirri similar to median antenna; small conical facial tubercle medial to ventral bases of tentaculophores (Figure 23A; Saint-Joseph, 1906, pl. 3: fig. 72). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri lateral to ventral mouth, similar to tentacular cirri and longer than following ventral cirri (Figure 23A). Extended pharynx with 9 pairs of border papillae and 2 pairs of hooked jaws.

Biramous parapodium with smaller notopodium rounded. with projecting acicular lobe on lower side; larger neuropodium with subconical presetal acicular lobe with small digitiform supraacicular process and shorter rounded postsetal lobe (Figure 23D,E). Notosetae numerous, forming radiating bundle, of several lengths, slightly stouter than neurosetae, with close-set spinose rows continuing to very short bare tips (Figure 23F). Neurosetae numerous, forming fan-shape bundle, upper ones with longer spinose regions and tapered tips with secondary tooth; middle and lower neurosetae with shorter spinose regions and more distinctly hooked tips, all with rather long secondary tooth (Figure 23G). Dorsal cirri with cylindrical cirrophores with bulbous glandular areas on posterior sides, and papillate styles extending beyond tips of neurosetae; dorsal tubercles nodular; ventral cirri short, papillate, tapered (Figure 23E). Pygidium with pair of long papillate anal cirri.

BIOLOGY.—Specimens from Santander, Bay of Biscay, Northern Spain, were reported by Alaejos y Sanz (1905) as commensal with balanoglossids, found on compact beaches near the surface of the sand under small rocks. The polynoids were encased in the mucus produced by the host. When the burrow was disturbed, the very lively polynoid was apt to escape.

DISTRIBUTION.—Eastern Atlantic Ocean, Mediterranean Sea. Depth 16–30 meters.

Malmgreniella darbouxi, new species

FIGURE 24

Harmothoe lunulata.—Katzmann, 1983:93 [not Delle Chiaje, 1830].

MATERIAL EXAMINED.—EASTERN ATLANTIC OCEAN: Mediterranean Sea: Near Zlorin Island, Yugoslavia, middle Adriatic Sea, 20–60 m, sand and mud, 1971, W. Katzmann, collector, holotype (USNM 74370).

DESCRIPTION.—Holotype, male with sperm, 9 mm long, 3.5 mm wide including setae, 34 segments. Ventral surface of posterior segments transversely banded with dark pigmentation (suggesting commensal relationship with unknown host). Elytra 15 pairs, on usual segments, large, oval, delicate, transparent; small group of microtubercles near middle of anterior border; dark pigmentation in form of transverse band over scar of attachment to elytrophore, to medial side, and continuing posteriorly along medial side (Figure 24B).

Bilobed prostomium with anterior lobes truncate, without peaks; anterior pair of eyes just anterior to greatest width, larger than posterodorsal pair; ceratophore of median antenna in anterior notch, with style small, regenerating; ceratophores of lateral antennae inserted terminoventrally, with styles short, subulate, with short papillae; palps long, tapered; tentaculophores lateral to prostomium, with single seta on inner side; dorsal and ventral tentacular cirri longer than prostomium, with short papillae (Figure 24A). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri, similar to tentacular cirri (Figure 24A).

Biramous parapodium with smaller notopodium rounded, with projecting acicular lobe on lower side; larger neuropodium with subconical projecting presetal acicular lobe with digitiform supraacicular process and shorter, rounded postsetal lobe (Figure 24C,D). Notosetae moderate in number, slender, of several lengths, with faint spinose rows, longest ones tapering to more slender tips (Figure 24E). Neurosetae numerous, slender, about same width as notosetae; upper ones with longer spinose regions and tapered bifid tips; middle and lower ones with shorter spinose regions and more distinct hooked tips, lower ones with entire tips (Figure 24F). Dorsal cirri with cylindrical cirrophores, inflated basally, with glandular areas on upper side; styles with short papillae, tapered and extending to about tips of neurosetae; dorsal tubercles nodular, ventral cirri short, tapered, with few short papillae (Figure 24D). Pygidium rounded, with pair of long anal cirri.

ETYMOLOGY.—The species is named for J. Gaston Darboux, in recognition of his basic research on the aphroditoid polychaetes (Darboux, 1900).

DISTRIBUTION.—Eastern Atlantic Ocean, Mediterranean Sea (Adriatic Sea), in 20–60 meters.

Key to the Species of Malmgreniella from the Western Atlantic Ocean (Virginia to Florida, Caribbean Sea, Belize, Panama, Gulf of Mexico, Brazil)

Malmgreniella maccraryae, new species

FIGURE 25

Harmothoe lunulata.—Taylor, 1971:70 [not Delle Chiaje, 1830].

? Malmgrenia lunulata. variety Gardiner, 1976:90, fig. 3a–e [part; not USNM 52850 = Lepidonopsis humilis (Augener)].

Malmgreniella sp. B.—Weston, 1984:21–20, figs. 21–15, 16a–g.

Malmgrenia lunulata.—Fox and Ruppert, 1985:127, 194, 254 [not Delle Chiaje, 1830].

MATERIAL EXAMINED.—NORTHWESTERN ATLANTIC OCEAN: North Carolina: Banks Channel, Wrightsville Sound, 1967, taken from brittle star, A.B. McCrary, collector, paratype (USNM 55065). Wrightsville Beach, intertidal, commensal on brittle star Amphipholis gracillima (Stimpson), Apr 1972, Mar 1973, S.L. Gardiner and T. Fox, collectors, holotype (USNM 52050) and 2 paratypes (USNM 52849, 55066, as ?Malmgrenia lunulata, variety by Gardiner, 1976). Georgia: Off Georgia, 31°55′N, 80°51′W, 8 m, Texas Instruments, R/V Pierce sta 4A, 20 Feb 1977, 3 specimens (USNM 59169). Florida: Off Florida, 29°27′N, 81°03′W, 20 m, R/V Pierce, sta 7A, 4 Mar 1977, 3 specimens (USNM 59171). Off Virginia Key: near Miami, 1 m, on brittle star Amphioplus sepultus Hendler, 23 Feb 1970, G. Hendler, collector, 2 paratypes (USNM 55067); 26 Feb 1970, 2 specimens (USNM 133560; LACM).

FIGURE 25.—Malmgreniella maccraryae, new species, holotype (USNM 52050): A, dorsal view of anterior end; B, right 1 st elytron, with detail of micropapillae; C, right middle elytron; D, right elytrigerous parapodium, anterior view, acicula dotted: E, right cirrigerous parapodium, posterior view; F, long and short notosetae; G, supraacicular neurosetae, with detail of tips; H, subacicular neurosetae. (Scales = 0.5 mm for A; 0.2 mm for B,C; 0.2 mm for D,E; 0.1 mm for F–H.)

GULF OF MEXICO: Florida: Tampa Bay, J.L. Taylor, collector, 7 paratypes (USNM 45513, as Harmothoe lunulata by Taylor, 1971). Hillsborough Bay, Tampa, sand and shell, 7 May 1975, S.L. Santos, collector, 9 paratypes (USNM 55068). Mississippi-Alabama-Florida (MAFLA) sta IV-2419, off Apalachicola River, Feb 1978, 29°47′N, 84°05′W, 10 m, medium-fine sand, 2 specimens (USNM 89685). Alabama: Mouth of Mobil Bay, 30°14′N, 88°03′W, 6.4 m, Mobile Oil sta Mc 151A,D, Jul 1978, 40 specimens (USNM 70075–6). About 3 miles SE of Dauphin Island, 30°11′N, 88°07′W, 13 m, Continental Shelf Assoc. sta 14, 18 Sep 1981, 6 specimens (USNM 71156). Texas: Bay City-Freeport, inner shelf, 28°53′N, 95°15′W, 15 m, T.R. Calnan, collector, 2 specimens (USNM 68008), as Malmgreniella sp. B. by Weston, 1984: Southwest Florida (SOFLA) sta 25B, Nov 1980, 24°47′N, 82°13′W, 24 m, silt-clay, 2 specimens (USNM 89684). South Texas Outer Continental Shelf (STOCS) sta II-1, off Port Aransas, Fall 1976, 27°40′N, 96°59′W, 22 m, sandy silty clay, 1 specimen (USNM 89688); sta IV-4, off Port Isabel, Fall 1976, 1977, 26°10′N, 97°08′W, 15 m, sand, 2 specimens (USNM 89686–7).

DESCRIPTION.—Holotype 7 mm long, 2.5 mm wide including setae, with 33 segments and 15 pairs of elytra. Paratype from Wrightsville Sound (USNM 55065, male with sperm) 5 mm long, 2.5 mm wide, with 31 segments and 14 pairs of elytra. Paratypes from Tampa Bay (USNM 45513) 4–6 mm long, 2–3 mm wide, with 27–34 segments and 13–15 pairs of elytra. Body short, depressed, tapering slightly anteriorly and posteriorly. Dorsum with transverse dark bands on posterior third of body; ventrum with wide longitudinal stripe along body or confined to posterior segments. Elytra up to 15 pairs, some sexually mature males with fewer than 15 pairs and fewer segments. Elytra oval, without tubercles, with scattered micropapillae along lateral and posterior borders and few on surface, with dark pigment spots on area of attachment to elytophores, on larger medial semilunar areas, and sometimes additional scattered spots (Figure 25B,C; Gardiner, 1976, fig. 3a,e; Weston, 1984, fig. 21–16a).

Bilobed prostomium with anterior lobes rounded to subtriangular, without distinct cephalic peaks; anterior pair of eyes larger than posterior pair, located just anterior to widest part of prostomium, median antenna with ceratophore in anterior notch, with rather short, subulate style; ceratophores of lateral antennae inserted ventrally, meeting midventrally, with styles about half as long as median antenna; palps long, smooth, tapered; tentaculophores each with 1–2 setae on inner side and pair of dorsal and ventral tentacular cirri, longer than median antenna and shorter than palps; antennae and tentacular cirri minutely papillated; light, net-like pigmentation on prostomium, tentaculophores, bases of styles of median antenna, and tentacular cirri (Figure 25A; Weston, 1984, fig. 21–16b). Segment 2 with biramous parapodia and long ventral buccal cirri similar to tentacular cirri. Pharynx with 9 pairs of border papillae and 2 pairs of hooked jaws.

Notopodium short, rounded, with projecting acicular lobe on lower side; neuropodium with subconical presetal acicular lobe, thick, digitiform supraacicular process, and shorter, rounded postsetal lobe (Figure 25D,E; Gardiner, 1976, fig. 3d). Notosetae numerous, forming radiating bundle, slightly stouter than neurosetae, short to long, with close-set spines along curved borders and tapered blunt tips (Figure 25F; Weston 1984, fig. 21–16d). Neurosetae moderate in number, forming fan-shape bundle; supraacicular neurosetae with longer spinose regions and secondary tooth rather far removed from slightly hooked tips (Figure 25G; Weston, 1984, fig. 21–16e,f); subacicular neurosetae with shorter spinose regions and bare, slightly hooked tips (Figure 25H; Weston, 1984, fig. 21–16g); neurosetae of segment 2 more slender than following, upper ones with bifid tips, middle and lower ones with slender, slightly clubbed, entire tips. Cirrophores of dorsal cirri cylindrical, wider basally, with papillate styles extending beyond tips of neurosetae; dorsal tubercles nodular; ventral cirri short, subulate, papillate, extending to tip of neuropodial postsetal lobe (Figure 25E). Pygidium medial to posterior, small parapodia, with pair of long anal cirri.

BIOLOGY.—Malmgreniella maccraryae was collected intertidally in Wrightsville Sound from a ophiuroid by A.B. McCrary and from the burrowing ophiuroid Amphipholis gracillima (Stimpson) by S.L. Gardiner and T. Fox. From Virginia Key near Miami, the species was collected subtidally on Amphioplus sepultus Hendler by G. Hendler.

ETYMOLOGY.—The species is named for Anne B. McCrary, who first collected one of the paratypes and who first noted its association with a brittle star.

DISTRIBUTION.—Northwestern Atlantic Ocean from North Carolina to southern Florida and Gulf of Mexico. Intertidal to 34 meters (see Weston, 1984).

Malmgreniella taylori, new species

FIGURES 26, 27

Hermadion sp.—Behre, 1950:14.

Polynoid B.—Taylor, 1971:89, fig. 2D–I.

Harmothoe, sp.—Wass, 1972:116.—Boesch, 1977:254.

Harmothoe lunulata.—Dauer, 1980:487 [not Delle Chiaje, 1830].

Malmgreniella sp. A.—Weston, 1984:21–20, figs. 21–13, 14a–g.

MATERIAL EXAMINED.—NORTHWESTERN ATLANTIC OCEAN: Virginia: York River, 9 m, mud, detritus, Nov-Dec 1960, M. Wass, collector, 11 specimens (USNM 55088). York River, mud, Jan-Mar 1961, M. Wass, collector, 16 specimens (USNM 55089). Rappahanock Shoals Channel, silty clay, 20 Nov 1963, M. Wass, collector, 1 specimen (USNM 55090). Chesapeake Bay and York River, 2–13 m, silty bottom, 1964, M. Wass, collector, 15 specimens (USNM 55091, as Harmothoe sp. by Wass, 1972, and Boesch, 1977). York River, 9 m, silty sand, on brittle star Amphiodia atra (Stimpson), Feb 1970, D. Boesch, collector, 6 paratypes (USNM 55078). Southern portion of Chesapeake Bay, 21 m, May 1976, D. Weston, collector, 1 specimen (USNM 55092). North Carolina: Wrightsville Sound, 23 Jun 1967, muddy sand, on oral disc of brittle star Amphiopholis gracillima (Stimpson), A.B. McCrary, collector, paratype (USNM 55076). Georgia: Sapelo Island, 31°23′N, 81°12′W, about 7 m, Apr 1970, A.S. Leiper, collector, 5 paratypes (USNM 55077).

GULF OF MEXICO: Florida: Tampa Bay, 1963, J.L. Taylor, collector, holotype (USNM 55073) and 5 paratypes (USNM 55074; LACM; as Polynoid B by Taylor, 1971). Hillsborough Bay, Tampa Bay, sand and shells, 7 May 1975, S.L. Santos, collector, 11 paratypes (USNM 55075). Alabama: Mouth of Mobile Bay, 30° 14′N, 88°03′W, Mobile Oil sta 051A, 154E,F, 6.1–8.2 m, sandy and grey mud, Jul-Aug 1978, 10 specimens (USNM 70077-8). About 3 miles SE of Dauphin Island, 30°11′N, 88°07′W, 13 m, Continental Shelf Assoc. sta 14, 18 Sep 1981, 1 specimen (USNM 71738). Louisiana: Grande Isle, 1934, 1 specimen (USNM 21379, as Hermadion sp. by Behre, 1950). Texas: Corpus Christi Bay, 27°45′ to 57′N, 97°00′ to 20′W, shelly clay and slightly shelly with fine sandy mud, sta 122, 127, 141, 142, 147, 152, Jun 1974 to Apr 1975, J.S. Holland and N.J. Maciolek, collectors, 202 specimens (USNM 55079-87). South Texas Outer Continental Shelf (STOCS) sta III-4, Spr 1976, 26°58′N, 97°20′W, 15 m, sand, 1 specimen (USNM 89682, as Malmgreniella sp. A by Weston, 1984). Off Texas, IXTOC Oil Spill sta N-40, 5 Dec 1979, 28°14′N, 96°29′W, 10 m, sand-silt-clay, 1 specimen (USNM 89683).

FIGURE 26.—Malmgreniella taylori, new species, holotype (USNM 55073): A, dorsal view of anterior end; B, right 1st elytron from segment 2, with detail of papillae; C, right 2nd elytron from segment 4; D, right middle elytron, with detail of papillae; E, right elytrigerous parapodium, anterior view, acicula dotted; F, right cirrigerous parapodium, posterior view, with dorsal view of cirrophore of dorsal cirrus and detail of papillae; G, long and short notosetae; H, lower and middle neurosetae; I, upper neuroseta. (Scales = 0.5 mm for A; 0.5 mm for B–D; 0.2 mm for E,F; 0.1 mm for G–I.)

SOUTHWESTERN ATLANTIC OCEAN: Brazil: Bay of Flamengo, São Paulo, 23°27′S, 45°06′W, 12 m, Jan 1963, L. Forneris, collector, 15 specimens (USNM 55093).

DESCRIPTION.—Holotype from Tampa Bay (USNM 55073) 9 mm long, 4 mm wide including setae, with 33 segments, and 15 pairs of elytra; three paratypes (USNM 55075) 6–7.5 mm long, 3 mm wide, with 31–32 segments and 14–15 pairs of elytra. Paratype from Wrightsville Sound (USNM 55076) 6 mm long, 2.5 mm wide, with 31 segments and 14 pairs of elytra. Two paratypes from Sapelo Island (USNM 55077) 10 mm long, 3.5–4 mm wide, with 34–35 segments and 15 pairs of elytra. Three paratypes from York River (USNM 55078) 5–9 mm long, 2–4 mm wide, with 29–32 segments and 14–15 pairs of elytra. Larger specimens from Corpus Christi Bay (USNM 55083) 5.5–7 mm long, 2.5–3 mm wide, with 29–30 segments and 14 pairs of elytra; small specimens 1.5–3 mm long, 1–2 mm wide, with 17–24 segments and 8–12 pairs of elytra. Specimens from Brazil (USNM 55093) 6–10 mm long, 3–5.5 mm wide, with 33–35 segments and 15 pairs of elytra.

Body short, flattened, tapering slightly anteriorly and posteriorly, colorless (rose color when living, according to M. Wass, in litt.), sometimes with greyish pigment on ventral side of posterior segments, with segments up to 35. Elytra up to 15 pairs, round, subreniform to oval, smooth, delicate, with prominent “veins,” without tubercles, with marginal fringe of clavate papillae on lateral and posterior borders (all around on first pair), without prominent color pattern but with opaque, denser area around place of attachment to elytrophore, sometimes faintly pigmented in this area and on additional crescent-shape area more medially (Figures 26B–D, 27A–D; Taylor, 1971, fig. 2E; Weston, 1984, fig. 21–14a). Bilobed prostomium with truncate anterior lobes and 2 pairs of minute eyes, anterior pair anterior to greatest width of prostomium and larger than posterior pair, ceratophore of median antenna in anterior notch, with style about as long as prostomium; ceratophores of lateral antennae inserted terminoventrally, with short subulate styles; palps stout, tapered, longer than median antenna; tentaculophores each with single seta on inner side and pair of dorsal and ventral tentacular cirri, similar to median antenna; styles of antennae and tentacular cirri with minute papillae (Figure 26A; Taylor, 1971, fig. 2D; Weston, 1984, fig. 21–14b). Segment 2 with first pair of elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri.

Parapodium with short, rounded notopodium with projecting acicular lobe on lower side; longer neuropodium with subconical presetal acicular lobe with bulbous supraacicular process, with shorter rounded postsetal lobe (Figure 26E,F). Notosetae numerous, forming radiating bundle, short to long, about as thick as neurosetae, tapering to blunt tips, with very faint spinose rows (Figure 26G; Weston, 1984, fig. 21–14d). Neurosetae forming fan-shape bundle, upper ones with longer spinose regions and more prominent spines, tapering to short, bare, entire tips (Figure 26I; Weston, 1984, fig. 21–14e); middle and lower neurosetae with shorter spinose regions and long, bare, slightly hooked tips (Figure 26H; Weston, 1984, fig. 21–14f,g). Cirrophores of dorsal cirri cylindrical, with bilobed glandular areas on anterior and posterior sides; styles extending to tips of neurosetae or beyond, with scattered clavate micropapillae; dorsal tubercles nodular; ventral cirri short, tapered (Figure 26F). Pygidium between last pair of parapodia, with pair of long anal cirri.

FIGURE 27.—Malmgreniella taylori, new species, paratype (USNM 55075): A, right 1st elytron from segment 2; B, right 2nd elytron from segment 4; C, right middle (7th) elytron from segment 13, with detail of papillae; D, right posterior (14th) elytron from last segment (29). (Scale = 0.5 mm.)

BIOLOGY.—In Tampa Bay, the type locality, Taylor (1971) reported the species (as Polynoid B) as the most common and widely distributed polynoid in the area. A specimen of M. taylori, filled with large yolky eggs, was taken from the oral disc of the brittle star Amphipholis gracillima (Stimpson) in Wrightsville Sound in June by Anne McCrary. The species was reported as commensal with the common burrowing brittle star Amphiodia atra (Stimpson) in the Chesapeake-York estuary by D.F. Boesch (1977) and in Mobile Bay, Alabama, by D. Weston (1984).

ETYMOLOGY.—The species is named for John L. Taylor, who first described the species (as Polynoid B) in his doctoral dissertation.

DISTRIBUTION.—Western Atlantic Ocean, from Chesapeake Bay to Georgia, Gulf of Mexico, Brazil. Low water to 21 meters.

Malmgreniella variegata (Treadwell, 1917), new combination

FIGURES 28–31, FRONTISPIECE A

Harmothoe variegata Treadwell, 1917:260, pl. 1: figs. 17, 18; pl. 2: figs. 1–3.—Hartman, 1956:251, 265, 273.

Paralepidonotus boholensis Gr. var. curacaoensis Horst, 1922:198.

Malmgrenia curacaoensis.—Augener, 1927:45, fig. 2A–D.—Kudenov, 1975b:79.

Harmothoe lunulata.—Millott, 1953:96, figs. 1, 2 [not Delle Chiaje, 1830].

Harmothoe sp.—Hanley, 1987:156.

MATERIAL EXAMINED.—NORTHWESTERN ATLANTIC OCEAN: Florida: Gulf of Mexico, Dry Tortugas, Jun-Jul 1909, holotype of Harmothoe variegata (AMNH 971). Straits of Florida, Looe Key, 24°33′N, 81°24′W, on brittle star Ophionereis reticulata (Say), G. Hendler, collector, by scuba: sta LK-1, 28 Aug 1984, fore reef spur and groove coral reef, 7 m, 1 specimen (USNM 133579; color photo by J.E. Miller); sta LK-33,12 May 1985, back reef, 4 m, 1 small specimen (USNM 133578); sta LK-53, 16 Aug 1985, near northeastern core marker, 2–3 m, 1 specimen (LACM).

CARIBBEAN SEA: Belize: Norval Cay, on Ophionereis reticulata, G. Hendler, J. Miller, collectors, 1.5 m, sta 89-11,4 Mar 1989, 2 specimens (USNM 133576). Curaçao: “Spaansch water,” in Porites porites, 10 Apr 1920, van der Horst, collector, 3 syntypes of Paralepidonotus boholensis var. curacaoensis (ZMA 1039; as Malmgrenia curacaoensis by Augener, 1927, and as Harmothoe sp. by Hanley, 1987). Piscadera Beach, adjacent to Caribbean Marine Biological Institute, sand and rubble, 25 Sep 1976, M.L. Jones, collector, 1 specimen (USNM 133580).

EAST PACIFIC OCEAN: Pamama: Gulf of Panama, Perlas Islands, Pedro Gonzales, on Ophionereis annulata (Le Contel), low water, under stones in coarse sand, 26 Apr 1976, S. Oldfield, collector, 2 specimens (USNM 133577).

DESCRIPTION.—Holotype of Harmothoe variegata from Dry Tortugas (AMNH 971) 26 mm long, 7 mm wide including setae, with 39 segments and 15 pairs of elytra. Complete syntype of Paralepidonotus boholensis var. curacaoensis from Curaçao (ZMA 1039) 18 mm long, 7 mm wide, with 39 segments and 15 pairs of elytra. Specimen from Curaçao (USNM 133580) 24 mm long, 7 mm wide, with 40 segments; largest specimen from Looe Key (USNM 133579) 24 mm long, 7 mm wide, with 40 segments. Young specimen from Looe Key (USNM 133578) 4 mm long, 2 mm wide, 22 segments with small posterior end, short anal cirri, and 11 pairs of elytra. Specimen on Ophionereis reticulata from Belize (USNM 133576) 23 mm long, 7 mm wide, with 38 segments. Two specimens on O. annulata from Gulf of Panama (USNM 133577), one incomplete with elytra, 11 mm long, 4 mm wide, with 32 segments; complete one with elytra missing, 13 mm long, 5 mm wide, with 40 segments, last one very small.

Body elongated, flattened, with sides nearly parallel, tapering posteriorly. Striking black or brown markings on elytra; on dorsum with small marks beginning on middle of body, becoming wider bands on posterior fourth (Figure 30B); on ventrum along midventral groove, on lateral bases of parapodia, and on ventral cirri (Figure 30C); and anteriorly on upper, lateral, and posterior lips of ventral mouth (Figure 29B). Transverse ciliated bands on dorsum, 2 per segment, between elytrophores and dorsal tubercles (Figures 28A, 29A). Fifteen pairs of elytra, covering dorsum except for few posterior segments. Elytra oval to subreniform, without papillae, with group of microtubercles near anterior borders (absent on first and some posterior elytra); pigmentation denser in some areas, consisting of wide funnel-shape area on posterior half of elytra, radiating from region of attachment to elytrophore to medial-posterior part, continuing as narrow band along posterior-lateral border, then to narrow transverse band completing the circle (Figures 28B,C, 30D–F; Treadwell, 1917, pl. 1: fig. 17; Augener, 1927, fig. 2A).

Bilobed prostomium with anterior lobes truncate or subtriangular, without distinct peaks, with 2 pairs of rather small eyes, anterior pair anterior to widest part of prostomium, posterior pair near posterior border, large ceratophore of median antenna in anterior notch, with long style; ceratophores of lateral antennae inserted terminoventrally, converging midventrally and with ciliated ridge on ventral side, and short styles; palps stout, tapered, with longitudinal rows of minute papillae; tentaculophores each with small projecting acicular lobe on inner side, without setae, and pair of dorsal and ventral tentacular cirri similar to median antenna; all appendages variously pigmented (Figures 28A, 29A,B, 30A, 31A; Treadwell, 1917, pl. 1: fig. 18). Segment 2 with first pair of elytrophores, biramous parapodia, and ventral buccal cirri with large cirrophores lateral to ventral mouth and long styles similar to tentacular cirri (Figures 28A,D, 29A,B, 30A). Notosetae similar to those of following segments; neurosetae slender, curved, tapering to fine blunt tips (Figure 28D–F).

FIGURE 28.—Malmgreniella variegata, holotype of Harmothoe variegata (AMNH 971): A, dorsal view of anterior end, styles of median antenna, right dorsal and left dorsal and ventral tentacular cirri missing, first left elytron partially shown; B, right 1st elytron from segment 2; C, right middle elytron, with detail of microtubercles; D, right elytrigerous parapodium from segment 2, posterior view; E, long and short notosetae from same; F, neurosetae from same; G, right middle elytrigerous parapodium, anterior view, acicula dotted; H, right middle cirrigerous parapodium, posterior view; I, short and long notosetae, with detail of tip; J, lower neuroseta; K, middle and upper neurosetae. (Scales = 0.5 mm for A; 1.0 mm for B,C; 0.5 mm for D,G,H; 0.1 mm for E,F,I–K.)

Parapodia biramous, with notopodia shorter than neuropodia; notopodium rounded with projecting acicular lobe on lower side; larger neuropodium with longer subconical, presetal acicular lobe with bulbous or digitiform supraacicular process, with shorter, rounded postsetal lobe (Figures 28G,H, 30G,H, 31B,C; Treadwell, 1917, pl. 2: fig. 2). Notosetae numerous, forming radiating bundle, stouter than neurosetae, with numerous, faint, close-set spinose rows, shorter notosetae with tapered tips, longer ones with blunt tips (Figures 28E,I, 30I, 31D; Augener, 1927, fig. 2B). Neurosetae numerous, forming fan-shape bundle, with numerous, faint, close-set spinose rows, tapering distally to slightly hooked tips, all, except for some lower shorter neurosetae, with secondary tooth (Figures 28J,K, 30J,K, 31E,F; Treadwell, 1917, pl. 2: fig. 3; Augener, 1927, fig. 2C, D). Dorsal cirri with cylindrical cirrophores and long styles extending beyond neurosetae, with 2 dark bands near tip; dorsal tubercles nodular, ventral cirri short, tapered (Figures 28H, 30H, 31C). Pygidium with anus between last pair of small parapodia, with pair of long anal cirri, color grading from darkly pigmented to colorless distal tips.

BIOLOGY.—Norman Millott (1953, figs. 1,2) described and published photographs of two polynoids, one wrapped around the aboral disc of the banded ophiuroid Ophionereis reticulata (Say), the other crawling along the arm of another individual, all found in sand beneath a submerged stone near Kingston, Jamaica. The polynoids showed a color pattern matching closely that of the brittle stars. Although the specimens were not available for study, the polynoids, identified as Harmothoe lunulata, are considered herein to be Malmgreniella variegata. On Looe Key, two adults and one young specimen of M. variegata were collected by scuba in 2–7 meters on specimens of O. reticulata by Gordon Hendler and John Miller. The adults showed the characteristic color pattern (Figure 30A–H; Frontispiece). The young polynoid was nearly colorless, only showing color markings on some anterior elytra and spots on the lower lip (Figure 31). On the Perlas Islands, Gulf of Panama, two specimens of M. variegata were collected on Ophionereis annulata (Le Conte) under stones in coarse sand by Susan Oldfield (in litt.). On Norval Cay, Belize, G. Hendler (in litt.) observed a good number of Ophionereis reticulata. When turning over a piece of coral rubble that was sheltering two ophiuroids, a polynoid, identified later as M. variegata, was found on the rock between them, suggesting that the polynoids move on and off their hosts. According to G. Hendler (in litt.), both O. annulata and O. reticulata, the hosts of M. variegata, have similar pigmentations and color patterns, both have banded arms. See Frontispiece.

FIGURE 29.—Malmgreniella variegata, two syntypes of Paralepidonotus boholensis var. curacaoensis (ZMA 1039): A, dorsal view of anterior end, styles of tentacular cirri missing; B, ventral view of anterior end, palps, styles of median antenna, right tentacular cirri, and ventral buccal cirri of segment 2 missing. (Scale = 0.5 mm.)

FIGURE 30.—Malmgreniella variegata, specimen from Looe Key (USNM 133579): A, dorsal view of anterior end. pharynx partially extended; B, middorsum of segment 36 showing pigmentation; C, ventral view of segments 21 and 22 showing pigmentation; D, left 1st elytron from segment 2; E, left 2nd elytron from segment 4, with detail of microtubercles; F, right 6th elytron from segment 11; G, right elytrigerous parapodium, anterior view, acicula dotted; H, right cirrigerous parapodium, posterior view; I, long and short notosetae, with detail of tips; J, middle neuroseta; K, lower neuroseta. (Scales = 0.5 mm for A,B; 1.0 mm for C; 1.0 mm for D–F; 0.5 mm for G,H; 0.1 mm for I–K.)

FIGURE 31.—Malmgreniella variegata, young specimen from Looe Key (USNM 133578): A, dorsal view of anterior end; B, right elytrigerous parapodium, anterior view, acicula dotted, style of ventral cirrus broken off; C, right cirrigerous parapodium, posterior view; D, long and short notosetae, with detail of tips; E, middle neuroseta; F, lower neuroseta. (Scales = 0.5 mm for A; 0.2 mm for B,C; 0.1 mm for D–F.)

DISTRIBUTION.—Northwestern Atlantic Ocean, off Florida; Gulf of Mexico; Caribbean Sea; East Pacific Ocean, Gulf of Panama. Intertidal to 7 meters.

Malmgreniella pierceae, new species

FIGURE 32

Malmgreniella sp. C.—Weston, 1984:21, 22, figs. 21-17,18a–g.

MATERIAL EXAMINED.—NORTHWESTERN ATLANTIC OCEAN: Georgia: Off Georgia, 31°03′N, 80°26′W, 34 m, Texas Instruments, R/V Pierce sta 5E, 16 May 1977, holotype (USNM 61750). 31°01′N, 80°17′W, 40 m, sta 5F, 16 May 1977, paratype (USNM 61762). Same, 30 Aug 1977, 3 paratypes (USNM 61763-4). 30°59′N, 80°08′W, 46 m, sta G, 25 Feb 1977, paratype (USNM 61304).

GULF OF MEXICO: Panama: Caribbean coast, Galeta Reef, 11 m, 4 Oct 1980, on brittle star Amphiura cf. fibulata Koëhler, G. Hendler, collector, 1 specimen (USNM 133561). Florida: as Malmgreniella sp. C by Weston, 1984: Southwest Florida (SOFLA) sta 20, Nov 1980, Jul 1981, 25°17′N, 82°09′W, 22 m, coarse sand, 2 specimens (USNM 89690-1). Mississippi-Alabama-Florida (MAFLA) sta 2422, off Apalachicola River, 29°30′N, 84°27′W, medium fine sand, 1 specimen (USNM 89689).

FIGURE 32.—Malmgreniella pierceae, new species, A, holotype (USNM 61750), B–I, paratype (USNM 61762): A, dorsal view of anterior end; B, same, prostomium defective, right eyes misplaced, ceratophore and style of median antenna missing; C, middle elytron; D, right elytrigerous parapodium, anterior view, acicula dotted; E, right cirrigerous parapodium, posterior view; F, long and short notosetae; G, upper neuroseta; H, middle neuroseta; I, lower neurosetae. (Scales = 0.5 mm for A,B; 0.5 mm for C–E; 0.1 mm for F–I.)

DESCRIPTION.—Holotype 10 mm long, 3 mm wide including setae, with 37 segments. Large paratype (USNM 61762) 13 mm long, 4 mm wide, with 37 segments. Two smaller paratypes 3–6 mm long, 2 mm wide, with 33 segments. Body flattened, tapering slightly anteriorly and more so posteriorly, without color. Fifteen pairs of oval elytra, without tubercles and papillae, colorless (Figure 32C),

Bilobed prostomium with subtriangular anterior lobes, without distinct peaks, with 2 pairs of rather large eyes on posterior half, and with some scattered pigment spots; ceratophore of median antenna in anterior notch, with style about length of prostomium; lateral antennae with distinct ceratophores inserted terminoventrally, converging midventrally, with short subulate styles; palps stout, tapered; tentaculophores without setae, dorsal and ventral tentacular cirri similar to median antenna; antennae and tentacular cirri minutely papillate (Figure 32A,B; Weston, 1984, fig. 21-18a). Segment 2 with long ventral buccal cirri similar to tentacular cirri.

Parapodium biramous, with notopodium shorter than neuropodium; notopodium rounded, with projecting acicular lobe on lower side; larger neuropodium with subconical presetal acicular lobe with very small bulbous supraacicular process, with shorter, rounded postsetal lobe (Figure 32D,E; Weston, 1984, fig. 21-18b). Notosetae numerous, forming radiating bundle, about same width as neurosetae, short to long, slightly curved, with tapering blunt tips, appearing smooth but finely spinose along convex border (Figure 32F; Weston, 1984, fig. 21-18c). Neurosetae moderate in number, forming fan-shape bundle; upper neurosetae with long spinose regions and entire, bare tips; some larger middle neurosetae with shorter spinose regions and bifid tips, with secondary tooth very long and slender; lower neurosetae shorter with less prominent spinose rows and entire, slightly hooked tips (Figure 32G–I; Weston, 1984, fig. 21-18d–g). Dorsal cirri with cylindrical cirrophores, wider basally and long styles extending beyond tips of neurosetae, with scattered micropapillae; dorsal tubercles nodular; ventral cirri short, tapered, with scattered micropapillae on upper side (Figure 32E). Pydigium small, rounded, between posterior pair of small parapodia.

BIOLOGY.—Malmgreniella pierceae was collected on the brittle star Amphiura cf. fibulata Koëhler on Galeta Reef, Panama, by G. Hendler.

ETYMOLOGY.—The species is named for the collecting ship R/V Pierce.

DISTRIBUTION.—Northwest Atlantic Ocean, off Georgia; Caribbean coast of Panama; Gulf of Mexico, off Florida. In 11–46 meters.

Malmgreniella hendleri, new species

FIGURE 33

MATERIAL EXAMINED.—NORTHWEST ATLANTIC OCEAN: Panama: Caribbean Sea, Galeta Reef, G. Hendler, collector, on burrowing brittle stars: 28 Oct 1977, on disc of Ophionephthys limicola Lütken, holotype (USNM 133567); 10 Oct 1977, 9 m, on mouth of Amphiura cf. fibulata Koëhler, paratype (USNM 133570); 13–14 Sep 1978, 9–11 m, on A. cf. fibulata, young paratype (USNM 133569); 2–5 Oct 1980, 11 m, sand, on O. limicola, 7 paratypes (USNM 133568; LACM).

DESCRIPTION.—Holotype 6 mm long, 3 mm wide including setae, with 34 segments; paratypes 6–9 mm long, 2.5–3 mm wide, with 31–34 segments; young paratype 1 mm long, 1 mm wide, with 13 segments and budding area, with 7 pairs of elytra. Posterior fourth of dorsum with black pigment spots medial to elytrophores and dorsal tubercles, and 4 groups of spots on posterior fourth of ventrum (Figure 33B). Color lacking on young paratype. Elytra 14–15 pairs, large, covering dorsum, oval to subreniform, without papillae or microtubercles; color pattern forming nearly complete circle on 1st and 2nd elytra, on following elytra, pattern is C-shape on medial part of elytra, including spot on place of attachment to elytrophore (Figure 33C–E). Bilobed prostomium with anterior lobes truncate, without peaks, with light pigmentation on middle of prostomium; 2 pairs of eyes, larger anterior pair slightly anterior to greatest width of prostomium, smaller posterior pair near posterior border; median antenna with ceratophore in anterior notch, long style with filamentous tip; lateral antennae with ceratophores inserted terminoventrally, with short, subulate styles; palps stout, tapered, about same length as median antenna; tentaculophores with single seta on inner side, dorsal and ventral tentacular cirri similar to median antenna (Figure 33A). Second segment with first pair of elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 33A). Extended pharynx with 9 pairs of border papillae and 2 pairs of hooked jaws.

Biramous parapodium with notopodium shorter than neuropodium, rounded, with projecting acicular lobe on lower side; neuropodium with subconical presetal acicular lobe, with bulbous supraacicular process; postsetal lobe shorter, rounded (Figure 33F,G). Notosetae numerous, forming radiating bundle, shorter and more slender than neurosetae, with faint spinose rows and slightly bulbous bare tips (Figure 33H). Neurosetae moderate in number (about 15), forming fan-shape bundle; supraacicular neurosetae with longer spinose regions, and more prominent spinose rows, with bifid, bare tips (Figure 33I); subacicular neurosetae with shorter spinose regions and entire, bare tips (Figure 33J). Dorsal cirri with cylindrical cirrophore, long styles with minute sensory papillae, tapering to slender tips and extending beyond tips of neurosetae; dorsal tubercles nodular; ventral cirri short, subulate (Figure 33G). Pygidium rounded, between posterior pair of small parapodia, with pair of long anal cirri.

BIOLOGY.—Malmgreniella hendleri was collected on two species of burrowing ophiuroids, Ophionephthys limicola Lütken and Amphiura cf. fibulata Koëhler, on Galeta Reef, Panama, by Gordon Hendler.

ETYMOLOGY.—The species is names for Gordon Hendler, specialist on the Ophiuroidea and collector of the polynoid commensals.

DISTRIBUTION.—Northwest Atlantic Ocean, Caribbean Sea, Panama. In 9–14 meters.

FIGURE 33.—Malmgreniella hendleri, new species, holotype (USNM 133567); A, dorsal view of anterior end, styles of right dorsal and ventral and left dorsal tentacular cirri missing; B, ventral view of posterior segments 27 and 28 showing pigmentation; C, right 1st elytron from segment 2; D, right 3rd elytron from segment 5; E, right middle elytron; F, right elytrigerous parapodium form segment 15, anterior view, acicula dotted; G, right cirrigerous parapodium form segment 16, posterior view; H, notoseta, with detail of tip; I, supraacicular neuroseta, with detail of tip; J, subacicular neurosetae. (Scales = 0.5 mm for A,B; 0.5 mm for C–E; 0.2 mm for F,G; 0.1 mm for H–J.)

Malmgreniella galetaensis, new species

FIGURE 34

MATERIAL EXAMINED.—NORTHWEST ATLANTIC OCEAN: Panama: Caribbean Sea, Galeta Reef, 11–12 m, sand, on burrowing ophiuroids, G. Hendler, collector: 12 Sep 1978, on Ophiophragmus septus Lütken, holotype (USNM 133571); 2 Oct 1980, on O. septus, paratype (USNM 133573); 3 and 5 Oct 1980, on O. septus, 3 paratypes (USNM 133574; LACM); 3 Oct 1980, on Amphiura cf. fibulata Koëhler, paratype (USNM 133575). Florida: Capron Shoals, 2 mi SE of Ft. Pierce Inlet, 14 m, shelly, silty sand, 11 Jul 1990, G. Hendler, collector, on disc of O. septus, paratype (USNM 133572).

FIGURE 34.—Malmgreniella galetaensis, new species, holotype (USNM 133571); A, dorsal view of anterior end, left dorsal tentacular cirrus missing; B, ventral view of 2 segments near posterior end showing pigmentation patter; C, left 1st elytron from segment 2; D, left middle elytron; E, right elytrigerous parapodium from segment 13, anterior view, acicula dotted; F, right cirrigerous parapodium from segment 12, posterior view; G, long and short notosetae, with detail of tip; H, supraacicular neurosetae; 1, subacicular neurosetae. (Scales = 0.5 mm for A,B; 0.5 mm for C,D; 0.2 mm for E,F; 0.1 mm for G–I.)

DESCRIPTION.—Holotype, female with eggs, 6 mm long, 2.5 mm wide including setae, with 32 segments, last one very small; complete paratype (USNM 133574), male with sperm, 5.2 mm long, 2 mm wide, with 30 segments; complete paratype (USNM 133572), male with sperm, 10 mm long, 4 mm wide, with 36 segments. Posterior one-half to two-thirds of body with black pigmentation in form of large oval midventral spots, smaller lateral spots on ventral bases of parapodia, and on basal two-thirds of dorsal cirri (Figure 34B,F). Elytra 14–15 pairs, oval, without papillae or microtubercles, with blackish pigmentation in spots over place of attachment to elytrophores, nearly complete ring on first elytra and semicircular ring on posterior half of following elytra (Figure 34C,D). Bilobed prostomium with subtriangular anterior lobes; 2 pairs of eyes, anterior pair in region of greatest width, posterior pair near posterior border; ceratophore of median antenna in anterior notch, with style about length of prostomium; lateral antennae with ceratophores inserted terminoventrally, with short subulate styles; palps stout, tapered; tentaculophores with single seta on inner side, dorsal and ventral tentacular cirri nearly as long as palps (Figure 34A). Segment 2 with first pair of elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 34A). Extended pharynx with 9 pairs of border papillae, some with dark pigment inside, and 2 pairs of hooked jaws.

Biramous parapodium with notopodium shorter than neuropodium, rounded, with projecting acicular lobe on lower side; neuropodium with subconical presetal acicular lobe and rounded supraacicular process; postsetal lobe shorter, rounded (Figure 34E,F). Notosetae numerous, forming radiating bundle, short to long, more slender than neurosetae, appearing smooth, but finely spinose with slightly bulbous bare tips (Figure 34G). Neurosetae moderate in number (about 14), forming fan-shape bundle; supraacicular neurosetae with long spinose regions, prominent spines, and bifid, bare tips (Figure 34H); subacicular neurosetae with shorter spinose regions, faint spinous rows, and slightly hooked, entire bare tips (Figure 34I). Dorsal cirri with long cylindrical cirrophores, styles long, with scattered micropapillae, tapering to slender tips and extending beyond neurosetae; dorsal tubercles nodular; ventral cirri short, tapered (Figure 34A,F). Pygidium between posterior pair of small parapodia, small, rounded, with ring of pigment, with pair of long anal cirri.

BIOLOGY.—Malmgreniella galetaensis was collected on two species of burrowing ophiuroids, Ophiophragmus septus (Lütken) and Amphiura cf. fibulata Koëhler, on Galeta Reef, Panama, by Gordon Hendler. At Ft. Pierce Inlet, Florida, the species was collected on the disc of O. septus by G. Hendler.

ETYMOLOGY.—The species is named for the collecting site, Galeta Reef, Panama.

DISTRIBUTION.—Northwest Atlantic Ocean, Caribbean Sea, West coast of Florida. In 9–14 meters.

Malmgreniella panamensis, new species

FIGURE 35

MATERIAL EXAMINED.—NORTHWESTERN ATLANTIC OCEAN: Panama: Caribbean Sea, Galeta Reef, 13–14 Sep 1978, 9–12 m, sand, on Ophiopsila cf. polysticta H.L. Clark, G. Hendler, collector, holotype (USNM 133562); 3 Oct 1980, paratype (USNM 133563).

DESCRIPTION.—Holotype, female with eggs, 7 mm long, 3.5 mm wide including setae, with 31 segments; paratype, female with eggs, 8 mm long, 3.2 mm wide, with 35 segments. Posterior two-thirds of body with black pigmentation in form of large oval midventral spots and smaller spots on ventral bases of parapodia, and on basal halves of dorsal cirri (Figure 35B). Elytra 14–15 pairs, oval to subreniform, delicate, transparent, without papillae or microtubercles, with blackish pigmentation in spots over place of attachment to elytrophores, in complete ring on first elytra, semicircular bands on following elytra, and nearly complete circle on posterior elytra (Figure 35C–E).

Bilobed prostomium with subtriangular anterior lobes, with slight indication of peaks; 2 pairs of eyes subequal in size, anterior pair in region of greatest width, posterior pair near posterior border; scattered pigmentation on prostomium and bases of antennae; ceratophore of median antenna in anterior notch, with style about as long as palps; lateral antennae with ceratophores inserted terminoventrally, with short subulate styles; palps stout, tapered; tentaculophores without setae, with dorsal and ventral tentacular cirri about same length as median antenna (Figure 35A). Segment 2 with first pair of elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 35A).

Biramous parapodium with notopodium shorter than neuropodium, rounded with projecting acicular lobe on lower side; neuropodium with subconical presetal acicular lobe and conical supraacicular process, not sharply set off; postsetal lobe shorter, rounded (Figure 35F,G). Notosetae numerous, forming radiating bundle, short to long, more slender than neurosetae, appearing smooth but finely spinose with short, bare, rounded tips (Figure 35H). Neurosetae moderate in number (about 20) forming fan-shape bundle; supraacicular neurosetae with long spinose regions, prominent spines, with entire, tapered tips (Figure 35I), or few with very small secondary tooth (on paratype); subacicular neurosetae with shorter spinose regions, entire, slightly hooked, bare tips (Figure 35J). Dorsal cirri with long cylindrical cirrophores, with styles long, tapering to filamentous tips, extending far beyond neurosetae; black pigmentation on basal third of styles; dorsal tubercles nodular; ventral cirri short, tapered (Figure 35G). Pygidium with pair of pigmented anal cirri.

BIOLOGY.—Malmgreniella panamensis was collected subtidally in sand on the burrowing ophiuroid Ophionpsila cf. polysticta H.L. Clark by Gordon Hendler.

ETYMOLOGY.—The species is named for Panama, the collection area.

DISTRIBUTION.—Northwestern Atlantic Ocean, Caribbean Sea, Panama. In 9–12 meters.

Malmgreniella puntotorensis, new species

FIGURES 36, 37, FRONTISPIECE B–D

MATERIAL EXAMINED.—NORTHWESTERN ATLANTIC OCEAN: Florida: Looe Key, back reef, 24°33′N, 81°24′W, sta LK-33,12 May 1985,4 m, sand, Thalassia, few sponges, on ophiuroid, G. Hendler, collector, paratype (USNM 133590).

CARIBBEAN SEA: Panama: Punto Toro, Fort Sherman, sta 147–1, 16 Apr 1973, sand bottom between rocks and Thalassia, M.L. Jones, collector, holotype (USNM 71760) and paratype (USNM 71761). Galeta Reef, 9–11 m, with burrowing ophiuroid Amphiodia trychna H.L. Clark, G. Hendler, collector: 27 Oct 1977, paratype (USNM 133588); 13 Sep 1978, paratype (USNM 133589); 3–5 Oct 1980, 3 paratypes (USNM 133586; LACM); 3 Oct 1980, on Ophiophragmus septus (Lütken), paratype (USNM 133587). Belize: Twin Cays, 16°50′N, 86°06′W, 0–2 m, 30 May 1983, on Ophionephthys limicola Lütken, G. Hendler, collector, paratype (USNM 133592). Twin Cays, Gator Creek, 1.5 m, 20 Jun 1985, on Ophiophragmus pulcher H.L. Clark, 3 paratypes (USNM 133596; LACM). Twin Cays, Hidden Creek, 1.5 mm, G. Hendler, collector, on Ophiophragmus pulcher, 30 May 1982, paratype (USNM 133591) and 23 Mar 1983, paratype (LACM); sta 85–11, 18 Jun 1985, 2 paratypes (USNM 133594); sta 85–12, 19 Jun 1985, 3 paratypes (USNM 133595, LACM); sta 86–7, 4 Apr 1986, 2 paratypes (USNM 133597); sta 89–14, 5 Mar 1989, paratype (USNM 133602); sta 89–22, 7 Mar 1989, paratype (USNM 133599); sta 89–25, 9 Mar 1989, paratype (USNM 133601); sta 89–31, 12 Mar 1989, 2 paratypes (USNM 133598); on Ophiophragmus cubanus (A.H. Clark), sta 85–12, 19 Jun 1985, paratype (LACM); sta 89–25, 9 Mar 1989, paratype (USNM 133600); on Amphiodia trychna, sta 85–11, 18 Jun 1985, paratype (USNM 133593).

FIGURE 35.—Malmgreniella panamensis, new species, holotype (USNM 133562): A, dorsal view of anterior end; B, ventral view of 2 segments near posterior end showing pigmented pattern, right parapodia not shown; C, left 1st elytron from segment 2; D, left middle elytron; E, left 12th elytron from segment 23; F, right elytrigerous parapodium from segment 13, anterior view, acicula dotted; G, right cirrigerous parapodium from segment 12, posterior view; H, long and short notosetae, with detail of tip; I, supraacicular neurosetae; J, subacicular neurosetae. (Scales = 0.5 mm for A,B; 0.5 mm for C–E; 0.2 mm for F,G; 0.1 mm for H–J.)

DESCRIPTION.—Holotype (USNM 71760), female with eggs, 7.5 mm long, 2.5 mm wide including setae, with 34 segments; paratype (USNM 71761), male with sperm, 6.5 mm long, 2.5 mm wide, with 34 segments. Additional paratypes from Panama and Belize with 25–36 segments, 4–11 mm long, and 2–3 mm wide.

Body of type specimens from Panama without color; on specimens commensal with ophiuroids from Belize, color variable: body and elytra without color; uniformly reddish brown, darker on antennae, tentacular cirri, and mouth area (Figure 37A,B); some with anterior half or two-thirds of body dark and posterior region colorless; some with additional chalky white transverse segmental patches on middorsum and scattered spots on dorsal tubercles, cirrophores of dorsal cirri, on midventral groove and ventral bases of parapodia, and cirrophores of ventral cirri. Elytra 15 pairs, large, oval, overlapping, covering dorsum, delicate, opaque, granular, denser around place of attachment to elytrophores, without papillae or tubercles (Figures 36B, 37C,D).

Bilobed prostomium with anterior lobes truncate, without peaks; 2 pairs of eyes, anterior pair slightly anterior to greatest width, posterior pair near posterior border; median antenna with ceratophore in anterior notch, with style about 1.5 times longer than prostomium; lateral antennae with ceratophores inserted terminoventrally, converging midventrally, with short subulate styles; palps stout, tapered; tentaculophores without setae, with dorsal and ventral tentacular cirri similar to median antenna (Figures 36A, 37A,B). Segment 2 with first pair of elytrophores, subbiramous parapodia and long ventral buccal cirri on cirrophores lateral to ventral mouth, with styles similar to tentacular cirri (Figures 36A, 37A,B). Extended pharynx with 9 pairs of border papillae and 2 pairs of jaws.

Parapodium subbiramous; notopodium small, digitiform, on anterodorsal side of larger neuropodium; neuropodium with conical presetal acicular lobe, without supraacicular process and shorter, rounded postsetal lobe (Figures 36C,D, 37E,F). Notosetae reduced to 1–2, short, smooth, acicular, with pointed tip, not extending beyond tip of notopodium and easily overlooked (Figures 36C,E, 37E–G). Neurosetae rather few (about 11), forming fan-shape bundle; upper and middle neurosetae with longer spinose regions and bifid tips, with long secondary tooth (Figures 36F, 37H); lower neurosetae with shorter spinose regions, with small secondary tooth, 3–4 lowest neurosetae with secondary tooth very small or absent (Figures 36G, 37I). Dorsal cirri with cylindrical cirrophores, bulbous basally, with styles smooth, tapering, and extending to tips of neurosetae or beyond; dorsal tubercles bulbous; ventral cirri short, tapered (Figures 36D, 37F). Small pygidium posterior to last segment, with pair of anal cirri.

FIGURE 36.—Malmgreniella punntotorensis, new species, holotype (USNM 71760): A, dorsal view of anterior end; B, left elytron: C, right elytrigerous parapodium, anterior view, acicula dotted, with detail of notopodium and notoseta from posterior view; D, right cirrigerous parapodium, posterior view; E, notosetae; F, upper and middle neurosetae; G, lower neuroseta. (Scales = 0.5 mm for A; 0.5 mm for B; 0.1 mm for C,D; 0.1 mm for E–G.)

FIGURE 37.—Malmgreniella puntotorensis, new species, specimen from Belize (USNM 133591): A, dorsal view of anterior end, turned slightly to right; B, ventral view of anterior end; C, left 2nd elytron from segment 4; D, right middle elytron; E, right elytrigerous parapodium, anterior view, acicula dotted; F, right cirrigerous parapodium, posterior view; G, notoseta; H, middle and upper neurosetae; I, lower neurosetae. (Scales = 0.5 mm for A,B; 0.5 mm for C,D; 0.2 mm for E,F; 0.1 mm for G–I.)

BIOLOGY.—Malmgreniella puntotorensis was collected by G. Hendler on Galeta Reef, Panama, and at Belize on five species of burrowing amphiurid ophiuroids: Amphiodia trychna H.L. Clark (7 specimens), Ophionephthys limicola Lütken) (1), Ophiophragmus cubanus (A.H. Clark) (2), Ophiophragmus pulcher H.L. Clark (17), and Ophiophragmus septus (Lütken) (1). The commensal polynoid was found by Hendler on the disc of a freshly excavated O. pulcher and, when the ophiuroid was allowed to burrow again in the sediment a couple of times, the polynoid remained attached to the disc of the brittle star. See color photos (Frontispiece B–D).

ETYMOLOGY.—The species is named for Punto Toro, the type locality.

DISTRIBUTION.—Northwestern Atlantic Ocean, Caribbean Sea, Florida (Looe Key), Panama, Belize. Intertidal to 11 meters.

FIGURE 38.—Malmgreniella lilianae, new species, paratype (USNM 55062): A, dorsal view of anterior end, left palp missing; B, right 1st elytron from segment 2; C, right 2nd elytron from segment 4; D, right middle elytron; E, left elytrigerous parapodium from segment 2, anterior view, acicula dotted; F, upper, middle, and lower neurosetae from same; G, right elytrigerous parapodium, anterior view, acicula doted, eggs visible in body cavity; H, right cirrigerous parapodium, posterior view; I, long and short notosetae; J, upper neuroseta; K, middle neuroseta; L, lower neurosetae. (Scales = 0.5 mm for A; 0.5 mm for B–D; 0.2 mm for E,G,H; 0.1 mm for F,I–L.)

Malmgreniella lilianae, new species

FIGURE 38

MATERIAL EXAMINED.—SOUTHWESTERN ATLANTIC OCEAN: Brazil: Bay of Flamengo, São Paulo, 23°27′S, 45°06′W, 12 m, Jan 1963, Liliana Forneris, collector, holotype (USNM 55061), 6 paratypes (USNM 55062).

DESCRIPTION.—Holotype, female with eggs, 8 mm long, 4 mm wide including setae, with 30 segments and 14 pairs of elytra. Paratypes (males and females with sperm and eggs) 6–7 mm long, 3–4 mm wide, with 29–30 segments and 13–14 pairs of elytra. Body flattened, tapering slightly anteriorly and posteriorly, with parapodia and setae longer than body width. Body transversely banded dorsally on posterior few segments and ventrally on posterior third to one half of body. Elytra oval, opaque, without tubercles, with some micropapillae along lateral and posterior borders and few on surface, with light grey to black pigmentation forming spots over place of attachment to elytrophore, complete circle on first elytra, and large C-shape area with larger portion on medial area on following elytra (Figure 38B–D).

Bilobed prostomium with anterior lobes rounded, without cephalic peaks, 2 pairs of small eyes, anterior pair anterior to greatest width of prostomium, posterior pair posterodorsal; ceratophore of median antenna in anterior notch with pigment spots on lateral sides, and style about as long as prostomium; ceratophores of lateral antennae barely visible dorsally, inserted ventrally and converging midventrally, with short styles less than half length of median antenna; palps stout, tapered; tentaculophores with single seta on inner side, dorsal and ventral tentacular cirri similar to and slightly longer than median antenna; antennae and tentacular cirri minutely papillate (Figure 38A). Segment 2 with first pair of elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri; notosetae similar to those of following parapodia; neurosetae more slender than those following, upper ones tapering to slender, sharp tip, middle and lower ones with long, bare, bulbous tip (Figure 38A,E,F).

Biramous parapodia with body wall delicate, transparent, with internal structures and eggs in coelom clearly visible; notopodium shorter than neuropodium, rounded, with projecting acicular lobe on lower side; neuropodium with subconical presetal acicular lobe, digitiform supraacicular process and postsetal lobe shorter, rounded (Figure 38G,H). Notosetae numerous, about 3 lengths, longest extending to about tips of neuropodia, slightly stouter than neurosetae, with prominent spinose rows, tapering to pointed bare tips (Figure 38G,I). Neurosetae moderate in number (about 17), forming fan-shape bundle, upper ones with longer spinose region and pointed bare tips, middle and lower ones with shorter spinose region and slightly hooked bare tips, all with entire tips (Figure 38G,J–L). Cylindrical cirrophores of dorsal cirri with wide glandular areas on posterior sides, with styles finely papillate, with filamentous tips extending to about tips of neurosetae; dorsal tubercles nodular; ventral cirri short, tapered, with few micropapillae (Figure 38H). Pygidium with anus medial to last pair of small parpodia, with pair of long anal cirri.

ETYMOLOGY.—The species is named for the collector, Liliana Forneris.

DISTRIBUTION.—South Atlantic Ocean, Brazil. Depth 12 meters.

Key to the Species of Malmgreniella from the Eastern Pacific Ocean (British Columbia, Washington, California, Galapagos, Panama)

Malmgreniella nigralba (E. Berkeley, 1923), new combination

FIGURES 39, 40

Malmgrenia nigralba E. Berkeley, 1923:213, pl. l: figs. 5–7; 1924:193.—Berkeley and Berkeley, 1941:21; 1942:187; 1948:10. fig. 8.—Pettibone, 1967:3.—Hartman. 1968:133, figs. 1–3.—Kudenov, 1975b:77, 79.

Harmothoe lunulata.—Hartman, 1944:244 [part]; 1968:81 [part]. [Not Delle Chiaje. 1830.]

Malmgrenia lunulata.—Pettibone, 1953:25, pl. 9: fig. 77; pl. 10: figs. 80–82, 85, 87: pl. 11: figs. 96–103 [part].—Banse et al., 1968:525 [part].—Lie, 1968:303, 370, 371 [part: sta 5]. [Not Delle Chiaje, 1830.]

Harmothoe nigralba.—Hanley, 1987:153, fig. 3F,H.

MATERIAL EXAMINED.—EASTERN PACIFIC OCEAN: British Columbia: Vancouver Island, Piper's Lagoon, near Nanaimo, very low tide, sandy, 20 Jun 1920, E. Berkeley, collector, 3 syntypes (USNM 32875–6). Pipers's Lagoon, 21 May 1924, coarse gravelly sand, commensal in vertical burrows of Leptosynapta clarki Heding. E. Berkeley, collector, 2 specimens (USNM 55040). Washington: Puget Sound, Seattle, Golden Gardens Park, shore, with Leptosynapta clarki Heding, 7 May 1949, M. Pettibone, collector, 2 specimens (USNM 55041). Olalla, shore, 22 Jun 1939, M. Pettibone, collector, 1 specimen (USNM 24931). Gig Harbor, shore, 3 Aug 1944, J.E. Lynch, collector, 2 specimens (USNM 24926). Near Dutchers Cove, Case Inlet, shore, gravelly, shelly sand, with L. clarki, 21 Jun 1947. 6 Jun and 17 Aug 1948. M. Pettibone, collector, 18 specimens (USNM 24925, as M. lunulata var. nigralba by Pettibone, 1953). Southwest of Johnson Point, Case Inlet, 47°10′N, 122°50′W, 15–36 m, coarse shelly sand mixed with rock and many L clarki, sta 5, 12 Feb 1963, U. Lie, collector, 3 specimens (USNM 36444, as M. lunulata form A by Banse et al., 1968). California: Tomales Bay, sandy-mud, with Lepto synapta clarki, summer 1941, F. Pitelka and R. Paulson, collectors, 2 specimens (USNM 24929; AHF 1202, as H. eurosetae with bifid tips [Figure 48H] H. lunulata by Hartman, 1944). Off Santa Cruz Island, 9–27 m, summer 1939, W.G. Hewatt, collector, 1 specimen (USNM 55042, as M. nigralba by Berkeley and Berkeley, 1941).

DESCRIPTION.—Syntypes all fragmented. Complete syntype, in 2 pieces (USNM 32876), 14 mm long, 5 mm wide including setae, and 40 segments, posterior segment very small. Adult specimens from Case Inlet (USNM 24925) 18–24 mm long, 5–7 mm wide, with 37–40 segments; small specimen 7.5 mm long, 3 mm wide, with 30 segments.

Body showing striking black and white pigmentation; both dorsal and ventral sides of posterior segments showing dark pigmentation. Elytra 15 pairs, on usual segments, circular to subreniform, with group of microtubercles on anterior medial part; microtubercles absent on 1st pair and some posterior elytra (Figure 39B–D). Pigmentation on elytra somewhat variable. On syntypes from Nanaimo and specimens from Tomales Bay, 1st pair of elytra with complete circle of dark pigmentation and large reticular area (Figure 39B); following elytra with reticular area confined to posterior two-thirds up to near border, with some scattered microtubercles, and complete or nearly complete oval ring of dark pigmentation (Figure 39C,D; Berkeley, 1923, pl. 1: figs. 6, 7). Melanistic specimens from Case Inlet dark gray on most of posterior two-thirds of elytra, including borders; reticular area with scattered microtubercles, circle of darker color around central colorless area, and group of microtubercles on anterior medial part (absent on first pair and more posterior elytra) (Figure 40A–D; Pettibone, 1953, pl. 11: figs. 96, 98–100).

FIGURE 39.—Malmgreniella nigralba, syntype of Malmgrenia nigralba (USNM 32875): A, dorsal view of anterior end, style of median antenna missing, anterior pair of eyes anteroventral (dotted); B, right 1st elytron from segment 2; C, right 2nd elytron from segment 4, with detail of microtubercles; D, right middle elytron; E, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; F, upper and middle neurosetae from same, with detail of tips; G, right elytrigerous parapodium, anterior view, acicula dotted; H, right cirrigerous parapodium, posterior view; I, long and short notosetae: J, upper, middle, and lower neurosetae, with detail of tips. (Scales = 0.5 mm for A; 0.5 mm for B–D; 0.2 mm for E,G,H; 0.1 mm for F,I,J.)

FIGURE 40.—Malmgreniella nigralba, melanistic specimen from Case Inlet (USNM 24925): A, right 1st elytron from segment 2; B, right 2nd elytron from segment 4, with detail of reticular area and microtubercles; C, right middle elytron; D, right 15th elytron from segment 32, with detail of reticular area and microtubercles. (Scale = 0.5 mm for A–D.)

Bilobed prostomium with anterior lobes truncate, without peaks; anteroventral pair of eyes slightly larger than posterodorsal pair, ceratophore of median antenna bulbous, in anterior notch, style with minute papillae, slightly longer than prostomium; ceratophores of lateral antennae inserted terminoventrally, converging midventrally, with styles short, subulate, papillate; palps stout, tapered; tentaculophores lateral to prostomium, with 0–1 seta on inner side; dorsal and ventral tentacular cirri similar to median antenna (Figure 39A; Berkeley, 1923, pl. 1: fig. 5; Pettibone, 1953, pl. 11: figs. 96, 97; Hanley, 1987, fig. 3F,H).

Segment 2 with 1st pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 39A,E); notosetae similar to following notosetae; neurosetae more slender than those following, with bulbous tips, upper and lower ones entire, middle ones with secondary tooth (Figure 39F). Extended pharynx with 9 pairs of border papillae and 2 pairs of hooked jaws.

Biramous parapodia with smaller notopodium rounded, with projecting acicular lobe on lower side; larger neuropodium with subconical presetal acicular lobe with digitiform or tapered supraacicular process and slightly shorter rounded postsetal lobe (Figure 39E,G,H; Pettibone, 1953, pl. 11: fig. 101). Notosetae moderate in number, about same width as neurosetae, short to longer, smooth or with faint spinose rows (Figure 39I). Neurosetae moderate in number, upper ones with longer spinose region, upper few with rounded blunt tip, some with small secondary tooth; middle ones with shorter spinose region, hooked tip with distinct secondary tooth; lower ones with entire rounded tip or with small tooth (Figure 39J; Pettibone, 1953, pl. 11: figs. 102, 103). Dorsal cirri with cylindrical cirrophore with glandular area on posterior side; style with short clavate papillae, tapered distally and extending to about tips of neurosetae; dorsal tubercles nodular; ventral cirri short, subulate, with small papillae (Figure 39H). Pygidium with anus medial to last pair of small parapodia, with pair of anal cirri.

BIOLOGY.—Malmgreniella nigralba was found in the Nanaimo area, Vancouver Island by Edith Berkeley in coarse gravelly sand living commensally in the vertical burrows of the synaptid Leptosynapta clarki Heding (as L. inhaerens) and by M. Pettibone in the Puget Sound area. In Tomales Bay, Central California, M. nigralba also was found associated with L. clarki (as L. albicans) by F. Pitelka and R. Paulson.

DISTRIBUTION.—Eastern Pacific Ocean from British Columbia to Central California, in low water to 36 meters.

Malmgreniella berkeleyorum, new species

FIGURE 41

Harmothoe lunulata.—Berkeley and Berkeley, 1945:321; 1948:11. [Not Delle Chiaje, 1830.]

MATERIAL EXAMINED.—EASTERN PACIFIC OCEAN: British Columbia: Gulf of Georgia, Mittelnacht Island off east coast of Vancouver Island, 183 m, 16 Jan 1929, E. and C. Berkeley, collectors, holotype (USNM 55039).

DESCRIPTION.—Holotype, female with eggs, with 30 segments, last 8 smaller, regenerating, 11 mm long, 4 mm wide including setae. Body dark. Elytra 14 pairs, on usual segments, last pair on segment 29 small (probably 15 pairs when complete). Elytra oval and subreniform, dark, opaque, darker around area of attachment, showing “veins,” with large group of microtubercles on anterior part, except first pair (Figure 41H–J).

Bilobed prostomium with anterior lobes wide, subtriangular, eyes rather small, anterior pair anterolateral, anterior to greatest width of prostomium, slightly larger than posterodorsal pair; median antenna with large ceratophore in anterior notch, style missing; ceratophores of lateral antennae inserted ventrally, converging midventrally, with styles short, subulate, papillate; palps stout, tapered; tentaculophores lateral to prostomium, each with 0–1 seta on inner side; tentacular cirri longer than prostomium, with short papillae and filamentous tips, dorsal ones longer than ventral ones (Figure 41A). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri, similar to tentacular cirri (Figure 41A,B); notosetae similar to those following; neurosetae more slender than those following, otherwise similar (Figure 41C).

Biramous parapodia with notopodium short, rounded, with projecting acicular lobe on lower side; longer neuropodium with subconical presetal acicular lobe continuing distally as subtriangular supraacicular process and shorter, rounded postsetal lobe (Figure 41B,D,E). Short and long notosetae forming radiating bundle, stouter than neurosetae, with double row of minute spines along one border and tapered blunt bare tip (Figure 41F). Neurosetae forming fan-shape bundle, rather slender, upper ones (5 or so) with longer spinose region and long, bare, slender, slightly knobbed tip; middle ones with shorter spinose region and bifid tip with extra long secondary tooth; lower ones (10 or so) with shorter spinose region and slightly knobbed bare tip (Figure 41C,G). Dorsal cirri with cylindrical cirrophore with style long, papillate, tapering distally and extending beyond tips of neurosetae; dorsal tubercles nodular; ventral cirri short, subulate, papillate (Figure 41E).

ETYMOLOGY.—The species is named for Edith and Cyril Berkeley, in recognition of their early work on the polychaetes of British Columbia.

DISTRIBUTION.—Eastern Pacific Ocean from British Columbia, in 183 meters.

Malmgreniella bansei, new species

FIGURE 42

Malmgrenia lunulata.—Pettibone, 1953:25 [part].—Banse et al., 1968:525 [part].—Lie, 1968:317, 370 [part]. [Not Delle Chiaje, 1830.]

MATERIAL EXAMINED.—EASTERN PACIFIC OCEAN: Washington: Puget Sound, Carr Inlet, off Green Point, 93 m, mud, 2 Aug 1938, M. Pettibone, collector, paratype (USNM 24932, as M. lunulata by Pettibone, 1953). Puget Sound, Carr Inlet, near Gertrude Island, between McNeil and Fox Islands, 128 m, mud, 25 Jun 1939, M. Pettibone, collector, holotype (USNM 24930). Puget Sound, Case Inlet, between Johnson Point and Heron Island, 47°13′N, 122°49′W, 47–89 m, sta 7, haul 4, 3 May 1963, U. Lie, collector, paratype (USNM 36445, as M. lunulata form C by Banse et al., 1968).

FIGURE 41.—Malmgreniella berkeleyorum, new species, holotype (USNM 55039): A, dorsal view of anterior end, style of median antenna missing; B, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; C, upper, middle, and lower neurosetae from same, with detail of tips; D, right elytrigerous parapodium, anterior view, acicula dotted; E, right cirrigerous parapodium, posterior view; F, long and short notosetae; G, upper, middle, and lower neurosetae, with detail of tips; H, right 1st elytron from segment 2; I, right 2nd elytron from segment 4; J, right middle elytron, with detail of microtubercles. (Scales = 0.5 mm for A; 0.5 mm for B,D,E; 0.1 mm for C,F,G; 0.5 mm for H–J.)

DESCRIPTION.—Holotype from Carr Inlet, male with sperm, 15.5 mm long, 4 mm wide including setae, and 34 segments. Paratype from Carr Inlet with posterior end regenerating, 28 segments, last 2 very small, 8 mm long, 4 mm wide. Paratype from Case Inlet, female with large eggs, 7 mm long, 3 mm wide, with 33 segments.

Body flattened, tapering slightly anteriorly and posteriorly. Elytra 15 pairs, on usual segments, subreniform, opaque, with branching “veins” and large group of microtubercles on anterior part; light brown pigmentation in form of spots over areas of attachment to elytrophores and crescent-shape areas near medial and posterior borders (Figure 42B,C).

Bilobed prostomium with anterior lobes wide, subtriangular, with slight indication of peaks; ceratophore of median antenna bulbous, in anterior notch; style slightly longer than prostomium, with short papillae and filamentous tip; ceratophores of lateral antennae inserted ventrally, converging midventrally, with styles short, subulate, papillate; palps stout, tapered; eyes moderate in size, anterior pair slightly anterior to greatest width of prostomium, slightly larger than posterodorsal pair; tentaculophores lateral to prostomium, each with 1–2 setae on inner side, dorsal and ventral tentacular cirri similar to median antenna; light pigmentation on ceratophores, tentaculophores, and styles of tentacular cirri (Figure 42A). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 42A,D); notosetae similar to following; neurosetae more slender than those following, all with secondary tooth (Figure 42E).

Biramous parapodia with short, rounded notopodium with projecting acicular lobe on lower side; longer neuropodium with subconical presetal acicular lobe with projecting supraacicular process tapering to curved tip, and shorter, rounded postsetal lobe (Figure 42D,F,G). Notosetae numerous, forming radiating bundle, of several lengths, slightly stouter than neurosetae, clear, with double row of minute spines along one border and blunt tapered tips (Figure 42H). Neurosetae numerous, forming long fan-shape bundle; upper ones with longer spinose region, upper few with entire bare tip or with small secondary tooth; middle ones with shorter spinose region and slightly hooked tip with secondary tooth; lower few with entire tip (Figure 42I). Dorsal cirri with cylindrical cirrophore and style extending slightly beyond tips of neurosetae, with scattered short papillae and filamentous tip; dorsal tubercles nodular; ventral cirri short, tapered, with short papillae (Figure 42G). Pygidium with anus medial to posterior pair of parapodia.

ETYMOLOGY.—The species is named for Karl Banse, in recognition of his many contributions to the study of polychaetes and who contributed specimens used in this study.

DISTRIBUTION.—Eastern Pacific Ocean, from Washington, Puget Sound, in 47–128 meters.

Malmgreniella liei, new species

FIGURE 43

Malmgrenia lunulata.—Banse et al., 1968:525 [part].—Lie, 1968:295, 370, 371 [part], [Not Delle Chiaje, 1830.]

MATERIAL EXAMINED.—EASTERN PACIFIC OCEAN: Washington: Puget Sound, Bainbridge Island, off Port Madison, 47°44′N, 122°32′W, 10–18 m, fine sand mixed with silt, sta 4, haul 10, 9 Jan 1963, U. Lie, collector, holotype (USNM 36446) and 2 paratypes (USNM 55058) (as M. lunulata form D by Banse et al., 1968).

DESCRIPTION.—Holotype, male with sperm, 10 mm long, 4 mm wide including setae, and 37 segments. Paratypes consisting of fragments of 2 anterior and one posterior ends. Body flattened, tapering slightly anteriorly and posteriorly, without color. Elytra 15 pairs, on usual segments, subreniform, delicate, opaque, denser around region of attachment, with group of microtubercles on anterior part of some elytra, absent on first and posterior elytra (Figure 43G).

Bilobed prostomium with anterior lobes wide, subtriangular, with distinct peaks; eyes not visible; ceratophore of median antenna large, bulbous, in anterior notch, with papillate style shorter than prostomium; ceratophores of lateral antennae inserted ventrally, converging midventrally, with papillate styles half as long as median antenna; palps stout, tapered; tentaculophores lateral to prostomium, each with 1–2 setae on inner side; tentacular cirri similar to and slightly longer than median antenna (Figure 43A). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 43A,B). Setae similar to those of following parapodia.

Biramous parapodia with notopodium shorter, rounded, with projecting acicular lobe on lower side; longer neuropodium with subconical presetal acicular lobe continuous with tapering supraacicular process and shorter rounded postsetal lobe (Figure 43B–D). Notosetae numerous, forming radiating bundle, slightly stouter than neurosetae, clear, with faint double row of spines along one side and blunt tapered tip (Figure 43E). Neurosetae numerous, forming fan-shape bundle, each with bifid split tip, upper ones with slightly longer spinose region and more prominent spines, lower ones with less prominent spines, appearing smooth (Figure 43F). Dorsal cirri with cirrophores stout, cylindrical, with papillate styles tapering distally and extending to tips of neurosetae or slightly beyond; dorsal tubercles nodular, ventral cirri short, subulate, papillate (Figure 43D).

FIGURE 42.—Malmgreniella bansei, new species, holotype (USNM 24930): A, dorsal view of anterior end; B, right 2nd elytron from segment 4, C, right middle elytron, with detail of microtubercles; D, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; E, upper and lower neurosetae from same, with detail of tips; F, right elytrigerous parapodium, anterior view, acicula dotted; G, right cirrigerous parapodium, posterior view; H, long and short notosetae; I, upper, middle, and lower neurosetae, with detail of tips. (Scales = 0.5 mm for A; 0.5 mm for B,C; 0.2 mm for D,F,G; 0.1 mm for E,H,I.)

FIGURE 43.—Malmgreniella liei, new species, holotype (USNM 36446): A, anterior end, dorsal view; B, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; C, right middle elytrigerous parapodium, anterior view, acicula dotted; D, right cirrigerous parapodium, posterior view; E, long and short notosetae; F, upper, middle, and lower neurosetae, with detail of tips; G, left 2nd elytron from segment 4, with detail of microtubercles. (Scales = 0.5 mm for A; 0.2 mm for B–D; 0.1 mm for E,F; 0.2 mm for G.)

ETYMOLOGY.—The species is named for Ulf Lie, who collected the specimens as a part of his detailed study on the benthic infauna in Puget Sound.

DISTRIBUTION.—Eastern Pacific Ocean, from Washington, Puget Sound, in 10–18 meters.

Malmgreniella scriptoria (Moore, 1910), new combination

FIGURE 44

Harmothoe scriptoria Moore, 1910:344, pl. 28: figs, 13–17.—Hartman, 1955:174; 1961:49; 1966:402; 1968:85, figs. 1–5.—Loi, 1980:124.

Malmgrenia lunulata.—Pettibone, 1953:25 [part], pl. 9: figs. 73–76; pl. 10: figs. 78, 79, 83, 84, 86; pl. 11: figs. 88–91.—Banse et al., 1968:525 [part].—Lie. 1968:271, 279, 370 [part; sta 2]. [Not Delle Chiaje, 1830.]

MATERIAL EXAMINED.—EASTERN PACIFIC OCEAN: California: Monterey Bay, off Point Pinos Light, Albatross sta 4460, 12 May 1904, 101–305 m, green mud and gravel, from ambulacral groove of spatangoid, holotype of Harmothoe scriptoria (USNM 17156). Washington: Puget Sound, off Skiff Point, Bainbridge Island, 254 m, mud with heart urchin Briaster townsendi, 17 Jul and 2 Aug 1938, M. Pettibone, collector, 14 specimens (USNM 24928, as M. lunulata by Pettibone, 1953). Puget Sound, Point Richmond, 73 m, gravel, mud and shell, 4 Jul 1940, M. Pettibone, collector, 7 specimens (USNM 24927, as M. lunulata by Pettibone, 1953). Puget Sound, off Port Madison, 47°42′N, 122°26′W, 172–216 m, fine soft mud, with many Briaster townsendi Agassiz, sta 2, haul 7, 21 May 1963, U. Lie, collector, 4 specimens (USNM 36443, as M. lunulata form B by Banse et al., 1968).

DESCRIPTION.—Holotype, female with eggs, in 2 pieces, 16 mm long, 5.5 mm wide including setae, and 39 segments, last one minute. Specimens from Puget Sound area off Skiff Point 10.5–14 mm long, 3.5–5 mm wide, with 34–38 segments. Specimens from Point Richmond 18–22 mm long, 5–6 mm wide, with 37–40 segments.

Body flattened, tapering slightly anteriorly and posteriorly. Elytra 15 pairs, on usual segments, oval to subreniform, thin, delicate, smooth, except for small group of microtubercles on anterior part (absent on 1st and some posterior elytra) and scattered round micropapillae (Figure 44B–E). Elytra opaque, with complete or incomplete circle of brown color on 1st elytral pair, following elytra with color confined to semicircular area on posterior and medial sides, plus pigmented spot over place of elytral attachment (Figure 44B–D; Moore, 1910, pl. 28: fig. 15; Pettibone, 1953, pl. 9: fig. 73; pl. 10: figs. 78, 79).

Bilobed prostomium with anterior lobes truncate or wide subtriangular, without distinct peaks; eyes rather small, anterolateral pair slightly larger than posterodorsal pair; ceratophore of median antenna large, bulbous, with style longer than prostomium; ceratophores of lateral antennae inserted terminoventrally, converging midventrally, with styles less than half as long as median antenna; palps stout, tapered; tentaculophores lateral to prostomium with 0–1 seta on inner side; dorsal and ventral tentacular cirri similar to median antenna; styles of antennae and tentacular cirri with minute papillae and pigmented on basal half (Figure 44A; Moore, 1910, pl. 28: fig. 13; Pettibone, 1953, pl. 9: figs. 75, 76). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 44A,F); notosetae similar to those following; neurosetae more slender than those following, with bulbous tips, upper ones each with secondary tooth (Figure 44G).

Biramous parapodia with notopodium short, rounded, with projecting acicular lobe on lower side; longer neuropodium with subconical presetal acicular lobe with wide supraacicular process and shorter, rounded postsetal lobe (Figure 44F,H,I; Moore, 1910, pl. 28: fig. 14; Pettibone, 1953, pl. 10: figs. 83, 84). Notosetae moderate in number, about as stout as neurosetae, short and longer, with faint spinose rows, tapering to blunt bare tip (Figure 44J; Moore, 1910, pl. 28: fig. 17; Pettibone, 1953, pl. 10: fig. 86). Neurosetae moderate in number, upper ones with longer spinose region, upper few with entire tip, rest with distinct secondary tooth; middle ones with shorter spinose region and bifid tip; some lower ones with or without small secondary tooth (Figure 44K; Moore, 1910, pl. 28: fig. 16; Pettibone, 1953, pl. 11: figs. 88–91). Dorsal cirri with cylindrical cirrophore and long style, extending beyond tips of neurosetae, with minute papillae and pigmentation on basal half; dorsal tubercles nodular; ventral cirri short, subulate (Figure 44I; Pettibone, 1953, pl. 10: fig. 83). Pygidium with anus medial to last pair of small parapodia, with pair of anal cirri.

BIOLOGY.—The type specimen of M. scriptoria from Monterey Bay was collected from the ambulacral groove of a spatangoid. In the Puget Sound area, specimens were found by M. Pettibone occupying the ambulacral groove of the heart urchin Briaster townsendi Agassiz, where they were well protected by the ambulacral spines of the host. In the revision of Briaster by Markov (1989:87), Briaster townsendi Agassiz, 1898 is referred to as B. latifrons Agassiz, 1898.

DISTRIBUTION.—Eastern Pacific Ocean, from Washington, Puget Sound to southern California, in 42–373 meters (see Hartman, 1968).

FIGURE 44.—Malmgreniella scriptoria, specimen from Puget Sound off Bainbridge Island (USNM 24928): A, dorsal view of anterior end; B, right 1st elytron from segment 2; C, right 2nd elytron from segment 4, with detail of microtubercles; D, right middle elytron, with detail of microtubercles; E, right 15th elytron from segment 32, with detail of micropapillae; F, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; G, upper neurosetae from same, with detail of tips of upper and lower neurosetae; H, right middle elytrigerous parapodium, anterior view, acicula dotted; I, right cirrigerous parapodium, posterior view; J, long and short notosetae; K, upper, middle, and lower neurosetae, with detail of tips. (Scales = 0.5 mm for A; 0.5 mm for B–E; 0.2 mm for F,H,I; 0.1 mm for G,J,K.)

Malmgreniella macginitiei, new species

FIGURE 45

Harmothoe lunulata.—Berkeley and Berkeley, 1941:21 [part; not Delle Chiaje, 1830].

Malmgeniella lunulata.—Pettibone, 1953:25 [part], pl. 9: fig. 77; pl. 10: figs. 80–82,85, 87; pl. 11: figs. 92–95 [not Delle Chiaje, 1830].

Harmothoe of. lunulata.—Kudenov, 1975a:42, fig. la–g [not Delle Chiaje, 1830].

MATERIAL EXAMINED.—EASTERN PACIFIC OCEAN: Central California: Tomales Bay, sandy mud flat, in burrow with ghost shrimp Callianassa californiensis Dana, 28 May 1941, F. Pitelka and R. Paulson, collectors, holotype (USNM 24933). Tomales Bay, Lawson's Flat, in tube of maldanid Axiothella rubrocincta (Johnson), 31 Jan 1970, J.K. Kudenov, collector, paratype (AHF 1119, as H. cf. lunulata by Kudenov, 1975a). Southern California: Newport Bay, Mar 1933, in burrow with Callianassa californiensis, G.E. MacGinitie, collector, paratype (USNM 55059, as H. lunulata by Berkeley and Berkeley, 1941). Santa Monica Bay, 60 m, on arm of Amphiodia urtica (Lyman), M/V Marine Surveyor, from G. Hendler, 1 specimen (USNM 133564).

DESCRIPTION.—Holotype from Tomales Bay, male with sperm, 13 mm long, 4.5 mm wide including setae, with 35 segments, last one minute. Paratype from same area (AHF 1119), female with eggs, 9 mm long, 3.5 mm wide, with 34 segments. Paratype from Newport Bay (USNM 55059), female with eggs, 14 mm long, 5 mm wide, with 37 segments. Body darkly pigmented or with dorsal and ventral transverse dark bands in posterior region. Elytra 15 pairs, on usual segments, rounded to subreniform, with anterior group of microtubercles on some anterior elytra; central part of elytra showing compartments, with or without pigment; pigmentation forming C-shape pattern on central, median, and posterior areas; some scattered round micropapillae on surface and border (Figure 45C,D); Pettibone, 1953, pl. 10: figs. 80–82; Kudenov, 1975a, fig. 1c).

Bilobed prostomium with anterior lobes wide, subconical, with distinct peaks; eyes moderate in size, anterior pair anterolateral, anterior to greatest width of prostomium, slightly larger than posterodorsal pair; ceratophore of median antenna large, bulbous, in anterior notch, style missing; ceratophores of lateral antennae inserted ventrally, converging midventrally, styles short, subulate, with short papillae; palps stout, tapered; tentaculophores lateral to prostomium, each with single seta on inner side; dorsal and ventral tentacular cirri longer than prostomium, papillate (Figure 45A,B; Pettibone, 1953, pl. 9: fig. 77; Kudenov, 1975a, fig. 1a). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri, similar to tentacular cirri (Figure 45A,B,E); notosetae similar to those following; neurosetae more slender than those following, upper and lower ones with slender entire tips, middle ones with bifid tips (Figure 45F).

Biramous parapodia with notopodium short, rounded, with projecting acicular lobe on lower side; larger neuropodium with subconical presetal acicular lobe with prominent subtriangular supraacicular process and shorter rounded postsetal lobe (Figure 45E,G,H; Pettibone, 1953, pl. 10: fig. 85; Kudenov, 1975a, fig. 1b). Notosetae numerous, forming radiating bundle, of several lengths, about as stout as neurosetae, clear, transparent, with double row of faint spines and blunt tapered tip (Figure 45I; Pettibone, 1953, pl. 10: fig. 87; Kudenov, 1975a, fig. 1d). Neurosetae numerous, forming fan-shape bundle, upper ones with longer spinose region, tapering to slender pointed tip, with or without slight indication of secondary tooth; middle ones with shorter spinose region and bifid hooked tip; lower ones with entire, slightly hooked tip (Figure 45J; Pettibone, 1953, pl. 11: figs. 92, 93; Kudenov, 1975, fig. 1e–g). Dorsal cirri with cylindrical cirrophore, style extending to about tips of neurosetae, with scattered short papillae; dorsal tubercles nodular; ventral cirri short, subulate, papillate (Figure 45H). Pygidium with anus medial to small parapodia of last segment, anal cirri lacking.

BIOLOGY.—Malmgreniella macginitiei was found living commensally in the burrows of the ghost shrimp Callianassa californiensis Dana by F. Pitelka and R. Paulson in Tomales Bay and by G. MacGinitie in Newport Bay. In Tomales Bay, J. Kudenov collected specimens occupying the tubes of the maldanid polychaete Axiothella rubrocincta (Johnson). From Santa Monica Bay, the specimen was found by G. Hendler on the arm of the ophiuroid Amphiodia urtica (Lyman).

ETYMOLOGY.—The species is named for George E. MacGinitie, collector of one of the paratypes and outstanding observer, as indicated by his book on the Natural History of Marine Animals (1949).

DISTRIBUTION.—Eastern Pacific Ocean, central and southern California. Intertidal to 60 meters.

Malmgreniella sanpedroensis, new species

FIGURE 46

Harmothoe lunulata.—Hartman, 1961:49 [part]; 1966:402 [part]. [Not Delle Chiaje, 1830.]

MATERIAL EXAMINED.—EASTERN PACIFIC OCEAN: California: San Pedro Basin, 6.2 mi south of Pt. Ferman Light, 33°36′N, 118°18′W, 393 m, clay and mud, Velero IV sta 2306, 25 Apr 1953, holotype (AHF 1305).

FIGURE 45.—Malmgreniella macginitiei, new species, A, paratype (AHF 1119); B–J, holotype (USNM 24933): A, dorsal view of anterior end, styles of median antenna, left dorsal and ventral and right dorsal tentacular cirri missing; B, dorsal view of anterior end, median and lateral antennae, palps, and tentacular cirri missing; C, left 2nd elytron from segment 4, with detail of microtubercles; D, left middle elytron, with detail of pigmented area and micropapillae along border and surface; E, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; F, upper, middle, and lower neurosetae from same, with detail of tips; G, right middle elytrigerous parapodium, anterior view, acicula dotted; H, right cirrigerous parapodium, posterior view; I, long and short notosetae; J, upper, middle, and lower neurosetae, with detail of tips. (Scales = 0.5 mm for A,B; 0.5 mm for C,D; 0.2 mm for E,G,H; 0.1 mm for F,I,J.)

DESCRIPTION.—Holotype, female with eggs, with posterior end incomplete, 14+ mm long, 5 mm wide including setae, and 33+ segments. Body without color. Elytra 15 pairs, on usual segments, oval and subreniform, delicate, with “veins”; large group of microtubercles on anterior part of elytra, absent on first pair, and smaller group on more posterior elytra; light brownish pigmentation in form of spots over area of attachment to elytrophores and C-shape masses on medial and posterior areas; exposed parts of elytra with scattered rusty red spots, giving speckled appearance (Figure 46B–E).

Bilobed prostomium with anterior lobes truncate, without distinct peaks; eyes moderate in size, anterior pair anteroventral, anterior to greatest width of prostomium, larger than posterodorsal pair; ceratophore of median antenna bulbous, in anterior notch, with papillate style about length of prostomium; ceratophores of lateral antennae inserted terminoventrally, with short, subulate, papillate styles; palps stout, tapered; tentaculophores lateral to prostomium, left one with up to 10 setae on inner side, 2 on slightly deformed right side (perhaps regenerating); tentacular cirri similar to median antenna, dorsal one slightly longer than ventral one; rusty red pigment spots on styles of lateral antennae and tentacular cirri (Figure 46A). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 46A,F); notosetae similar to those following; neurosetae more slender than those following, lower ones with rather long bare bulbous tips.

Biramous parapodia with notopodium short, rounded, with projecting acicular lobe on lower side; longer neuropodium with subconical presetal acicular lobe with subtriangular supraacicular process and shorter rounded postsetal lobe (Figure 46F–H). Notosetae numerous, forming radiating bundle, of several lengths, much stouter than neurosetae, clear, with faint, close-set spinose rows along one border, and short tapered bare tips (Figure 46I). Neurosetae numerous, forming fan-shape bundle; upper ones with longer spinose region, tapering to sharply pointed bare tips; middle ones with shorter spinose region and bifid tips; lower few with entire tips (Figure 46J). Cirrophore of dorsal cirri cylindrical, with bulbous glandular area on posterior side; style with short clavate papillae, rusty red pigment spots, and filamentous tip extending to tip of neuropodia; dorsal tubercles nodular; ventral cirri short, subulate, papillate (Figure 46G).

ETYMOLOGY.—The species is named for the collecting site, the San Pedro Basin.

DISTRIBUTION.—Eastern Pacific Ocean, southern California, San Pedro Basin, in 393 meters.

Malmgreniella baschi, new species

FIGURE 47

MATERIAL EXAMINED.—EASTERN PACIFIC OCEAN: California: Santa Catalina Island, Big Fisherman Cove, 33°27′N, 118°29′W, sand, 8–30 m, associated with ophiuroid Ophiopsila californica A.H. Clark, L.W. Basch, collector: no 5C, 20 Jul 1982, paratype (USNM 133582); no 6 and 7, 28 Jul 1982, 2 paratypes (USNM 133583; no 8, 24 Nov 1982, paratype (LACM); no 10, 9 Apr 1983, 2 paratypes (USNM 133584); no 11, 10 Apr 1983, 2 paratypes (USNM 133585); no 12, 16 Apr 1983, holotype (USNM 133581).

DESCRIPTION.—Holotype, male with sperm, 14 mm long, 4.5 mm wide including setae, with 38 segments and 15 pairs of elytra; figured paratype (USNM 133582), incomplete male with sperm. 9+ mm long, 5 mm wide, with 28+ segments; paratype (LACM), female with eggs, 9 mm long, 3.5 mm wide, with 37 segments.

Black pigmented bands on posterior 4 segments of dorsum and posterior 8 segments of ventrum. Elytra 15 pairs, oval to subreniform, delicate, opaque, with micropapillae on posterior and lateral borders and small group of microtubercles on anterior part of few elytra; some light to dark mottled pigmentation forming spots over place of attachment to elytrophores and larger C-shape areas more posteriorly (Figure 47C,D). Bilobed prostomium with anterior lobes subtriangular, without distinct peaks, with mottled pigmentation; 2 pairs of eyes, anterior pair larger, anteroventral, anterior to widest part of prostomium and not visible dorsally, smaller posterior pair near posterior border; ceratophore of median antenna in anterior notch, with style about twice as long as prostomium, with slender tip; lateral antennae with ceratophores inserted ventrally, with short subulate styles; palps long, tapered; tentaculophores without setae, with brown pigment spots; dorsal and ventral tentacular cirri similar to median antenna (Figure 47A,B). Second segment with biramous parapodia and long ventral buccal cirri similar to tentacular cirri.

Biramous parapodia with notopodia shorter than neuropodia, rounded, with projecting acicular lobe on lower side; neuropodium with presetal acicular lobe subconical, with digitiform supraacicular process; postsetal lobe shorter, rounded (Figure 47E,F). Notosetae numerous, forming radiating bundle, short to long, about as stout as neurosetae, tapering to short bare tips, with faint close-set spinose rows (Figure 47G). Neurosetae moderate in number, forming fan-shape bundle; supraacicular neurosetae with long spinose region with prominent spines, upper few tapering to pointed tips, some lower ones with slight indication of secondary tooth (Figure 47H); subacicular neurosetae with shorter spinose region, less prominent spines, and slightly hooked, bare tips (Figure 47I). Dorsal cirri with prominent cylindrical cirrophore, ciliated on upper side, with black pigment on lower side; style long, flattened, with scattered micropapillae, and slender tip extending beyond tips of neurosetae; dorsal tubercles nodular; ventral cirri short, tapered, with micropapillae (Figure 47F). Pygidium with dorsal anus medial to posterior parapodia, with pair of long anal cirri.

FIGURE 46.—Malmgreniella sanpedroensis, new species, holotype (AHF 1305):—A, dorsal view of anterior end, right side slightly deformed, anterior pair of eyes not visible dorsally (dotted); B, right 1st elytron from segment 2; C, right 2nd elytron from segment 4, with detail of microtubercles; D, right middle elytron; E, right posterior elytron; F, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; G, right cirrigerous parapodium, posterior view; H, right elytrigerous parapodium, anterior view, acicula dotted; I, long and short notosetae; J, upper, middle, and lower neurosetae, with detail of tips. (Scales = 0.5 mm for A; 0.5 mm for B–E; 0.2 mm for F–H; 0.1 mm for I,J.)

FIGURE 47.—Malmgreniella baschi, new species, A, holotype (USNM 133581), B–I, paratype (USNM 133582): A, dorsal view of anterior end, anterior pair of eyes anteroventral (dotted), left ventral tentacular cirrus missing; B, same, left palp short, regenerating; C, left elytron with few marginal micropapillae; D, left elytron, with detail of microtubercles; E, right elytrigerous parapodium, anterior view, acicula dotted; F, right cirrigerous parapodium, posterior view; G, long and short notosetae; H, supraacicular neurosetae, with detail of tip; I, subacicular neurosetae. (Scales = 0.5 mm for A,B; 0.5 mm for C,D; 0.5 mm for E,F; 0.1 mm for G–I.)

BIOLOGY.—The specimens were collected at night by L.W. Basch from the discs of the very large and brilliantly bioluminescent ophiuroid Ophiopsila californica A.H. Clark, in 8–30 meters off Santa Catalina Island. According to the collector, the ophiuroids dwell in sand and in tubes of Chaetopterus variopedatus (Renier).

ETYMOLOGY.—The species is named for Lawrence V. Basch, the collector and donor of the specimens.

DISTRIBUTION.—East Pacific Ocean, off southern California, in 8–30 meters.

Malmgreniella pacifica (Monro, 1928), new combination

FIGURE 48

Harmothoe lunulata var. pacifica Monro, 1928a:559 [part; see Fauchald, 1977:5, 16].

Harmothoe lunulata.—Hartman, 1944:246 [part; H. lunulata var. pacifica Monro in synonymy]. [Not Delle Chiaje, 1830.]

Harmothoe lunulata.—Pettibone, 1953:25 [part; H. lunulata var. pacifica Monro in synonymy]. [Not Delle Chiaje, 1830.]

MATERIAL EXAMINED.—EASTERN PACIFIC OCEAN: Galapagos: James Island, James Bay, 9–11 m, clean sand with several species of “weed,” C. Crossland, collector, “St. George” Pacific and Panamanian Expedition 1923–1924, 4 syntypes (BMNH 1928.9.14.73–75; USNM 54748). Fernandina Island, along shore of Espinosa Point, 00°15′S, 91°26′W, 1–3 m, associated with algal-covered rocks, R/V Anton Bruun cruise 16, 25 May 1966, S.A. Earle, collector, 2 specimens (USNM 71735). Nameless Island, intertidal, 22 Feb 1978, W.D. Hope, collector, 1 specimen (USNM 75368).Panama: Panama City, Paitilla Beach, 08°58′N, 79°30′W, sta 25–1, 26 Apr 1971, M.L. Jones, collector, 1 specimen (USNM 71736). Recife de Casa de Putas, hard rock substrate at low tide, sta 92–1, 18 Apr 1972, M.L. Jones, collector, 1 specimen (USNM 71737).

REMARKS.—According to Fauchald (1977:5, 16), the syntype of H. lunulata var. pacifica from Taboga Island, Panama, deposited in the Zoological Museum, University of Copenhagen, is an unidentifiable hesionid species of Gyptis.

FIGURE 48.—Malmgreniella pacifica, syntype of Harmothoe lunulata var. pacifica from the Galapagos Islands (BMNH 1928.9.14.73–75): A, dorsal view of anterior end, anterior pair of eyes not visible dorsally (dotted); B, right 2nd elytron from segment 4, with detail of microtubercles; C, right 5th elytron from segment 9; D, right 13th elytron from segment 26, with detail of micropapillae; E, right elytrigerous parapodium, anterior view, acicula dotted; F, right cirrigerous parapodium, posterior view; G, long and short notosetae; H, upper and middle neurosetae, with detail of tips. (Scales = 0.5 mm for A; 0.5 mm for B–D; 0.2 mm for E,F; 0.1 mm for G,H.)

DESCRIPTION.—Four syntypes from Galapagos 9–13 mm long, 2–2.5 mm wide including setae, with 36–37 segments. Body elongated, slender, subrectangular. Elytra 15 pairs, on usual segments, rounded, subrectangular, and elongate oval, with group of microtubercles on anterior part (absent on first and more posterior elytra), and scattered rounded micropapillae; mottled pigmentation in form of spot on scar of attachment to elytrophore and irregular mass on medial part (Figure 48B–D).

Bilobed prostomium with anterior lobes wide, subtriangular, with cephalic peaks; eyes moderate in size, anterior pair anteroventral, anterior to greatest width, slightly larger than posterodorsal pair; ceratophore of median antenna in anterior notch, with papillate style longer than prostomium; ceratophores of lateral antennae inserted ventrally, with short subulate papillate styles; palps stout, tapered, minutely papillate; tentaculophores lateral to prostomium, each with 2 setae on inner side; tentacular cirri similar to median antenna; scattered pigmentation on prostomium and appendages (Figure 48A). Segment 2 with 1st pair of large elytrophores, biramous parapodia, and long ventral buccal cirri, similar to tentacular cirri (Figure 48A).

Biramous parapodia with small rounded notopodium with short projecting acicular lobe on lower side; larger neuropodium with subconical presetal acicular lobe, with digitiform supraacicular process and shorter rounded postsetal lobe (Figure 48E,F). Notosetae not extending to tip of neuropodium, of several lengths, subequal in width to neurosetae, with rather prominent spinose rows and tapered blunt tip (Figure 48G). Neurosetae with prominent spinose rows, somewhat longer spinose region on upper neurosetae, with bifid split tip; lower neurosetae with slender secondary tooth or entire (Figure 48H). Dorsal cirri with cirrophore long, cylindrical, with papillate style extending beyond tips of neurosetae; dorsal tubercles low, nodular; ventral cirri short, subulate, papillate (Figure 48F). Pygidium with anus medial to last pair of small parapodia, anal cirri missing.

DISTRIBUTION.—Central Eastern Pacific Ocean, Galapagos, Panama, intertidal to 11 meters.

Paragattyana, new genus

TYPE SPECIES.—Malmgrenia micropoides Augener, 1918. Gender: feminine.

DIAGNOSIS.—Segments up to 40. Elytra and elytrophores 15 pairs, on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, and 32. Elytra with or without anterior group of microtubercles, and micropapillae. Prostomium bilobed, anterior lobes subtriangular, without distinct cephalic peaks, with 3 antennae, 2 palps, and 2 pairs of eyes; lateral antennae with distinct ceratophores inserted ventrally. Tentaculophores of first segment lateral to prostomium, each with aciculum, with or without few setae on inner side, and pair of dorsal and ventral tentacular cirri. Second or buccal segment with first pair of elytrophores, biramous parapodia, and long ventral buccal cirri lateral to ventral mouth. Biramous parapodia with smaller notopodia and larger neuropodia, both rami with projecting acicular lobes; presetal acicular lobe of neuropodium longer, with supraacicular process; postsetal lobe shorter, rounded. Notosetae with spinose rows, of 2 types; shorter, with blunt tips and longer, with capillary tips. Neurosetae with spinose rows and bifid hooked tips or some lower ones with entire tips. Dorsal cirri on nonelytrigerous segments with cylindrical cirrophores and long styles; dorsal tubercles slightly developed; ventral cirri short. Pygidium with pair of anal cirri.

ETYMOLOGY.—From the Greek Para (besides) plus the genus Gattyana McIntosh.

COMPARISONS.—Among the genera of Harmothoinae, Paragattyana agrees with Gattyana in having two types of notosetae; upper shorter ones with blunt tips and lower longer ones ending in capillary tips. The latter genus differs from the new genus in having neurosetae with entire tips, not bifid; the elytral surfaces are covered with numerous microtubercles and some macrotubercles; and the prostomium has distinct cephalic peaks.

The genus is established for two species, both from West Africa: P. micropoides (Augener), new combination and P. intesi, new species.

Paragattyana micropoides (Augener, 1918), new combination

FIGURE 49

Malmgrenia micropoides Augener, 1918:146, pI. 2: fig. 21; pI. 3: fig. 43,text-fig. 8a–c.—Intes and Le Loeuff, 1975:271.—Kudenov, 1975b:79; 1977:95.—Hanley, 1987:160 [under incertae sedis].

MATERIAL EXAMINED.—NORTH ATLANTIC OCEAN: West Africa: Spanish Guinea, Bata, A. Hupfer, collector, holotype of Malmgrenia micropoides (ZMH V-595).

DESCRIPTION.—Holotype, female with eggs, with 36 segments, 12 mm long, and 4 mm wide including setae. Body delicate, flattened, tapering slightly anteriorly and more so posteriorly. Elytra 15 pairs, on usual segments, large, covering dorsum, round to oval; some anterior elytra with group of microtubercles on anterior part; scattered delicate, rather long papillae along external border and on surface; some darker pigmentation on central part (Figure 49B–D; Augener, 1918, pl. 2: fig. 21).

FIGURE 49 (opposite page).—Paragattyana micropoides, holotype of Malmgrenia micropoides (ZMH V-595): A, dorsal view of anterior end, style of median antenna missing; B, right 1st elytron from segment 2; C, right 2nd elytron from segment 4, with detail of microtubercles and papillae; D, right middle elytron; E, right elytrigerous parapodium from segment 2, anterior view, acicular dotted; F, right middle cirrigerous parapodium, posterior view; G, right middle elytrigerous parapodium, anterior view, acicula dotted; H, long and short notosetae, with detail of tip; I, upper, middle, and lower neurosetae, with detail of tips. (Scales = 0.5 mm for A; 0.5 mm for B–D; 0.2 mm for E–G; 0.1 mm for H,I.)

Bilobed prostomium with anterior lobes subtriangular, without peaks; eyes rather small, anterodorsal pair anterior to widest part of prostomium, larger than posterolateral pair; ceratophore of medial antenna oval, in anterior notch of prostomium, style missing (about twice as long as lateral antennae, according to Augener); ceratophores of lateral antennae inserted ventrally, with short, tapered styles; palps stout, long, tapered; tentaculophores lateral to prostomium, each with 0–1 seta on inner side; tentacular cirri almost as long as palps, dorsal ones slightly longer than ventral ones (Figure 49A). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri similar to tentacular cirri (Figure 49A,E); notosetae similar to those following; neurosetae more slender than those following, with long spinose region, tapering to slender sharp tip.

Biramous parapodium with notopodium much shorter than neuropodium, conical, with projecting acicular lobe on lower side; larger neuropodium with subconical presetal acicular lobe with digitiform supraacicular process; postsetal lobe shorter, rounded (Figure 49E–G; Augener, 1918, pl. 3: fig. 43). Notosetae rather numerous, subequal in width to neurosetae, with numerous spinose rows; upper shorter and longer ones ending in blunt, slightly pointed tips; longest lower ones tapering distally to long capillary tips (Figure 49H; Augener, 1918, text-fig. 8a,b). Upper neurosetae with longer spinose regions and bifid hooked tips; middle and lower neurosetae with shorter spinose regions and bifid tips or secondary tooth lacking on few lower ones (Figure 49I; Augener, 1918, text-fig. 8C). Dorsal cirri with cylindrical cirrophores, wider basally; styles long, extending beyond tips of neurosetae, with scattered minute papillae; dorsal tubercles indistinct; ventral cirri short, tapered, with minute papillae (Figure 49F). Pygidium with anus medial to posterior pair of small parapodia, with pair of anal cirri.

DISTRIBUTION.—North Atlantic Ocean, West Africa.

REMARKS.—The specimen from off Annobon in the Gulf of Guinea, questionably referred to this species by Monro (1930:45), was examined (BMNH 1930.10.8.225); it cannot be positively identified because it consists of a posterior fragment only, lacking elytra.

Paragattyana intesi, new species

FIGURE 50

Harmothoe lunulata.—Intes and Le Loeuff, 1975:275 [not Delle Chiaje, 1830].

MATERIAL EXAMINED.—NORTH ATLANTIC OCEAN: West Africa: Ivory Coast, 05°12′N, 04°02′W, 69 m, Reine Pokou sta SD59, 21 Nov 1969, A. Intes, collector, holotype (MNHN).

DESCRIPTION.—Holotype with 36 segments, last one minute, 12.5 mm long, and 3 mm wide including setae. Body subrectangular, flattened dorsoventrally, without color dorsally; ventral half of body with segmental brown spots lateral to midventral groove and brownish areas on ventral bases of parapodia on posterior fourth of body and pygidium. Elytra 15 pairs, on usual segments, large, oval, covering dorsum, delicate, with small group of microtubercles on anterolateral part, and some scattered micropapillae on border and surface; brownish pigmentation in form of spots over places of attachment to elytrophores and crescent-shape area on medial and posterior sides, forming almost complete circle on some posterior elytra (Figure 50C,D).

Bilobed prostomium with anterior lobes subtriangular, without peaks; anterior pair of eyes anteroventral, larger than posterodorsal pair; ceratophore of median antenna oval, in anterior notch of prostomium, style longer than prostomium; ceratophores of lateral antennae inserted ventrally, with short tapered styles; palps stout, long, tapered; tentaculophores lateral to prostomium, each with aciculum, 2 setae on inner side, and pair of dorsal and ventral tentacular cirri, similar to median antenna; styles of antennae and tentacular cirri with small papillae; brownish pigmentation on prostomium, ceratophores of antennae, tentaculophores, and area between palps (Figure 50A,B,F). Segment 2 with first pair of large elytrophores, biramous parapodia and long ventral buccal cirri similar to tentacular cirri, and lateral to ventral mouth (Figure 50A,B,G); notosetae similar to those of following parapodia; neurosetae more slender than those following, ending in slender sharp tips, upper ones with longer spinose regions (Figure 50H).

Biramous parapodia with notopodium much shorter than neuropodium, conical, with projecting acicular lobe on lower side; larger neuropodium with subconical presetal acicular lobe with digitiform supraacicular process; postsetal lobe shorter, rounded (Figure 50G,I,J). Notosetae numerous, subequal in width to neurosetae, with numerous spinose rows, of 3 lengths: shorter and longer upper ones ending in blunt tips; longest lower ones tapering distally to long capillary tips (Figure 50K). Upper neurosetae with longer spinose regions and bifid hooked tips; middle and lower ones with shorter spinose regions and bifid tips, few lower ones with entire tips (Figure 50L). Dorsal cirri with cylindrical cirrophores and styles extending to tips of neurosetae or beyond, with scattered minute papillae; dorsal tubercles indistinct; ventral cirri short, tapered, with minute papillae (Figure 50J). Pygidium small lobe with anus medial to posterior small parapodia, with pair of small anal cirri.

FIGURE 50 (opposite page).—Paragattyana intesi, new species, holotype (MNHNP): A, dorsal view of anterior end, right lateral antenna small, regenerating, left palp and left dorsal tentacular cirrus missing, anterior pair of eyes anteroventral (dotted), first pair of elytra removed; B, same, ventral view; C, right 1st elytron from segment 2; D, right 7th elytron from segment 13, with detail of microtubercles; E, right 15th elytron from segment 32, with detail of microtubercles; F, right tentaculophore of segment 1, outer view (dorsal side left); G, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; H, upper and lower neurosetae from same; I, right middle elytrigerous parapodium, anterior view, acicula dotted; J, right cirrigerous parapodium, posterior view; K, long capillary and 2 shorter notosetae; L, upper, middle, and lower neurosetae, with detail of tips. (Scales = 0.5 mm for A,B; 0.5 mm for C–E; 0.2 mm for F,G,I,J; 0.1 mm for H,K,L.)

ETYMOLOGY.—The species is named for André Intes, the collector of the holotype.

DISTRIBUTION.—North Atlantic Ocean, West Africa, in 69 meters.

REMARKS.—Paragattana intesi differs from P. micropoides by the position of the anterior pair of eyes; anteroventral in the former and anterodorsal in the latter. The elytra of P. intesi have only scattered micropapillae and a marked color pattern, whereas the elytra of P. micropoides have some long delicate papillae on the border and surface and only an indistinct color pattern.

Rullieriella, new genus

TYPE SPECIES.—Malmgrenia monoechinata Rullier, 1965. Gender: feminine.

DIAGNOSIS.—Segments up to 44. Elytra and elytrophores 15 pairs, on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, and 32. Elytra with single recurved boss or tubercle, with or without anterior group of microtubercles, and micropapillae. Prostomium bilobed, with 3 antennae, 2 palps, and 2 pairs of eyes; ceratophore of median antenna in anterior notch; ceratophores of lateral antennae indistinct, inserted terminoventrally, separated from prostomium by faint line or continuous with prostomium (as in Halosydna). Tentaculophores of first segment lateral to prostomium, each with small acicular lobe on inner side, without setae, and pair of dorsal and ventral tentacular cirri. Second or buccal segment with first pair of elytrophores, biramous parapodia, and long ventral buccal cirri lateral to ventral mouth. Biramous parapodia with smaller notopodia and larger neuropodia, both rami with projecting acicular lobes; presetal acicular lobe of neuropodium longer, notched distally, with supraacicular process; postsetal lobe shorter, rounded. Notosetae more slender than neurosetae, finely spinose, all tapering to slender capillary tips, except on segments 2 and 3, shorter ones with blunt tips. Neurosetae with spinose regions and variable tips: slender, pointed; entire, blunt; or bifid. Dorsal cirri on nonelytrigerous segments, with cylindrical cirrophores and long styles; dorsal tubercles nodular; ventral cirri short. Pygidium with pair of anal cirri.

ETYMOLOGY.—The genus is named for the late François Rullier, in recognition of his many contributions to the biology of polychaetes.

COMPARISONS.—Among the genera of Harmothoinae, Rullieriella differs by having the elytra each with a large central recurved boss or tubercle; lateral antennae with indistinct ceratophores inserted terminoventrally, resembling Halosydna; notosetae more slender than neurosetae, finely serrated, all tapering to capillary tips (except on segments 2 and 3, some with blunt tips); and variable neurosetae with either tips tapering to slender pointed tips, entire blunt tips, or some with bifid tips.

The genus is established for a single species, Rullieriella monoechinata (Rullier, 1965), new combination.

Rullieriella monoechinata (Rullier, 1965), new combination

FIGURES 51–53

Malmgrenia monoechinata Rullier, 1965:166, fig. 1A–J.—Kudenov, 1977:95 [under incertae sedis, “probably represents a new genus”].—Day and Hutchings, 1979:9.—Hanley, 1987:161, fig. 3C [under incertae sedis, includes Parahalosydna chrysostichtus in synonymy].

Parahalosydna chrysostichtus Hutchings and Rainer, 1979:746, fig. 1A–D; 1980:36.—Hutchings, 1982:188; 1984:49, fig. 66a–c.—Hutchings and Murray, 1984:15.

MATERIAL EXAMINED.—SOUTH PACIFIC OCEAN: Australia: Queensland, Dunwich, Stradbroke Island, Moreton Bay, midshore in Zostera, 3 May 1962, F.C.V., collector, holotype of Malmgrenia monoechinata (AMS W3792). New South Wales, types of Parahalosydna chrysostichtus: Careel Bay, Pittwater, 33°37′S, 151°20′E, 16 Apr 1972, sheltered estuary area with Zostera, holotype (AMS W10570); paratype (AMS W10571). Port Hacking, Mainbar, N.S.W. State Fisheries sta 1051, 4 Nov 1975, Posidonia core, paratype (BMNH ZB1977:110).

TYPE MATERIAL.—Holotype of Malmgrenia monoechinata with 43 segments, last one very small, 18 mm long and 6 mm wide including setae; all prostomial appendages, except palps, tentacular and dorsal cirri missing; only 2 elytra remaining in vial. Holotype of Parahalosydna chrysostichtus in 2 pieces, with 34 segments, last 3 small, regenerating, 23 mm long and 9 mm wide including setae; paratype (AMS W10571) with anterior 15 segments missing, posterior fragment of 29 segments (segments 16–44), 13 mm long and 9 mm wide; paratype (BMNH ZB1977:110) with 38 segments, posterior 8 segments smaller, regenerating, 20 mm long and 7 mm wide.

DESCRIPTION.—Body flattened dorsoventrally, tapering posteriorly, with parapodia as long as body width; scattered brownish or light yellow pigmentation on prostomium, body, and parapodia. Elytra 15 pairs, on usual segments, large, oval, imbricated, covering dorsum, except for posterior 5–6 segments; large erect conical tubercle or boss near place of attachment to elytrophore, indistinct basally and directed posteriorly; group of microtubercles on anterior region (absent on first and some posterior elytra); globular papillae on lateral and posterior areas (Figures 51C,D, 52C,D; Rullier, 1965, fig. 1A–D; Hutchings and Rainer, 1979, fig. 1a).

FIGURE 51 (opposite page).—Rullieriella monoechinata, holotype of Malmgrenia monoechinata (AMS W-3792): A, dorsal view of anterior end, styles of median and lateral antennae, right and left dorsal and ventral tentacular cirri missing; B, same, ventral view, only bases of palps shown, styles of ventral buccal cirri of segment 2 missing; C, left anterior elytron, with detail of microtubercles and globular papillae; D, left middle elytron, with detail of papillae; E, right elytrigerous parapodium from segment 2, anterior view, acicula dotted, style of ventral buccal cirrus missing; F, long and short notosetae from same; G, neurosetae from same; H, right cirrigerous parapodium from segment 18, posterior view, style of dorsal cirrus missing; I, right elytrigerous parapodium from segment 19, anterior view, acicula dotted; J, long and short notosetae; K, upper, middle (2), and lower neurosetae. (Scales = 0.5 mm for A,B; 0.5 mm for C,D; 0.2 mm for E,H,I; 0.1 mm for F,G,J,K.)

FIGURE 52.—Rullieriella monoechinata holotype of Parahalosydna chrysostichtus (AMS W-10570): A, dorsal view of anterior end, right palp smaller, regenerating; B, ventral view of anterior end, median antenna bent dorsally; C, right 5th elytron from segment 9, with detail of microtubercles and papillae; D, right elytron from posterior fragment, with detail of microtubercles and papillae; E, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; F, shorter notosetae from same; G, upper, middle, and lower neurosetae from same. (Sclaes = 0.5 mm for A,B; 0.5 mm for C–E; 0.1 mm for F,G.)

FIGURE 53.—Rullieriella monoechinata, holotype of Parahalosydna chrysostichtus (AMS W-10570): A, right elytrigerous parapodium from segment 15, anterior view, acicula dotted; B, right cirrigerous parapodium from segment 16, posterior view; C, long and short notosetae from same; D, upper, middle, and lower neurosetae from same; E, right cirrigerous parapodium from posterior fragment, posterior view; F, upper, middle, and lower neurosetae from same. (Scales = 0.5 mm for A,B,E; 0.1 mm for C,D,F.)

Bilobed prostomium with anterior lobes without distinct peaks; ceratophore of median antenna in anterior notch, with style longer than prostomium; ceratophores of lateral antennae inserted terminoventrally, separated from prostomium dorsally by faint line (Figure 51A,B; Rullier, 1965, fig. 1A) or continuous with prostomium, as in Halosydna (Figure 52A,B; Hutchings and Rainer, 1979, fig. 1a); styles of lateral antennae short, subulate; palps stout, tapered, minutely papillate; eyes rather small, anterolateral pair in region of greatest width of prostomium, slightly larger than posterodorsal pair, tentaculophores lateral to prostomium, each with small acicular lobe on inner side, without setae, and pair of dorsal and ventral tentacular cirri, similar to median antenna; antennae and tentacular cirri with scattered micropapillae (Figures 51A,B, 52A,B; Rullier, 1965, fig. 1A; Hutchings and Rainer, 1979, fig. 1a; Hanley, 1987, fig. 3C). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri lateral to ventral mouth, similar to tentacular cirri (Figures 51A,B,E, 52A,B,E; Hutchings and Rainer, 1979, fig. la); notosetae of 2 types: shorter ones with blunt tips and longer ones tapering to fine tips (Figures 51F, 52F); neurosetae all tapering to slender sharp tips (Figures 51G, 52G).

Biramous parapodium with notopodium much shorter than neuropodium, subconical, with projecting digitiform acicular lobe on lower side; larger neuropodium subconical, with presetal acicular lobe notched distally and slightly longer supraacicular process; postsetal lobe shorter, rounded (Figures 51E,H,I, 52E, 53A,B,E; Rullier, 1965, fig. 1E; Hutchings and Rainer, 1979, fig. 1b). Notosetae moderate in number, more slender than neurosetae, finely spinose, short and longer, all tapering to slender capillary tips (Figures 51J, 53C; Rullier, 1965, fig. 1F), except on segments 2 and 3, with shorter notosetae ending in blunt tips (Figures 51F, 52F). Neurosetae numerous, forming fan–shape bundle, of 2–3 types, upper ones with long spinose regions, tapering to slender pointed tips; middle ones with shorter spinose regions and entire blunt tips, some with slight indication of secondary tooth, and some with distinct secondary tooth; lowest ones with slender sharp tip (Figures 51K, 53D,F; Rullier, 1965, fig. 1G–I; Hutchings and Rainer, 1979, fig. 1c,d). Dorsal cirri with cylindrical cirrophores, inflated basally, and long tapered styles extending to near tips of neurosetae, with scattered micropapillae; dorsal tubercles nodular, ventral cirri short, tapered (Figures 51H, 53B,E; Rullier, 1965, fig. 1E; Hutchings and Rainer, 1979, fig. 1b). Pygidium small lobe with dorsal anus and pair of long anal cirri.

DISTRIBUTION.—South Pacific Ocean, Australia.

Wilsoniella, new genus

TYPE SPECIES: Malmgrenia furcosetosa Loshamn, 1981. Gender: feminine.

DIAGNOSIS.—Segments up to 42. Elytra and elytrophores 15 pairs, on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, and 32. Elytra with or without anterior group of microtubercles and micropapillae. Prostomium bilobed, without cephalic peaks, with 3 antennae, 2 palps, and 2 pairs of eyes; ceratophore of median antenna in anterior notch; ceratophores of lateral antennae distinct, inserted terminoventrally. Tentaculophores of first segment lateral to prostomium, each with small acicular lobe on inner side, 0–2 setae, and pair of dorsal and ventral tentacular cirri. Second or buccal segment with first pair of elytrophores, biramous parapodia, and long ventral buccal cirri lateral to ventral mouth. Biramous parapodia with smaller notopodia and larger neuropodia, both with projecting acicular lobes; presetal acicular lobe of neuropodium longer, with large subtriangular supraacicular process; postsetal lobe shorter, rounded. Notosetae slightly stouter than neurosetae, with numerous spinose rows, of 2 types: shorter with blunt tips and longer, tapering to slender bifid split tip. Neurosetae with spinose regions and bifid hooked tips or lower ones entire. Dorsal cirri on nonelytrigerous segments, with cylindrical cirrophores and long styles; dorsal tubercles nodular; ventral cirri short. Pygidium with pair of anal cirri.

ETYMOLOGY.—The genus is named for Douglas P. Wilson, in recognition of his many contributions in the study of polychaetes including descriptions of numerous polychaete species.

COMPARISONS.—Among the genera of Harmothoinae, Wilsoniella differs by having notosetae of two types: shorter ones with blunt tips and long ones tapering to slender split tips. The general characters of the prostomium, parapodia, and elytra agree with Malmgreniella.

The genus is established for a single species, Wilsoniella furcosetosa (Loshamn, 1981), new combination, from the west coast of Sweden and the Plymouth area, England.

Wilsoniella furcosetosa (Loshamn, 1981), new combination

FIGURES 54, 55

Harmothoe lunulata.—Orton, 1923:861 [part; large specimens with Amphitrite edwardsii]. [Not Delle Chiaje, 1830.]

Harmothoe sp. nov.—G.M. Spooner, in Plymouth Marine Fauna, 1957:xxxvi, 112 [with Amphitrite edwardsii].

Malmgrenia furcosetosa Loshamn, 1981:5, fig. 1A–I.

Harmothoe furcosetosa.—Tebble and Chambers, 1982:42, fig. 1c, 13a–c, 39.—Hanley, 1987:151, fig. 3G.

MATERIAL EXAMINED.—NORTH ATLANTIC OCEAN: England: Salcombe Estuary near Plymouth, in burrows of Amphitrite edwardsii (Quatrefages), 29 Jul 1956, 26 Mar 1959, 18 Mar 1961, G.M. Spooner, collector, 4 specimens (LCHP S-18–20; USNM 55053). Salcombe, west side, 27 Mar 1971, P.E. Gibbs, collector, 3 specimens (LCHP G-1; USNM 55054). Sweden: Boluslän, Säcken, Tränsholmen, 125 m, L. Almkvist, collector, 6 Apr 1974, holotype of Malmgrenia furcosetosa (NRS 3173). Skagerrak, 58°21′N, 09°11′E, 366 m, clay, Gunnild Expedition sta 14, 1879, paratype of M. furcosetosa (NRS 3174).

FIGURE 54 (opposite page).—Wilsoniella furcosetosa, specimen from Plymouth (USNM 55053): A, dorsal view of anterior end; B, ventral view of anterior end, only bases of median antenna, palps, and tentacular cirri shown; C, right elytrigerous parapodium from segment 2, anterior view, acicula dotted; D, upper and lower neurosetae from same, with detail of tips; E, right 1st elytron from segment 2; F, right 2nd elytron from segment 4; G, right 10th elytron from segment 19, with detail of microtubercles; H, right 15th elytron from segment 32; I, dorsal view of right side of segments 13 and 14, dorsal cirrus flattened and curled dorsally; J, ventral view of left side of 2 segments near posterior end. (Scales = 1.0 mm for A,I,J; 0.5 mm for B,C; 0.1 mm for D; 1.0 mm for E–H.)

FIGURE 55.—Wilsoniella furcosetosa, specimen from Plymouth (USNM 55053): A, right elytrigerous parapodium from segment 19, anterior view, acicula dotted: B, right cirrigerous parapodium from segment 20, posterior view; C, long and short notosetae, with detail of tips; D, upper and lower supraacicular neurosetae, with detail of tips; E, upper and lower subacicular neurosetae. (Scales = 0.5 mm for A,B; 0.1 mm for C–E.)

TYPE MATERIAL.—Holotype, male with sperm, in 3 pieces, with 39 segments, 38 mm long and 10 mm wide including setae. Paratype, female with eggs, in 3 pieces, with 38 segments, 36 mm long and 9 mm wide. Specimens from Plymouth area with 40–42 segments, 46–56 mm long and 11–13 mm wide.

DESCRIPTION.—Body elongate, flattened ventrally, arched dorsally, tapering slightly anteriorly and more so posteriorly. Body showing brownish pigmentation on posterior regions: dorsally in form of spindle-shape bands between dorsal tubercles and additional smaller intersegmental spots; ventrally in form of narrow anterior transverse bands and wide posterior transverse bands, joining laterally and extending onto parapodia, with spots on cirrophores and styles of ventral cirri (Figure 54J). Elytra 15 pairs, on usual segments, large, covering dorsum, except for posterior 5 or so segments; round, subreniform, and elongate oval, with mottled pigmentation, tending to be darker near place of attachment to elytrophore and large area posterior to it without color; also colorless on anterior part; wide band of microtubercles on anterior part of elytra; except for first pair and some posterior elytra; without papillae, except for some scattered micropapillae (Figure 54E–H; Loshamn, 1981, fig. 1I,J; Tebble and Chambers, 1982, fig. 39).

Bilobed prostomium with anterior lobes without distinct peaks; ceratophore of median antenna large, cylindrical, in anterior notch, with style about 1.5 times length of prostomium; ceratophores of lateral antennae inserted terminoventrally, with styles about half as long as median antenna; eyes rather small, anterior pair anterolateral, barely visible dorsally, slightly larger than posterodorsal pair, palps long, tapered, longer than median antenna; tentaculophores lateral to prostomium, each with small acicular lobe and 0–2 setae on inner side; dorsal and ventral tentacular cirri similar to median antenna; antennae and tentacular cirri with minute papillae; facial tubercle bulbous (Figure 54A,B; Loshamn, 1981, fig. 1A,L; Tebble and Chambers, 1982, fig. 1c; Hanley, 1987, fig. 3G). Segment 2 with first pair of large elytrophores, biramous parapodia, and long ventral buccal cirri lateral to ventral mouth, similar to tentacular cirri (Figure 54A–C); notosetae similar to following segments, only few longer ones tapering to slender tips; neurosetae more slender than following, lower ones with bulbous entire tips, some upper ones with slender secondary tooth (Figure 54D); extended pharynx with 9 pairs of border papillae and 2 pairs of hooked jaws.

Biramous parapodium with notopodium much shorter than neuropodium, rounded, with projecting acicular lobe on lower side; neuropodium with longer subconical presetal acicular lobe, with prominent subtriangular supraacicular process, and shorter rounded postsetal lobe (Figure 55A,B; Loshamn, 1981, fig. 1K). Notosetae very numerous, slightly stouter than neurosetae, with numerous spinose rows, of 2 types: shorter ones, of 3 lengths, tapering to blunt tips; longer ones, extending almost to tips of neurosetae, tapering distally to slender bifid tips, one tooth slightly larger than other, or some with tips entire (Figure 55C; Loshamn, 1981, fig. 1B–D,H; Tebble and Chambers, 1982, fig. 13a–c). Neurosetae very numerous, supraacicular ones with longer spinose regions and bifid tips; upper few with longer secondary tooth (Figure 55D; Loshamn, 1981, fig. 1E); subacicular neurosetae with shorter spinose regions and bifid hooked tips, lower few with entire hooked tips (Figure 55E; Loshamn, 1981, fig. 1F,G). Cirrophores of dorsal cirri cylindrical, bulbous basally, with anterior and posterior orange colored glandular areas, with styles flattened dorsoventrally, tending to curl dorsally and extending beyond neurosetae, with scattered micropapillae; dorsal tubercles nodular, with orange colored glandular areas; ventral cirri short, subulate, with scattered micropapillae (Figures 54A,I, 55B). Pygidium with anus medial to last pair of parapodia, with pair of long anal cirri.

BIOLOGY.—Wilsoniella furcosetosa was collected by G.M. Spooner and P.E. Gibbs in the Plymouth area in the burrows of the terebellid polychaete Amphitrite edwardsii (Quatrefages).

DISTRIBUTION.—North Atlantic Ocean, Great Britain and west coast of Sweden, up to 366 meters.

Genus Lobopelma Hanley, 1987

Lobopelma microscala (Kudenov, 1977)

Malmgrenia microscala Kudenov, 1977:90, pl. 2a–m.

Lobopelma microscala.—Hanley, 1987:157, figs. 3J, 4A–H.

REMARKS.—Harmothoinae with up to 38 segments, 16–18 pairs of elytra (15 pairs on usual segments, and 1–3 posterior pairs variable, on segments 35–38). The type material from Port Phillip Bay, Victoria, Australia, in 2–10 meters (NMV, AMS) was not examined but is well described by Kudenov (1977) and Hanley (1987). Lobopelma microscala agrees with the general characters of Malmgreniella, except for the more numerous pairs of elytra and the unique character on the ventral side of the parapodia: a tripalmate lobe and small cylindrical lobe between the ventral cirri and nephridial papillae (Kudenov, 1977, pl. 2b; Hanley, 1987, fig. 4C,D).

Malmgrenia perspicua Intes and Le Loeuff, 1975, incertae sedis

Malmgrenia perspicua Intes and Le Loeuff, 1975:271–273, fig. 1j–q.

REMARKS.—Harmothoinae, with 36 segments and 15 pairs of elytra. The species, described from material from the Ivory Coast, was not available for study. Based on the original description and figures, it differs from the genera covered herein. The attachment of the lateral antennae on the prostomium is not clear (said to be subterminal); notosetae are stout and smooth; neurosetae are slender with entire tips, the lower ones with capillary tips.

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Devaney, D.M., and J.H. Bailey-Brock

1987.   Polychaetes of Enewetak Atoll. In D.M. Devaney, E.S. Reese, B.L. Burch, and P. Helfrich, editors, The Natural History of Enewetak Atoll, II: Biogeography and Systematics, 13:97–103. Oakridge, Tennessee: U.S. Department of Energy, Office of Scientific and Technical Information.

Ditlevsen, H.

1917.   Annelides, I. In The Danish Ingolf-Expedition Copenhagen, 4(4):1–71, 6 plates. Copenhagen: Bianco Luno.

1929.   Polychaeta. Zoology of the Faroes, Copenhagen, 16:1–83.

Ehlers, E.

1864.   Die Borstenwürmer (Annelida Chaetopoda) nach systematischen und anatomischen Untersuchungen, Ersten Band. Pages 1–268, 11 plates. Leipzig: Wilhelm Engelmann.

Fauchald, K.

1977.   Polychates from Intertidal Areas in Panama, with a Review of Previous Shallow-water Records. Smithsonian Contributions to Zoology, 221:1–81, 13 figures.

Fauchald, K., and A.A. Reimer

1975.   Clave de Poliquetos Panamenos con la inclusion de una clave para todas la Familias del Mundo. Boletin del Instituto Oceanografico, 14(1):71–94, 9 figures.

Fauvel, P.

1913.   Quatrième note préliminaire sur les polychètes provenant des campagnes de l'Hirondelle et de la Princesse-Alice, ou déposées dans le Musée Océanographique de Monaco. Bulletin de l'Institut Océanographique de Monaco, 270:1–80, 13 figures.

1914.   Annélides polychétes non pélagiques provenant des campagnes de l'Hirondelle et de la Princesse-Alice (1885–1910). Résultats des Campagnes Scientifiques Accomplies par le Prince Albert I, Monaco, 46:1–432, 31 plates.

1923.   Polychètes Errantes. Faune de France, 5:1–488, 181 figures.

1927.   Polychètes sédentaires; Addenda aux Errantes, Archiannélides, Myzostomaires. Faune de France, 16:1–494, 152 figures.

1953.   Annelida Polychaeta. In K.B. Seymour Sewell, editor, The Fauna of India Including Pakistan, Ceylon, Burma and Malaya. 507 pages, 250 figures. Allahabad: The Indian Press.

Fox, R.S., and E.E. Ruppert

1985.   Shallow-water Marine Benthic Macroinvertebrates of South Carolina, Species Identification, Community Composition and Symbiotic Associations. The Belle W. Baruch Library in Marine Science, 14:1–330.

Gardiner, S.L.

1976 (“1975”). Errant Polychaete Annelids from North Carolina. Journal of the Elisha Mitchell Scientific Society, 91(3):77–220, 29 figures. [Date on title page is 1975. Actually published in 1976.]

Giard, A.

1886.   Sur quelques Polynoidiens. Bulletin Biologique de la France et de la Belgique, series 2, 9:1–18, 6 figures.

Gibbs, P.E.

1969.   Aspects of Polychaete Ecology with Particular Reference to Commensalism. Philosphical Transactions of the Royal Society of London, B255:443–459, figures 130–138, plates 69, 70.

1971.   The Polychaete Fauna of the Solomon Islands. Bulletin of the British Museum (Natural History), Zoology, 21(5):99–211, 17 figures.

1972.   Polychaete Annelids from the Cook Islands. Journal of Zoology, London, 168:199–220, 8 figures.

Grube, A-E.

1856.   Annulata Örstediana: Enumeratio Annulatorum, quae in itinere per Indiam occidentalem et Americam centralem annis 1845–1848 suspepto legit cl. A.S. Örsted, adjectis speciebus nonnullis a cl. H. Kröyero in itinere ad Americam meridionalem collectis. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjøbenhavn, 1856:44–62.

Guille, A.

1965.   Contribution à l'étude de la systématique et de l'écologie d'Ophiothrix quinquemaculata d.Ch. Vie et Milieu, 15:243–308, 6 plates.

Hamond, R.

1966.   The Polychaeta of the Coast of Norfolk. Cahiers de Biologie Marine, 7:383–436.

Hanley, J.R.

1987.   Taxonomic Status of Some Species Formerly Referred to Malmgrenia MacIntosh 1874, with the Description of a New Genus Lobopelma (Polychaeta: Polynoidae). The Beagle, Records of the Northern Territory Museum of Arts and Sciences, 4(1):147–163, 4 figures.

Hartman, O.

1939.   The Polychaetous Annelids Collected by the Presidential Cruise of 1938. Smithsonian Miscellaneous Collections, 98:1–22, 3 figures.

1944.   Polychaetous Annelids from California Including the Descriptions of Two New Genera and Nine New Species. Allan Hancock Pacific Expeditions, 10:239–307, plates 19–26.

1955.   Quantitative Survey of the Benthos of San Pedro Basin, Southern California, Part I: Preliminary Results. Allan Hancock Foundation Pacific Expeditions, 19:1–185, 7 plates, 2 charts.

1956.   Polychaetous Annelids Errected by Treadwell, 1891 to 1948, Together with a Brief Chronolgy. Bulletin of the American Museum of Natural History, 109(2):243–310, plate 2.

1959.   Catalogue of the Polychaetous Annelids of the World, Part 1. Allan Hancock Foundation Publications Occasional Paper, 23:1–353.

1961.   Polychaetous Annelids from California. Allan Hancock Pacific Expeditions, 25:1–226, 34 plates.

1966.   Quantitative Survey of the Benthos of San Pedro Basin, Southern California, Part II: Final Results and Conclusions. Allan Hancock Pacific Expeditions, 19:187–456, 13 plates.

1967.   Polychaetous Annelids Collected by the USNS Eltanin and Staten Island Cruises, Chiefly from Antarctic Seas. Allan Hancock Monographs in Marine Biology, 2:1–387, 51 plates.

1968.   Atlas of the Errantiate Polychaetous Annelids from California. 828 pages. Los Angeles, California: Allan Hancock Foundation University of Southern California.

Hartmann-Schröder, G.

1971.   Annelida, Borstenwürmer, Polychaeta. Die Tierwelt Deutschlands und der Angrenzenden Meeresteile, 58:1–594, 191 figures.

1977.   Polychaeten aus dem Sublitoral und Bathyal vor der portugiesischen und marokkanischen Küste Auswertung der Fahrt 8 (1967) von F.S. “Meteor.” “Meteor” Forschungsergebnisse, Reihe D, 26:65–99, 83 figures, 1 plate.

1984.   Zwei neue kommensalische Polychaeten der Gattung Hololepidella Willey (Polynoidae) von der Philippine. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 81:63–70, 17 figures.

Hendler, G., and D.L. Meyer

1982.   An Association of a Polychaete, Branchiosyllis exilis with an Ophiuroid, Ophiocoma echinata, in Panama. Bulletin of Marine Science, 32(3):736–744, 3 figures.

Hornell, J.

1891.   Report on the Polychaetous Annelids of the L.M.B.C. District. Proceedings of the Liverpool Biological Society, 5:223–268, plates 13–15.

Horst, R.

1915.   On New and Little-known Species of Polynoinae from the Netherlands' East Indies. Zoologische Mededeelingen, Leiden, 1:2–30.

1917.   Polychaeta Errantia of the Siboga-Expedition, Part 2: Aphroditidae and Chrysopetalidae. Siboga-Expeditie (Leiden), 24b:1–140, 5 figures, plates 11–29.

1922.   On Some Polychaetous Annelids from Curaçao. Bijdragen tot de Dierkunde (Amsterdam), Feest-nummer: 193–201.

Hutchings, P.

1982 (“1981”). The Fauna of Australian Seagrass Beds. Proceedings of the Linnean Society of New South Wales, 106(2):181–200. [Date on title page is 1981. Actually published in 1982.]

1984.   An Illustrated Guide to the Estuarine Polychaete Worms of New South Wales. 160 pages, 155 figures. Sydney: Coast and Wetlands Society, Sydney South 2000.

Hutchings, P., and A. Murray

1984.   Taxonomy of Polychaetes from the Hawkesbury River and the Southern Estuaries of New South Wales, Australia. Records of the Australian Museum, 36(3):1–118, 32 figures.

Hutchings, P., and S. Rainer

1979.   The Polychaete Fauna of Careel Bay, Pittwater, New South Wales, Australia. Journal of Natural History, 13(6):745–796, 10 figures.

1980 (“1979”). A Key to Estuarine Polychaetes in New South Wales. Proceedings of the Linnean Society of New South Wales, 104(1): 35–48. [Date on title page is 1979. Actually published in 1980.]

Intes, A., and P. Le Loeuff

1975.   Les Annélides polychètes de Côte d'Ivoire, I: Polychètes Errantes—Compte rendu systématique. Cahiers de l'Office de la Recherche Scientifique et Technique Outre-Mer, série Océanographie, 13(4):267–321, 13 figures.

Katzmann, W.

1983.   Bermerkungen zur Systematik, Ökologie und Tiergeographie der mitteladriatischen Weichbodenpolychaeten. Annalen des Naturhistorischen (Hof) Museums, Wien, 84/B:87–122, 2 figures, 1 plate.

Kinberg, J.G.H.

1856 (“1855”). Nya slägten och arter af Annelider. Ōfversigtaf Konglia Vetenskaps-Akademiens Förhandlingar, Stockholm, 12:381–388. [Date on title page is 1855. Actually published in 1856.]

Kirkegaard, J.B.

1969.   A Quantitative Investigation of the Central North Sea Polychaeta. Spolia Zoologica Musei Hauniensis, 29:1–285, 11 figures.

1983.   The Polychaeta of West Africa, Part II: Errant Species, I: Aphroditidae to Nereididae. Atlantide Report, 13:181–240.

Knox, G.A., and D.B. Cameron

1971.   Port Phillip Bay Survey Part 2, 4: Polychaeta. Memoirs of the National Museum of Victoria, 32:21–41, 35 figures.

Koehler, R.

1894.   Echinodermes recueillis à La Ciotat pendant l'été 1894. Mémoires de la Société Zoologique de France, 7:405–426.

Kudenov, J.D.

1975a. The Occurrence of the Polynoid Harmothoe cf. lunulata from the Tube of the Maldanid Axiothella rubrocincta (Polychaeta). Bulletin of the Southern California Academy of Sciences, 74(1):42–43, 1 figure.

1975b. Two New Species of Errant Polychaetes from the Gulf of California, Mexico. Bulletin of the Southern California Academy of Sciences, 74(2):75–80, 2 figures.

1977.   Polychaeta from Southeastern Australia, 2: Malmgrenia spp. (Polynoidae) from Port Phillip Bay, Victoria. In D.J. Reish and K. Fauchald, editors, Essays on Polychaetous Annelids in Memory of Dr. Olga Hartman, pages 85–102, plates 1, 2. Los Angeles, California: Allan Hancock Foundation, University of Southern California.

Lie, U.

1968.   A Quantitative Study of Benthic Infauna in Puget Sound, Washington, U.S.A., in 1963–1964, with a Section on Polychaetes by Karl Banse, Katharine D. Hobson, and Frederic H. Nichols. Fiskeridirektoratets Skrifter, serie Havundersøkelser, 14(5):230–556, 70 figures.

Loi, T.

1980.   Catalogue of the Types of Polychaete Species Erected by J. Percy Moore. Proceedings of the Academy of Natural Sciences of Philadelphia, 132:121–149.

Loshamn, A.-A.

1981.   Descriptions of Five Polynoid Species (Polychaeta) from the Coasts of Norway and Sweden, Including Three New Species, One New Genus and One New Generic Name. Zoologica Scripta, 10(1):5–13, 3 figures.

MacGinitie, G.E., and N. MacGinitie

1949.   Natural History of Marine Animals. 473 pages, 282 figures. New York: McGraw-Hill Book Company, Inc.

Macnae, W., and M. Kalk

1962.   The Fauna and Flora of Sand Flats at Inhaca Island, Moçambique. The Journal of Animal Ecology, 31:93–128, 5 figures.

Malaquin, A.

1890.   Les Annélides polychètes des côtes du Boulonnais. Revue Biologique du Nord de la France, Lille, 2(1b):275–285.

Malmgren, A.L.

1865.   Nordiska Hafs-Annulater. Översigt af Konglia Vetenskaps-Akademiens Förhandlingar, Stockholm, 21(1):51–110, plates 8–15.

Markov, A.V.

1989.   The Species Composition of the Genus Brisaster (Echinoidea). Zoologischesky Zhurnal, 68(11):86–93, 1 figure.

McIntosh, W.C.

1868.   Report on the Annelids Dredged off the Shetlands by Mr. Gywn Jeffreys, 1867–68. Report of the British Association for the Advancement of Sciences, for 1868, pages 336–340.

1874a. On the Annelida of the Gulf of St. Lawrence, Canada: Family 1, Euphrosynidae, to Family 6, Sigalionidae. Annals and Magazine of Natural History, series 4, 13(76):261–270, plates 9, 10.

1874b. On the Invertebrate Marine Fauna and Fishes of St. Andrews: Annulata, Discophora, Oligochaeta and Polychaeta. Annals and Magazine of Natural History, series 4, 14:192–207.

1876a. On British Annelida, Part 1. Transactions of the Zoological Society of London, 9:371–394, plates 67–70.

1876b. On the Annelida of the Porcupine Expeditions of 1869–1870, Part I: Euphrosynidae, Amphinomidae, Aphroditidae, Polynoidae, Acoetidae and Sigalionidae. Transactions of the Zoological Society of London, 9(7):395–416, plates 71–73.

1885.   Annelida Polychaeta. In Report on the Scientific Results of the Voyage of H.M.S. Challenger…1873–76…Zoology, 12(34):1–554, plates 1–55, 1A–39A.

1900.   A Monograph of the British Annelids, Part 2: Polychaeta, Amphinomidae to Sigalionidae. Ray Society, London, pages 215–442, plates 24–42.

1919.   Number 42, Notes from the Gatty Marine Laboratory, St. Andrews, 1: Preliminary Studies on Filograna; 2: On Harmothoe watsoni M'I., and var. H. marphysae, M'I. Annals and Magazine of Natural History, series 9, 3:125–164

1923.   The British Marine Annelids, Polychaeta: Sabellidae to Serpulidae and Additional Species. Ray Society, London, 4(2):251–539, plates 115–117, 128–138.

Millott, N.

1953.   A Remarkable Association between Ophionereis reticulata (Say) and Harmothoe lunulata (Delle Chiaje). Bulletin of Marine Science of the Gulf and Caribbean, 3(2):96–99, 2 figures.

Monro, C.C.A.

1928a. Polychaeta of the Families Polynoidae and Acoetidae from the Vicinity of the Panama Canal, Collected by Dr. C. Crossland and Dr. Th. Mortensen. Journal of the Linnean Society, London, 36:553–576, 30 figures.

1928b. On Some Polychaeta of the Family Polynoidae from Tahiti and the Marquesas. Annals and Magazine of Natural History, London, series 10, 3:467–473, 4 figures.

1930.   Polychaete Worms. Discovery Reports, 2:1–222, 91 figures.

1937.   Polychaeta. In The John Murray Expedition 1933–34. Scientific Reports, 4(8):243–321, 28 figures.

Moore, J.P.

1910.   The Polychaetous Annelids Dredged by the U.S.S. “Albatross” off the Coast of Southern California in 1904, II: Polynoidae, Aphroditidae and Segaleonidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 62:328–402, plates 28–33.

Orton, J.H.

1923.   Some New Commensals from the Plymouth District. Nature, London, 112:861, 1 figure.

Pérès, J.M.

1954.   Contribution à l'étude des annélides polychae`tes benthiques de la Mediterranée occidentale. Recueil des Travaux de la Station Marine d'Endoume, 8(13):83–155, 1 plate.

Pettibone, M.H.

1953.   Some Scale-bearing Polychaetes of Puget Sound and Adjacent Waters. 89 pages, 40 plates. Seattle: University of Washington Press.

1967.   Type-specimens of Polychaetes Described by Edith and Cyril Berkeley (1923–1964). Proceedings of the United States National Museum, 119:1–23.

1969a. Review of Some Species Referred to Scalisetosus McIntosh (Polychaeta, Polynoidae). Proceedings of the Biological Society of Washington, 82:1–30, 12 figures.

1969b. The Genera Polyeunoa McIntosh, Hololepidella Willey, and Three New Genera (Polychaeta, Polynoidae). Proceedings of the Biological Society of Washington, 82:43–62, 6 figures.

1975.   Review of the Genus Hermenia, with a Description of a New Species (Polychaeta: Polynoidae: Lepidonotinae). Proceedings of the Biological Society of Washington, 88(2):233–248, 6 figures.

1977.   Review of Halosydnopsis and Related Genera (Polychaeta Polynoidae: Lepidonotinae). In D.J. Reish and K. Fauchald, editors, Essays on Polychaetous Annelids in Memory of Dr. Olga Hartman, pages 39–62, 6 figures. Los Angeles, California: Allan Hancock Foundation, University of Southern California.

1989.   A New Species of Benhamipolynoe (Polychaeta: Polynoidae: Lepidastheniinae) from Australia, Associated with the Unattached Stylasterid Coral Conopora adeta. Proceedings of the Biological Society of Washington, 102(2):300–304, 2 figures.

Picard, J.

1965.   Recherches qualitatives sur les biocoenoses marines des substrats meubles dragables de la région marseillaise. Recueil des Travaux de la Station Marine d'Endoume, 36(52):1–160.

Plymouth Marine Fauna

1957.   Plymouth Marine Fauna. Third edition, 487 pages. Plymouth, England: Marine Biological Association of the United Kingdom. [Polychaetes revised by D.P. Wilson and N. Tebble.]

Potts, F.A.

1910.   Polychaeta of the Indian Ocean, Part 2: The Palmyridae, Aphroditidae, Polynoidae, Acoetidae and Sigalionidae. The Transactions of the Linnean Society of London, series 2, 13:325–353, plates 18–21.

Rullier, F.

1965.   Contribution à la faune des Annélides polychètes de l'Australie. Papers, Department of Zoology, University Queensland, 2(9):163–201, 9 figures.

Saint-Joseph, A. de

1888.   Les Annélides polychètes de côtes de Dinard, Part 2. Annales des Sciences Naturelles, Zoologie (Paris), series 7, 5:141–338, plates 6–13.

1898.   Les Annélides polychètes des côtes de France (manche et ocèan). Annales des Sciences Naturelles, Zoologie (Paris), series 8, 5:209–464, plates 13–23.

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Sloan, N.A., A.M. Clark, and J.D. Taylor

1979.   The Echinoderms of Aldabra and Their Habitats. Bulletin of the British Museum (Natural History), Zoology, 37(2):81–128.

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1914.   Clare Island Survey, Part 47: Archiannelida and Polychaeta. Proceedings of the Royal Irish Academy, 31(2):1–160, 15 plates.

Spooner, G.M., D.P. Wilson, and N. Tebble

1957.   Phylum Annelida. In Plymouth Marine Fauna, third edition, pages 109–149. Plymouth, England: Marine Biological Association of the United Kingdom.

Støp-Bowitz, C.

1948.   Polychaeta from the Michael Sars North Atlantic Deep-sea Expedition 1910. Report on the Scientific Results of the “Michael Sars” North Atlantic Deep-sea Expedition 1910, Bergen Museum, Norway, 5(8):1–91, 51 figures.

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1971.   Polychaetous Annelids and Benthic Environments in Tampa Bay, Florida. 1332 pages. Doctoral dissertation, University of Florida, Gainesville.

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1982.   Polychaetes from Scottish Waters, Part I: Family Polynoidae. In Royal Scottish Museum Studies, 73 pages, 58 figures. Edinburgh: Royal Scottish Museum.

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1917.   Polychaetous Annelids from Florida, Porto Rico, Bermuda, and the Bahamas. Papers from the Department of Marine Biology of the Carnegie Institution of Washington, 11(10)251:255–272, 3 plates.

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1975.   Ectocommensal Polynoids (Annelida, Polychaeta) Associated with the Echinoderms. Bulletin of the Marine Park Research Stations, 1(1):19–30, 6 figures.

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1902.   Polychaeta. In Report on the Collections of Natural History Made in the Antarctic Regions during the Voyage of the “Southern Cross,” 12:262–283, plates 41–46. London.

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