Blake, 1983_Spionidae S America and Antarctica

Biology of the Antarctic Seas XIV Antarctic Research Series, Voluale 39, Paper 3, Pages 205-288

POLYCHAETES OF THE FAMILY SPIONIDAE FROM SOUTH AMERICA, ANTARCTICA, AND ADJACENT SEAS AND ISLANDS

James A. Blake

Battelle New England Marine Research Laboratory Duxbury, Massachusetts 02332

Abstract. Fifty-nine species in 18 genera of Splonidae are recorded from South America and Antarctica. Eight additional species are also represented but cannot be specifically identified due to the fragmentary nature of the specimens. Two new genera are erected: Pygospiopsis for Pygospio dubia Monro and Amphipolydora for Polydora ~chiata Hartman. Twelve species are new to science: Scolelepis eltaninae, Dispio brachychaeta, Scolecolepides uncin~Prionospio (Prionospio) orensanzi, Spio ~setosa, Microspio hartmanae:-M. paradoxa, Boccardiella occipitalis,-Polydora antonbruunae, P. ecuadoriana, P. magellanica, and Pseudopolydora primigenia. Revisionary data is presented for 12 species. This includes several new synonyms and redescriptions of Prionospio (Minuspio) patagonica Augener and Spiophanes soederstroemi Hartman. The distribution and relationships of the spionid faunas from various regions of South America and Antarctic seas are discussed. Only 4 species are common in the high Antarctic region, with 1 species, Laonice antarcticae, being a probable immigrant from the deep Atlantic Ocean. More endemic species occur on the eastern side of South America south of 300s latitude than on the western side. The faunas of both shores, however, show more zoogeographic affinities to the species of the Pacific Ocean than of the Atlantic Ocean.

Contents

Introduction •••.•••.••••••••••••••••••••• 205 Systematic account of Spionidae •••••••••• 206 Discussion ••.••.•.•••..•..••••••.•••••.•• 267 Appendix: Station 10cations •.•••.••••.•.• 271

Introduction

Spionid polychaetes are among the more familiar components of marine benthic communities along most continental shorelines. The best studied spionid faunas in the southern oceans are from South Africa [Day, 1967], Australia [Blake and Kudenov, 1978], and New Zealand [Ranier, 1973; Read, 1975], where collectively, over 100 species have been reported. Approximately 28 species of Spionidae are currently known from the seas

Copyr ight 1983 by the American Geophysical Union.

around South America and Antarctica [Ehlers, 1897, 1901a; Augener, 1923a, b; Hartman, 1953, 1966, 1967, 1978; Hartmann-Schroder, 1962a, b, 1965; Orensanz and Estivariz, 1971; Blake and Woodwick, 1971; Carrasco, 1974; Blake, 1979a].

The present report describes new collections of Spionidae from the east and west coasts of South America, the seas around Antarctica, and adjacent island groups. Some species described in previous publications have been reexamined in order to clarify taxonomic problems.

The new materials come from several sources. Intertidal and subtidal spionids from Uruguay and Argentina were provided by J. M. Orensanz, formerly of the Instituto de Biologia Marina, Mar del Plata, Argentina (IBM), and include specimens collected by a variety of oceanographic cruises (see appendix Table AI). The Smithsonian Oceanographic Sorting Center (SOSC) provided numerous lots of unidentified specimens collected by the USNS Eltanin (Table A3) and R/V Hero (Table A4) as part of the U. S. Antarctic Research Program (USARP) and by the R/V Anton Bruun (Table A2) as part of the Southeastern Pacific Biological and Oceanographic Program (SEPBOP). The Spionidae collected by the Deep Freeze I-IV cruises of 1956-1959 (Table A5) and some miscellaneous unworked Antarctic materials were provided by the National Museum of Natural History, Smithsonian Institution (USNM). A small portion of the spionid collections taken during the Lund University Chile Expedition in 1948-1949 (Table A6) was made available through the courtesy of K. Fauchald while he was wi th the Allan Hancock Foundation. Some of this latter material has already been treated [Blake, 1971; Blake and Woodwick, 1971]. Additional unidentified spionids from Chile were provided by D. Stewart, Old Dominion University, Norfolk, Virginia, and C. Bertran, Universidad Austral de Chile. Older collections were acquired from the following institutions: Allan Hancock Foundation, Los Angeles, California (AHF); British Museum of Natural History, London (BMNH); Swedish Museum of Natural History, Stockholm (SMNH); National Museum of Natural History, Smithsonian Institution (USNM); Zoological Museum of Hamburg (ZMH); and the Zoological Museum of East Berlin (ZMB). Type specimens of the

205

206 BIOLOGY OF THE ANTARCTIC SEAS XIV

new taxa and most of the collections are deposited in the National Museum of Natural History, Smithsonian Institution (USNM). A few duplicate type specimens are deposited in the Allan Hancock Foundation (AHF). When available, museum catalogue numbers are included with the lists of materials examined.

Based upon the study of these new materials and reexamination of older ones, the total number of named spionid species from the region is increased to 59. Another 8 species are identified but not named either due to fragmentary specimens or to difficult taxonomic problems yet to be resolved. Two new genera are erected, and 12 species are new to science. Revisionary comments, including new synonyms, are presented for another 12 species. There has been no effort to provide fauna 1 keys since the collections encompass such diverse zoogeographic areas. Instead, the reader is referred to Blake and Kudenov [1978] for keys and diagnoses to the genera. Each species is provided with descriptive details or descriptive remarks to enable readers to use this paper independent of other sources. Synonymies are complete for poorly known species, with the better known species including major references. The following species are included in this report (new synonyms are in brackets):

Scolelepis aitutaki Gibbs, 1972 Scolelepls chllensIs (Hartmann-Schroder,

1962), new combination Scolelepis eltaninae, n. sp. Scolelepis quinquedentata (Hartmann

Schroder, 1962), new combination Scolelepis gaucha (Orensanz and Gianuca,

1974), ~mbination Scolelepis sp. A Dispio brachychaeta, n. sp. Dispio uncinata Hartman, 1951 ~s paucibranchiata Southern, 1914 Scolecolepides uncinatus, n. sp. ?Scolecolepides sp. Malacoceros indicus (Fauvel, 1928)

[Spio cf. obtusa: Hartmann-Schroder l~--not Ehlers, 1913] ,

Rhynchospio glutaea (Ehlers, 1897) Laonice antarcticae Hartman, 1953 Laonice weddellia Hartman, 1978 Laonice cf. cirrata (Sars, 1851) Laonice sp. A Paraprionospio pinnata (Ehlers, 1901) Prionospio (Prionospio) orensanzi, n.sp. Prionospio (P.) cf. ehlersi Fauvel, 1928 Prionospio (P.) pygmaea Hartman, 1961 •. Prionospio (P.) steenstrupi Malmgren, 1867 Prionospio (P.) sp. A Prionospio (P.) sp. B Prionospio (Aquilaspio) peruana Hartmann

Schroder, 1962 Prionospio (Minuspio) sp.--cirrifera

complex

Prionospio (Minuspio) patagonica Augener,1923 r po. cirrifera chilensis HartmannSchroder, 1962]

Spiophanes bombyx (Claparede, 1870) Spiophanes ~stroemi Hartman, 1953

[So chilensis Hartmann-Schroder, 1965] Spiophanes kroeyeri Grube, 1860 Spiophanes tcherniai Fauvel, 1951 Spio quadrisetosa, n. sp. Spio sp. A M1Crospio hartmanae, n. sp. Microspio paradoxa, n. sp. Microspio minuta (Hartmann-Schroder, 1962) Microspio moorei (Gravier, 1911) Microspio gracIlis (Hartmann-Schroder,

1962), new combination Pygospiopsis dubia (Monro, 1930), n. g.,

new combination Boccardia acus (Rainer, 1973) Boccardia ~hthalma (Rioja, 1962) Boccardia chilensis Blake and Woodwick,

1971 Boccardia natrix (Soderstrom, 1920) Boccardia ~anchia (Haswell, 1885)

[Polydora paucibranchus Ehlers, 1913] [B. (Boccardia) wellingtonensis Read, 1975]

Boccardia tricuspa (Hartman, 1939) Boccardiella ligerica (Ferronniere, 1898) Boccardiella occipitalis, n. sp. Boccardiella sp. A Carazziella carrascoi Blake, 1979 Carazziella patagonica Blake, 1979 Amphipolydora abranchiata (Hartman,

1953),n. g., new combination Polydora colonia Moore, 1907 Polydora cirrosa Rioja, 1943 Polydora ligni Webster, 1879 Polydora CiIIita (Johnston, 1838) Polydora websteri Hartman, 1943 Polydora biocciptalis Blake and Woodwick,

1972 Polydora rickettsi Woodwick, 1961 Polydora ecuadoriana, n. sp. Polydora armata Langerhans, 1881 Po1ydora ~runnae, n. sp. Polydora magel1anica, n. sp. Po1ydora giardi Mesni1, 1896 Po1ydora ~C1aparede, 1870 Po1ydora SOCIi1is (Schmarda, 1861) Po1ydora sp. A Pseudopo1ydora primigenia, n. sp.

Systematic Account of Spionidae

Genus Sco1e1epis B1ainvi11e, 1828 Sco1e1epis aitutaki Gibbs, 1972 Emended

Fig. 1

Sco1e1epis (Sco1e1epis) aitutakii Gibbs, 1972, pp. 211-212, fig. 6.

Material examined. Uruguay: IBM Sta. N-244, 128 m (3, USNM 67559).--Argentina,

BLAKE: SOUTH AMERICAN AND ANTARCTIC SPIONIDAE 207

subtidal: IBM Sta. N-I054, 58-65 m (1, USNM 67560); Sta. N-I066, 72-86 m (1, USNM 67561). --Argentina, intertidal in sand, collected by J. M. Orensanz: Caleta de los Loros, February 1972 (20+, USNM 67563); Las Crutas, July 26, 1971 (1, USNM 67562), January 14, 1973 (1, USNM 67564); Puerto Lobos, January 21, 1973 (2, USNM 58991); Colfo San Jose, San Ramon, February 17, 1975 (20+, USNM 58989); Colfo Nuevo, Puerto Madryn, November 15, 1975 (2, USNM 58990).--Cook Islands, Aitutaki Lagoon, September 1969, collected by P. E. Cibbs, New Zealand Royal Society Expedition, Sta. AL16, 4.5-6 m, silty sand (holotype, BMNH ZB 1972:6).

Description. A moderate-sized species, up to 19 mm long and 1.5 mm wide for up to 60 setigers. Color in alcohol: pink, with yellow tipped branchiae.

Prostomium acutely pointed on anterior margin, extending posteriorly as caruncle to middle of setiger 1 (Figure lA); without occipital tentacle; 2 pairs of inconspicuous, subcuticular eyes; palps extending posteriorly for 15 or more setigers (not shown on Figure lA). Peristomium well developed, with low lateral wings.

Setiger 1 reduced, with long, fingerlike notopodial lamellae, lacking notosetae; neuropodial lamellae small, rounded, with fascicle of delicate capillary setae. Notopodial lamellae of setigers 2 to 15-17 entirely fused to branchiae (Figure IB), separation occuring more posteriorly as notches near branchial tips, and finally progressing to complete separation of lamellae and branchiae by setiger 25 with gill having expanded, flaglike tip bearing yellow pigment and lamella reduced to lateral earlike lobes (Figure lC); dorsal ridges present, connecting bases of branchiae (Figure lA). Neuropodial lamellae of se tigers 2-25 entire, rounded (Figure IB), thereafter elongating dorsally, becoming less rounded and eventually dividing into small ventral lamellae and larger triangular interramal lamellae (Figure lC).

Notosetae all capillaries, arranged in 2 rows in anterior setigers and in single rows in posterior setigers; usually with 3-4 additional long capillaries located near superior end of anterior row; setae of anterior row and superior group with granulated shafts and clear, narrow sheaths (Figure IH); posterior row of capillaries lacking granulations. Neurosetae of anterior se tigers arranged in 2 rows with granulated capillaries in anterior row and nongranulated capillaries in posterior row; no inferior sabre setae; bidentate or tridentate hooded hooks present from setiger 22; hooks numbering 1-2 at first, then increasing to 12-16 and accompanied by 1-5 delicate capillaries randomly dispersed among hooks (Figure lC), hooks with

apical tooth appearing single or partially divided into 2 separate teeth (Figures IDIF), hood inflated, with smooth opening, about 10 times longer than main fang.

Pygidium an irregular disc, with large dorsal anal opening (Figure lC).

Remarks. A comparison of the holotype of Scolelepis aitutaki from the Cook Islands with specimens from Uruguay and Argentina establishes that they are the same species. There are tridentate hooded hooks in addition to the bidentate ones described by Cibbs [1972]. By having branchiae with flaglike expansions of the tips, S. aitutaki resembles S. gilchristi (Day) from-South Africa. Scolelepis gilchristi, however, has notosetae of setiger 1, an occipital tentacle and multidentate hooded hooks while S. aitutaki lacks notosetae on setiger 1, has no occipital tentacle and has bidentate and tridentate hooded hooks. Nerinides vexillatus Hutchings and Rainer [1979] from Australia is probably referrable to S. aitutaki. Minor differences emphasized by the authors in the point at which the fused branchiae and dorsal lamellae separate and in the number of capillary setae accompanying the hooded hooks do not appear to be very significant, especially since only a single anterior fragment was examined.

Distribution. Cook Islands, 4-6 m in silty sand; Argentina and Uruguay, intertidal to 65 m in sand; ? Australia.

Scolelepis chilensis (Hartmann-Schroder, 1962)

new combination Fig. 2

Nerine cirratulus chilensis Hartmann--schroder, 1962b, pp. 142-144, figs. 179-

182. Neriides sp. [sic] Cuiler, 1959, p. 179.

Material examined. Chile, Mehuin, midintertidal in sand, March 30, 1975, collected by C. Bertran (3, AHF): Montemar, Cuchoa Beach in sand, February 1955, collected by E. R. Cuiler (5, AHF); Huasco, intertidal in sand, July 28, 1960, collected by C. Hartmann-Schroder (holotype, ZMH P-14925).

Description. A large species, with 1 incomplete specimen from Mehuin measuring 40 mm long and 6 mm wide for 43 setigers, 2 smaller, incomplete specimens from same locality measuring 26 and 20 mm long for 48 and 39 setigers, respectively; holotype complete, measuring 50 mm long and 2.5 mm wide for 135 setigers. Color in alcohol: dark gray, light brown, or opaque white.

Prostomium anteriorly elongated, distally pointed, continuing posteriorly as caruncle to end of setiger 1 (Figure 2A); 2 pair of eyes arranged in transverse line: medial

~H E F

500pm

T 300JIID

1

r 20Jlm

L D

Fig. 1. Scolelepis aitutaki (USNM 67563). A, anterior end in dorsal view; B, right setiger 7 in anterior view; D, right setiger 30 in anterior view; D, hooded hook in lateral view; E, F, hooded hooks in frontal view with hoods omitted; G, pygidium and 2 posterior setigers in dorsal view; H, capillary notoseta from anterior parapodium (inset not to scale).

"

500Jlm

Fig. 2. Scolelepis chilensis (Mehiun specimen AHF). A, anterior end in dorsal view; B, posterior end in dorsal view; C, left setiger 30 in anterior view; E, F, capillary notosetae from anterior row of anterior setiger; G, inset of F (not to scale); H, I, capillary notosetae from posterior row of anterior setiger; J, neuropodial hooded hook from middle of body in lateral view; K, L, same in frontal view.


pair oval, lateral pair cup-shaped. Peristomium well developed, forming moderate lateral wings, not overlapping prostomium. Palps thickened, with basal inflation, each palp extending posteriorly for 11-13 setigers.

Setiger I reduced, with short postsetal lamellae; dorsal lamellae longer than ventral (Figure 2A). Dorsal lamellae and branchiae fused from setiger 2, being free only at their tips (Figure 2C), then becoming distinctly separated by setiger 25 and retaining only basal fusions in far posterior setigers lFigure 2D). Neuropodial lamellae elliptical in anterior setigers (Figure 2C), divided in middle setigers forming small ventral lamel-' lae and elongated, thickened interramal lamellae, continuing through posterior se tigers (Figure 2D).

Anterior notosetae and neurosetae all cap" illaries. arranged in 2 rows; most setae short, thickened, numbering 30,-32 per ramus, with 4-5 longer notosetae in superior position near anterior row; capillaries with minute granulations along internal striations of shafts (Figures 2E 21), sheath transpar" ent, with fine striations apparent at 1000X (Figure 2G); hooded hooks begining in neuropodia from se tigers 25-32, numbering 4-5, accompanied by 8··9 capillaries lacking granulations and 3-4 granulated inferior sabre setae. Hooks beginning in notopodia from setiger 32-60, numbering 2-3 among longer capillaries. Hooks bidentate (Figures 2J-2L) with unique apical tooth appearing to be formed of 2 incomplete paired teeth joined by connecting ridge, having prominent notch; shaft prominently bent; hood with faint striations.

Pygidium forming ventral cushion (Figure 2B). Remarks. Scolelepis chilensis, originally

described as a subspecies of Nerine cirratuIus [>Scolelepis squamata (Muller)] is herein elevated to full species status. The species is most closely related to S. squamata in having bidentate hooded hooks ~ posterior segments and notosetae on setiger 1. S. chilensis has the apical teeth of the hooded hooks fused by a ridge, whereas .§.. squamata is reported to have unidentate, bidentate, and tridentate hooded hooks with distinct apical teeth which are frequently worn [Foster, 1971a; Light, 1978j. In S. chilensis, the anterior dorsal lamellae-are separated from the gills only at their tips, while in S. squamata the separation is more complete.- The fine striations observed on the hoods of the hooded hooks and sheaths of capillaries of S. chilensis have not been reported for S.-squamata. Further, the ventral lamellae-of posterior neuropodia are poorly developed in.§.. chilensis and f<lrm prominent papillae in S. squamata.

Distribution. Chile, intertidal in sand.

Scolelepis eltaninae, n. sp. Figs. 3, 4

Material examined. Ross Sea, Robertson Bay, Eltanin Sta. 2126, 143 m (holotype and 10 paratypes USNM 60428-9); Pennell Bank, Eltanin Sta. 5761, 388-399 m (3 paratypes, USNM 69401).--Bransfield Strait, Eltanin Sta. 410, 220-240 m (1 specimen, USNM 61961).

Description. All specimens anterior fragments, with largest measuring 17 mm long and 2.6 mm wide for 38 setigerous segments. Color in alcohol: light tan to flesh colored, no body pigment.

Prostomium pointed on anterior margin, flaring subterminally, continuing posteriorly to bases of palps, terminating in large occipital tentacle (Figure 3A); 2 pairs of light red eyes on anterior face of tentacle (Figure 4A), posterior pair oval, anterior pair cup-shaped; palps with length equivalent to 7-8 setigers, thickened basally, with conspicuous sheath (Figure 3B). Peristomium with well-developed lateral wings.

Setiger 1 ,"ell developed, wi th conical notopodial lamellae bearing long capillary notosetae and elliptical neuropodial lamellae with spreading fascicles of capillary neurosetae (Figures 3A and 4A). Notopodial lamellae from setigers 2-20 fused to branchiae, being free only at their tips (Figures 4B-4C); nature of fusion in far posterior notopodia unknown; lower portion of notopodial lamellae directed ventrally toward neuropodium from setiger 25, forming interramal channel (Figure 4D). Neuropodial lamellae of setigers 2-17 somewhat elongated dorsally (Figure 4B), becoming divided by setiger 18-23 into large, triangular and dorsally directed interramal lamellae and smaller, elliptical neuropodial lamellae (Figure 4D).

Notosetae all capillaries (at least to setiger 38), arranged into 2 rows in anterior setigers and single rows more posteriorly; 3-4 longer, superior capillaries present at dorsal end of posterior row, anterior capillaries with finely granulated shafts and clear, narrow sheaths (Figure 3C); granulations diminished or inconspicuous on capillaries by about setiger 25. Neurosetae on anterior se tigers similar in arrangement and structure to notosetae except for lower group of 3-4 long granulated capillaries (Figures 4B-4C), these setae gradually thickening over successive setigers, becoming sabre setae with commencement of neuropodial hooks from setiger 20-24; hooks numbering up to 7-8 per neuropodium, accompanied by 4-6 thin, simple capillaries and 7-8 granulated sabre setae (Figure 4D); hooks unidentate, acicular, some exhibiting remnants of closely adhering hood (Figure 3D).

Nature of posterior setigers and pygidium not known.

BLAKE: SOUTH AMERICAN AND ANTARCTIC SPIONIDAE

(

\ i ( l

~ , /' ~ T I t :: ~ j) .:~

~~m ( ~

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Fig. 3. Scolelepis eltaninae n. sp. (paratype, USNM 60429). A, anterior end in dorsal view; B, basal part of palp; C, capillary notoseta from anterior segment; D, unidentate neuropodial hooks and bases of accompanying capillaries.

211


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T O.5mm

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Fig. 4. Scolelepis eltaninae n. sp. (paratype, USNM 60429). A, prostomium and right setiger 1 in anterior view, showing occipital tentacle and position of eyes; B, right setiger 5 in anterior view; C, right se tiger 20 in anterior view; D, right se tiger 30 in anterior view.


Remarks. Scolelepis eltaninae is most closely related to S. foliosa (Audouin and Milne-Edwards) from-the North Atlantic in having an occipital tentacle and neuropodial unidentate acicular hooks. S. eltaninae, however, has only hood fragm~nts, while S. foliosa has conspicuous hoods surrounding the ends of the hooks. In S. eltaninae. the anterior branchiae and dorsal lamellae are free at their tips, while in S. foliosa, they are completely fused along their lengths. Furthermore, S. eltaninae has a conspicuous basal palpal sheath and inconspicuous eyes at the base of the occipital tentacle, while S. foliosa lacks palpal sheaths and has conspicuous eyes on the prostomium anterior to the occipital tentacle.

Etymology. This species is named for the USNS Eltanin, the former research USARP research vessel.

Distribution. Antarctica, the Ross Sea and Bransfield Strait, 143-399 m.

Scolelepis quinquedentata (Hartmann-Schroder, 1965)

new combination

Nerinides quinquedentata Hartmann-Schroder, 1965, pp. 203-204, figs. 187-190.--Carrasco, 1974, pp. 189-190, figs. 10-14; 1976, pp. 18-21, figs. 5-6.

Material examined. Chile, Isla Mocha, 26 m, dark fine sand, March 11, 1960, collected by G. Hartmann-Schroder (holotype and 4 paratypes, ZMH P-14928-9).

Remarks. Only fragmentary specimens of S. quinquedentata have been described. The oc= cipital tentacle, not reported by HartmannSchroder [1965], but clearly depicted by Carrasco [1974, Fig. 11] is definitely present on the holotype and paratypes.

Scolelepis quinquedentata belongs to a group of closely related species including S. papillosa (Okuda) and S. yamaguchi (Imajima) from Japan, S. trid~ntata (Southern) from the North Atlantic (and reported from California [Light, 1977, 1978]), S. bousfieldi Pettibone from eastern United States and S. texana Foster from the Gulf of Mexico and eastern United States. These species all have multidentate hooded hooks with a long main fang, an occipital tentacle, and a basal fringe of papillae around the palps. Scolelepis quinquedentata, S. texana, and S. papillosa lack notosetae on-se tiger 1. Scolelepis papillosa is known from only a single, posteriorly incomplete specimen [Okuda, 1937J in which the branchiae remain fused to the dorsal lamellae throughout the fragment. In S. quinquedentata and S. texana, the branchiae of posterior n~topodia are separated from an earlike postsetal lamella. The posterior neuropodia of

~. ~ are distinctly divided into an interramal lamella and ventral papilla, S. quinquedentata has only the interramal lamella, while the neuropodia of S. papillosa are undivided. These data suggest that S. texana and S. quinquedentata are so similar-that the form~r may be no more than a subspecies of the latter. Additional specimens of all 3 species are needed in order to clarify these relationships.

Distribution. Chile, shallow subtidal sandy substrata.

Scolelepis gaucha (Orensanz and Gianuca, 1974) new combination

Spio (Microspio) gaucha Orensanz and Gianuca, --r974, pp. 16-17~ 10.

Remarks. Specimens of this species have not been examined, but the description and figures published by Orensanz and Gianuca [1974J clearly indicate that it belongs to the genus Scolelepis rather than Spio (Microspio). The prostomium is pointed on-rhe anterior margin and the branchiae, which begin on setiger 2, are fused to the dorsal lamellae in anterior setigers. By having notosetae on setiger 1 and neuropodial bidentate hooded hooks, S. gaucha appears to be closely related to S. ~ta except that notopodial hooks are lacking. The species is unusual in that the pygidium is terminally divided into 4 cirriform appendages.

Distribution. Southern Brazil and Uruguay.

Scolelepis sp. A

Material examined. Weddell Sea, Glacier Sta. 68-1, 650 m (1 specimen, USNM 61963).

Remarks. The specimen is a small, anterior fragment which lacks notosetae on setiger 1 and has no occipital tentacle or hooded hooks. It differs from others reported from the region but is too fragmentary to be further identified.

Distribution. Weddell Sea, 650 m.

Genus Dispio Hartman, 1951 Emended

Type species: Dispio uncinata Hartman, 1951, by monotypy.

Diagnosis. Prostomium fusiform, anteriorly pointed, with narrow caruncle extending posteriorly. Eyes present or absent. Peristomium moderately developed, forming low lateral wings. Anterior parapodial lamellae lobed or entire. Presetal notopodial and neuropodial lamellae present or absent. Branchiae present from setiger 1, fused to notopodial lamellae for half or more of their length. With or without accessory branchiae on posterior face of middle and posterior no-


topodia. With capillary notosetae only. Neurosetae include capillaries, hooded hooks, and sabre setae. Some capillaries of anterior rows with distinct transverse barring pattern, composed of partitions and chambers in shaft, giving setae cracked appearance; sometimes reticulations interspersed with dense groups of granules. Pygidium with midventral flap and prominent anal cirri.

Remarks. Dispio is closely related to Scolelepis and differs chiefly in having branchiae from setiger 1 instead of 2 and some capillary setae with distinct reticulated barring patterns instead of all capillaries with uniformly arranged granules or no granules at all. Most species of Dispio have the postsetal lamellae of anteriormost parapodia divided into separate lobes or serrations, have accessory branchiae in middle and posterior notopodia, and have prolonged notosetae on setiger 1. The absence of 1 or more of the latter 3 characters occurs among various species and all 3 are lacking in D. brachychaeta (see below). For this reason, they should be used only as species level characters.

Dispio brachychaeta, n. sp. Fig. 5

Material examined. Argentina, off the Rio de la Plata, IBM Sta. Comp V-79, 31 m (holotype, USNM 67512).

Description. Holotype large, incomplete, measuring 35 mm long and 3 mm wide for 100 segments. Color in alcohol: yellow to orange.

Prostomium pointed on anterior margin; caruncle short, merging with setiger 1; eyes and occipital tentacle absent, but with scar suggesting possible tentacle location, prominent semicircular nuchal area across setigers 2-3 (Figure 5A).

Setiger 1 reduced, with dorsal lamellae about half as large as those of subsequent setigers (Figure 5A). Branchiae from setiger 1, entirely fused to notopodial postsetal lamellae in anterior setigers (Figure 5B), with branchial tips becoming free at setiger 20 (Figure 5C) but not entirely free from lamellae until about setiger 80 (Figure 5D), notopodial postsetal lamellae longest and pointed in anterior setigers, becoming shorter and blunt-tipped in posterior setigers. Neuropodial lamellae entire, rounded on setigers 1-30 (Figure 5B), becoming dorsally pointed and elongated more posteriorly; interramal channel between notopodial and neuropodial lamellae with lateral ciliated organs developing in posterior setigers.

Notosetae all capillaries, arranged in 2 rows in anterior se tigers and in single rows in middle and posterior setigers, setae of setiger 1 not elongated as in other species of genus, being similar in length to follow-

ing setigers; anterior row of capillaries heavier, bearing granulations and distinct transverse bars on shaft (Figure 5E), sheath clear; separate group of longer, thinner capillaries present dorsal to anterior row; capillaries of posterior row more delicate, lacking granulations and transverse bars. Neurosetae of anterior setigers lacking granulations and transverse bars, arranged in 2 rows; 4-6 unilimbate inferior sabre setae with granulations on distal end of shaft (Fjgure 51); 2-3 unidentate hooded hooks replacing anterior row of capillaries from setiger 32; hooks with closely applied hoods (Figures 5F-5H), frequently torn from shaft; unilimbate capillaries accompanying hooks in posterior row.

Nature of pygidium unknown. Remarks. Dispio brachychaeta differs from

other species~e genus in lacking distinct lobes or serrations on the dorsal lamellae and accessory branchiae on the posterior setigers and having short notosetae on se tiger 1 rather than long prominent ones. The species is referred to Dispio because branchiae occur from setige~d transverse bars are present on capillary notosetae of the anterior row. The latter character has been observed in 2 species of Dispio and appears typical for the genus [Light, 1978; Blake and Kudenov, 1978].

Etymology. Brachychaeta is derived from the Greek brachy, for short and chaeta, for seta. The-uame-is suggested by the short notosetae on setiger 1.

Distribution. Argentina, in sand, 31 m.

Dispio uncinata Hartman, 1951

Dispio uncinata Hartman, 1951, pp. 87-90, pIs. 22-23; 1961, p. 88; 1969, pp. 105-106, figs. 1-4.--Bellan, 1968, pp. 49-55, figs. 1-2.--Foster, 1971a, pp. 73-78, figs. 161-174 [synonymyJ.--Fauchald, 1973, pp. 24, 28.--Light, 1977, p. 78; 1978, pp. 113-116, figs. 113, 115.--Ibanez and Vieitez, 1973, pp. 5-8, fig. l.--Hartmann-Schroder, 1979, pp. 133-134, figs. 291-298.

Material examined. Argentina, Puerto Lobos, intertidal in sand, January 21, 1973, collected by J. M. Orensanz (1, USNM 67530).

Remarks. Dispio uncinata has been well described by Bellan [1968J, Foster [1971aJ, and Light [1978J. The species is characterized by having long notosetae on setiger 1, anterior notopodial postsetal lamellae divided into a variable number of fingerlike lobes or serrations, accessory branchiae located basally on the posterior faces of notopodia from setiger 18-28 and anterior capillaries with distinct reticulations (barring patterns) on the shaft. The present specimen appears to be typical.

I

2mm

T 500)1 m

1 F G

T 5()C

I c

Fig. 5. Dispio brachychaeta n. sp. (holotype, USNM 67512). A, anterior end in dorsal view; B, right se tiger 13 in anterior view; C, right setiger 70 in anterior view; D, left setiger 100 in anterior view; E, capillary notoseta from an anterior setiger; F-H, hooded hooks in various orientations and degrees of separation of the hood; I, inferior sabre seta.


Distribution. East and west coasts of North America, Gulf of Mexico, Spain, northwestern Australia, Argentina. Intertidal to shallow subtidal depths, in sand.

Genus Aonides Claparede, 1864 Aonides paucibranchiata Southern, 1914

Fig. 6

Aonides paucibranchiata Southern, 1914, pp. 100-101, pI. 11, figs. 24A-24E.--Fauvel, 1927, p. 40, fig. 13f-13k.--Friedrich, 1938, p. 128.--Hartmann-SchrHder and Stripp, 1968, pp. 16-17, figs. 7f-7k. --Hartmann-SchrHder, 1971, pp. 329-331, fig. 113.

Material examined. Argentina, IBM Sta. SAO-I-52, 29 m (1, USNM 58960); Sta. SAO 1-24, 15 m (1, USNM, 67525); Sta. SAO I-A, 15 m (3, USNM 67524); Golfo San Mat{as, Las Grutas, intertidal, February 9, 1971, collected by J. M. Orensanz (1, USNM 67526).

Description. Argentinian specimens small, up to 2.2 mm long and 0.12 mm wide for 30 setigers. Color in alcohol: brown or light green.

Prostomium fusiform, narrowed anteriorly, extending posteriorly to middle of setiger 1; without eyes or occipital tentacle (Figures 6A-6B). Peristomium reduced, with weakly developed lateral wings; palps not observed.

Setiger 1 well developed, with short postsetal lamellae and prominent setal fascicles; notopodial lamellae of setiger 2 and subsequent setigers leaflike, becoming smaller, conical in posterior setigers; neuropodial lamellae short, fingerlike.

Notosetal and neurosetal capillaries with narrow sheaths, granulations lacking; neuropodia with hooded hooks from setiger 11-13 and notopodia from setiger 18; hooks bidentate or tridentate with main fang surmounted by 1 or 2 apical teeth in a vertical series, hood short, with finely serrated opening and shaft weakly constricted (Figure 6C); neuropodial inferior sabre setae from setiger 18 lacking granulations or sheath; hooks accompanied by capillaries throughout.

Branchiae from setiger 2, numbering 3-4 pairs. Pygidium with 2 short lobes and 3 longer, fingerlike cirri (Figure 60).

Remarks. In all previous accounts of Aonides paucibranchiata, the hooded hooks were said to be tridentate. This is the first report of the co-occurence of bidentate and tridentate hooks and of serrations on the hood opening. The possibility that A. paucibranchiata is a juvenile of A. oxycephala cannot be ruled out at the present time. The latter species is apparently cosmopolitan in distribution. It has bidentate hooded hooks and is considerably larger. A thorough study of the larval and ju-

venile morphology of A. oxycephala is needed in order to determine-if there is an overlap in characters of the 2 species.

Distribution. Northeastern Atlantic, North Sea, Skagerrak, Black Sea, Argentina in shallow sandy substrata.

Genus Scolelcolepides Ehlers, 1907 Scolecolepides uncinatus, n. sp.

Fig. 7

Material examined. Argentina, intertidal, collected by J. M. Orensanz: Riacho jabali, sand and mud, with planL debris, October 4, 1968 (holotype and 13 paratypes, USNM 67518-9); Riacho jabali, north of the bridge, April 17, 1970 (39 paratypes, AHF Poly 1294); same locality, October 17, 1970 (15 paratypes, AHF Poly 1293); Caleta de los Loros, midintertidal, in sand, February 1972 (paratype, USNM 58980); Bahia de San Antonio, in sand, February 1971 (10, paratypes, USNM 58979).

Description. A large species, up to 60 mm long and 3 mm wide for about 160 setigers. Color in alcohol: brown to light tan.

Prostomium truncate on anterior margin, with short lateral horns (Figure 7); prostomial ridge elevated between palp bases, then tapering, with caruncle merging with se tiger 2; 2 pairs of indistinct, deeply imbedded eyes (not shown on Figure 7A); palps basally thickened, extending to setiger 15.

Branchiae from se tiger 1, small at first, then increasing to full size on setigers 2-4; branchiae fused basally to dorsal lamellae throughout, absent from posterior one third of body; branchiae with cilia on inner and outer margins. Notopodial lamellae elongate on anterior setigers, tapering to fine tips (Figure 7C), becoming smaller, leaflike in posterior setigers (Figure 70). Neuropodial lamellae of setiger 1 triangular in shape (Figure 7B), becoming elliptical more posteriorly (Figure 7C), with those of middle and posterior se tigers bearing a small nipplelike projection (Figure 70).

Capillary notosetae of anterior se tigers unilimbate, lacking granulations (Figure 7E), arranged in 2 rows. Neurosetae of setigers 1-8 similar in form and arrangement to notosetae except capillaries of posterior row becoming thicker and darker and replaced by dark, thickened acicular spines on setigers 9-18 (Figures 7F-7H); these acicula numbering 6-7 per neuropodium and accompanied by numerous smaller capillaries, 3-4 curved inferior sabre setae in neuropodia from about setiger 40. Unidentate hooded hooks beginning in neuropodia from about setiger 60 and in notopodia from setiger 90-100; up to 6-7 in neuropodia and 3-4 in notopodia; hooks interspersed throughout with fine capillaries (Figure 70), hooks with narrow tips surround-


30pm

Fig. 6. Aonides paucibranchiata (USNM 67524). A, anterior end in dorsal view; B, prostomium from another specimen showing shape variability; C, hooded hooks in lateral view; DJ pygidium in dorsal view.

ed by long, delicate striated hoods having serrated openings (Figures 71-7J).

ures 57 and 58]. Scolecolepides uncinatus is unique in having unidentate hooded hooks.

217

Pygidium forming thickened ring with numerous cirri.

Remarks. Scolecolepides currently includes 4 species: S. viridis (Verrill, 1873) from eastern North-America and Alaska, S. benhami Ehlers, 1907 from New Zealand, S. aciculatus Blake and Kudenov, 1978 from-Australia, and S. uncinatus from Argentina. The 3 southern h;mispheric species differ significantly from S. viridis in having short, lateral subte~inal prostomial horns instead of nearly T-shaped frontal horns [George, 1966, Figures la and Ib; Foster, 1971a, Fig-

Etymology. Uncinatus is Latin for hooked or barbed, and refers to the unidentate hooded hooks.

Distribution. Argentina, intertidal in sand.

?Scolecolepides sp.

Material examined. Argentina, Tierra del Fuego, Hero Sta. 71-2-6 (6,USNM 60427). Rema~ These specimens are anterior

fragments superficially resembling Scolecolepides uncinatus (see above). Neuropodial

Imm

I J

Fig. 7. Scolecolepides uncinatus n. sp_ (paratype, USNM 67519). A, anterior end in dorsal view; B, left setiger 1 in anterior view; C, right setiger la in anterior view; D, right setiger 100 in anterior view; E, anterior capillary notoseta from setiger 5; F-H, thickened neuropodial spines from setiger 12, showing various degrees of wear; I, J, hooded hooks in frontal (1) and lateral (J) views.


hooded hooks beginning in setigers 30-32 are bidentate, with a small and frequently worn apical tooth which resembles that of S. aciculatus Blake and Kudenov [1978J from-AuS=tralia. Notopodial hooks were not present in these fragments precluding generic confirmation.

Distribution. Tierra del Fuego, intertidal.

Genus Malacoceros Quatrefages, 1843 Malacoceros indicus (Fauvel, 1928)

Scolelepis indica Fauvel, 1928a, p. 90. Malacoceros~coceros) indicus: Foster,

1971a, pp. 50-53, figs. 93 99 [synonymy]; 1971b, pp. 1455-1457, figs. 1-6.

Malacoceros indicus: Blake and Kudenov, 1978, p. 195.

Spio cf. obtusa: Hartmann-Schroder, 1965, -p. 214, figs. 205-207. [not Ehlers, 1913]

Material examined. Chile, Galera bis Quedal, 80-84 m, in fine sand and debris, March 1960, collected by G. Hartmann-Schroder (6, ZMH P-14943 as Spio cf. obtusa).

Remarks. These specimens exhibit all of the characters of Malacoceros indicus as reported by Foster [1971a, b]. The outer border of the neuropodial lamellae bears a distinct nipple, and the hooded hooks include both tridentate and quadridentate types.

Distribution. Circumtropical; Chile, 80 m in fine sands.

Genus Rhynchospio Hartman, 1936 Rhynchospio glutaea (Ehlers, 1897)

Scolecolepis glutaea Ehlers, 1897, pp. 83-85, pI. 5, figs. 129-132, pl.6, figs. 133-135; 1901a, p. 165

Scolecolepis cornifera Ehlers, 1913, pp. 509-510, pI. 36, fig. 5.

Microspio glutaea: Soderstrom, 1920, p. 252, figs. 162-163.--Augener, 1932a, p. 108.

Rhynchospio arenincola Hartman, 1936, p. 51, figs. 20 22; 1969, p. 171, figs. 1-3. --Wieser, 1959, pp. 105-106.--Banse, 1963, pp. 203-204.--Blake, 1975, pp. 211, 215, fig. 242.

Scolelepis cornifera: Monro, 1939, pp. 125-126, fig. 14.--Fauvel, 1953a; 1953b, p. 308.

Scolelepis cornigera [sic]: Fauvel, 1952, p. 297.

Rhynchospio glutaea: Hartman, 1953, p. 42; 1966, pp. 21-22, pI. 5, figs. 8-10.--Day, 1961, p. 491; 1967, p. 478, fig. 18.6a-18. 6c.--Hartmann-Schroder, 1962b, p. 138; 1965, pp. 213-214.--Carrasco, 1974, pp. 196-197, figs. 33-36, 1976, pp. 45-48, figs. 18, 21-22.--Blake and Kudenov, 1978, p. 199.--Bellan, 1975, p. 790.

Rhynchospio arenicola [sic] asiatica Khlebo-

vitsch, 1959, pp. 175-176, fig. 6; 1961, p. 196. Fide Foster, 1971a.

Malacoceros-cRhynchospio) glutaeus: Pettibone, 1963, p. 90.--Foster, 1971a, pp. 53-56, figs. 100-111 [synonymy].

Malacoceros (Rhynchospio) arenincolus: Pettibone, 1963, p. 90. Fide Foster, 1971a.

Material examined. Argentina, intertidal, collected by J. M. Orensanz: Santa Elena, among rocks, December 23, 1968 (12, USNM 67558); Mar del Plata, rocky area (20, USNM 67557); Golfo Nuevo, Puerto Madryn, sand beach, 1975 (3, USNM 58962).--Argentina, Tierra del Fuego, Hero Sta. 656, 18 m (1, USNM 60355); Sta. 658, 13-34 m (4, USNM 60358); Sta 664, 29 m (1, USNM 60359); Sta. 71-2-8, intertidal (4, USNM 60353); Sta. 71-2-14, intertidal (3, USNM 60354).--Argentina, Staten Island, off Tierra del Fuego, Hero Sta. 71-2-16, intertidal (9, USNM 60356); Sta. 71-2-41, intertidal to 1 m (1, USNM 60360).--Falkland Islands, April 22, 1927, Sta. 115, collected by W. L. Schmitt (1, USNM 24437); Eltanin Sta. 344, 119 m (2, USNM 60352).--off Cape Horn, Eltanin Sta. 453, 31 m (2, USNM 61960).

Remarks. All specimens are small and agree well with the numerous descriptions of Rhynchospio glutaea. Hooded hooks are bidentate. The frontal horns sometimes project forward, forming a distinct notch on the anterior end of the prostomium. Rhynchospio is considered to be distinct from Malacoceros because the branchiae begin on setiger 2 instead of setiger 1 (see Blake and Kudenov, [1978, pp. 198-199] for discussion).

Distribution. West coast of North America; Florida; Kurile Islands; South Africa; Chile, Argentina, MacQuarie Island; South Georgia; Falkland Islands; Kerguelen Islands. Intertidal to 120 m.

Genus Laonice Malmgren, 1867 Laonice antarcticae Hartman, 1953

Fig. 8

Laonice cirrata antarcticae Hartman, 1953, pp. 40-41.

Laonice antarcticae: Hartman, 1965, pp. 147-148; 1978, p. 61 [in part].--Banse and Hobson, 1968, p. 24.--Hartman and Fauchald, 1971, p. 104.

?Laonice cirrata: Hartman, 1967, p. 112 [in part [not Sars, 1851] spionids, Hart-man, 1967, p. 114 [in part].

Material examined. Uruguay, 3300'S; 51°10'W, 80 m, dark gray mud, December 12, 1901, collected by the Swedish Antarctic Expedition, 1901-1903 (2 syntypes, SMNH 613).--Argentina, IBM Sta. N-I055, 92-96 m (2, USNM 67531); Sta. N-I066, 72-86 m (2, USNM 67532); Sta. A 11-60-282, 200-215 m (3


':.: ..

",:

Fig. 8. Laonice antarcticae (syntype, SMNH 613). A, anterior end in dorsal view; B, hooded hook in lateral view; C, inferior sabre seta; D, detail of C (not to scale).

juveniles, USNM 67533 .--South Shetland Islands, Eltanin Sta. 410, 220-240 m (5, USNM 60538; 20, USNM 61948); Sta. 418, 311-426 m (6, USNM 60540); Hero Sta. 726, 64-82 m (1, USNM 60544).--Bransfield Strait, Eltanin Sta. 412, 1180 m (1, USNM 60539); Sta. 997, 769 m (7, USNM 61950).--Antarctic Peninsula, Eltanin Sta. 272, 412 m (1, USNM 56390); Sta. 439, 128-165 m (1, USNM 60541).--Ross Sea, Eltanin Sta. 2031, 535 m (1, USNM 60543); off Cape Adare, Eltanin Sta. 1870, 659-714 m (1, USNM 60542); Relay Bay, Deep Freeze I, Edisto Sta. 4, 400 m (1, USNM 58992).--Weddell Sea, Glacier Sta. 68-1, 650 m (3, USNM 46691); Sta. 69-4, 743 m (2, USNM 46693); Sta. 69-23, 3697 m (3, USNM 46697).--South Georgia, Islas Orcadas Sta. 21 (cruise 575), May 16, 1975, 53 057.5'S; 37 020.7'W, 27-40 m, Blake trawl (1, USNM 69391).--South Sandwich Islands, Islas Orcadas Sta. 76 (cruise 575), June 3, 1975, 56015.6'S; 27 035.0·W, 110-155 m, Blake trawl (3, USNM 69392).

Description. A moderate-sized species, up to 13 mm long and 2 mm wide for 60 set igers. Col or in alcohol: dark brown.

Prostomium truncate to weakly rounded on anterior margin, with frontal horns develop-

ed to varying degrees on some specimens (Figure 8A); caruncle bordered by nuchal ciliary tracts, continuing posteriorly for 5-12 setigers; occipital tentacle present; eyes present or absent; if present, a single pair, with each elongated, reddish in color.

Setiger 1 well developed, with flattened postsetal lamellae. Notopodial lamellae of setiger 2 and subsequent setigers larger, triangular (Figure 8A); posterior se tigers with smaller lamellae, rounded.

Capillary notosetae arranged in 2 rows with about 35-40 setae per fascicle in anterior se tigers and about 25-30 in posterior setigers; capillaries with narrow, clear sheaths, shafts with fine granulations along internal striae. Anterior capillary neuro-setae similar to notosetae in arrangement, number and form; neuropodial tridentate hooded hooks from setiger 35-40, numbering 9-12 per fascicle; hooks with main fang surmounted by paired apical teeth (Figure 8B); a single inferior sabre seta from setiger 23-24, strongly curved with fine granulations along internal striae (Figure 8C-8D).

Branchiae from setiger 2, continuing posteriorly to about setiger 30; branchiae short, triangular, basally free from dorsal


lamellae. Genital pouches beginning between setigers 4 and 5. Nature of pygidium not known.

Remarks. Laonice antarcticae was originally described as a subspecies of L. cirrata (Sars) and was distinguished f;om-rhe stem species by having genital pouches from setigers 4 and 5 instead of 28 to 35. The subspecies was later considered to be a full species [Hartman, 1965; Hartman and Fauchald, 1971J. Foster [197IaJ, however, defined L. cirrata in a broad sinse and did not consIder the first occurrence of genital pouches to be taxonomically important and therefore lumped all Laonice species and subspecies into L. cirrata. Blake and Kudenov [1978J, Tn a study of Australian spionids, were able to distinguish 3 distinct froms of Laonice and described them all as new species after carefullY comparing them with one another and with Foster's defin\tion of L. cirrata.

In the present study, L. antarcticae is considered to be a valid species. The species appears to most closely related to L. bassensis Blake and Kudenov from AustralIa in having a double nuchal ridge. It differs from L. bassensis in having an entire rather than an incised prostomium, in having rather than lacking eyes, in having tridentate rather than bidentate hooded hooks and in having the genital pouches first present from between setigers 4 and 5 instead of 2 and 3. Furthermore, L. antarcticae differs from L. bassensis and-all other species of the genus in the development of horns on the frontal margin of the prostomium.

All of the specimens from the Weddell Sea which Hartman [1978J referred to L. antarcticae have been reexamined. Most h~ve been referred to L. weddellia (see below). Many of the specimens are poorly preserved or damaged, and this probably accounts for their misidentification by Hartman. A brief inspection of some deep-water specimens of ~. antarcticae identified by Hartman and Fauchald [1971J from the North Atlantic (Gay Head-Bermuda Transect: Chain Sta. 87, 39 048.7'N; 70040.8'W, 1102 m, deposited AHF) confirms that they are morphologically indistinguishable from the southern hemispheric specimens recorded in the present study.

Distribution. North Atlantic Ocean, slope and abyssal depths to 2864 m off eastern North America; South America: off Surinam and Brazil in depths of 520-1500 m; off Argentina and Uruguay, 70-96 m; Antarctic and subantarctic seas, 27-3697 m.

Laonice weddellia Hartman, 1978 Fig. 9

?Laonice cirrata: Hartman, 1967, p. 112 [in partJ. [not Sars, 1851J.

nerinid Hartman, Prionospio spp.

partJ. Laonice weddellia

fig. 16.

1967, p. 112. Hartman, 1967, p. 113 [in

Hartman, 1978, pp. 161-163,

Material examined. Off southwestern Chile, Eltanin Sta. 72, 805-970 m, (1, USNM). --Straits of Magellan, Eltanin Sta. 962, 256-320 m (1, USNM 60519).--Falkland Islands, Eltanin Sta. 344, 119 m (4, USNM 60514); Sta. 363 (2, USNM 60515).--Drake Passage. Eltanin Sta. 1089, 641 m (1, USNM 56387.--South Orkney Islands, Eltanin Sta. 500, 489-490 m (2, USID1 56395); Sta. 1079, 593-598 m (5, USNM 56385); Sta. 1082, 298-302 m (2, USNM 56386).--South Shetland Islands, Eltanin Sta. 418, 311-426 m (1, USNM 56391); Sta. 419, 509-549 m (3, USNM 60518); Deep Freeze Ill, West-wind Sta. 9, 170 m (2, USNM 58998); Hero Sta. 726, 64-82 m (3, USNM 60531); Sta. 1060, 44 m (3, USNM 60529); Sta. 1063, 44 m (2, USNM 60530).--Bransfield Strait, Eltanin Sta. 410, 220-246 m (26, USNM 61952); Sta. 412, 1180 m (18, USNM 56394, 60517, 61953); Sta. 413, 1113-1153 m (3, USNM 61954); Sta. 416, 494-507 (2, USNM 61955); Sta. 437, 267-311 m (9, USNM 56394).--Antarctic Peninsula, Eltanin Sta. 272, 412 m (10, USNM 61951); Hero Sta. 767, off Anvers Island, diving (I:1USNM 60532); Sta. 843, off Anvers Island, 107 m (1, USNM 60533); Sta. 970, Bismark Straits, 102 m (1, USNM 60528); Sta. 1066, Anvers Island (1, USNM 60534); Sta. 1075, 91-110 m (2, USNM 60535); Sta. 1937, Bismark Straits, 96-133 m (1, USNM 60536); Anvers Island, Hero Inlet, Palmer Station, March 17, 1968; Glacier Sta. 68-Palmer 11, 40 m (9, USNM 46704).--Ross Sea, Eltanin Sta. 1885, 311-328 m (2, USNM 60520); Sta. 1930, 831-836 m (2, USNM 60521); Sta. 2005, 864-870 m (4, USNM 60522); Sta. 2012, 589-608 m (3, 60523); Sta. 2029, 335-338 m (I, USNM 60524); Sta. 2035, 876 m (I, USNM 60525); Sta. 2050, 909-923 m (1, USNM 60526); Sta. 2073, 503 m (1, USNM 60527); Moubray Bay, off Cape Hallett, Deep Freeze IV, Northwind Sta. 8, 135 m (6, USNM 58998); Robertson Bay, west of Cape Adare, Deep Freeze IV, Staten Island Sta. 10, 128 m (1, USNM 58997)~Edward VII Peninsula, Cape Colbeck, Staten Island Sta. dredge 1, consecutive 14, December 24, 1960, 77006'S; 158°17'W, 100 m (1, USNM 58997).--Ross Sea, McMurdo Sound, Deep Freeze I, Edisto Sta. 8, 321 m (2, USNM 58994); Deep Freeze 11, Glacier Sta. 1, 400 m (2, USNM 58995).-Weddell Sea, Deep Freeze 11, Staten Island Sta. 19, 430 m (2, USNM 58996); Glacier Sta. 68-1, 650 m (4, USNM 61956); Sta. 68-5, 400 m (2, USNM 46702); Sta. 68-9, 526 m (1, USNM 46703); Sta. 68-18, 1664 m (3, USNM 46692); Sta. 69-1, 513 m (3, USNM 46705), Sta. 69-2, 412 m (5, USNM 46706); Sta. 69-6, 513 m (3,


USNM 46707); Sta. 69-7, 512 m (15, USNM 46708); Sta. 69-8, 585 m (holotype and 10 paratypes, USNM 46700-1), Sta. 69-10, 659 m (3, USNM 46709); Sta. 69-22, 3111 m (1, USNM 46696).--W of South Georgia, Eltanin Sta. 1536, 659--686 m (5, USNM 69393).

Description. A large species, up to 34 mm long and 2.5 mm wide for approximately 130 setigerous segments. Color in alcohol: light tan to dark brown.

Prostomium longer than wide, rounded on anterior margin, with small apical incision or notch evident on some specimens (Figure 9A); narrowing posteriorly and continuing to end of setiger 1, then merging indistinctly with nuchal ridge and continuing posteriorly for 12-20 setigers; nuchal ridge bordered laterally by ciliated grooves; prostomium with a single fingerlike occipital tentacle at posterior end and with 2-3 pairs of red eyes: anterior pair indistinct, deeply imbedded within cuticle; middle pair largest, composed of up to 6 individual pigment cups, these sometimes dispersed or scattered obscuring overall structure; posterior pair small, oval, consisting of single pigment cup. Peristomium well developed, with moderately high lateral wings.

Setiger 1 well developed, with conical notopodial lamellae and subtriangular neuropodial lamellae. Parapodial lamellae of sub-sequent setigers larger, triangular, somewhat auriculate (Figure 9B); posterior notopodial lamellae subtriangular, earlike; posterior neuropodial lamellae more or less elliptical (Figure 9C).

Capillary notosetae and neurosetae numerous, with about 100 setae arranged in 4-5 rows per fascicle; shafts of setae of first row bearing fine granulations (Figure 9D) and clear sheaths, capillaries of rows 2-5 lacking granulations; notosetae fewer by about setiger 15, with fewer rows and loss of granulations; posterior neurosetae longer, giving spinous appearance to posterior end; neuropodia with 1-3 heavily granulated inferior sabre setae by about setiger 12 and bidentate and tridentate hooded hooks from setiger 20-21; hooks with main fang surmounted by 1-2 apical teeth; when present, second apical tooth located in tandem with first (Figures 9E-9G) •

Branchiae from setiger 2, thin and short at first, then increasing in size and thickness on subsequent setigers (Figure 9A); each with ciliated inner border, branchiae continuing posteriorly for 20-50 setigers, absent from posterior region. Genital pouches present from between setigers 8-9 or 10-11. Pygidium with 7-8 anal cirri, with ventralmost being thickest.

Remarks. Laonice weddellia is endemic to Antarctic and subantarctic seas and differs

from other species of the genus in having an elongated, narrow prostomium with 3 pairs of eyes. The prostomium of some specimens (Eltanin Sta. 412) superficially resembles Scolelepis species, although it is not acutely pointed on the anterior margin.

Laonice weddellia differs from L. antarcticae, with which it frequently occur~ having genital pouches from setigers 8-11 instead of 4-5, 5 rows of capillary setae in anterior parapodia instead of 2 and by having the apical teeth of bidentate hooded hooks in a tandem arrangement instead of paired. Furthermore, the anterior margin of the prostomium is narrow and entire, sometimes with a weakly developed notch, instead of broad, truncate, and with short lateral horns.

Distribution. Off southern South America, Straits of Magellan; Falkland Islands; South Orkney Islands; South Shetland Islands; widespread in Antarctic seas; 44-3111 m.

Laonice cirrata (Sars, 1851)

Laonice cirrata: Soderstrom, 1920, p. 220.-Fauvel, 1927, p. 38, figs. 12a-12e; 1936a, p. 29.--Monro, 1939, p. 125.--HartmannSchroder,1965, pp. 204-206, figs. 191-192. --Hartman, 1966, pp. 15-16, pI. 3, figs. 13-15; 1969, pp. 107-108, figs. 1-4.-Foster, 1971a, pp. 69-72, figs. 155-160 [in part].--Carrasco, 1974, pp. 187-188, figs. 5-9.

?Laonice cirrata: Hartman, 1967, p. 112 [in part].

Laonice cirrata postcirrata HartmannSchroder, 1965, p. 206, figs. 193-194. Fide Foster, 1971.

Laonice cirrata praecirrata HartmannSchroder, 1965, p. 207, figs. 195-196. Fide Foster, 1971.

Material examined. field Strait, Eltanin (1, USNM 56392); Sta. 56393).

Antarctica, BransSta. 428, 662-1120 m 432, 884-935 m (1, USNM

Remarks. According to Fauvel [1927] and Hartman [1969] Laonice cirrata has a broadly rounded prostomium with 2 pairs of eyes, the posterior of which is the larger. The hooded hooks are bidentate and begin on setiger 40-50. Genital pouches occur from about setiger 25. Foster [1971a] lumped all known species with L. cirrata and included forms having bidentate and tridentate hooded hooks from setiger 29-60 and genital pouches from setiger 4-50. Because at least 1 of the species she lumped with L. cirrata has been found to be valid (L. antarcticae, see above), it would seem best to follow the earlier diagnoses of Fauvel [1927] and Hartman [1969] until a larger comparative study can be undertaken. Carrasco [1974] observed

O.5mm E F

20J.lm

L D

O.5mm

L Fig. 9. Laonice weddellia (paratype, USNM 46701). A, anterior end in dorsal view; B, left setiger 5 in anterior view; C, left setiger 100 in anterior view; D, detail of anterior capillary notoseta showing granulations of shaft; E-G, hooded hooks in lateral (E, G) and frontal (F) views.


that shallow water L. cirrata from Chile tended to agree with the earlier descriptions, for he noted bidentate hooded hooks from se tiger 35-36 and genital pouches from se tiger 21-36. Hartmann-Schroder [1965] described typical ~. cirrata and 2 separate subspecies, L. cirrata postcirrata and L. cirrata prae~irrata from Chile. The 3 forms were separated on the basis of caruncle length and placement of the occipital tentacle. Foster [1971a) was unable to separate these forms following an examination of Hartmann--Schroder's specimens.

The majority of specimens which Hartman (1967) referred to ?Laonice cirrata from South America and Antarctic seas have been examined and are referred to L. antarcticae and L. weddellia. Two anteri;r fragments agree-well in overall appearance to L. cirrata. Neither genital pouches nor hooded hooks were observed since the largest fragment consisted of 20 setigers and it can be assumed that if present, they began in the middle and posterior body segments. Unlike L. cirrata, however, the prostomium is in~ised on the anterior margin.

Because of the widespread occurrence of L. antarcticae and L. weddellia in Antarctic seas, the records of L. cirrata by Fauvel [1936] and Monro [1939] should be reexamined.

Distribution. Widespread in the northern hemisphere; Chile, 7-240 m; Antarctica: Bransfield Strait, 662-1120 m; ?Alexander Island [Fauvel, 1936a], ?MacRobertson Land, 219 m, Ingrid Christensen Coast, 1266 m, Adelie Coast, 640 m [Monro, 1939].

Laonice sp. A

?Laonice cirrata; Hartman, 1967, p. 112 [in part]. Not Sars, 1851.

Material examined. Off southwestern Chile, Eltanin Sta. 208, 957 m (1, USNM 56381).

Remarks. This specimen is unusual in having genital pouches from setiger 1-2. The prostomium is broadly rounded on the anterior margin and lacks eyes. The caruncle extends posteriorly for approximately 50 segments. The specimen is an anterior fragment which resembles L. hermaphroditica Blake and Kudenov from Qu;ensland, Australia.

Distribution. Off southwestern Chile, 957 m.

Genus Paraprionospio Caullery, 1914 Paraprionospio pinnata (Ehlers, 1901)

Prionospio pinnata Ehlers, 1901a, pp. 163-164.--0kuda, 1937, p. 247, fig. 22. --Fauvel, 1953c, pp. 323-324, fig. 174e. --Hartmann-Schroder, 1965, p. 211.--Day,

1967, p. 488, fig. 18.8.i-18.8.1.--Hartman, 1967, p. 113.

Paraprionospio pinnata: Foster, 1971a, pp. 102-104, figs. 237-246 [synonymy).--Carrasco, 1974, pp. 190-193, figs. 15-19. --Blake and Kudenov, 1978, pp. 209-210.

Prionospio spp. Hartman, 1967, p. 113 [in part].

Material examined. Uruguay, IBM Sta. N-242, 63 m (13, USNM 67549).--Argentina, IBM Sta. N-250, 83 m (3, USNM 67552); Sta. N--263, 39 m (2, USNM 67551); Sta. N-I054, 58--65 m (4, USNM 67552); Sta. 1059, 72-80 m (1, USNM 67553); Sta. Mej-5, 55 m (6, USNM 67542); Sta. Mej-ll, 47 m (6, USNM 67543); Sta. Mej-18, 48 m (1, USNM 67544); Sta. Mej-22, 40 m (3, USNM 67545); Sta. G n--3, 16.5 m (1, USNM 67546); Sta. G n--8, 20 m (1, USNM 67547); Sta. G 11-14 (2, USNM 67548); Sta. H-15, 70-80 m (25+, USNM 67554).--Near South Georgia, Shag Rocks, Eltanin Sta. 463, 3403-3484 m (17, USNM 56384, 60593).--Off western Chile, Eltanin Sta. 752, 209 m (10, USNM 56388); Sta. 758, 156 m (21, USNM 60592); Sta. 194 (cruise 21), 137-141 m (394, USNM 69394). --South Pacific Ocean, west of Juan Fernandez Islands, Eltanin Sta. 204 (cruise 21), 183-283 m (5, USNM 69395); Sta. 205, 128-183 m (1, USNM 69396).

Description. Prostomium spindle-shaped, without caruncle; eyes lacking. Palps with unique basal sheath. Peristomium fused to achaetous first segment, forming large, elevated peristomial wings partially enclosing prostomium. Setiger 1 distinctly separated from achaetous segment. Notopodial lamellae large, foliose on setigers 1-5, smaller on subsequent setigers. Three pairs of pinnate branchiae occurring on setigers 1-3; each branchia with large, platelike pinnules; individual branchiae sometimes resembling plumes. Distinct dorsal ridge connecting branchial bases of setiger 1. Anterior setae all capillaries; multidentate hooded hooks from setiger 9 in neuropodia and setiger 20 in notopodia; with 1-3 neuropodial inferior sabre setae from setiger 9. Pygidium with 1 long medial cirrus and 2 short lateral cirri.

Remarks. Paraprionospio pinnata is a widespread and apparently cosmopolitan species occuring in shallow subtidal sediments along continental coasts to moderate depths. The record from Eltanin Sta. 463 from the southwestern Atlantic represents an unusual occurrence from abyssal depths. The specimens all agree with published accounts of the morphology of the species.

Distribution. Cosmopolitan; off Uruguay and Argentina on hard bottoms of mud and sand to 83 m; off western Chile and beyond the Juan Fernandez Islands, 128-283 m; Shag rocks, southwestern Atlantic, 3403-3484 m.


Genus Prionospio Malmgren, 1867

Type species: Prionospio steenstrupi Malmgren, 1867, by monotypy.

Remarks. Blake and Kudenov [1978] revised the genera of the Prionospio complex to include Paraprionospio Caullery, Orthoprionospio Blake and Kudenov, Streblospio Webster, and Prionospio Malmgren sensu lato. To the latter genus were referred Aquilaspio Foster and Minuspio Foster as subgenera. Apoprionospio Foster was considered to be a synonym of Prionospio (Prionospio) because it was based only on a particular ordered arrangement pinnate and nonpinnate branchiae. Recently, 2 new genera, Aurospio Maciolek [1981aJ and Laubierellus Maciolek [1981b] have been added as full genera to the Prionospio complex. In the present study, the concept of Prionospio and its sub-genera follows Blake and Kudenov [1978]: Prionospio (Prionospio) includes species having smooth (nonpinnate) branchiae and pinnate branchiae in various combinations; Prionospio (Aquilaspio) includes species having all pinnate branchiae; and Prionospio (Minuspio) includes species having all nonpinnate branchiae.

Prionospio (Prionospio) orensanzi, n. sp. Figs. 10, 11

Material examined. Argentina, IBM Sta. Mej-12, 24 m (paratype, USNM 67517); Sta. SAO I-C, 15 m in sand (paratype, USNM 67516); Sta. SAO V-31, 21 m in sand (holotype, USNM 67515) .

Description. Holotype complete, measuring 15 mm long and 0.75 mm wide for 65 setigers. Color in alcohol: light tan.

Prostomium broadly rounded on anterior margin, with weak medial incision (Figure lOA); caruncle extending posteriorly as low ridge to middle of setiger 2; ciliated nuchal grooves occurring lateral to caruncle; 2 pairs of eyes: posterior pair larger, irregular in shape, composed of numerous ocelli, anterior pair smaller. Peristomium reduced, lacking lateral wings.

Setiger 1 well developed, separated from peristomium, with broad notopodial lamellae and smaller, leaflike neuropodial lamellae (Figure lOA); notopodial lamellae of setigers 2-5 large, triangular (Figure llA); lamellae of setigers 6-19 continued across dorsum forming dorsal crests (Figure lIB); neuropodial lamellae of anterior middle setigers rounded, slightly wider than long, becoming leaflike in posterior setigers.

Capillary notosetae of anterior and middle setigers unilimbate, arranged in 2 rows, those of first row shorter, with wider

sheaths (Figure lOB) than those of second row (Figure 10C); both with granulations on distal half of shaft. Capillary neurosetae bilimbate, arranged in 2 rows, also with granulations on shaft; neuropodia with 1-2 inferior sabre setae from setiger 10 having granulations on shaft and clear, narrow sheath (Figure 100); multidentate hooded hooks in neuropodia from setiger 20 and in notopodia from setiger 36; hooks with 10-12 apical teeth arranged in 2 ro\.-s above main fang (Figures 10E-IOG); internal hood inconspicuous.

Branchiae present on setigers 2-5 (Figure lOA); first and fourth pairs pinnate, second and third pairs cirriform. Pygidium typical, with 2 short lateral cirri and 1 long medial cirrus.

Remarks. Prionospio orensanzi belongs to the K. steenstrupi group in having the first and fourth pairs of branchiae pinnate and the second and third pairs cirriform and lacking pinnules. By lacking peristomial wings, P. orensanzi most closely resembles 3 Australian species: P. australiensis Blake and Kudenov, 1978; K~ queenslandica Blake and Kudenov, 1978; and P. lanceolata HartmannSchroder, 1979. P.-orensanzi differs from these 3 species by having prominent dorsal crests from setiger 6 through setiger 19. In P. australiensis a dorsal fold occurs on setiger 5 and low, inconspicuous dorsal crests are present from setigers 9-13; P. lanceolata lacks dorsal crests. P. orensanzi further differs in having a distinct separation between the peristomium and first setigerous segment. These 2 segments are normally fused in other species of the genus, and this character was used by Blake and Kudenov [1978] as a generic character for Prionospio.

Etymology. The species is named in honor of Jose M. Orensanz in recognition of his studies on the Polychaeta of Argentina.

Distribution. Argentina, subtidal, 15-24 m in sand.

Prionospio (Prionospio) cf. ehlersi Fauvel, 1928

Prionospio ehlersi Fauvel, 1928b, pp.lO-12, fig. 1; 1936b, pp. 61-62, fig. 4.--Day, 1967, p. 490, fig. 18.9.d-18.9.f.

Prionospio (Prionospio) ehlersi: Blake and Kudenov, 1978, p. 217, fig. 20m [synonymy].

Spiophanes spp. Hartman, 1967, p. 114 [in part].

Material examined. Chile, off Valparaiso, Eltanin Sta. 750, 624 m (1, USN}j 60596) .---Drake Passage, Eltanin Sta. 265, 3691-3695 m (1, USNM 61958); Sta. 303, 4077-4176 m (1, USNM 60625).--Off King George V


BeD T

300)1 m

1

',.' . .', ),::' ," .... :~., ','

" .,',

T 2 0)1 m

1 T

JO)Jrn

'F 1

G

~' );

.' ,;r-"

"~;'.p ""H

.:,:', ",<

Fig. 10. Prionospio orensanzi n. sp. (holotype, USNM 67515). A, anterior end in dorsal view; B, capillary notoseta from the anterior row of setiger 7; C, capillary no toseta from the posterior row of setiger 7; D, inferior sabre seta; E, hooded hook in lateral view; F, G, enlargements of hooded hook to show teeth arrangement, hood omitted of F; H, posterior end in dorsal view.

Coast, Eltanin Sta. 7 Gr 7, 3543 m (2, USNM 69399).--Off Peter I Island, Eltanin Sta. 939, 3848-3980 m (1, USNM 56373). "South Pacific Ocean, Albatross Cordillera, Eltanin Sta. 1248, 3386-3495 m (1, USNM 60626).

Remarks. Various branchial pairs are missing from all of the specimens. They

generaly agee with the overall character of the species in having the first pair of branchiae long and pinnate, the second and third pair short and cirriform and the fourth pair long, thin and cirriform. In contrast, lateral genital pouches, usually considered to be characteristic of P. ehlersi have not been


T 300)1 m

1

/

B

\ \

Fig. 11. Prionospio orensanzi n. sp. (holotype, USNM 67515). A, left setiger 5 in anterior view; B, right setiger 7 in anterior view.

observed. Similar appearing specimens have been found with pouch bearing examples in North Atlantic localities (N. J. Maciolek, personal communication, July 1981).

Distribution. Eastern North America; Africa; Solomon Islnds; Korea; Australia; off Chile; South Pacific Ocean; Antarctic seas. A predominantly deep-sea species, Eltanin specimens occurring from 624-4176 m.

Prionospio (Prionospio) pygmaea Hartman, 1961

Prionospio pygmaeus Hartman, 1961, pp. 93-94, 1969, pp. 163-164, figs. 1-3.

Apoprionospio pygmaea: Foster, 1971a, pp. 94-97, figs. 213-225 [synonymy].

Prionospio (Apoprionospio) pygmaea: Light,

1977:82; 1978:84-87, figs 84-86 [synonymy].

Material examined. Argentina, IBM Sta. G 11-3, 16.5 m, muddy sand (9, USNM 67555); Sta. SAO I-C, 15 m, sand (1, USNM 67556). --Juan Fernandez Islands, Anton Bruun Sta. 65240, 24-27 m (1, USNM 60597).

Remarks. These specimens agree well with ~ublished accounts. The prostomium is truncate on the anterior margin. The first 3 branchial pairs are cirriform, and the fourth is pinnate. Eyes are absent. Hooded hooks are multidentate. Thin dorsal membranous crests described by Light [1978] were not observed.

Distribution. California; Gulf of Mexico; Argentina; Juan Fernandez Islands. In fine sands, shallow depths of 15-30 m.


Prionospio (Prionospio) steenstrupi Malmgren, 1867

Prionospio steenstrupi Malmgren, 1867, pp. 202-203, pI. 9, fig. 55.--Foster, 1971a, pp. 84-87, figs. 175-185 [synonymy].

Prionospio (Prionospio) steenstrupi; Light, 1978, pp. 88-92, figs. 89-90 [synonymy]. --Blake and Kudenov, 1978, p. 213, fig 20a [synonymy]. Material examined. Juan Fernandez Is

lands, Anton Bruun Sta. Drab 130, 46 m (6, USNM 60627).

Remarks. These specimens generally agree with the descriptions of various authors. Branchiae are pinnate on se tigers 2 and 5 and cirriform on se tigers 3 and 4. Lateral peristomial wings are well developed, and low dorsal crests are present from setiger 6.

Distribution. Cosmopolitan; Chile, Juan Fernandez Islands, 46 m.

Prionospio (Prionospio) sp. A

Material examined. Argentina, off the Rio de la Plata, IBM Sta. N-250, 83 m (5, USNM).

Remarks. These specimens have broad cirriform branchiae on setigers 2-4 but lack branchiae on setiger 5 precluding specific identification. They differ from P. pygmaea by having the prostomium rounded on the ante rior margin rather than truncate.

Distribution. Argentina, subtidal in 83 m.

Prionospio (Prionospio) sp. B

Laonice antarcticae: Hartman, 1978 [in part]. Not Hartman, 1953.

Material examined. Weddell Sea, Glacier Sta. 68-1, 650 m (12, USNM 61959).

Remarks. These specimens are all small. Minute cirriform branchiae, present on setigers 3-4, are obscured by the large triangular shaped dorsal lamellae. There is no evidence of branchiae on setigers 2 or 5. The specimens resemble others seen from deepsea samples in the North Atlantic which are considered to be damaged (N. J. Maciolek, personal communication, 1980).

Distribution. Weddell Sea, 650 m.

Prionospio (Aquilaspio) peruana Hartmann-Schroder, 1962

Prionospio peruana Hartmann-Schroder, 1962a, pp. 138-139, figs. 101-104; 1965, pp. 212-213 •

Aquilaspio peruana: Foster, 1971a, p. 106.

Remarks. No specimens of this species were encountered. The species is known from

both Peru and Chile [Hartmann-Schroder, 1962a, 1965] and has recently been identified from among the collections of the Lund University Chile Expedition, 1948-1949 (K. Fauchald, personal communication, 1978).

The species is characterized by having a truncated prostomium, 4 pairs of pinnate branchiae from setiger 2, well-developed dorsal crests from setiger 6--7, neuropodial multidentate hooded hooks from setiger 16-17, and notopodial hooks from setiger 27.

Distribution. Peru; Chile, in fine sands, 15-71 m.

Prionospio (Minuspio) sp.--cirrifera complex

Material examined. Galapagos Islands, Anton Bruun Sta. 66122, 17-18 m (1, USNM 60595).--Ecuador, Anton Bruun Sta. 6670, 8-9 m (2, USNM 60594).

Remarks. Prionospio (minuspio) cirrifera Wiren, 1883 is an especially difficult species to define owing to a wide range of morphological variability attributed to it. Recent investigations have shown that various reports of ~. cirrifera from North America and other areas of the North Atlantic refer to at least 7 species (N. J. Maciolek, personal communication, August 1981). The southern hemispheric forms are yet to be sorted out. The present specimens may be referred to P. cirrifera in a broad sense by having all clrriform branchiae from setiger 2 and well-developed notopodial lamellae on se tiger 1. It will be necessary, however, to compare details of prostomial shape, hooded hooks, and parapodial form with the northern hemispheric species before they can be specifically identified.

Distribution. Ecuador, Galapagos Islands, 8-18 m.

Prionospio (Minuspio) patagonica Augener, 1923 Revised

Fig. 12

Prionospio patagonica Augener, 1923a, pp. 3-· 5.

Prionospio cirrifera chilensis HartmannSchr6der, 1962b, pp. 138-140, figs. 169-173; 1965, pp. 213, 302. New Synonymy.

Minuspio chilensis: Foster, 1971a, pp. 107, 111.

Minuspio cirrifera: Orensanz, 1974, p. 43. Not Wiren, 1883.

Material examined. Chile, western Patagonia, near Straits of Magellan, Estero Kelly (5 syntypes of P. patagonica, ZMH-9438); Estero lenga, i;tertidal in fine sand with debris, October 31, 1959, collected by G. Hartmann (holotype of P. cirrifera chilensis, ZMH P-14984).


500pm

00 00 00

QC

30pm Fig. 12. Prionospio (Minuspio) patagonica (syntype, ZMH 9438). A, anterior end in dorsal view; B, hooded hook in lateral view; C, diagram of tooth arrangement of hook in frontal view.

Description. A large species, up to 19 mm long and 1.5 mm wide for 100+ segments. Color in alcohol: light tan with no pigment spots.

Prostomium entire, blunt on anterior margin, with short caruncle, extending to anterior margin of setiger 2, without eyes (Figure 12A). Peristomium inflated, with low lateral wings.

Setiger 1 reduced, lacking postsetal lamellae (Figure 12A); anterior notopodial lamellae triangular; neuropodial lamellae rounded in anterior and middle segments, with notopodia reduced to short lobes and neuropodial lamellae lacking.

Capillary notosetae and neurosetae of anterior setigers with narrow sheath bearing fine punctations; hooded hooks in neuropodia from setiger 22 and in notopodia from setiger 46; neuropodia with up to 4 hooks, 1-2 capillaries and single inferior sabre seta; notopodia with 1-2 hooks and 5-6 capillaries; hooded hooks with 6 apical teeth arranged in 2 rows, with distinct internal hood (Figures 12B-12C).

Branchiae from setiger 2, numbering 10-12 pairs, all cirriform (Figure 12A). Pygidium with 2 thin lateral lappets and single long medial cirrus.

Remarks. Foster [1971a] referred P. patagonica to the cosmopolitan P. cirrifera-,-based on the original description. An examination of the syntypes of P. patagonica, deposited in the collections-of the Zoological Museum of Hamburg, clearly reveal it to be a distinct species. P. patagonica is much larger than reports-of P. cirrifera and differs significantly in the reduction of setiger 1. P. cirrifera has prominent notopodial lamellae and long capillary notosetae on setiger 1, while the lamellae are lacking and the notosetae are short on P. patagonica. No specimens of P. cirrifera have been found which exceeded 15 mm in length or have more than 80 setigers (N. J. Maciolek, personal communicaion, July 1981). Specimens of P. patagonica are up to 19 mm long and hav; over 100 setigers. The heavily granulated anterior capillary setae found in P. cirrifera do not occur in P. patagonica.


Prionospio cirrifera chilensis Hartmann-Schroder was raised to full species status by Foster [1971a]. An examination of the holotype, however, reveals the species to be the same as P. patagonica.

Distribution. Southern Chile and Patagonia, intertidal in fine sands.

Genus Spiophanes Grube, 1860

Type species: Spiophanes kroeyeri Grube, 1860, by monotypy.

Remarks. Indentification of the new collections of Spiophanes has proven to be particularly difficult, and it has been necessary to examine the types of some species previously described from South American waters. Results of these observations are presented separately for each species considered below. Four species are present: (1) S. bombyx (Claparede), (2) S. soederstraemi Hartman, (3) s. kroeyeri Grube, and (4) ~. tcherniai-Fauvel.

Spiophanes bombyx (Claparede, 1870)

Spio bombyx Claparede, 1870, pp. 485-487, pI. ----rr,~2. Spiophanes bombyx: Mesnil, 1896, pp. 249-

257, pI. 15.--Fauvel, 1916, p. 439; 1927, p. 41, figs. 14a-14i.--Soderstrom, 1920, pp. 243-244, fig. 153.--Augener, 1932b, p. 39.--Hartman, 1966, p. 22, pI. 5, figs. 14-16.--Day, 1967, p. 474, fig. 18.5.a-18.5.e. --Hartmann-Schroder, 1971, pp. 327-328, fig. 112.--Foster, 1971a, pp. 40-43, figs. 66-75 (synonymy].--Carrasco, 1974, pp. 197-198, figs. 37-41.--Bellan, 1975, p. 790. --Light, 1978, pp. 60-62, figs. 60-61. --Blake and Kudenov, 1978, p. 224.

Material examined. Argentina, IBM Sta. 1055, 92-96 m (1, USNM 67566); Sta. Mej-12, 24 m (1, USNM 67565).--Falkland Islands, Eltanin Sta. 344, 119 m (4, USNM 60350); St~ 363 (1, USNM 60351).

Remarks. The specimens agree well with previous descriptions of S. bombyx. The prostomium has long, tapering frontal horns and the neuropodial hooks have a partial hood.

Distribution. Cosmopolitan in sandy substrata, ranging from intertidal to shallow sub tidal depths; Argentina, in mud, sand, and hard bottoms, 47-92 m; Falkland Islands, 119 m, Kerguelen Islands, intertidal to 95 m.

Spiophanes soederstroemi Hartman, 1953 Revised Fig. 13

Spophanes kroyeri: S6derstrom, 1920, p. 240 [in part]. Not Grube, 1860.

Spiophanes soderstromi Hartman, 1953, p. 41, fig. 14 [in part, Sta. 1 only]. Not Hartman, 1966, p. 23, pI. 6, figs. 1-3.

Not Spiophanes s6derstromi [sic]: Day, 1961, pp. 484-485.

Not Spiophanes soederstromi (sic]: Day, 1967,p.474, fig. 18.1.a-18.1.e.

Spiophanes chilensis Hartmann-Schroder, 1965, pp. 215-218, figs. 208-211.

Spiophanes soederstroemi: Orensanz and Gianuca, 1974, pp. 17-18.

Material examined. Off Uruguay, December 12, 1901, 33000'S; 51010'W, 80 m, bottom of dark gray mud, collected by Swedish Antarctic Expedition, 1901-1903, Sta. 1 (2 syntypes of Spiophanes soderstromi, SMNH 3380); IBM Sta. N-242, 163 m (4, USNM 69362).--Off Argentina, IBM Sta. N-248, 170 m (1, USNM 69363); Sta. N-258, 50 m (10, USNM 69364); Sta. N-I054, 58-65 m (2, USNM 69366); Sta. N-I055, 92-96 m (6, USNM 69367); Sta. N-I059, 72-80 m (1, USNM 69368); Sta. N-I066, 72-86 m (1, USNM 69365); Sta. N-I075, 68 m (44, USNM 69369); Sta. Mej-5, 55 m (1, USNM 69353); Sta. Mej-ll, 47 m (5, USNM 69354-5); Sta. Mej-15, 28 m (3, USNM 69356); Sta. Mej-22, 40 m (1, USNM 69357); Sta. SAO I-IS, 28 m (3, USNM 69358); Sta. SAO I-53, 38 m (4, USNM 69359); Sta. G 11-3, 16.5 m (1, USNM 69360); Sta. G 11-8, 20 m (1, USNM 69361); Sta. A 11-60-282, 200-250 m (22, USNM 69370); Sta. A 11-60-284, 200-250 m (6, USNM 69371). --Argentina, intertidal: Albufera Mar Chiquita de embocadura, January 15, 1969, collected by J. M. Orensanz (1, USNM 69352). --Chile, Puerto Tortuga bei Coquimbo, 29057.4'S; 71 022.9'W, February 21, 1960, collected by G. Hartmann-Schroder (holotype of Spiophanes chilensis, ZMH P-14946).--Juan Fernandez Islands, Anton Bruun Sta. Drab 130, 46 m (1, USNM 60431); Sta. MV65-IV-47 (1, USNM 60432); Sta. MV65-IV-63 (1, USNM 60433).

Description. A moderate-sized species, up to 14 mm long and 0.7 mm wide for about 65 setigers. Color in alcohol: opaque white to brown.

Prostomium triangular to bell-shaped, broad on anterior margin, sometimes with slight medial incision, with short, laterally directed processes (Figure 13A, a); caruncle short, terminating in middle of setiger 1; without occipital tentacle; 2 pairs of eyes present or absent, when present arranged trapezoidally, with anterior pair more widely spaced and cup-shaped; nuch-al organs present or absent. Peristomium moderately developed as 2 lateral bulges, not forming wings.

Setigers 1-4 dorsally elevated, with long cirriform postsetal notopodial lamellae, those of subsequent setigers shorter, becoming stubby in appearance (Figure 13A) and reduced to fingerlike lobes by setiger 9.


E F G H

Fig. 13. Spiophanes soederstroemi (IBM Sta. N-I075, USNM 69369). A, anterior end in dorsal view; a, prostomium of another specimen; B, posterior end in dorsal view; C, posterior notopodium in dorsal view (not to scale); D, enlarged curved notoseta from posterior notopodium; E, inferior sabre seta; F, capillary neuroseta from anterior parapodium; G, H, neuropodial hook in lateral (G) and frontal (H) views.

Neuropodia of setigers 1-2 fingerlike, those of subsequent segments smaller, rounded. Ciliated mid-dorsal ridges from setiger 19-20, continuing to posterior end. Setigers 9-14 with glandular material, sometimes darkly pigmented.

Anterior capillary notosetae long, non-

granulated, with narrow sheaths, notopodia of posterior setigers with 1 or 2 unusually long, flat, distally curved capillaries among normal capillaries (Figures 13B-13C); each long capillary thickened, nongranulated, pointed with narrow sheath (Figure 13D). Anterior neuropodia with capillaries short,

231


broad, bilimbate, with finely granulated shafts (Figure 13F); fascicles arranged in 2 rows; setiger 1 with 1-2 large curved spines in addition to capillaries; single inferior sabre seta from setiger 4-6, each heavily granulated near tip (Figure 13E); neuropodial hooks from setiger 15, numbering 5-6 per neuropodium, accompanied by inferior sabre seta; hooks quadridentate with main fang surmounted by single unpaired tooth and pair of smaller teeth (Figure 13 H), appearing tridentae in lateral view (Figure 13G); hoods lacking. Bacillary setae not apparent, but glandular segments 9-14 with 1 or 2 small pointed setae visible upon dissection.

Pygidium with 2 anal cirri (Figure 13B). Remarks. Soderstrom [1920] included 2

specimens from off Uruguay in his redescription of specimens of S. kroeyeri. These specimens were subsequently examined by Hartman [1953] and determined to represent a distinct species, which she named S. soederstroemi. She also referred 4 specimens from South Georgia to the same species. All 6 of these specimens were obtained from the Swedish Museum of Natural History, Stockholm, and compared with the new Spiophanes collections from South America and Antarctica. The holotype of S. chilensis Hartmann-Schr6der, 1965 from the Zoological Museum, Hamburg, was also obtained. Following an examination of all of the available specimens, it is apparent that 2 species are included: (1) Spiophanes soederstroemi Hartman included the 2 specimens from off Uruguay from Hartman [1953, Sta. 1], numerous specimens from Argentina, the holotype of ~. chilensis Hartmann-Schroder and specimens from the Juan Fernandez Islands; (2) Spiophanes kroeyeri Grube included the 4 specimens from South Georgia from Hartman [1953, Sta. 34] and several specimens from the Falkland Islands, western Chile, and Ross Sea collected by the Eltanin.

Although not originally designated as types by Hartman [1953], the 2 specimens from Uruguay (Sta. 1), should be considered syntypes of S. soederstroemi Hartman and Uruguay the type locality. Hartman's figures 14a-14c agree with ~. soederstroemi, but the description is probably a mixture of the Uruguay and South Georgia (=S. kroeyeri) materials, despite her indication that it was based on the Uruguayan specimens. Subsequent confusion was introduced by Hartman [1966, p. 23] in her review of Antarctic polychaetes when she cited only South Georgia as the locality for S. soederstroemi. The figures [Hartman, 1966, pI. 6, figs. 1-3], however, were taken from the 1953 paper. Because of the brevity of the original description, S. soederstroemi was stated to be 'insufficiently characterized' by Pettibone [1962, p. 85] and 'indeterminable' by Foster [1971a, p. 40]. Both

incorrectly listed the type locality as 'Antarctic.' Records of S. soederstroemi by

Day [1961, pp. 484-485, 1967, p. 474, fig. 18.5.a-18.5.e] represent a different species, perhaps S. kroeyeri, due to the presence of an occipItal tentacle [Day, 1967, fig. 18. 5.1.]

The prostomium of S. soederstroemi has been observed to be variable in shape, triangular to bell-shaped, and degree of development of the lateral processes on its frontal margin. It is possible that this variation is a preservation artifact. The presence of enlarged notosetae in posterior segments has not been previously reported for the species. These spines are curved and have distinct limbations near the tip. Similar setae discovered in S. tcherniai and S. kroeyeri are not as thi~k and lack limbarions.

Spiophanes soederstroemi is closely related to S. missionensis Hartman, 1941 from California. The 2 differ in that the cirriform notopodial postsetal lamellae of setigers 1-4 of ~. missionensis are shifted to a more medial location than S. soederstroemi. Spiophanes missionensis lacks the enlarged and flattened notopodial setae in the posterior setigers, and its pygidium has a terminal cone with 2 lateral cirri, while S. soederstroemi has enlarged posterior notopodial setae and a ventromedial enlargement on the pygidium and 2 dorsolateral cirri.

Distribution. Argentina and Uruguay, intertidal to 250 m in muddy sands; Chile, intertidal; Juan Fernandez Islands, 46 m.

Spiophanes kroeyeri Grube, 1860

Spiophanes kroeyeri Grube, 1860, p. 88. -Pettibone, 1962, p. 85 [synonymy].

Spiophanes fimbriata Moore, 1923, p. 179. Fide Pettibone, 1962.

Spr;phanes soderstromi Hartman, 1953, p. 41 [in part, Sta. 34 only].

Spiophanes spp. Hartman, 1967, p. 114 [in part].

Spiophanes kroeyeri: Fauchald, 1972, p. 99, figs. 4C-4D. -Light, 1977, pp. 79-80, fig. 5d [synonymy].

Material examined. Off Chile, Eltanin Sta. 203, 436 m (1, USNM 56371); Sta. 208, 957 m (2, USNM 56372); Sta. 748, 1025 m (2, USNM 60435); Sta. 749, 1007 m (1, USNM 60436).--Falkland Islands, Eltanin Sta. 338, 587-595 m (1, USNM 60434).--South Georgia, outside of Cumberland Bay, 54011'S; 36 0 18'W, June 5, 1902, 252-310 m, on bottom of gray mud with stones, collected by Swedish Antarctic Expedition, 1901-1903, Sta. 34 (1, SMNH 3306; 2, SMNH 3304-k).--Ross Sea, off Moubray Bay, Victoria Land, Eltanin Sta. 1996, 143 m (1, USNM 60429).

Remarks. These specimens agree well with


published descriptions of S. kroeyeri. An occipital tentacle is present. Bacillary setae are feathery and conspicuous. Anterior neuropodial capillary setae are broad with narrow sheaths and bear heavy granulations on the shaft. Posterior notopodia bear enlarged, curved capillary setae but, unlike those of S. soederstroemi, lack limbations. Notopodial postsetal lamellae of se tigers 1-2 are long, cirriform with those of subsequent segments being shorter and broader. Genital pouches are present.

Distribution. Cosmopolitan in northern hemisphere; Australia; South Georgia; Falkland Islands; off Chile; Antarctica, Ross Sea. Subtidal, 252-1025 m in South American waters.

Spiophanes tcherniai Fauvel, 1951 Fig. 14

Spiophanes tcherniai Fauvel, 1951, pp. 762-764, figs. la-lg; 1953a, p. 10.--Hartman, 1966, p. 23, pI. 6, figs. 4-6; 1967, pp. 113-114; 1978, p. l63.--Be11an, 1975, p. 790.

Material examined. South Orkney Islands, Eltanin Sta. 500, 489-490 m (1, AHF; 1, USNM 56379); Sta. 1084, 298-403 m (10, USNM 56375).--South Shetland Islands, Eltanin Sta. 408, 223-225 m (4, USNM 60361); Sta. 418, 311-426 m (3, USNM 56377, 60352); Hero Sta. 819, Deception Island, 30 m (30, USNM 60372).--Bransfield Strait, Eltanin Sta. 437, 267- 311 m (3, USNM 56378); Sta. 1001, 238 m (4, USNM 60363); Sta. 1003, 210-230 m (5, USNM 56380, 60364).--Antarctic Peninsula, near Anvers Island, Hero Sta. 848, south of Arthur Harbor, 94-165 m (3, USNM 60373); Sta. 970, 102 m (2, USNM 60370); Sta. 5444, 34-40 m (2, USNM 60374); Sta. 1897, 22 m (I, USNM 60375); Sta. 983, Grandidier Channel, 38 m (1, USNM 60371); Marguerite Bay, Eastwind Sta. 4A, January 24, 1966, 67 0 53'S; 69 0 l0.5'W, 300 m, collected by Pawson and Squires (1, USNM 59886).--South Pacific Ocean, north of Ross Sea near Scott Island, Eltanin Sta. 1944, 516-595 m (1, USNM 60365).--Ross Sea, Eltanin [Sta. 2016, 581-586 m (1, USNM 60366); Sta. 2020, 256 m (3, USNM 60367).--Ross Sea, off Edward VII Peninsula: Staten Island Sta. 1-14, 77 0 06'S; l580 l7'W, December 24, 1960, 100 m (1, USNM 59889).--Ross Sea, off Victoria Land; Robertson Bay, Deep Freeze I, Edisto Sta. 3, 27 m (55, USNM 59893); Relay Bay, SW of Robertson Bay, Deep Freeze I, Edisto Sta. 4, 400 m (355, USNM 59891); Sta. 6, 100 m (16, USNM 59892); W of Cape Adare, Deep Freeze IV, Staten Island Sta. 10, 128 m (6, USNM 59894); Eltanin Sta. 2126, 143 m (3, USNM 60369); off Cape Hallett, Deep Freeze IV, Northwind Sta. 8, 135 m (22, USNM 59896).--Off Ross Ice Shelf, Eltanin Sta.

2065, 473-475 m (1, USNM 60368).--Budd and Knox Coasts, Antarctica, Deep Freeze Ill, Atka Sta. 24, off Vincennes Bay, near Wilkes Station, 45 m (1, USNM 59890).--Weddell Sea, Glacier Sta.: 69-1, 69-2, 69-4, 69-6, 69-7, 69 8, 69-10, 69-11, 412-851 m (290+, USNM 467l4-2l).--Weddell Sea, Coats Land, off Vahsel Bay, Deep Freeze Ill, Westwind Sta. 7, no. 91, off Filchener Shelf Ice, 225 m (1, USNM 59887); Deep Freeze IV, Edisto Sta. 20, Tr. 5, 384 m (2, USNM 59895).---

Description. Large specimen from Eltanin Sta. 500 16 mm long and 0.7 mm wide for 91 setigers; most specimens smaller, incomplete. Color in alcohol: light tan or opaque white.

Prostomium triangular narrowing posteriorly, truncate on anterior margin, with distinct frontal horns, caruncle short, extending to end of setiger 1; 2 pairs of eyes: anterior pair cup-shaped, larger, widely spaced; posterior pair oval, medial in location (Figure l4A). Peristomium well developed, forming thickened ridges lateral to prostomium.

Setiger 1 with a single, long curved neuropodial spine bearing granulations in addition to normal capillaries. Notosetal and neurosetal capillaries of anterior setigers long, with narrow nongranulated sheaths (Figure l4C). Setigers 15-18 with special broadly sheathed notosetae with fine venations in addition to normal capillaries (Figure l4D). Neuropodia with tridentate partially hooded hooks from setiger 16; each hook with main fang surmounted by 2 apical teeth (Figure l4B); a single inferior sabre setae in neuropodia from se tiger 16. Fourfive posterior setigers with 1-2 enlarged, curved granulated notosetae in addition to long capillaries (Figures l4E-14F).

Pygidium with 2 short cirri (Figure l4E). Remarks. Spiophanes tcherniai is most

closely related to S. bombyx in having a partial hood on the ne~ropodial hooks. It differs from other species of the genus in having special broadly sheathed notosetae on setigers 15-18 and enlarged, curved notopodial setae in far posterior segments.

Distribution. Endemic and widespread in Antarctic seas; Kerguelen Islands. Intertidal to 851 m.

Genus Spio Fabricius, 1785 Spio ~isetosa, n. sp.

Fig. 15

Material examined. Argentina, intertidal in sand, collected by J. M. Orensanz: Las Grutas, January 14, 1973 (holotype, USNM 67520); Golfo Nuevo, Puerto Madryn, February 15, 1975 (28 paratypes, USNM 67522); Golfo Nuevo, San Ramon, November 17, 1975 (150+ paratypes, AHF Poly 1295; USNM 58963, 67523); Los Chanares, February 1972 (1 paratype, USNM 67521).

200)Jrn

B T

10)1 m 1

Fig. 14. Spiophanes tcherniai (Eltanin Sta. 500, AHF). A, anterior end in dorsal view; C, capillary notoseta from anterior se tiger; DJ broad-sheathed notoseta from setiger 16; D, posterior end in dorsal view; F, curved notoseta from posterior setiger.


Description. A moderate-sized species, up to ID mm long and 1 mm wide for 45 setigerous segments. Color in alcohol: light tan, with body anteriorly pigmented, with prominent black ventrolateral intersegmental stripes; brown pigment on peristomium and dorsal surface (Figure 15A).

Prostomium entire on anterior margin; caruncle short, merging with setiger 1 (Figure 15A); without nuchal organs or occipital tentacle; 2 pairs of eyes: anterior pair cup-shaped, posterior pair oval.

Setiger 1 well developed, with prominent fascicles of capillary setae, notosetae especially long (Figure 15A). Sheathed capillaries of anterior setigers arranged in 2 rows, those of anterior row strongly granulated (Figure 15F), those of posterior row finely granulated or lacking granulations. Hooded hooks replacing posterior row of neuropodial capillaries from setiger 11, with anterior row remaining intact throughout body; third row of very fine, delicate, nongranulated capillaries also beginning from setiger 11 (Figure 15G) and 4-6 granulated inferior sabre setae from setiger 12 (Figure 15E); hooded hooks bidentate or tridentate with large apical tooth over main fang, sometimes surmounted by very small tooth (Figures 15C-15D); hooks numbering 10.-12 per row.

Parapodial lamellae low, rounded, inconspicuous throughout; branchiae from setiger 1, continuing to near end of body, fused basally to notopodial lamellae (Figure 15B). Pygidium unknown.

Remarks. Spio quadrisetosa resembles ~. pettiboneae Foster, 1971 from the Gulf of Mexico and S. pacifica Blake and Kudenov, 1978 from Australia in having tridentate hooded hooks and an entire prostomium. S. quadrisetosa, however, differs from these species in having unusually prolonged notosetae on setiger 1. The very unusual arrangement of the neurosetae, in which the capillaries and hooded hooks are arranged in 3 distinct rows, joined by a fourth group of inferior sabre setae further distinguishes S. quadrisetosa from other Spio species.

Etymology. Derived from the Latin, quadri from quattuor, four; ~, from seta, bristle. The name quadrisetosa is suggested from the arrangement of 4 distinct setal types in the neuropodia.

Distribution. Argentina, intertidal in sand.

Spio sp. A

Material examined. Ecuador, Salinas, north side of Punta Santa Elena, 20 13'S; 8D0 57'W, March 4, 1967, collected by D. W. Kirtley (1, USNM).--Juan Fernandez Islands, Anton Bruun Sta. 65260., 26 m (1, USNM 61253); Sta. 65256, 9-12 m (1, USNM 61252).

Description. Both specimens from Juan Fernandez Islands posteriorly incomplete, measuring 3.2 mm long and 0..75 mm wide for 18 setigers, and 4.0. mm long and 0..5 mm wide for 22 setigers. Color: richly pigmented on anterior setigers as typical for several other Spio species.

Pro~ium entire on anterior margin, with caruncle extending to posterior border to setiger 3, and surrounded by prominent lateral nuchal organs; occipital tentacle lacking; 2 pairs of eyes; posterior pair oval, more medially situated than cup-shaped anterior pair.

Notosetae of se tigers 1-6 long, giving spinous appearance to anterior end of body. Setal fascicles arranged in 2 rows; anterior row of capillaries bilimbate with granulated shafts and clear sheaths; capillaries of posterior row lacking granulations; neuropodial hooded hooks replacing posterior row capillaries from setiger 11; 6-7 per row, accompanied by 1-2 inferior sabre setae from setiger 17; hooks bidentate and tridentate with main fang surmounted by small apical tooth sometimes bearing a minute additional tooth; angle between main fang and first apical tooth reduced; hood long, extending well down shaft.

Branchiae from setiger 1, long, straplike, and overlapping at midline. Posterior segments and pygidum unknown.

Specimen from Ecuador posteriorly incomplete, poorly preserved, measuring 4.8 mm long and 1. 4 mm wide for 20. segments. Body pigment lacking. Hooded hooks numbering only 4-5 in neuropodia, instead of 6-7. In other respects, similar to Juan Fernandez Islands specimens and probably representing same species.

Remarks. Spio sp. A differs from S. quadrisetosa from Argentina in lacking a third row of capillaries in segments having hooded hooks. The species cannot be determined due to the fragmentary nature of the specimens.

Distribution. Ecuador; Juan Fernandez Islands. Intertidal to 26 m.

Genus Microspio Mesnil, 1896 Microspio hartmanae, n. sp.

Fig. 16

Material examined. Argentina, IBM Sta. N-ID64, 20.-26 m (paratype, USNM 58975); Sta. SAD I-I, 18 m (holotype and paratype, USNM 67513-4); Sta. SAD 1-7, 18-24 m (2 paratypes, USNM 58974).

Description. A small species, holotype largest, measuring 3.5 mm long and 0..3 mm wide for 34 setigerous segments. Color in alcohol: tan with pigment spots scattered over anterior dorsal surface.

Prostomium broadly rounded on anterior

l 500}Jrn

J

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'. ~

~'. :: , ....

~':: ' :',

c 'D F

..... ..., ~ ,

................ " ~

....... ~ " ... :: ...... :~" -. ..... .... ... ,

~.' ~ :::. ~ --~--:.-:: ~ -~ -:~~ ;'~ , - -- - .~

G Fig. 15. Spio quadrisetosa n. sp. (holotype, USNM 67520). A, anterior end in dorsal view; B, right se tiger 9 in anterior view; C, D, hooded hooks in lateral (C) and frontal (D) views; E, inferior sabre seta; F, capillary neuroseta from anterior row of setiger 11; G, same from third row.


margin (Figure 16A); caruncle short, terminating at posterior margin of setiger 1; curved lateral ciliated nuchal ridges extending posteriorly from caruncle, 2 pairs of eyes, anterior pair cup-shaped, widely spaced and posterior pair rounded, more closely spaced; no occipital tentacle. Peristomium well developed, lacking peristomial wings; palps missing.

Setiger 1 well developed, with conical postsetal lamellae and fascicles of capillary setae. Anterior 8-9 se tigers with capillary notosetae arranged in 2 rows; setae of anterior row shorter, thicker, with granulated shaft and clear sheath (Figure 16C); those of posterior row longer, thinner, lacking granulations (Figure 16D); capillaries from setiger 9-10 on lacking granulations, fewer in number and gradually increasing in length in far posterior setigers (Figure 16B). Neurosetae of setigers 1-10 arranged similarly to notosetae, with anterior row of shorter, granulated capillaries and posterior row of longer, nongranulated capillaries; posterior row replaced by hooded hooks from setiger 11; anterior row of granulated capillaries remaining intact for 7-8 setigers, thereafter becoming smaller, with less distinct granulations; single inferior sabre seta from setiger 12-13 (Figure 16E), increasing to 2 per neuropodium in middle setigers; tridentate hooded hooks numbering 4-5 in a row and bearing prominent main fang surmounted by large apical tooth and very small one (Figure 16F).

Branchiae from setiger 2, continuing to near posterior end; each branchia appearing glandular, extending across middorsal line (Figure 16A). Dorsal ciliary bands extending across segments from bases of branchiae on setigers 3-6, with cilia of setiger 3 merging to some extent with large dorsal field of nuchal cilia on setigers 2 and 3. Pygidium with 4 fingerlike cirri, with bacillary glands, dorsal pair slightly longer and thinner than ventral pair (Figure 16B).

Remarks. Microspio hartmanae resembles M. maculata (Hartman) and M. pigmentata (Reish) from southern California in having the anterior margin of the prostomium entire and hooded hooks from setiger 11. It differs in having paired nuchal organs posterior to the caruncle. No such structure is reported for the California species. Furthermore, the tridentate hooded hooks of M. hartmanae number 4-5 in a fascicle, while-numbering 9-12 in the California species. Microspio pigmentata has quadridentate hoo~ hooks according to Hartman [1961]. Both California species are heavily banded with pigment anteriorly, with the arrangement of their pigment being specifically distinct. Such pigment is limited to small indistinct markings in M. hartmanae.

Etymology. It is a pleasure to name this species for the late Olga Hartman in recognition of her major contributions to polychaete systematics of the seas of South America and Antarctica.

Distribution. Argentina, subtidal on rocky and sandy bottoms, 18-26 m.

Microspio paradoxa, n. sp. Fig. 17

Material examined. Galapagos Islands, Anton Bruun Sta. 66122, 17-18 m (holotype, USNM 60513).

Description. Holotype posteriorly incomplete, measuring 3 mm long and 0.3 mm wide for 20 setigerous segments. Color in alcohol: opaque white without distinct pigmentation.

Prostomium deeply incised anteriorly, forming distince lobes (Figure 17A); caruncle indistinct; without occipital tentacle; with 2 pairs of eyes arranged trapezoidally.

Setiger 1 reduced, lacking notopodial lobes and notosetae; neuropodium short, conical, with fascicle of 4 slender capillaries. Setigers 2-3 with notosetal and neurosetal capillaries arranged in 2 more or less dorsoventral rows, with setae of anterior row shorter, thicker, and granulated and those of posterior row longer, thinner, and nongranulated. Setigers 4-6 slightly modified with notosetae thicker, and darker than on setigers 3-7 and displaced into U-shaped arrangement (Figure 17B); anterior row of granlulated capillaries (Figure 17D) forming anterior and ventralmost portion of U and posterior row on nongranulated capillaries (Figure 17E) composing posterior portion of U; 3 longer, thinner, nongranulated capillaries (Figure 17C) located at dorsalmost end of U; neurosetae of setigers 4-6 also thicker than those of 3 and 7. Notosetae of setigers 7-10 arranged in 2 rows, with anterior row bearing short, thick, granulated bilimbate capillaries; posterior row with 3-4 peculiar capillaries having notch or opening in sheath (Figure 17F) and 3 long, finely bristled capillaries (Figure 17G) in superior position; bristled capillaries occurring on some following setigers. Notopodia subsequent to setiger 10 with capillaries as on setigers 2-3. Neurosetae of setigers 7-8 arranged in typical pattern of 2 rows; hooded hooks replacing posterior row on setiger 9, continuing to end of fragment; granulated setae of anterior row persisting for a few setigers; inferior sabre setae present on some segments; tridentate hooded hooks with main fang surmounted by 2 small apical teeth (Figure 17H).

Branchiae from setiger 2, each broad, with numerous bacillary glands.

Remarks. Despite the fragmentary nature

T

r-0 .2mm ----t

Fig. 16. Microspio n. sp. (holotype, USNM 67513). A, anterior end in dorsal Vlew; B, posterior end in dorsal view; C, anterior row capillary notoseta from setiger 5; D, same from posterior row; E, inferior sabre seta; F, hooded hook in lateral view.

o


. :./

": . • • ~I

o F G

I I

Fig. 17. Microspio paradoxa n. sp. (holotype, USNM 60513). A, anterior end in dorsal view; B, diagram illustrating arrangement of capillary notosetae on setiger 5, with dorsalmost setae on left, anterior row on bottom and posterior row toward top (arrows identify respective setal types); C, superior capillary notosetae from setiger 5; D, same from anterior row; E, same from posterior row; F, 'notched' capillary notoseta from anterior row of setiger 8; G, bristled capillary notoseta from posterior row of setiger 8; H, hooded hook in lateral view.

239


T 20~m

1

D

Fig. 18. Microspio minuta (holotype, ZMH P-14930). A, anterior end in dorsal view; B, capillary notoseta from an anterior setigerj C, hooded hook in lateral view; D, inferior sabre seta.

of the holotype, Microspio paradoxa is clearly an unusual and significant species. By having thickened capillaries arranged in a U shape on setigers 4-5, the species exhibits a tendency toward modification of setiger 5, which is typical for species of the Polydora complex. Blake (1979c] has proposed that the Polydora complex may have originated form an ancestral form similar to Microspio and noted general similarities between Tripolydora spinosa and species of Microspio. The discovery of M. paradoxa exhibiting a trend toward modifIcation of se tiger 5 lends strong support to this hypothesis (see also Blake and Woodwick, 1981]. The species itself is unique among spionids in the nature of the peculiar notched notopodial capillaries and the bristled capillaries occurring on setigers 7-10.

Etymology. Paradoxus, Latin for strange or contrary to expectation.

Distribution. Galapagos Islands, 17-18 m.

Microspio minuta (Hartmann-Schroder, 1962) Fig. 18

Paraspio minuta Hartmann-Schroder, 1962b, p. 144, figs 183-186.

Spio (Microspio) ~: Foster, 1971a, p.35.

Material examined. Galapagos Islands, Anton Bruun Sta. 66122, 17-18 m, (1, USNM 60537).--Chile, 4 km north of Taltal, July 26, 1960, on rhizoids of Macrocystis, collected by G. Hartmann (holotype, ZMH P-14930).

Description. Holotype small, incomplete, measuring 2.5 mm long for 26 setigers. Specimen from Galapagos Islands 2.6 mm long with 18 setigers. Color in alcohol: brown, with medial dorsal pigment spots on holotype, suggesting it to be postlarval. Bacillary glands prominent on body, especially on parapodial lobes; each bacillary gland with terminal sensory cilium.

Prostomium deeply incised on anterior margin, forming prominent lobes (Figure 18A); 2 pairs of cup-shaped eyes; without caruncle and occipital tentacle; 2 prominent curved nuchal organs lateral to prostomium and nearly meeting at posterior margin of setiger 2. Dorsal sensory organs beginning on setiger 3 as double rows of ciliary patches (Figure I8A).

Setiger 1 reduced, lacking notopodium and notosetae. Notosetae and neurosetae of an-


terior se tigers arranged in 2 rows of unilimbate capillaries, with those of anterior row having granulated shafts (Figure 18B) and those of posterior row nongranulated. Tridentate hooded hooks (Figure 180) from setiger 9, accompanied by capillaries and single, inferior, granulated sabre seta throughout (Figure 180).

Branchiae from se tiger 2. Posterior end of both specimens damaged precluding observation of pygidial structure.

Remarks. The nuchal and dorsal sense organs were not noted in the original description. Microspio minuta is similar to M. moorei (Gravier) from Antarctic seas and M. ~kowianus (Claparede) from European -waters in having an incised prostomium and hooded hooks from setiger 9. M. minuta would appear to differ from those species in details relative to the nuchal and sensory organs, but since a global review of this genus is needed, a final discussion of specific differences must await that study.

Distribution. Galapagos Islands; Chile. Intertidal to 18 m.

Microspio ~ (Gravier, 1911)

Mesospio moorei Gravier, 1911, pp. 100-105, pI. 7, figs. 80-83, pI. 8, figs. 84-86. --Augener, 1932b, pp. 39-40.--Hartman, 1966, p. 17, pI. 4, figs. 1-3.--Bellan, 1975, p. 789.

Microspio ~: Foster, 1971a, p. 35.

Remarks. No specimens of Microspio moorei were encountered in the present study. The species was referred to Microspio by Foster [1971a]. This referral was supported by Blake and Kudenov [1978J but not by Fauchald [1977, p. 24J. As originally described, Gravier [1911] depicts a moderateized species having about 50 setigerous segments and measuring about 16 mm long. His illustrations of the anterior end [Gravier, 1911, pI. 7, fig. 80] are confusing in that the peristomium is incised on its anterior margin and extends beyond the dorsally located prostomium. The latter is entire and truncate on its anterior margin, not incised, and there are no frontal horns. It is probable that Hartman [1966J and Fauchald [1977J confused the incised anterior margin of the peristomium with the prostomium when they stated that frontal horns were present. According to Gravier, the caruncle extends to the anterior border of setiger 2 and lacks an occipital tentacle. Four eyes are present. Branchiae begin on setiger 2 and are free from dorsal lamellae. Neuropodial hooded hooks are bidentate and begin on se tiger 15. The pygidium bears 4 anal cirri.

It should be possible to identify M. moorei in future collections. No other described species to Microspio has an entire

prostomium and bidentate hooded hooks from setiger 15.

Distribution. South Shetland Islands, 6-30 m; Kerguelen Islands, intertidal to 15 m.

Microspio gracilis (Hartmann-Schroder, 1962) new combination

Fig. 19

Prionospio gracilis Hartmann-Schroder, 1962b, pp. 141-142, figs. 174-178.

Minuspio gracilis: Foster, 1971a, p. 107. --Orensanz, 1974, p. 43.

Material examined. Argentina, Santa Cruz, east of Patagonia, between hillocks of Salicornia, February 12, 1960, collected by-C:Hartmann-Schroder (holotype, ZMH P-14987).

Description. Holotype small, incomplete, measuring 3.0 mm long and 0.35 mm wide for 37 setigers. Color in alcohol: light tan, with small flecks of dorsal pigment.

Prostomium broadly rounded on anterior margin, with caruncle extending posterior to setiger 2 (Figure 19A); 2 pairs of eyes, with medial pair cup-shaped and lateral pair rounded. Peristomium reduced, not forming lateral wings; palps extending posteriorly for 6 setigers.

Parapodia with thin, rounded notopodial and neuropodial lamellae throughout, with interramal ciliary tuft (Figure 19B). Anterior capillary notosetae and neurosetae arranged in spreading fascicles; setae unilimbate, with narrow sheaths, granulations lacking. Bidentate hooded hooks beginning in neuropodia from setiger 18 and in notopodia from setiger 25; neuropodia with 2-3 hooks, 4-5 capillaries and 2 inferior sabre setae; notopodia with 1-2 hooks and 4-5 capillaries; hooks with small teeth barely wider than shaft (Figures 19C-19D).

Branchiae from setiger 2, numbering 14 pairs on holotype; free from notopodial lamellae (Figure 19A). Pygidium lacking on holotype, but described as being composed of 2 short lobes and 4 longer cirri [HartmannSchroder, 1962b].

Remarks. Prionospio gracilis differs from the Prionospio complex of genera in the structure of its parapodia, hooded hooks, and pygidium. Except for having notopodial hooded hooks as well as neuropodial, the species is well accommodated in the genus Microspio. All other known species of Microspio have only neuropodial hooded hooks. Another species of Prionospio related to K. gracilis, and also generically misplaced, is P. cirrobranchiata Day, 1961. It is possible that P. cirrobranchiata and P. gracilis may event~ally be assigned to a new genus. The present referral of K. gracilis to Microspio reflects this author's present opinion of which spionid genus best accommodates the features of the species.


I ]OJl

L

/301!_m __ Fig. 19. Microspio gracilis (holotype, ZMH P-14987). A, anterior end in dorsal view; B, right setiger 19 in anterior view; C, hooded hook in lateral view; D, detail of C (not to scale).

Distribution. Argentina, intertidal.

Pygospiopsis, n.g.

Type species: Pygospio dubia Monro, 1930. Gender feminine.

Diagnosis. Prostomium truncate on anterior margin, laterally flaring into divergent lobes but lacking distinct frontal horns; occipital tentacle present. Branchiae occurring on se tigers 2-3 and from setiger 7 to about 16, then absent from middle and posterior segments; branchiae of setigers 2-3 cirriform, free from notopodial lamellae; other branchiae fused to membranous notopodial lamellae. Notosetae all capillaries; neurosetae including capillaries and bristled acicular hooks lacking hoods. Pygidium surrounded by ring of anal ci rri.

Remarks. Pygospiopsis n. g. is similar to Pygospio Claparede in having an interrupted branchial pattern. Pygospiopsis differs most significantly from Pygospio and other spionids in lacking hooded hooks. Their presence is considered to be characteristic for the Spionidae.

Pygospiopsis ~ (Monro, 1930) new combination

Fig. 20

Pygospio dubia Monro, 1930, pp. 146-147, fig. 55a-55i.--Hartman, 1966, p. 21, pI. 5, figs. 4-7.--Richardson and Hedgpeth, 1977, p. 186.

Material examined. South Georgia, west of Cumberland Bay, 9.5 km SW of Jason Light, R/V Discovery Sta. 29, March 16, 1926, dredged on mud and stone bottom, 25 m (25 syntypes, BMNH ZB 1930.10.8.1576-1600).

Description. Largest complete syntypes up to 12 mm long and 1.0 mm wide for 54 segments. Color in alcohol: light tan.

Prostomium truncate on anterior margin, weakly incised, forming lateral lobes; caruncle short, extending to middle of setiger 1, bearing short occipital tentacle; eyes lacking (Figure 20A). Peristomium inflated; palps extending posteriorly to about setiger 7.

Setigers 1-7 with elliptical notopodial lamellae; setigers 7-16 with thin, transparent, rounded lamellae, and from se tiger 16 to end of body somewhat elongated, cirriform

300)1 m

Fig. 20. Pygospiopsis ~ (syntype, BMNH ZB 1930. 10.8.1576-1600). A, anterior end in dorsal view; B, posterior end in dorsal view; C, neuropodial hook in lateral view with inset showing detail of bristled portion (not to scale); D, left setiger 3 in anterior view; E, left setiger 8 in anterior view.


lamellae; neuropodial lamellae rounded throughout body. Branchiae on setigers 2-3 and 7 to about 16, absent from middle and posterior setigers; branchiae of setigers 2-3 cirriform (Figure 20D), free from notopodial lamellae; branchiae from setiger 7 fused to notopodial lamellae (Figure 20E).

Capillary notosetae and neurosetae unilimbate, thin, and nongranulated; setal fascicles arranged in 2 rows in anterior setigers and in single row in posterior setigersj 1-2 acicular hooks accompanying neurosetal capillaries from se tiger 17-20, lightly bristled near tips (Figure 20C).

Pygidium of 1 specimen observed with circle of 10-12 anal cirri (Figure 20B), but see Monro [1930, fig. 55h and 55iJ, who observed' ... a distinct pygidial sucker consisting of two vertical halves, which under the microscope have the appearance of oyster shells ••• ' [Monro, 1930, p. 146J.

Distribution. South Georgia; Antarctic Peninsula, Anvers Island. Shallow subtidal depths.

Genus Boccardia Carazzi, 1893 Boccardia acus (Rainer, 1973)

Fig. 21

Paraboccardia acus Rainer, 1973, pp. 550-553, fig. 3.

Boccardia (Paraboccardia) acus: Read, 1975, pp. 406-407, fig. 5a.

Material examined. New Zealand, Auckland, Chevelier Park Beach, Eltanin Sta. 91 (cruise 20), intertidal (6, USNM 69387).

Description. A moderate-sized species, up to 7 mm long and 0.4 mm wide for 60 setigerous segments. Color in alcohol: light tan.

Prostomium weakly incised on anterior margin; caruncle broad, extending to posterior border of se tiger 2; occipital tentacle lacking; with 2 pair of eyes, with anterior pair more widely separated than posterior pair (Figure 2lA).

Setiger 1 approximately one half as large as se tiger 2, with both notosetal and neurosetal capillaries present (Figure 21A). Capillary notosetae of setigers 2-4, 6 and subsequent se tigers limbate, numbering 14-16 per fascicle, arranged in 2 rows; posterior notopodia with 7-10 acicular spines and 3-4 slender capillaries (Figures 21B-21C). Neurosetae of se tigers 3-4 and 6-7 similar to notosetaej bidentate hooded hooks replacing all but 1-2 capillaries from setiger 8; hooks with reduced angle between main fang and apical tooth and with acute angle between main fang and shaft (Figure 21D).

Modified setiger 5 with 2-3 falcate, smooth, nonbristled spines, 3-4 bristletopped spines (Figure 21E) and 2-3 short capillary notosetae.

Branchiae present on setigers 2-4, 6 and following segments (Figure 21A) to about setigers 20-25, absent from posterior two thirds of body. Pygidium a broad, dorsally open disc (Figure 21B).

Remarks. Boccardia acus is only known previously from the north and south islands of New Zealand, where it is a common borer in mollusc shells, coralline algae, soft wood and stone.

Distribution. New Zealand. Intertidal to shallow subtidal.

Boccardia anophthalma (Rioja, 1962)

Polydora anophthalma Rioja, 1962, pp. 185-188, figs. 89-93.

Boccardia anophthalma: Blake, 1981a, pp. 957-959, fig. 5.

Material examined. Ecuador, Bah{a de Santa Elena, Anton Bruun Sta. 6670, 8-9 m (7, USNM 60479).

Remarks. In a recent investigation of polychaetes from the Gulf of California, Blake [1981aJ has determined that Polydora anophthalma Rioja [1962J is actually a species of Boccardia. The species is unique in the genus in having an accessory flange on the unbristled, smooth falcate spines of setiger 5. All other species have only simple spines lacking accessory flanges. The species is further unusual in that the prostomium and first 5 setigers are dorsoventrally compressed, appearing shrunken against the larger, oval-shaped setigers which follow. The Ecuadorian specimens exhibit all of these features and agree in other respects with the Mexican forms.

Distribution. Western Mexico; Ecuador. Bores into calcareous substrata.

Boccardia chilensis Blake and Woodwick, 1971

Polydora polybranchia: Fauvel, 1916, p. 441 [not Haswell, 1885J.

Boccardia sp. Hartman, 1948, p. 109. Boccardia chilensis Blake and Woodwick, 1971,

p. 36-37, fig. 3.--Read, 1975, pp. 398-399. --Carrasco, 1976, pp. 11-15, figs. 3, 8B. --Blake and Kudenov, 1978, pp. 238-240, fig. 33d-33e.

Boccardia jubata Rainer, 1973, pp. 547-548, fig. 1. Fide Read, 1975.

Material examined. Peru, Anton Bruun Sta. 65215, 0-5 m (2, USNM).--Chile, Lota, June 16, 1927, collected by W. L. Schmidt (1, USNM 61947).--Straits of Magellan, Hero Sta. 71-2-8, intertidal (5, USNM 60481).~ Argentina, Tierra del Fuego, Hero Sta. 71-2-8, intertidal (1, USNM 60482).

Remarks. Boccardia chilensis is widely distributed in the southern hemisphere [Blake and Kudenov, 1978J. The notosetae of seti-


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.. :' ... :.:

.. i '.'

" ..

Fig. 21. Boccardia acus (Eltanin Sta. 91, USNM 69387). A, anterior end in dorsal view; B, posterior end in dorsal view; e, posterior notopodium in dorsal view; D, hooded hook; E, 2 groups of bristled and nonbristled major spines from setiger 5.

245


ger 1 are variable in length. Some are as long as those of setiger 2 while they are much longer on other specimens. The species is characterized by having a deeply incised prostomium with a prominent occipital tentacle and the caruncle extending posteriorly to the end of se tiger 2. There is an additional middorsal swelling on setigers 5-8. The major setae of setiger 5 include simple falcate spines and spines with a distal concavity and central cone. Branchiae occur on setigers 3-4, 6, and subsequent setigers. The pygidium is a weakly divided, fleshy pad. The Peruvian specimens represent a northward range extension for the species.

Distribution. Peru; Chile; Australia, New South Wales, Victoria; New Zealand; Falkland Islands; Macquarie Island. Intertidal to 5-7 m in various microhabitats associated with bivalves and encrusting organisms.

Boccardia natrix (Soderstrom, 1920) Fig. 22

Polydora natrix Soderstrom, 1920, pp. 254-256, figs. 165-166.

Boccardia natrix: Hartman, 1948, pp. 107-108, pl.~igs. 6-9; 1966, p. 15, pI. 3, figs. 7-10.

Polydora polybranchia: Ehlers, 1901a, pp. 164-165.--Pratt, 1901, p. 16 [not Haswell, 1885].

Material examined. Argentina, IBM Sta. WH-389 , in sponge, 200 m (1, USNM 67527). --Chile, Straits of Magellan, between Bah1a Inutil and Admiralty Sound, Puerto Condor (now El Condor), (syntype, SMNH 530).--Near Falkland Islands, Burwood Bank, Eltanin Sta. 1596, 124 m (2, USNM 69388).--South Georgia, Eltanin Sta. 220-320 m (1, USNM 60483). --South Pacific Ocean, SW of Campbell Island, Eltanin Sta. 1989, 589-594 m (2, USNM 60484) •

Description. A moderate-sized species, measuring up to 13 mm long and 0.75 mm wide for 52 setigers.

Prostomium incised on anterior margin, without dorsal furrow; without occipital ten-tacle; 4 eyes present or eyes lacking; caruncle extending posteriorly to near posterior margin of setiger 2; nuchal grooves located lateral to caruncle (Figure 22A). Peristomium not inflated; palps thickened, extending posteriorly for 7-8 setigers.

Anterior border of setiger 1 delimited by distinct ridge, separating it from peristomium (Figure 22A); setiger 1 well developed, with parapodia displaced laterally rather than elevated dorsally as in related species; notopodium oblong, fingerlike; neuropodium rounded; both fascicles with thin capillary setae. Notosetae of setigers 2-4, 6, and subsequent setigers arranged in 3 rows, those

of anterior row shortest and those of posterior row longest; capillaries unilimbate, with narrow transparent sheath; notosetae of posterior setigers reduced to simple fascicle of 7-8 capillaries of various lengths; posterior end of body appearing somewhat spinous, but special posterior needles or spines lacking. Neurosetae of setigers 2-4 and 6 similar in form and arrangement to notosetae; bidentate hooded hooks beginning in neuropodia from setiger 7, numbering 4-5 per neuropodium; hooks with reduced angle between apical tooth and main fang and wide angle between main fang and shaft (Figure 22E); hooks accompanied by 2-3 thin inferior capillaries throughout.

Setiger 5 heavily modified, twice as large as setiger 6 (Figure 22A); modified setae including 3 large bristle-topped spines lying ventral to 1-2 smaller, simple falcate spines (Figure '!aJ,; bristle-topped spilleis with rounded apex bearing large and small smooth bosses, with a tall cone of bristles between them (Figure 22B); these spines appearing falcate in some views (Figures 22C-22D); unworn falcate spines with a terminal mucron (Figure 22B); notopodial capillaries lacking; with neuropodial fascicle of 4-5 unilimbate capillaries.

Branchiae present on se tigers 2-3, 6 and subsequent se tigers to near posterior end of body. Pygidium consisting of 4 equal lobes.

Remarks. Boccardia natrix is apparently commensal with sponges in South American waters. The records of Polydora (Boccardia) natrix by Berkeley and Berkeley [1936, 1952] and Banse et al. [1968] from British Columbia and the Puget Sound have been referred to!. pugettensis Blake [1979b]. The retord from the Kurile Islands by Khlebovitsch [1961] needs to be reevaluated on the basis of these findings.

Among the specimens used by Soderstrom [1920, p. 256] as the basis for the original description of Polydora ~ were some which had previously been identified as P. polybranchia by Ehlers [1901a, pp. 164-165] from Puerto Condor in southern South America. Those specimens as well as others from the Burwood Bank near the Falkland Islands were later re examined by Hartman [1948, pp. 107-108], who noted that 'The original description was probably based on the find from Puerto Condor.' When the type material was requested from the Swedish Museum by the present investigator, L. Orrhage forwarded the Puerto Condor specimens from their type collections. The Burwood Bank specimens, however, should also be considered as syntypes since Hartman [1948] stated that they were the same species. The exact locality of the Puerto Condor collection is not given by either Soderstrom [1920] or Hartman [1948]. Ehlers [1901a, p. 5], however, provides the


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500pJD I 50)lJD I 120pm] Fig. 22. Boccardia natrix (IBM Sta. WH-389, USNM 67527). A, anterior end in dorsal view; B, major spines and companion setae from se tiger 5; C, D, bristle-topped major spines; E, hooded hook in lateral view.

following details: 'Magellanisches Gebiet --Bahia Inutil and Admiralty Sound: Rio Condor 54 0 0'Br., 700 8'L.' This locality is approximately 100 km southwest of Punta Arenas, Chile, off the main avenue of the Strait of Magellan. There is no indication of habitat or depth. The Burwood Bank specimens are listed as being from 137-150 m by Soderstrom [1920, p. 256J. This is station 59 of the Swedish Antarctic Expedition 1901-1903 [Hart-

man, 1953, p. 6J. Two newly collected specimens of Boccardia natrix are from Burwood Bank (Eltanin Sta. l~

Boccardia natrix is most closely related to!. androgyna Read [1975J, a hermaphroditic species inhabiting shallow subtidal sponges in New Zealand. Both species have notosetae on setiger 1 and similar major spines in setiger 5. Boccardia androgyna, however, has only a weakly notched prostomium, acicular


notopodial spines in posterior setigers, and a 4-10bed pygidium with 2 dorsal lobes being much smaller than the ventral ones. ~dia natrix, on the other hand, has a distinctly notched prostomium and lacks posterior notopodial spines, and the 4 lobes of the pygidium are of equal size. Moreover, B. natrix reportedly broods the larvae within the body of the female [Soderstrom, 1920], while Read [1975] indicates that ~. androgyna deposits its eggs into strings of capsules, with some containing nurse eggs. Boccardia natrix is also related to B. otakouica ~, 1973 from New Zeal~nd. The latter species, however, is a shell-boring form and has dorsal capillaries on setiger 5. Boccardia natrix is not known to bore into shells ~cks dorsal capillaries on setiger 5. Additional information on B. otakouica is presented by Read [1975]. - --------Distribution. Southern South America, including Straits of Magellan, subtidal to 200 m; near Falkland Islands, 124-150 m; South Georgia, 220-320 m; South Pacific Ocean, 589-594 m.

Boccardia polybranchia (Haswell, 1885)

Perialla claparedei Kinberg, 1866, p. 253; 1910, pp. 63-64, pI. 24, fig. 9.

Polydora (Leucodore) polybranchia Haswell, 1885, p. 275.

Polydora polybranchia: LoBianco, 1893, p. 28.--Ehlers, 1900, p. 217.--Soderstrom, 1920, pp. 256-257, fig. 167.--Augener, 1923b, pp. 70-71; 1926, pp. 157-158.

Polydora (Boccardia) polybranchia: Carazzi, 1893, pp. 16-17, pI. 2, figs. 1-3.--Mesnil, 1893, p. 645; 1896, pp. 221-227, pI. 14, figs. 9-21.--Ehlers, 1897, p. 87; 1901b, p. 265.--Augener, 1918, pp. 410-411. --Fauvel, 1927, pp. 58-59, fig. 20a-20i; 1941, pp. 287-288.--Monro, 1933, p. 1047; 1939, p. 125.--Day, 1934, p. 61.--0kuda, 1937, p. 238.--Berkeley and Berkeley, 1950, p. 52; 1952, pp. 16-17, figs. 24-25. --Rullier, 1964, p. 1092.--HartmannSchroder, 1974, pp. 171-172.

Polydora paucibranchus Ehlers, 1913, pp. 510-511, pI. 36, fig. 6.--Hartman, 1966, p. 18, pI. 4, fig. 13. New Synonymy.

Boccardia polybranchia: Chamberlin, 1919, p. 369. -Hartman, 1948, pp. 108-109, pI. 16, figs. 1-5, 13; 1966, p. 15, pI. 3, figs 11-12; 1967, pp. 111-112.--Hartmann-Schroder, 1962b, pp. 134-136, figs. 165-166; 1965, pp. 202-203, 302.--Imajima and Hartman, 1964, p. 279.--Day, 1967, pp. 463-464, fig. 18.1.a-18.1.h; 1971, p. 386.--Blake and Kudenov, 1978, pp. 236-238, fig. 32.

Polydora euryhalina Hartmann-Schroder, 1960, pp. 33-34, figs. 76-77. Fide HartmannSchroder, 1962b, p. 136.

Boccardia (Boccardia) wellingtonensis Read, 1975, pp. 404-406, fig. 4. New Synonymy.

Material examined. Brazil, Rio de Janeiro, 7-9 m, Eugenie Expedition, circa December 1851 (6 syntypes of Perialla claparedei, SMNH 742).--Argentina, intertidal collected by J. M. Orensanz: Santa Clara, March 1965 (4, USNM 67583); March 6, 1965 (35+, USNM 67584); January 1966 (7, USNM 67585); February 22, 1966 (100+, USNM 67586-67587); June 22, 1967 (25+, USNM 67590); June 1968 (3, USNM 67592); Santa Elena, May 5, 1967 (6, USNM 67588); May 7, 1967 (15, USNM 67589); Puerto Mar del Plata, Club Nautico, June 19, 1978 (28, USNM 67591); Miramar, with Ulva, May 2, 1969 (3, USNM 67594); Bahla de San BIas, north of Riacho Jabali, October 6, 1978 (13, USNM 67593); NE of Isla Jabali, April 20, 1970 (1, USNM 67595); Bahia de San Antonio, with Brachydontes, December 11, 1971 (7, USNM 67597); Golfo San Matlas, Las Grutas, December 9, 1971 (1, USNM 67596); Golfo San Jose, February 1973 (3, USNM 67598); Isla de las Fijacon, sandy beach, January 18, 1975 (13, USNM 67600); Isla de las Pajaros, in limestone with Cianophyra, January 18, 1975 (4, USNM 67599); Golfo Nuevo, Puerto Madryn, sandy beach, February 15, 1975 (71, USNM 67601); November 28, 1975 (35, USNM 67602). --Straits of Magellan, Hero Sta. 71-2-2, intertidal (19, USNM 6048~S of Punta Arenas, Chile, SOSC Sta. 69-6A, 53030.48'S, 70050.33'W, 0-1 m (6, USNM 69390).--Tierra Del Fuego, Hero Sta. 71-2-8, intertidal (19, USNM 60488).--Staten Island, off Tierra del Fuego, Hero Sta. 71-2-16, 18, 19, 21, 30, 31, 32,~ 37, 39, 40, 41, intertidal to 1 m (189, USNM 60489-60501).--Chile, Lund University Chile Expedition (1948-1949): Sta. M-56, 60, 113, 115, 116, 121, 123, 131, 133, intertidal (23, SMNH).--Chile, Montemar, Eltanin Sta. 300 (cruise 25), intertidal (10, USNM 69389); Mehuin, estuarine, intertidal, May 20, 1974 and in sand, February 20, 1975, collected by C. Bertran (14, AHF).--Peru, Punta Chira, August 9, 1955, collected by G. Hartmann (25, ZMH P-14974); Playa de Cascadas, below cliffs at Chorillos, 12°10'S, 77 002.4'W, March 10, 1967, collected by D. W. Kirtley (100+, USNM 59912); Playa Conchas, S of Playa Herradura, 12°12'S, 77 003'W, March 10, 1967, collected by D. W. Kirtley (125, USNM 59913); SE of Punta Hermosa, 12012.5'S, 77003'W, sand and rocks on beach, March 10, 1967, collected by D. W. Kirtley (60, USNM 59914). --Ecuador, Bahla de Santa Elena, ~ Bruun Sta. 6671, 3 m (1, USNM 60485).--South Pacific Ocean, SW Campbell Island, Eltanin Sta. 1989, 589-594 m (34, USNM 60486). Kerguelen Islands, intertidal in algal zone (holotype of Polydora paucibranchus, 2MB 5857).

Description. A large species, up 15 mm long for about 80 segments. Color in alcohol: body light tan, with conspicuous dark pigment on prostomium and palps. Prostomium incised on anterior margin; caruncle extend-


ing to end of setiger 3; without occipital tentacle; with 2 pairs of eyes. Se tiger 1 reduced, lacking notosetae. Setiger 5 with simple falcate, nonbristled spines, bristletopped spines, and neuropodial fascicle of capillaries. Bidentate neuropodial hooded hooks from setiger 7. Branchiae on setigers 2-4, 6 and subsequent setigers, missing from last one fourth of body. Pygidium with 4 equal lobes.

Remarks. A redescription of Boccardia polybranchia from near the type locality in New South Wales, Australia, was provided by Blake and Kudenov [1978]. The Australian specimens generally agreed with the more widely accepted definition of the species as established by Carazzi [1893] from Naples and by Mesnil [1896] from France. The extensive materials now available from South American waters agree very well with those from Australia and Macquarie Island reported by Blake and Kudenov [1978]. One feature of these forms from the southern hemisphere which differs from the European forms is the structure of the pygidium: it is divided into 4 nearly equal lobes. Both Carazzi [1893, fig. 18] and Mesnil [1896] described a tubular, undivided pygidium. It will be necessary to examine specimens from Europe before any conclusions can be reached about this difference.

Perialla claparedei Kinberg, 1866 from Rio de Janeiro is definitely the same species as !. polybranchia (Haswell, 1885). Kinberg's specimens, deposited in the Swedish Museum of Natural History, are in excellent condition and exhibit all of the diagnostic characters of !. polybranchia. The nomenclatural history of the species, however, is confused. Perialla claparedei was first named by Kinberg in 1866 with a short, inadequate description and without figures. The same short description but with very good figures was published 44 years later [Kinberg, 1910]. During this interval, Polydora polybranchia Haswell, 1885 was described. The description was supplemented and figured by Carazzi [1893] and placed in a new subgenus Boccardia. This was followed by LoBianco [1893], Mesnil [1893, 1896], and others. In his monograph on the Spionidae, Soderstrom [1920] examined Kinberg's types and referred Perialla claparedei Kinberg, 1910 (he overlooked the earlier reference of Kinberg, 1866, p. 253) to Polydora polybranchia Haswell, 1885 and included the record of these syntypes from Brazil as a new locality for ~. polybranchia. Hartman [1948] also examined Kinberg's specimens and referred them to !. polybranchia despite the formers priority ' ..• because of the incompletely known characters of the older one.' [Hartman, 1948, p. 109]. It is likely that Hartman did not examine all of the syntypes,

as she indicated only 1 specimen from Rio de Janeiro, Brazil (Eugenie Expedition). While the name Perialla claparedei Kinberg, 1866 clearly has priority, no effort is made here to replace Boccardia polybranchia, as it upsets the well-established names of Boccardia Carazzi and !. polybranchia (Haswell). Instead, application will be made to the International Commission of Zoological Nomenclature to place Perialla claparedei on the Official Index of Rejected Names.

Polydora paucibranchus Ehlers, 1913, described fro~ the Kerguelen Islands and deposited with the Zoological Museum, Berlin, was examined and referred to Boccardia ~branchia. Boccardia wellingtonensis Read, 1975 was described as lacking both notosetae and notopodial lamellae on setiger 1. Many of the South American specimens exhibit the same features, or a minute lobe may occasionally be present on se tiger 1. Blake and Kudenov [1978] reported only very small notopodial lobes on their Australian specimens. Since they agree in all other characteristics, they are here considered to by synonymous with!. polybranchia.

Distribution. Widespread in the southern hemisphere: Australia; New Zealand; subantarctic islands; Chile, Peru; Argentina, Brazil; South Africa. Central America; western Canada; Mediterranean. Intertidal to shallow subtidal depths and occasional reports to 500+ m in South Pacific.

Boccardia tricuspa (Hartman, 1939)

Polydora tricuspa Hartman, 1939, pp. 16-17, figs. 3c-3k; 1961, p. 29.--Rioja, 1939, pp. 304-308, figs. 22-31; 1941, p. 727; 1943, p. 238; 1962, p. 185.

Boccardia tricuspa: Woodwick, 1963, pp. 209-212, fig. l.--Hartman, 1969, pp. 99-100, figs. 1-5.--Blake, 1981a, pp. 959-960.

Boccardia proboscidea: Carrasco, 1974, pp. 185-187, figs. 1-4; 1976, pp. 8-11, figs. 2, 8A, 13H-C [not Hartman, 1940]. Fide Blake and Kudenov, 1978.

Material examined. Ecuador, Bahia de Santa Elena, Anton Bruun Sta. 6670, 8-9 m (5, USNM 60502).

Description. Prostomium entire on anterior margin; caruncle extending to end of setiger 3; no occipital tentacle; with 4 small eyes. Setiger 1 reduced, lacking notosetae. Setiger 5 large, with simple falcate spines, tridentate spines, and neuropodial fascicle of capillaries. Neuropodial bidentate hooded hooks from setiger 7. No specialized setae in posterior notopodia. Branchiae very small on setigers 2-4 and 6, becoming large in subsequent setigers. Pygidium with 4 small lobes.


Remarks. These specimens agree with previous descriptions.

Distribution. California; Mexico; Ecuador; Galap~gos Islands; Chile. Intertidal to shallow subtidal; bores in calcareous substrata.

Genus Boccardiella Blake and Kudenov, 1978 Boccardiella ligerica (Ferronniere, 1898)

Boccardia ligerica Ferronniere, 1898, pp.l09-Ill, pI. 6, figs. a-i.--Blake and Woodwick, 1971, pp. 32-34, fig. l.--Light, 1977, pp. 67-68; 1978, pp. 144-146, fig. 145.

Polydora redeki Horst, 1920, p. Ill. Fide Blake and Woodwick, 1971.

Polydora (Boccardia) redeki: Fauvel, 1927, pp. 58, fig. 19 t,u.--Augener, 1939, pp. 141-142, fig. 2.--Rullier, 1960, pp. 231-243, figs. 1-31.--Eliason and Haahtela, 1969, pp. 215-218, fig. 1. [Not Okuda, 1937, fide Blake and Woodwick, 1971].

Polydora~ccardia) ligerica: Fauvel, 1927, pp. 57-58, figs. 19n-19s.

Polydora uncatiformis Monro, 1938, pp. 311-313, figs. 1-3. Fide Blake and Kudenov, 1978.

?Boccardia redeki: Hartman, 1941, pp. 304-305, pI. 48, figs. 44-45. Fide Light, 1977 •

Polydora nr. uncata: Hartman, 1954, p. 9 [not Berkeley, 1927, fide Light, 1977].

Boccardia nr. ligerica: Day, 1955, p. 415; 1967, p. 463, fig. 18.1.j.

Polydora~: Filice, 1958, p. 170 [not Berkeley, 1927, fide Light, 1977].

Boccardia redeki: Hartman, 1961, p. 28; 1969, p.~igs. 1-3.

Boccardia hamata: Orensanz and Estivariz, 1971, pp. 102-104, figs. 32-34 [not Webster,1879].

Boccardiella ligerica: Blake and Kudenov, 1978, p. 265.

Polydora (Boccardiella) nahe ligerica: Hartmann-Schroder, 1980, p. 400.

Material examined. Uruguay, Canelones, Arroyo de las Brujas, May 30, 1938, from brackish water among tubes of Ficopomatus enigmaticus (6, syntypes of Polydora uncatiformis, BMNH ZB 1938:5:30:1 4).--Argentina, ~ra de Mar Chiquita, Puerto Oridina, February 1965 (1, USNM 67528); among valves of molluscs at Club Nautico, January 25, 1968 (7, USNM 67529); in canal No. 7 with Ficopo~ enigmaticus, October 12, 1968 (55, USNM 58964; 20, AHF).

Description. Prostomium entire to slightly incised on anterior margin; caruncle eXtending posteriorly to se tiger 2-3; with 4 eyes; no occipital tentacle. Branchiae present on se tigers 2, 3, 7 and subsequent 15-18 setigers; absent on setigers 4-6. Setiger 1 reduced, lacking notosetae. Mod-

ified fifth setiger with notopodial and neuropodial capillaries and curved row of large, nonfalcate simple spines alternating with lanceolate, pennoned companion setae. Neuropodial bidentate hooded hooks from setiger 7. Posterior notopodial with 1-2 large, curved boat hooks projecting toward middorsal line. Pygidium formed by an undivided ventral plate bearing 2 posterolateral cirri.

Remarks. The synonymy of Polydora ~iformis Monro with Boccardia hamata (Webster), proposed by Blake [1966] was incorrect. In that earlier study, only part of the type series of P. uncatiformis was examined. Following a-new examination of all of the syntypes, the synonymy of P. uncati~ and Boccardiella ligerica Ts readily apparent.

Boccardiella ligerica occurs in brackish waters and has possibly been introduced into such areas in various parts of the world. The species was recently discovered in San Francisco Bay and its tributaries by Light [1977, 1978] and may be expected to occur in similar localities subjected to heavy ship traffic. The species is frequently associated with Ficopomatus (=Mercierella) enigmati~ (Fauvel).

Distribution. Western Europe; California; Uruguay and Argentina; West Indies; South Africa. Intertidal to shallow depths in brackish to near freshwater environments.

Boccardiella occipitalis, n. sp. Fig. 23

Material examined. Argentina, intertidal, collected by J. M. Orensanz: Bahia San BIas, N of Riacho Jabali, June 10, 1968 (holotype, USNM 67509), October 6, 1968 (paratype, USNM 67510); NE of Isla Jabali, April 1970 (2 paratypes, USNM 67511).

Description. A moderate-sized species, up to 12.5 mm long and 1.5 mm wide for 75 setigers. Color in alcohol: brown with black pigment on prostomium, caruncle, and dorsal and ventral surface.

Prostomium deeply incised on anterior margin, forming divergent lobes; caruncle narrow, coiled, continuing posteriorly to middle of setiger 4; occipitial tentacle present at level of setiger 1; 2 pairs of eyes: anterior pair cup-shaped, posterior pair oval, lateral to occipital tentacle (Figure 23A). Peristomium with irregular folding; palps short, extending posteriorly for 7-8 setigers.

Setiger 1 well developed, with long, fingerlike notopodial lamellae (Figure 23A); notosetae lacking; neuropodium elliptical, with fascicle of bristled capillaries. Setigers 2-4, 6 and subsequent segments with spreading fascicles of capillary notosetae arranged in 3 rows; capillaries with fine bristles

1 3 4

H I J K L

• m

-/ • !l1II111 I. ~ -, -

N Fig. 23. Boccardiella occipitalis n. sp. (holotype, USNM 67509). A, anterior end in dorsal view; B, capillary notoseta from setiger 6, with inset showing detail of shaft (not to scale); C, boathook from posterior notopodium; D, curved spines from posterior notopodium; E-G, series of hooded hooks from anterior (E), middle (F), and posterior (G) neuropodia showing reduction and loss of apical tooth and hood; H-J, companion setae from setiger 5, showing progressive degrees of wear; K, L, major spines of setiger 5; M, ventral fimbriated capillary from setiger 5; N, detail of M (not to scale); 0, left posterior parapodium in anterior view; P, pygidium in dorsal view. Scale: 1 = 500 ~m (Figure 23A); 2 = 200 ~m (Figures 230-23P); 3 = 100 ~m (Figures 23C-23D); 4 = 40 ~m (Figures 23B, 23E-23M).


arranged in transverse rows (Figure 23B); far posterior notopodia with single large, recurved boathooks (Figure 23C), 3-4 shorter curved spines (Figure 23D) and fascicle of long bristled capillaries; notosetae projecting medially toward channel formed between dorsally elevated notopodia (Figure 230). Neurosetae of se tigers 2-3 and 6 with capillaries similar in arrangement and structure to notosetae; hooded hooks from setiger 7, up to 8-9 per neuropodium with 1-2 inferior capillaries in anterior setigers; bidentate hooks with reduced angle between main fang and apical tooth in anterior setigers (Figure 23E), with apical tooth gradually becoming smaller in middle segments (Figure 23F) and lost entirely, along with hoods in far posterior setigers (Figure 23G).

Setiger 5 modified, with dorsal masculature overlapping se tiger 6 (Figure 23A); setae including curved row of 7-8 simple, blunttipped spines (Figures 23K-23L) alternating with hastate companion setae showing different degrees of wear (Figures 23H-23J) and neuropodial fascicle of bristled capillaries (Figures 23M-23N); dorsal fascicle lacking.

Branchiae on setigers 2, 3, 6 and continuing (Figure 23A) for about one half of body; branchiae short, never extending more than one half distance to midline.

Pygidium with 2 lappets, each bearing long cirrus on lateral corners (Figure 23P).

Remarks. Boccardiella occipitalis is similar to ~. hamata (Webster), !. ligerica (Ferronniere), and B. bihamata Blake and Kudenov in having r;curved boathooks projecting medially into a channel between posterior notopodia. The species is closest to B.bihamata in having 2 types of posterior ~otopodial spines and unidentate hooded and unhooded hooks in posterior neuropodia. Boccardiella occipitalis differs from!. bihamata and other species of the genus in having an occipital tentacle, unusual bristled capillary setae, and a coiled, narrow caruncle and by lacking dorsal capillaries on setiger 5.

Etymology. Occipital, Latin referring to the back of the head. The name is suggested by the presence of an occipital tent-acle on this species.

Distribution. Argentina, intertidal.

Boccardiella sp. A

Material examined. Ecuador, Bah{a de Santa Elena, Anton Bruun Sta. 6669, Sta. 6670, 8-9 m (2 specimens, USNM 60503-60504).

Remarks. Both specimens are posteriorly incomplete, precluding specific identification due to the critical importance of the pygidial structure. The specimen from station 6670 has enough of the posterior end to

see posterior notopodial spines, including a single large recurved boathook and 2-3 straight acicular spines. There is no evidence of the occipital tentacle which characterizes !. occipitalis, and both specimens possess a fascicle of dorsal capillaries on setiger 5 which are lacking on!. occipitalis (see above). The apical tooth is retained on the hooded hooks in posterior neuropodia. The specimens appear to most closely resemble B. hamata (Webster) from North America [Blake, 1966] and B. bihamata Blake and Kudenov from Australi~. Final identification must await additional collections.

Distribution. Ecuador.

Genus Carazziella Blake and Kudenov, 1978 Carazziella carrascoi Blake, 1979

Polydora citrona: Carrasco, 1974, pp. 193-194, figs. 20-26; 1976, pp. 25-28, figs. 9, 13G, 21A [not Hartman, 1941]. Fide Blake, 1979a.

Carazziella carrascoi Blake, 1979a, pp. 473-475, fig. 5.

Remarks. Species of the genus Carazziel~ have 2 types of major spines on setiger 5 and branchiae beginning posterior to setiger 5. They lack constrictions on the shafts of the hooded hooks. Carazziella carrascoi is characterized by having well-developed notopodial and neuropodial lobes on se tiger 1, both of which bear fascicles of capillary setae. The major spines of setiger 5 include 1 type having an expanded bristled top bearing 2 distinct knobs and a second bristled type with a narrow, falcate tip. Details concerning this species and its relationship to other species are given by Blake [1979a]. .

Distribution. Chile, Bahfa de Concepcion, 2-14 m in mixed sediments.

Carazziella patagonica Blake, 1979

Carazziella patagonica Blake, 1979a, pp. 475-477, fig. 6.

Remarks. Carazziella patagonica is similar to ~. carrascoi in the general configuration of the major spines of se tiger 5 but differs from all species of the genus in having a well-developed notopodium with setae on setiger 1 while entirely lacking a neuropodium. Details of this species are given by Blake [1979a].

Distribution. Argentina, Golfo San Matfas, 36 m in mixed sediments.

Amphipolydora, n.g.

Type species: Polydora abranchiata Hartman, 1953. Gender feminine.


Diagnosis. Prostomium entire on anterior margin, extending posteriorly as caruncle; eyes and occipital tentacle absent. Setiger 1 without notosetae. Modified setiger 5 with 2 types of major spines: (1) first type having an expanded end with terminal tooth and surrounding collar; (2) second type simple acicular in form, dorsal to first type; notopodial and neuropodial capillary fascicles present. Hooded hooks bidentate, from setiger 7, with conspicuous angle between teeth, no constriction on shaft. Branchiae completely lacking. Prominent dor-sal bacillary glands on anterior setigers. Pygidium reduced, with 2-4 indistinct knobs.

Remarks. Amphipolydora is monotypic and unusual among the Polydora complex in lacking branchiae. Prominent bundles of dorsal bacillary glands occur on some setigers in the usual location of branchiae. Amphipolydora resembles Boccardia Carazzi and Caraz ziella Blake and Kudenov in having 2 types of major spines on setiger 5. The form of the expanded spines appears to be closer to some Boccardia species rather than Carazziella. An affinity to Boccardia is also suggested by the form of the hooded hooks, in which there is a prominent angle between the teeth; this angle is reduced in most Carazziella species. Amphipolydora exhibit-s--a--greater degree of reduction of the pygidium than in any other polydorid.

Etymology. Amphi, Greek for around; Polydora, mythical Greek boy and name of the reIat;d genus, Polydora Bosc. The name is suggested by the close affinities Amphipoly~ shares with related genera.

Amphipolydora abranchiata (Hartman, 1953) new combination

Fig. 24

Polydora abranchiata, Hartman, 1953 p. 44, fig. 13c-13d.

Material examined. Off Argentina, 370

50'S, 56011'W, 100 m, mixed sand and gravel, collected by Swedish Southpolar Expedition 1901-1903, Sta. 2, December 23, 1901 (5 and 21 syntypes, SMNH 618, 621).

Description. A small species, measuring up to 2.3 mm long and 0.27 mm wide for 25 setigerous segments. Color in alcohol: light tan, body pigment lacking. First 11 segments narrow, remaining segments longer, appearing moniliform (Figure 24A). Some specimens exhibiting regenerating anterior and/or posterior ends (Figure 241).

Prostomium entire on anterior margin, continuing posteriorly as broad caruncle to posterior border of setiger 2 (Figure 24A). Setiger 2-4, 6-11 with capillary bilimbate notosetae arranged in 2 rows, 2-3 short, thickly sheathed in first row and 4-5 long,

thin capillaries, without special posterior spines. Neurosetae of setigers 2-4 and 6 with fascicles of unilimbate capillaries; bidentate hooded hooks from setiger 7, numbering 2-3 per neuropodium, accompanied by single inferior capillary seta; hooks with prominent angle between main fang and apical tooth and about 900 angle between main fang and shaft (Figure 24H).

Setiger 5 with 2 notopodial bilimbate capillaries having inflated sheaths (Figure 24B), 2-3 slender acicular spines (Figures 24C-24D) located dorsal to 2-3 enlarged, distally expanded major spines and ventral fascicle of 4-5 unilimbate capillaries; major spines with single large, smooth terminal tooth surrounded by subterminal collar (Figure 24E-24G).

Branchie completely lacking. Bundles of prominent bacillary glands present dorsally on setigers 3 and 5-11 (Figure 23A). Pygidium with 2-4 small, indistinct knobs (Figure 24J).

Remarks. The original description of Polydora abranchiata by Hartman [1953] was deficient in several respects: the smaller acicular spines of setiger 5 were not mentioned; superior dorsal capillary setae are present on setiger 5, not absent; the pygidium has small lobes; and prominent bundles of bacillary glands are present. The presence of enlarged moniliform body segments in all specimens and the presence of regenerating fragments suggest that architomic asexual reproduction occurs in this species.

Distribution. Argentina, subtidal in sand and gravel, 100 m.

Genus Polydora Bosc, 1802 Polydora colonia Moore, 1907

Polydora colonia Moore, 1907, pp. 199-201, pI. 15, figs. 18-23.--Hartman, 1945, pp. 32-33.--Blake, 1971, pp. 15-16, fig. 10 [synonymy].--Day, 1973, pp. 69.

Polydora ancistrata Jones, 1962, pp. 185-187, figs. 55-65. Fide Blake, 1971.

Polydora hoplura inhaca Day, 1957, p. 99, fig. 6k-l; 1967~68, fig. 18.2.n. Fide Day, 1967 [footnote, p. 468]; 1973, p.~

Material examined. Argentina, IBM Sta. Comp-l, 18-30 m in sponge (2, USNM 67534).

Description. Prostomium entire on anterior margin, with caruncle extending posteriorly to setiger 2; eyes and occipital tentacle absent. Notosetae absent on setiger 1. Modified fifth setiger with notopodial and neuropodial fascicles of capillaries and 3-4 heavy spines alternating with pennoned companion setae; major spines with 2 terminal teeth surrounded by subterminal collar. Bidentate hooded hooks from se tiger 7. Posterior notopodia with 1-2 large recurved boat-

.' :.~

\ \\ F\ , , \ \

B \t

T IOOJlm

1

Fig. 24. Amphipolydora abranchiata (syntype, SMNH 621). A, anterior end in dorsal view; B, capillary notoseta from setiger 5; C, D, acicular spines from same: E-G, major cusped spines from same; H, hooded hook in lateral view; I, regenerating anterior end; J, posterior end in lateral view.


hooks. Branchiae from se tiger 7, absent from posterior one half of body. Pygidium disclike, flattened with wide dorsal gap.

Remarks. This is the first record of Polydora colonia from South America. The species has been recorded from South Africa [Day, 1967] and is well known from the east coast of North America and Jamaica [Blake, 1971; Day, 1973]. The species is known to be limited to shorelines bordering the Atlantic Ocean. The records of P. colonia from the northwestern Pacific by~nnenkova [1938] and Uschakov [1955, 1965] need to be reevaluated after the recent description of P. uschakovi by Buzhinskaja [1971]. The latter species is larger than~. colonia, has similar appearing spines on setiger 5, but is not associated with sponges, as is typical for P. colonia.

Distribution. Eastern North America; Jamaica; South Africa; Argentina.

Polydora cirrosa Rioja, 1943 Fig. 25

Polydora cirrosa Rioja, 1943, pp. 233-238, figs. 8-25.--Reish, 1959, pp. 38, 67, 78. --Light, 1978, p. 177.

Material examined. Ecuador, Bahia de Santa Elena, Anton Bruun Sta. 6669, 8-9 m (1, USNM 60547).

Description. Specimen posteriorly incomplete, measuring 2.8 mm long and 0.5 mm wide for 28 setigerous segments. Color in alcohol: light tan.

Prostomium incised on anterior margin; caruncle continuing posteriorly to setiger 3 (Figure 25A); occipital tentacle present at about level of setiger 1; 2 pairs of eyes present: anterior pair cup-shaped, posterior pair oval.

Setiger 1 with long fingerlike notopodial lamellae, without notosetae (Figure 25A); neuropodial lamellae reduced, conical, with capillary neurosetae. Setigers 2-4, 6 and subsequent setigers with fascicles of unilimbate or bilimbate capillary notosetae arranged in 2 rows; from se tiger 17 setal fascicles reduced to 5-6 simple capillaries. Anterior neurosetae similar to notosetae; bidentate hooded hooks from setiger 7, numbering 12-17 per ramus; without accompanying capillaries; hooks with a distinct manubrium on shaft, angle between apical tooth and main fang reduced and hood covered with minute bristles (Figures 25E-25F).

Setiger 5 heavily modified, with dorsal musculature overlapping setiger 6 and part of 7 (Figure 25A); setae include curved row of 7-8 major spines alternating with plumose companion setae (Figure 25D), without notopodial or neuropodial capillaries; major spines falcate, lacking accessory structures

but with distinct subterminal expansion (Figures 25B-25C).

Branchiae from setiger 7, small at first, becoming full size by setiger 11 and overlapping at midline.

Remarks. Aside from 2 southern California records [Reish, 1959, p. 38; Light, 1978, p. 177 footnote], Polydora cirrosa has remained unreported since its original description by Rioja [1943] from Mazatlan and Guaymas, Mexico. The present specimen from Ecuador is small and posteriorly incomplete but agrees very well with the original account. Polydora cirrosa, ~. ligni Webster, and P. nuchalis (Woodwick) form a distinct specTes group. Each bears an occipital tentacle and has distinctly bristled companion setae on setiger 5 and a manubrium on the hooded hooks. Polydora ligni has a distinct accessory tooth on the major spines of setiger 5, while it is lacking in P. nuchalis and P. cirrosa. Polydora cirrosa Ts distinguished from P. nuchalis by the distinctly falcate nature of the major spines and the feathery companion setae on setiger 5 and by the long, fingerlike notopodial lamellae on setiger 1. Polydora nuchalis, on the other hand, bears nearly acicular major spines and bristled but not feathery companion setae on setiger 5 and has only short, conical notopodial lamellae on setiger 1.

Distribution. Southern California; western Mexico; Ecuador. Intertidal to 9 m.

Polydora ligni Webster, 1879

Polydora ligni Webster, 1879, p. 119, pI. 5, figs. 45-47.--Blake, 1971, pp. 5-6, figs. 1-2 [synonymy].--Orensanz and Estivariz, 1971, pp. 104-105, figs. 35-37.

Material examined. Argentina, intertidal, collected by J. M. Orensanz: Albufera de Mar Chicquita, Club Nautico, August 16, 1967 (2, USNM 67578); Desembocadura, January 15, 1970 (24, USNM 67580); Puerto de Mar del Plata, August 15, 1967 (100+, USNM 67577); January 19, 1968 (11, USNM 67579); April 1970 (6, USNM 67581); June 27, 1979, collected by M. T. de Mandri (12, USNM 67582).

Remarks. Polydora ligni is well known, and the present specimens agree very well with previous descriptions. The prostomium is bilobed on the anterior margin, and the caruncle bears an occipital tentacle just behind 4 eyes. Branchiae begin on setiger 7. The pygidium is a large, flaring disc. Notosetae are lacking on se tiger 1. The modified fifth setiger lacks both notopodial and neuropodial capillaries but has a curved row of large spines and closely adhering, featherlike companion setae. The major spines are slightly falcate with a short accessory tooth. J. D. Kudenov (personal com-


l O.3mm

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d it if ~~ W ,~w

~!.

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I . .'

<I -.':

:.~

.,

B C D L50Jlm~

Fig. 25. Polydora cirrosa (USNM 60547). A, anterior end in dorsal view; B-C, major spines from setiger 5; D, companion seta from setiger 5; E, hooded hook in lateral view; F, enlargement of E showing bristles on hood (not to scale).

munication, May 1980) has observed that unworn major spines of setiger 5 of ~. ligni bear a small flange in addition to the accessory tooth. As part of another study, I have observed this same structure with the scanning electron microscope. Rather than being a flange, the structure is a thin, membranous extension of the tooth which extends apically across the falcate curvature of the spine. Since it does not actually curve around the shaft, it should be considered as a keel rather than a flange. It is similar to, but not identical to, the tooth and flange arrangements of Polydora haswelli Blake and Kudenov [1978] and P. bioccipitalis Blake and Woodwick [1972]. The structure definitely occurs in the Argentinian specimens and has been observed in materials from New England, South Carolina, and California. Since the keel is readily worn away once the major spine is emergent, its value as a diagnostic character is limited.

Distribution. East, west, and gulf coasts of North America, Europe, Mexico, Argentina, Australia. Intertidal to shallow estuarine localities. In Argentina, the species is

frequently associated with the serpulid Ficopomatus (,Mercierella) enigmaticus (Fauvel) in areas of reduced salinities.

Polydora ciliata (Johnston, 1838)

Leucodore ciliatus Johnston, 1838, pp. 67-68, pI. 3, figs. 1-6.

Polydora ciliata: Soderstrom, 1920, pp. 261-262, fig. 169. Fauvel, 1916, p. 440: 1927, pp. 49-50, fig. 16i-16p.--HartmannSchroder, 1971, pp. 312-314, fig. 106.

Material examined. Uruguay, Jose Ignacio, La Pedrera, October 30, 1970, collected by S. Olivier (2, USNM 67574-5).--Argentina, intertidal, collected by J. M. Orensanz: Mar de Cobo, October 12, 1968, endolithic, with Bryopsis (4, JAB); Santa Clara, endolithic, March 1965 (7, JAB); January 21, 1966 (1, JAB); January 18, 1968, with Corallina (1, USNM 67570); June 1968 (10, USNM 67573); Mar del Plata, Cabo Corrientes, January 24, 1968 and February 1968, high intertidal, with bryozoans and sponges (11, USNM 67571-2); Miramar, with algae (7, USNM 67476).--South


Shetland Islands, Eltanin Sta. 993, 300 m (USNM 56451).

Remarks. These specimens agree well with published accounts. The prostomium is weakly incised. There is no occipital tentacle. Notosetae are absent from setiger 1. The major spines of setiger 5 are falcate and have a lateral accessory tooth rather than a flange as in P. websteri (see below). Branchiae begin on setiger 7. The pygidium is a flaring disc, with a dorsal notch.

The distribution of P. ciliata is probably more restricted than generally believed. Many records have proven to belong to related species such as P. websteri [see Blake, 1971; Blake and Woodwi~k, 1972; Blake and Kudenov, 1978]. The specimens from Uruguay and Argentina have been compared to specimens from Britain (Kent) and agree in all particulars. The species may prove to be limited to the Atlantic Ocean.

Distribution. Europe; South Africa; Uruguay and Argentina; Falkland Islands; South Shetland Islands. A borer in calcareous rock, molluscan shells and coralline algae. Intertidal to 300 m.

Polydora websteri Hartman, 1943

Polydora websteri Hartman, 1943, pp. 70-72, fig. l.--Blake, 1971, pp. 6-8, fig. 3 (synonymy]; 1981a, pp. 954-955.--Foster, 1971a, pp. 26-27, figs. 30-36 (synonymy].

Material examined. Peru, island near Pucusana, S of Callao, Anton Brunn Sta. 65215, 0-5 m (5, USNM 60564).--Ecuador, Bah!a de Santa Elena, Anton Brunn Sta. 6670, 8-9 m (1, USNM 60565).

Remarks. Polydora websteri differs from K. ciliata chiefly in having a distinct accessory flange on the major spines of setiger 5 instead of a tooth.

Distribution. East, west, and gulf coasts of North America; Mexico; Peru and Ecuador; Australia. A borer in calcareous substrata. Intertidal to ca. 10 m.

Polydora bioccipitalis Blake and Woodwick, 1972

Polydora biocci,italis Blake and Woodwick, 1972, pp. 75- 7, figs. 3-4.

Material examined. Chile, La Playa Aguilla (Eagle Beach), approximately 100 km S of Iquique, November 10, 1977, in mud blister of shells of living Mesodesma donacium, I-m depth, collected by D. Stewart (12, USNM 61957).

Remarks. Polydora bioccipitalis is unique in several respects. The prostomium is deeply notched on the anterior margin, continuing posteriorly over many segments as a low nuch-

al ridge. Two occipital tentacles are present in tandem arrangement immediately following 4 oval eyes. Palps are short for a Polydora, and extend posteriorly for only 4-5 setigers. Notosetae are absent on setiger 1. Bidentate hooded hooks first appear in neuropodia of se tiger 9-14. Notosetal capillaries are absent on setiger 5 and neuropodial capillaries are present only on small specimens. The major spines of setiger 5 alternate with pennoned companion setae, with each spine having an accessory tooth and flange. Branchiae begin on setiger 7 and continue to near the posterior end. The pygidium is disclike, with a dorsal gap. These specimens from Chile agree completely with the original account from California.

Distribution. California; Chile. A borer in mollusc shells. Intertidal.

Polydora rickettsi Woodwick, 1961

Polydora rickettsi Woodwick, 1961, pp. 78-81, figs. 1-7.

Material examined. Chile, Montemar, in shell of Concholepis concholepis, collected by R. Riveros Junigo (9, AHF).--Mexico, Cape San Lucas, attached to tube of Spirobranchus, intertidal, collected by E. F. Ricketts, March 17, 1940 (holotype and paratype, AHF Poly 620-1).

Remarks. Polydora rickettsi is similar to P. ciliata in having an accessory tooth on the major spines of se tiger 5, branchiae from setiger 7, and a disclike pygidium. Polydora rickettsi, however, has an entire instead of an incised prostomium, and the caruncle extends to the posterior margin of setiger 4 instead of 2. The hooded hooks of P. rickettsi were described as lacking a constriction on their shafts. An examination of the holotype and paratype and a series of slides prepared from them confirms most of Woodwick's [1961] description except that the hooded hooks do have a weakly developed constriction on the shaft. The Chilean specimens have more pronounced constrictions and distinct manubriums than are found on the type series from Mexico.

Distribution. Mexico; Chile. A borer in molluscan shells and an associate of serpulid masses. Intertidal.

Polydora ecuadoriana, n. sp. Fig. 26

Material examined. Ecuador, Bah!a de Santa Elena, Anton Bruun Sta. 6670, 8-9 m (holotype and paratype, USNM 60548-9; paratype, AHF Poly 1298); Sta. 6669, 200 m offshore, 8-10 m in shell fragments (33 paratypes, USNM 59920-2).

Description. A moderate-sized species,


up to 13.2 mm long and 0.32 mm wide for about 95 setigerous segments. Color in alcohol: light tan, with paired black spots dorsally on setigers 2-4 and transverse black bars on palps (Figure 26A).

Prostomium entire to incised on anterior margin; caruncle broad, extending posteriorly to middle of setiger 3; no occipital tentacle; 2 pairs of eyes present: posterior pair cup-shaped, anterior pair oval (Figure 26A). Peristomium broad, not inflated.

Setiger 1 with short notopodial and neuropodial lobes, without notosetae. Setigers 2-4, 6 and subsequent se tigers with 2 rows of notosetae formed of bilimbate fimbriated capillaries having finely granulated shafts (Figure 26C); posterior se tigers with 4-5 long, thin capillaries and 3-4 short pointed spines (Figure 26G). Neurosetae of se tigers 1-4, and 6 with unilimbate capillaries arranged in 2 rows; bidentate hooded hooks from setiger 7, completely replacing capillaries, numbering 5-6 per ramus initially, then increasing to 10-11 in middle body segments, and decreasing to 4-5 in far posterior setigers; hooks with distinct angle between main fang and apical tooth and reduced angle between main fang and shaft, with constriction and manubrium on shaft (Figure 26D).

Modified setiger 5 with dorsal fascicle of 3-4 fimbriated capillaries, 4-5 major spines with hastate companion setae (Figure 26B) and ventral fascicle of 5-6 unilimbate capillaries. Major spines falcate, with large accessory tooth and smaller accessory flange; flange very delicate and worn on older spines.

Branchiae from se tiger 7, continuing posteriorly for about three fourths of body. Pydigium a large disc, appearing scoop-shaped (Figures 26E-26F).

Remarks. Polydora ecuadoriana belongs to the P. ciliata-websteri group in having hooded hooks from se tiger 7 with a manubrium on the shaft of the hooded hooks, by lacking notosetae on setiger 1, and by having accessory flanges or teeth on the major spines of se tiger 5. Polydora ecuadoriana most closely resembles ~. pygidialis Blake and Woodwick from California and British Columbia and P. haswelli Blake and Kudenov from Australia. Both~. ecuadoriana and ~. pygidialis have large scoop-shaped pygidia and are borers in coralline habitats. Polydora pygidialis, however, has only a single tooth on the major spines of setiger 5, while K. ecuadoriana has both Po tooth and flange. In P. haswelli, the tooth and flange on the major-spines are much smaller in proportion, and the pygidium is disclike, not scoop-shaped. Another related species having both an accessory tooth and flange on the major spines is ~. bioccipitalis Blake and Woodwick, which occurs in Chile (see above). This species, however, has 2 occipital tentacles, branchiae extend

to the posterior end of the body, the pygidium is disclike, and most importantly, the hooded hooks begin on setiger 10-14 instead of 7. Polydora ecuadoriana differs from the above-mentioned species in having short, pointed spines in posterior notopodia.

Distribution. Ecuador, shallow subtidal sandy bottoms, boring into coralline structures and mollusc shells, 8-10 m.

Polydora ~ Langerhans, 1881

Polydora armata Langerhans, 1881, pp. 93-94, pI. 4, fig. 5a-5c.--Fauvel, 1927, pp. 55-56, fig. 19a-1ge.--Hartman, 1941, p. 306; 1969, p. 127, figs. 1-5.--Woodwick, 1964, p. 14, fig. 2 (1-6).--Day, 1967, pp. 466-468, fig. 18.2.i-18.2.j.--Rainer, 1973, pp. 557-558, fig. 7.--Read, 1975, pp. 412-413. --Blake and Kudenov, 1978, pp. 255-256, fig. 43a-43g.--Hartmann-Schroder, 1979, p. 134, figs. 299-302.

Polydora monilaris Ehlers, 1905, pp. 43-44, pI. 6, fig. 5-14. Fide Day, 1954.

Material examined. Ecuador, Bah{a de Santa Elena, Anton Bruun Sta. 6670, 8-9 m (2, USNM).--Chile, off SW coast, Archipelago Reino Adelaida, Eltanin Sta. 958, 92-101 m (20, USNM 61992); Sta. 960, 64 m (13, USNM 61993).--Juan Fernandez Islands, Anton Bruun Sta. Drab 135, 160-180 m (2 fragments, USNM 62000); Sta. MV65IV-55 (3, USNM 61999); Sta. MV65IV-57 (1, USNM 62001); Sta. 65235, 3-11 m (1, USNM 61997); Sta. 65239, 2-8 m (1, USNM 61998); Sta. 65240, 26-29 m (20+, USNM 62004).--Argentina, off Tierra del Fuego, Eltanin Sta. 222, 79-80 m (1, USNM 61991); Staten Island, Hero Sta. 672, 50 m (1, USNM 62002); Sta. 67~0 m (6, USNM 62003); Sta. 71-2-16, intertidal (1, USNM 61990).--Near Falkland Islands, Burwood Bank, Eltanin Sta. 344, 119 m (1, USNM 61989).--Drake Passage, near Powell Islands, Eltanin Sta. 993, 300 m (1, USNM 61994).--MacQuarie Island, Eltanin Sta. 1418, 80-101 m (8, USNM 61995).--New Zealand, South Island, off eastern coast, N of Dunedin, Eltanin Sta. 1431, 51 m (1, USNM 61996).

Remarks. There is evidence that the present material represents 2 distinct species. The specimens from Ecuador and along the coast of Chile are similar to those observed from southern California (J. A. Blake, unpublished data, 1965) and Australia [Blake and Kudenov, 1978]. The major spines of setiger 5 have 2 teeth connected by a hood or cowling that is cloaked with bristles. The posterior notopodial spines are very thick, frequently blunted, and arranged in a rosette which is sometimes deeply imbedded within the body. Specimens from the Juan Fernandez Islands and southeastern Argentina have major spines with a large, curved, and flattened tip. Each spine bears a somewhat


~ O.3rnrn----1

--------B C D 30prn IIIOJlrn I

Fig. 26. Polydora ecuadoriana n. sp. (holotype, USNM 60548). A, anterior end in dorsal view; B, major spines and companion setae from setiger 5; C, bilimbate fimbriated capillary notosetae from setiger 4; D, hooded hook in lateral view; E, posterior end in lateral view; F, same in dorsal view; G, left posterior notopodium in dorsal view (not to scale).

259


flattened, lateral accessory tooth, with the tip emerging from a cloak of bristles. The shape of the spines resemble those of Polydora caulleryi Mesnil [see Blake, 1971, fig. 8c-8d]. The posterior notopodial spines are thinner and more delicate, with the rosettes sometimes spread fanlike.

The specimen from South Island, New Zealand, is typical P. armata with very thick posterior spines,-while the specimen from Macquarie Island resembles the form with thin posterior spines.

These observations suggest that 2 species are involved. Typical P. armata with thickened posterior spines appear to be most common in warmer subtropical and temperate seas, ranging from near 38 0 N to 47 0 S latitudes. The form with thin posterior spines, on the other hand, appears to be most common in colder and perhaps deeper waters, with Eltanin records being between 520 -620 S lati----tude. This latter form appears to be transitional between the northern f. caulleryi on the one hand and typical P. armata on the other. To approach the problem more thoroughly, specimens of f. armata are being acquired from various parts of its reported range. A comprehensive analysis of its morphology, habitat preference, and temperature requirements are needed. A search is also underway for the location of the type of P. monilaris Ehlers, from New Zealand (refer;ed to P. armata by Day [1954]). Until these studie~completed, f. armata remains a broadly defined species.

Distribution. Australia; New Zealand; South America; Europe; northeastern Pacific; Japan.

Polydora antonbruunae, n. sp. Fig. 27

Material examined. Peru, Callao, Isla San Lorenzo, at Naval Base, Anton Bruun Sta. 65211, 2-6 m, in gastropod shells (holotype and 13 paratypes, USNM 60545-60546).

Description. A moderate-sized species, measuring up to 8.5 mm long and 0.4 mm wide for about 90 setigerous segments. Color in alcohol: light tan, body pigment lacking.

Prostomium conical, narrowed anteriorly, appearing almost pointed (Figure 27A); caruncle extending posteriorly to middle of setiger 3; occipital tentacle absent; eyes 0-4, when present, anterior pair oval and widely spaced, posterior pair cup-shaped, more closely spaced, often hidden by overlapping fold of caruncle; nuchal organs present lateral to caruncle (Figure 27A). Peristomium truncate anteriorly, with distinct prominences on some specimens (Figure 27A).

Setiger 1 reduced and hidden by overlapping setiger 2; with conical notopodial postsetal lamellae and 3-4 long, erect capillary

setae; neuropodia 1 lamellae elliptical with small fascicle of short capillaries. Setigers 2-4, 6 and subsequent segments with capillary notosetae arranged in 2 rows, with those setae of first row shortest and broadest; in middle and posterior setigers notosetae fewer, including 1-2 curved acicular spines (Figure 27C) and 4-5 longer, thinner capillaries; acicular spines tending to curve toward midline on some specimens. Neurosetae of setigers 2-4 and 6 with fascicles of unilimbate capillaries arranged in 2 rami; bidentate hooded hooks from setiger 7, numbering 4-5 per ramus and accompanied initially by 1-2 inferior capillaries; hooks with reduced angle between main fang and apical tooth and acute angle between main fang and shaft in anterior segments (Figure 27D); apical tooth reduced in middle and posterior segments becoming nearly lost in far posterior segments (Figure 27E).

Setiger 5 about twice size of adjacent setigers, overlapping setigers 6 and 7 with heavy dorsal musculature (Figure 27A). Setae including a curved row of 4-5 major spines accompanied by hastate companion setae (Figure 27B) and ventral fascicle of unilimbate capillaries; with dorsal fascicle lacking; major spines falcate, with large, prominent accessory flange (Figure 27B).

Branchiae from se tiger 7 (rarely 6), broad, straplike, meeting at midline by setiger 9 (Figure 27A), absent from posterior one third of body. Dorsal sense organs from setiger 7. Pygidium formed of 4 small glandular lobes, dorsal pair smaller (Figures 27F- 27G).

Remarks. Polydora antonbruunae appears to be closely related to the P. commensalis group by having notosetae ~n setiger 1, large flanges on the major spines of setiger 5, and

hooded hooks which lack constrictions on their shafts. Polydora antonbruunae, however has a unique conical prostomium which is nearly pointed on the anterior end, an unusual truncated appearance to the anterior margin of the peristomium, a greatly reduced setiger 1, and posterior acicular spines in the notopodia. It is highly unlikely that this unusual species will be confused with related forms.

Etymology. The species is named after the former research vessel, R/V Anton Bruun, on whose cruise to western South America this species was collected.

Distribution. Peru, boring into gastropod shells, 2-6 m.

Polydora magellanica, n. sp. Fig. 28

Material examined. Southwestern Chile, Eltanin Sta. 960, 64 m, boring into calcareous substrata (holotype and paratype, USNM 60556-60557).

, T·.


' .. :

.. :

I.

r--------B c

F

Fig. 27. Polydora antonbruunae n. sp. (holotype, USNM 60545). A, anterior end in dorsal view; B, major spines and companion setae from setiger 5; C, acicular notopodial spine from posterior se tiger; D, hooded hook from anterior setiger in lateral view; E, same from posterior setiger in lateral view; F, posterior end in dorsal view; G, pygidium and last 6 setigers in dorsal view.

261

" ..

.,

.'

., .. ' ., ..

:',

:.:


Description. A small species, holotype up to 5.4 mm long and 0.3 mm wide for approximately 85 setigers. Col or in alcohol: light tan.

Prostomium deeply incised on anterior margin, forming 2 lobes; caruncle continuing posteriorly to middle of se tiger 4, without occipital tentacle or eyes (Figure 28A).

Setiger 1 with conical notopodial postsetal lamellae and slender capillary notosetae and neurosetae. Notopodia of setigers 2-4, 6 and subsequent setigers with unilimbate capillaries arranged into 2 rows; posterior notopodia with several short acicular spines in addition to longer capillaries (Figure 28H). Neurosetae of setigers 2-4 and 6 unilimbate capillaries arranged in 2 rows. Bidentate hooded hooks from se tiger 7, 3-4 per ramus accompanied initially by single inferior capillary seta; hooks with wide angle between main fang and shaft and reduced angle between teeth (Figure 28B); apical tooth gradually reduced in size in middle setigers (Figure 28C) and finally lost altogether in posterior setigers (Figure 28D).

Modified setiger 5 with dorsal fascicle of 3-4 bilimbate capillaries having granulated shafts (Figure 28E), curved row of 4 major spines (Figure 28F), alternating with bilimbate capillary companion setae (Figure 28G) and ventral fascicle of unilimbate capillaries; major spines falcate, with 2 lateral accessory teeth (Figure 28F).

Branchiae from se tiger 10 (Figure 28A), continuing through middle of body, absent from posterior one third. Pygidium divided into 4 lobes, with deep dorsal and ventral notches (Figure 281).

Remarks. Polydora magellanica belongs to the ~. giardi group and is most closely related to~. bifurcata Blake [1981b] from California in having 2 accessory teeth on the major spines of setiger 5 and posterior notopodial spines. Polydora magellanica has branchiae from setiger 10, posterior notopodial spines not conspicuously wider than the accompanying capillaries, and the apical tooth of the hooded hooks is lost in posterior neuropodia. In contrast, P. bifurcata has branchiae from setiger 8, p;sterior notopodial spines much thicker than the capillaries and hooded hooks which retain the apical tooth throughout.

Distribution. Southwestern Chile boring into calcareous substrata, associated with Polydora~, 64 m.

Polydora giardi Mesnil, 1896

Polydora giardi Mesnil, 1896, pp. 195-202, pI. 13, figs. 1-12.--Blake and Kudenov, 1978, p. 252, fig. 3'8i-k [synonymy].--Day and Blake, 1979, pp. 20-30, figs. 1-2. --Blake, 1981a, p. 951, figs. 2A-B.

Material examined. Ecuador, Bahia de Santa Elena, Anton Bruun Sta. 6670 8-9 m (1 USNM 60555).--Chile, Montemar, Elt~nin Sta. ' 300 (cruise 25), intertidal (6, USNM 69397); in shell of Concholepis concholepis, collected by F. Riveros Junigo, intertidal (41, AHF).--Juan Fernandez Islands, Anton Bruun Sta. Drab 126, 1 m (1, USNM 60551); Sta. Drab 134 (30+, USNM 60553); Sta. Drab 135, 160-180 m (10+, USNM); Sta. 65235, 3-11 m (10+, USNM 59918); Sta. 65239, 2-8 m (2, USNM 60550); Sta. 65240, 24-27 m (70+, USNM 62005); Sta. 65243, 9 m (1, USNM 60552); Sta. 65253, 80-125 m (20+, USNM 60554); Sta. MV65IV-58 (30+, USNM).

Description. Prostomium deeply incised on anterior margin, with some specimens having exceptionally long prostomial lobes; caruncle extending posteriorly to setiger 3; without occipital tentacle or eyes. Setiger 1 well developed, biramous. Neuropodial bidentate hooded hooks from se tiger 7. Without specialized posterior notosetae. Modified setiger 5 with fascicle of geniculate notosetae, a row of heavy spines and hastate companion setae, and small fascicle of neuropodial capillaries; major spines falcate, with large accessory tooth on concave side and small accessory spur on convex side. Branchiae from setiger 9, absent from posterior one third of body. Pygidium small disclike. '

Remarks. These specimens agree well with previous descriptions. Polydora giardi differs from ~. magellanica (see above) in lacking instead of having specialized posterior notopodial spines. Further, P. giardi has a single large accessory tooth and very small accessory spur on the major spines of setiger 5, while ~. magellanica has 2 very prominent accessory teeth.

Distribution. Ecuador; Chile; cosmopolitan in calcareous substrata. Associated with !. rickettsi and P. armata in Chilean gastropods. Intertidal~O m.

Polydora ~ Claparede, 1870

Polydora flava Claparede, 1870, pp. 487-488.--Mesnil, 1896, pp. 182-191, pI. 11, figs. 18-26, pI. 12, figs. 1-22.--Fauvel, 1927, pp. 52-54, figs. 17n-17u.--Day, 1967, pp. 468-469, fig. 18.3.a-18.3.d. --Hartmann-Schroder, 1971, p. 305. --Blake and Kudenov, 1978, p. 248, fig. 38a-38c.

Material examined. Uruguay, IBM Sta. N-248, 170 m (2, USNM 67535).--Argentina, collected by J. M. Orensanz, intertidal, 1975: Golfo de San Matias, Isla de las Jijacan, sandy beach, January 18 (2, USNM); Golfo Nuevo, Puerto Madryn, sandy beach, February 15 (32, USNM 67536); November 28 (50+, USNM 67537) .

, i; ::

,~ ~

" ; .: .~ ~

~

C ,D~ i ~ F

Fig. 28. Polydora magellanica n. sp. (holotype, USNM 60565). A, anterior end in dorsal view; B-D, hooded hooks in lateral view, showing progressive sequence in reduction of apical tooth from anterior (B), middle (C) to posterior (D) neuropodia; E, dorsal bilimbate capillary from setiger 5; F, major spine from se tiger 5; G, bilimbate capillary companion seta from setiger H, notopodium from posterior segment in dorsal view; I, posterior end in dorsal view. Scale: 1 = 200 #m (Figures 28A, 281); 2 = 100 #m (Figure 28H); 3 = 20 #m (Figures 28B-28G).


Description. Prostomium incised on anterior margin, with caruncle extending to setiger 3; without occipital tentacle or eyes. Setiger 1 well developed, biramous, with long notopodial postsetal lamella. Posterior notopodia with dense packets of fine needles; these needles glisten by reflected light. Major spines of setiger 5 simple, falcate, lacking accessory structures. Branchiae from setiger 8. Pygidium disclike, entire or usuually divided into 3 lobes.

Remarks. These specimens agree well with those recently described from Australia [Blake and Kudenov, 1978] and recently seen by this author from European waters.

Distribution. Argentina and Uruguay; Australia; New Zealand; Europe; Ceylon; Sumatra; Japan. Intertidal to 170 m.

Polydora socialis (Schmarda, 1861)

Leucodore socialis Schmarda, 1861, pp. 64-65, figs. a-c, pI. 26, figs. 209, 209a.

Polydora socialis: Hartmann-Schroder, 1962b, pp. 137-138, figs. 167-168; 1965, pp. 209-

211, figs. 200-203.--Blake, 1971, pp. 20-23, figs. 13-14 [synonymy]; 1979b, pp. 607-609 [synonymy].--Carrasco, 1974, pp. 194-196, figs. 27-32.--Light, 1977, p. 71; 1978, pp. 179-181, fig. 180.

Material examined. Ecuador, Bahia de Santa Elena, Anton Bruun Sta. 6669, 8-10 m (3, USNM 59917, 60561); Sta. 6670, 8-9 m (1, USNM 60562); 6671, 8-9 m (4, USNM 60563).--Peru, Callao, Isla San Lorenzo, Anton Bruun Sta. 65211, 2-6 m (1, USNM 60559); island near Pucusano, S of Callao, 0-5 m (13, USNM 60560).--Chile, off Valparaiso, Eltanin Sta. 753, 192 m (1, USNM 60558).--Juan Fernandez Islands, Anton Bruun Sta. 65239, 2-8 m (2, USNM 60550).--Argentina, IBM Sta. GII-12 (2, JAB); Mar del Plata, November 1970, collected by J. M. Orensanz (1, USNM 67537); Puerto Ignacio White, collected by Valentinuzzi (2, USNM 67538); Bahia San BIas, NE of Isla Jabali, April 20, 1970, collected by J. M. Orensanz (2, JAB); Golfo San Matias, IBM Sta. SAO 111-1041, off Lte. Lobos, midintertidal, collected by J. M. Orensanz (10, USNM 67541); IBM Sta. SAO 111-1113, Caleta de los Loros, February 1972 (12, USNM 67540); Golfo San Jose, Isla de las Faijacon, sandy beach, intertidal, January 18, 1971 (1, JAB).

Remarks. Polydora socialis is widespread and very well known. It is closely related to!. flava Claparede in having notosetae on setiger 1, an incised prostomium, branchiae from setiger 8, and hooded hooks without a constriction or manubrium on the shaft. Polydora socialis differs from!. flava in having an internal gizzard present and visible through the body wall at about se tigers 18-19. Polydora socialis has only capil-

laries in posterior notopodia instead of specialized bundles of needles and the dorsal surface of the body frequently bears a distinct pattern of black pigment spots. Detailed descriptions of !. socialis and complete synonymies are provided by Blake [1971 1979b].

Distribution. East, west and gulf coasts of North America; east and west coasts of South America; Australia; New Zealand. Intertidal to about 200 m.

Polydora sp. A

Material examined. Ecuador, Bah{a de Santa Elena, Anton Bruun Sta. 6670, 8-9 m (3, USNM 60567).

Remarks. These specimens are anterior fragments resembling Polydora convexa Blake and Woodwick [1972] and P. barbilla Blake [1981a], in having a distinct collar on the convex side of the major spines of se tiger 5. Both species are known from Mexico and may be expected to occur in Ecuador. Since the form of the posterior notopodial setae are diagnostic, their absence precludes specific identification.

Distribution. Ecuador.

Genus Pseudopolydora Czerniavsky, 1881 Pseudopolydora primigenia, n. sp.

Figs. 29-30

Material examined. Ecuador, Bahia de Santa Elena, Anton Bruun Sta. 6669, 8-9 m (holotype, 3 paratypes, USNM 59924-5).

Description. A small species, holotype complete, 3.2 mm long and 0.2 mm wide for 26 setigers; largest paratype incomplete, 3.8 mm long and 0.35 mm wide for 30 setigers. Color in alcohol: light tan to opaque white; holotype and smaller paratypes with small black pigment markings on dorsal surface; largest paratype without pigment. Two paratypes with oocytes in coelom (Figure 29D).

Prostomium incised on anterior margin, forming 2 rounded lobes (Figures 29A-29B); caruncle continuing posteriorly to end of setiger 1; without occipital tentacle; nuchal cilia forming 2 lateral diagonal rows from termination of caruncle to bases of notopodial lamellae of setiger 1, resulting in V-shaped pattern (Figure 29A); 2 eyes present. Peristomium narrow, longer than wide; palps short, with length up to 5-6 anterior setigers; with thickened bases, tapering distally.

Se tiger 1 well developed, with fingerlike notopodial and neuropodial lamellae and fascicles of long capillary notoand neurosetae. Setigers 2-10 with short fingerlike notopodial lamellae; subsequent setigers with shorter, less distinct lamellae, fused basally with broad membranous branchiae by


Fig. 29. Pseudopolydora primigenia n. sp. (paratypes, USNM 59925). A, B, anterior ends of female and male in dorsal views; C, posterior end of female in dorsal view; D, oocytes from female.

265


\ \

/ f ~ ~ ,

I

/ \

) ~

50~m

I A B C D E F G

20~m

Fig. 30. Pseudpolydora primigenia n. sp. (holotype, USNM 59924). A, lateral view of right setiger 5 showing positions and form of setal types; B, modified spine from same; C, enlarged neuroseta from same; D, enlarged notoseta from same; E, smaller seta from posterior row of same; F, granulated neuroseta fom posterior row of se tiger 3; G, hooded hook in lateral view.

setiger 11. Neuropodial lamellae of setigers 2-7 short, fingerlike, similar to notopodia; lacking on subsequent setigers.

Notosetae and neurosetae of se tigers 1-3 and 7 similar in form and arranged in 2 rows with capillaries of anterior row shorter and thicker than those of posterior row; posterior row of neurosetae of se tigers 3 and 7 and notosetae of setiger 7 with distinct granulations on sheath (Figure 30F), rest of capillaries of those setigers lacking granulations, or with very fine granulations on

shaft. Notosetae and neurosetae of setigers 4-6 enlarged and modified to varying degrees, with those of setiger 5 being most modified, especially in holotype and smallest paratype; neurosetae of se tiger 5 with anterior and posterior rows of capillaries formed into closed ring, anterior row composed of enlarged and thickened capillaries having distinct ribbing visible along one edge (Figure 30C); posterior row of capillaries less modified, smaller, with distinct granulations present on shaft (Figure 30E); notosetae of setigers


4-6 similar to neurosetae, with anterior row of short, thickened ribbed setae (Figure 30D) and posterior row of granulated capillaries with additional group of 2-4 long, thin capillaries located dorsally (Figure 30A); setiger 5 of holotype and 1 paratype with 4 pointed spines instead of thickened capillaries (Figures 30A-30B); spines with narrow transverse ridge below pointed tip; 2 larger paratypes with thickened capillaries instead of spines; notosetae of setigers 8 and subsequent setigers all capillaries, but fewer in number and arranged irregularly. Neuropodial bidentate hooded hooks replacing capillaries from setiger 8, numbering 4-5 per ramus in smaller specimens and 6-8 in larger specimens; hooks with distinct manubrium on shaft; with apical tooth closely applied to main fang (Figure 30G).

Branchiae from se tiger 11, each broad, basally fused to notopodial lamellae and continuing through middle body segments (Figures 29A-29B); holotype and 1 paratype with extra pair of cirriform branchiae on se tiger 2 (Figure 29B). Dorsal transverse row of cilia between notopodia and branchiae where present.

Pygidium with 4 separate lobes, dorsal pair smaller and erect, with bacillary glands (Figure 29C).

Remarks. Pseudopolydora primigenia differs from other species of Pseudopolydora in having a well-developed first setiger with notosetae, modified notosetae and neurosetae on setigers 4-6, branchiae from setiger 11 and sometimes with an extra pair of branchiae on setiger 2. This latter feature is known to occur on males of Pygospio elegans and P. californica [Light, 1978] and may also be a sexual character on Pseudopolydora primigenia. The 2 specimens having the extra branchiae had dense reflective granules in the coelom, possibly representing sperm packets. The 2 specimens lacking extra branchiae contained oocytes. The occurence of this type of sexual dimorphism in a species of Pseudopolydora indicates a close relationship to Pygospio. Such a generic similarity between postlarval forms has already been suggested [Blake, 1969]. The absence of modified spines in setiger 5 in 2 of the paratypes is difficult to explain. Since these specimens are the largest, it is possible that the presence of the spines is a juvenile character in Pseudopolydora primigenia. Replacement of 1 type of major spine by another type is well documented during development of species of Polydora [Blake, 1969], but there is no prior record for a complete loss of spines during development. It will be necessary to obtain additional specimens before commenting further on this phenomenon.

Etymology. Primigenius, Latin for first of its kind. The name is suggested by close affinities this new species of Pseudopolydora exhibits with the genu~spio.

Distribution. Ecuador, 8-10 m.

Discussion

The Spionidae are one of the dominant polychaete families in the southern oceans with numerous endemic species described from Australia [Blake and Kudenov, 1978], South Africa [Day, 1967], New Zealand [Rainer, 1973; Read, 1975; J. A. Blake, manuscript in preparation, 1982], and South America (this paper). Spionids are poorly known from the Indian Ocean, but preliminary studies suggest that new species await description (J. A. Blake, unpublished data, 1980). Despite the diversity of spionid polychaetes throughout the southern oceans, they are surprisingly poorly represented in Antarctic seas.

In an effort to understand differences between spionid faunas of Antarctic seas and adjacent waters of South America, the geographic categories of Knox [1977] are employed. According to Knox [1977] there is (1) an Antarctic area which includes the entire Antarctic coastline, the islands of the Scotia Arc, Kerguelen Island and Heard Island; (2) a Subantarctic Island group; and (3) the Magellanic Region including southwestern Chile, the Straits of Magellan, Tierra del Fuego, and the Falkland Islands. In the present discussion, the Magellanic Region is incorporated into a consideration of the South American continent as a whole. The Juan Fernandez Islands, off Chile, and the Galapagos Islands, off Ecuador, are treated separately here as an Eastern Pacific Island category.

It is beyond the scope of the present paper to provide detailed zoogeographical analyses of spionid distributions and patterns of evolution and dispersal. Such analyses will follow completion of other studies in progress on southern ocean spionids.

Antarctic Area

The diverse polychaete fauna of Antarctica has been summarized by Hartman [1964, 1966]. From those works and more recent studies by Hartman [1967, 1978], Averincev [1972], and Knox [1977] it appears that the number of Antarctic polychaetes may eventually exceed 800 species. The most diverse families are the Polynoidae (27 genera, 68 species), Syllidae (20 genera, 50 species), and Terebellidae (18 genera, 29 species). Overall, 57% of the Antarctic polychaete species are endemic. In contrast, the Spionidae of Antarctica are represented by only 10 genera and 16 species (Table 1), of


TABLE 1. Distribution of Spionidae in South American Waters and Antarctic Seas

Regions

Western East Eastern Antarctic Subantarctic Magellanic South Pacific South

Species Seas Islands Region America Islands America

Scolelepis aitutaki A,U S. chilensis C S. eltaninae RS,BS S. quinguedentata C S. gaucha B,U Dispio brachychaeta A D. uncinata A Aonides paucibranchiata A Scolecolepides uncinatus A Malacoceros indicus C Rhynchospio glutaea SG,KI MI TF,SI,FI C A Laonice antarcticae AP ,~BS, SG, SS, - FI A,U

SSI, WS 1- weddellia AP,RS,SG,SO,- C,DP,SM

WS L. cf. cirrata BS Paraprionospio pinnata SG FI C JFI A,U Prionospio orensanzi A P. cL ehlersi KG,PI DP C P. Eygmaea JFI A P. steenstrupi JFI P. Eeruana C,P P. cf. cirrifera E GI P. pata~onica SM,C C Spiophanes bombyx KI FI C JFI S. kroeyeri RS,SG FI C S. soederstroemi C A,U S. tcherniai AC,AP,BC,BS,-

SO,SS,WS,KI Microspio hartmanae A M. paradoxa GI M. minuta C GI M. iiiCiOreI" KI,SS SEio quadrisetosa A Pygospiopsis dubia AP ,SG Boccardia anoEhthalma E B: ch11ensis MI FI,TF C,P B. natrix SG Cl SM,FI A B. polybranchia KI CI,MI TF,SI,SM C,P,E A,B B. tricusEa E Boccardiella ligerica A,u B. occi,Eitalis A Carazziella carrascoi C C. Eata~onica A Amphi,Eolzdora abranchiata A Polydora colonia A P. cirro sa E P. ligni A P. cH ia ta SS FI A,U P. websteri P,E P. bioccipitalis C P. rickettsi C P. ecuadoriana E


TABLE 1. (cont inued)

Regions

Species Antarctic

Seas Subantarctic

Islands Magellanic

Region

Western South

America

East Pacific Islands

Eastern South

America

P. armata

P. antonbruunae P. magellanica P. giardi P. flaVa P. ~lis Pseudopolydora primigenia

SS MI DP,C,FI, SI, TF

C

E

P

C,E

C,P,E E

JFI

JFI A A

Code: Antarctic seas: AC, Adelie Coast; AP, Antarctic Peninsula; BC, Budd-Knox Coasts; BS, Bransfield Strait; KG, King George V Coast; KI, Kerguelen Islands; PI, Peter I Island; RS, Ross Sea; SG, South Georgia; SO, South Orkney Islands; SS, South Shetland Islands; WS, Weddell Sea. Subantarctic islands: Cl, Campbell Island; MI, Macquarie Island. Magellanic Region: C, Chile; DP, Drake Passage; FI, Falkland Islands; SI, Staten Island; SM, Straits of Magellan; TF, Tierra del Fuego. Western South America: C, Chile; P, Peru; E, Ecuador. Eastern Pacific Islands: GI, Galapagos Islands; JFI, Juan Fernandez Islands. Eastern South America: A, Argentina; U, Uruguay; B, Brazil.

which only 4 species are endemic: Scolelepis eltaninae, Spiophanes tcherniai, Microspio moorei, and Pygospiopsis~. Of these, only S'. tcherniai is relatively common and widespread around Antarctica with the other 3 species being known from only 2 localities each. Laonice antarcticae and L. weddellia are the only nonendemic spionids to be widely distributed and common in Antarctica. Laonice antarcticae is known elsewhere from deep waters off eastern North America and eastern South America, while L. weddellia occurs to the north in the Magellanic region. The remaining 10 species listed in Table 1 are more common in northern localities. Two of these species, Boccardia polybranchia and Rhynchospio glutaea, are abundant on some subantarctic islands and in the Magellanic area.

Among the endemic species, Scolelepis eltaninae, Spiophanes tcherniai, and ~. moorei are closely related to congeners in southern hemispheric localities north of the Antarctic convergence. Pygospiopsis ~ is the only endemic genus in Antarctic seas.

According to Knox [1977J, the Antarctic fauna has been isolated from northern continental land masses for over 20 m.y. This long period of isolation has been maintained by southern ocean circulation patterns and temperature-salinity barriers which have effectively blocked intrusions by northern shallow water faunas and allowed a prolonged period of uninterrupted evolution. It is noteworthy in this context to observe that

no species of either the Polydora complex or Prionospio complex are endemic to Antarctic seas. Both generic groups have radiated into numerous species on other shorelines. Their absence in Antarctica suggests that current success in other oceans is the result of adaptive radiation during the 20 m.y. isolation of Antarctica. While Antarctica appears to be effectively isolated from migration from other oceans by temperature and salinity barriers in shallower waters, Knox [1977J suggests that deepsea species are already preadapted to low temperatures and, if not depth limited, should be capable of invading the Antarctic shelf. Averincev [1970J has found evidence supporting such a hypothesis with errant polychaetes, and Hessler et al. [1979J suggest a similar shallow water Antarctic emergence for deep-sea isopods. Among the spionids, Laonice antarcticae may be an example of a species which entered Antarctica via such a deep-sea route. The species is known from the deep Atlantic [Hartman, 1965; Hartman and Fauchald, 1971J. A pathway from the Atlantic to Antarctica via the Scotia Arc might be envisioned for this species. A similar route might be proposed for L. wed~, which occurs in both Antarctica and the Magellanic region. The latter species, however,is not as yet known from the Atlantic.

Subantarctic Islands

Five species of Spionidae occur in various subantarctic islands groups outside of the


Scotia Arc (Table 1). No genera or species are endemic to these islands. Auckland Island polychaetes are believed to have affinities with New Zealand [Knox, 1958], but to date there is no evidence with the Spionidae. The 5 species are more widely distributed in the southern hemisphere, with Rhynchospio glutaea and Boccardia polybranchia also being known from the northern hemisphere.

South America

A total of 50 species of Spionidae occur in the seas around South America. Eighteen species or 36% are endemic. It is beyond the scope of this paper to analyze the zoogeography of this fauna.

Magellanic Region. This area includes the southern tip of South America and the Falkland Islands. Fourteen species of Spionidae are recorded from this area (Table 1), with Polydora magellanica being the only endemic species present. The records of Laonice weddellia are the northernmost for a predominantly Antarctic species. Prionspio patagonica was described from the Straits of Magellan, but the species occurs elsewhere along the Chilean coast. Boccardia chilensis is known from Australia, New Zealand, MacQuarie Island, and along the west coast of South America. The remaining la species are all common elsewhere in South America and are recorded from a few Antarctic localities.

Western South America. Twenty-eight species occur from Chile to Ecuador, including 9 which are endemic. Boccardia polybranchia, Polydora giardi, and~. socialis occur along the entire western coast and are well known in other geographic areas as well. A distinct faunal break occurs between Chile and Peru/Ecuador. Eight species are limited to Peru/Ecuador, 1 is only known from Peru, and 14 are limited to Chile. Only 2 species occur in Peru and Chile. The basis for this separation is most certainly related to the oceanic current patterns along the western coast of South America. A northern branch of the cold West Wind Drift, the Peru Current, carries cold subantarctic water northward along the west coast. This water grad-

ually warms and eventually meets a warm south flowing current and turns westward at about 60 S to merge with the South Equatorial Current [Ekman, 1953]. The southern limit of the tropical-subtropical water is on the Peruvian coast and is farther north in this regard than anywhere else in the world [Ekman, 1953J.

Eastern South America. Twenty-six species are recorded from Argentina to southern Brazil (Table 1). The spionid fauna of most of Brazil remains largely unknown. Nine of the endemic species occur in Argentina and Uruguay. Four species, Rhynchospio glutaea, Paraprionospio pinnata, Spiophanes bombyx, and S. soederstroemi, also occur on the west coast. For the most part, however, the 2 faunas are very different in species composition. Scolelepis aitutaki appears to be distributed across the Pacific Ocean and eastern South America but has not been found on the west coast. Ten species are either Atlantic in distribution or well known from North America and elsewhere. Ekman [1953] noted that the fauna between Patagonia and Rio de Janeiro is poorly known but speculated that the greatest change in faunal composition possibly occurs at the mouth of the Rio de la Plata, where the water temperature rises abruptly. In this regard, he is probably correct, as studies of Brazilian spionids indicate a totally different species composition to that of Argentina and Uruguay (J. A. Blake, unpublished data, 1982). A preliminary analysis of the Brazilian spionids suggests a close affinity with the Caribbean Sea and Gulf of Mexico.

Eastern Pacific Islands

Five species of Spionidae are recorded from the Juan Fernandez Islands. All are widely distributed elsewhere. Three species were reported from the Galapagos Islands, of which Microspio paradoxa is endemic. An undescribed species of Boccardia from the Galapagos is in the author's collection. The record of Microspio minuta is the only known locality for that species outside of Chile.


Appendix

TABLE Al. List of Oceanographic Stations Taken by the Instituto de Biologia de Mar del Plata (IBM)

Station Latitude Longitude Depth, Bottom Area S W m Type

N-242 off Uruguay 63 sand Uruguay N-244 36024.5 ' 53051. 7' 128 Uruguay N-248 35036' 52 043 ' 170 muddy sand Uruguay N-250 34051 ' 52 035 ' 83 Uruguay N-258 34034' 53014 ' 50 Uruguay N-263 34051' 54004 ' 39 Uruguay N-I054 35056.5 ' 54015.7' 58-65 Argentina N-I055 36016' 54 001.5 ' 92-96 Argentina N-I056 36 030.5' 53055' 155-192 Argentina N-I058 35 057' 53032 ' 150-156 muddy sand Argentina N-I059 35 025.9' 53027.9' 72-80 Argentina N-I064 34 024.5' 53 027.2' 20-26 Uruguay N-I066 34 029.2 ' 52 020 ' 72-86 muddy sand Uruguay N-I070 33 050' 53 003 ' 20-23 Uruguay N-I073 35010.5' 52042.5 ' 115-117 muddy sand Uruguay N-I075 35036.5 ' 53 032 ' 68 muddy sand Argentina Mej-5 37040 ' 56028 ' 55 hard bottom Argentina Mej-ll 37 030' 56033' 47 Argentina Mej-12 37030' 56041' 24 Argentina Mej-15 37010' 56 015 ' 28 Argentina Mej-18 37016' 56 009 ' 48 Argentina Mej-22 37 027' 56029' 40 Argentina Mej-27 37033' 56 024 ' 51 Argentina G II-3 37056.6' 57 028.1' 16.5 muddy sand Argentina G II-8 38001' 57026.7' 20 muddy sand Argentina G II-12 38003.9' 57022 ' sand and shells Argentina G II-13 38004.3' 57027.2' 21 sand and stones Argentina G II-14 38003.5 ' 57030.9 ' muddy sand Argentina SAO I-A 4 km off 15 sand Argentina

Las Grutas SAO I-C off Las Grutas 15 sand Argentina SAO I-I 40055 ' 64054 ' 18 sand Argentina SAO 1-5 40054' 64031' 20-24 sand Argentina SAO 1-7 40057.5' 64020.5 ' 18-24 stones Argentina SAO 1-15 41001' 64 015 ' 33-36 mud and sand Argentina SAO 1-24 41 002' 65 008.5 ' 15 shell bottom Argentina SAO 1-49 41016 ' 65 004 ' 36 sand and mud Argentina SAO I-52 41005.5 ' 65007.5' 29 sand Argentina SAO I-53 41000' 65006' 38 sand and shells Argentina SAO II-54 40054 ' 65 002' 13-15 fine sand Argentina SAO IlI-I113 Golfo San sand Argentina

Madas SAO V-31 off Bajo 21 sand Argentina

Oliviera, Golfo San Madas

272

Station Latitude

S

H-15 37037' A II 60-282 off the Rio

de la Plata AIl 60-284 off the Rio

de la Plata Comp-I 38000' Comp-V-79 37 038' WH-389 38056'

BIOLOGY OF THE ANTARCTIC SEAS XIV

TABLE Al. (continued)

Longitude W

57027.5' 56 045 ' 55 0 31'

Depth, m

70-80 200-250

200-250

18-30 31

200

Bottom Type

sand sand

Area

Argentina Argentina

Argentina

Argentina Argentina Argentina

Cruise and ship identification codes: N: Oceanographic cruises of the Soviet research vessel, A. Knipovich, along the coasts of Argentina and Uruguay (1966 and 1968); V. Scarabino, collector. Mej: Fisheries research cruise of the Mejil16n I, Penchaszdeh, collector. G II: Oceanographic cruise of the Goyena II; S. R. Olivier, R. Bastida, and R. Capitoli, collectors. SAO I-V: Exploratory cr~in the Golfo San Matias; cruises I-II under supervision of IBM and FAO fisheries of Argentina; cruises Ill-IV included cooperative investigations of the federal council of the Province Rio Negro and the IBM; cruise V was fisheries investigations of selected areas in the Golfo San Matias. H: Oceanographic cruise of the American research vessel, R/V Hero (1971); R. Bastida, collector. A II 60: Cruise 60 of the R/V Atlantis II of the Woods Hole Oceanographic Institution (1971) involving studies off the mouth of the Rio de la Plata; R. Bastida, collector. Comp I-V: Oceanographic cruises of the IBM in cooperation with the Argentine navy; R. Olivier and Salanouve, collectors. WH: Fisheries cruise of the Walther Herwig (1966).

TABLE A2. List of Stations Taken by the R/V Anton Bruun (November 1965 to May 1966) Western South America

Station Date Latitude Longitude Depth, Gear Area S W m Type

65211 Nov. 26 2-6 SCUBA Peru, Callao, Isla San Lorenzo Naval Base*

65215 Nov. 29 0-5 SCUBA Peru, island near Pucusana, S of Callao

65235 Dec. 11 33 038.20' 78048.50' 3-11 SCUBA Juan Fernandez Islands, Mas a Tierra, Cumberland Bay

65239 Dec. 11 33 038.20' 78049' 2-8 SCUBA Juan Fernandez Islands, Mas a Tierra, Cumberland Bay

65240 Dec. 12 33 037.18' 78050.20 ' 24-27 SCUBA Juan Fernandez Islands, shell fragments

65243 Dec. 12 33 037' 78050.50' 0-9 SCUBA Juan Fernandez Islands

65253 Dec. 14 33038.12 ' 78056.36' 80-125 Otter trawl Juan Fernandez Islands, off shore Mas a Tierra

65256 Dec. 15 9-12 SCUBA Juan Fernandez Islands, West Carvajal Bay


TABLE A2. (continued)

Latitude Longitude Depth, Gear Station Date S W m Type Area

65260 Dec. 16 26 SCUBA Juan Fernandez Islands, SE of Bacalao Point

MV65-IV-47 Dec. 12 33037.5' 78°49.7' Otter trawl Juan Fernandez Islands

MV65-IV-55 Dec. 13 33 036.4' 78°47.8' Otter trawl Juan Fernandez 33 037.3' 78°48.0' Islands

MV65-IV-57 Dec. 14 33039' 78059' Otter trawl Juan Fernandez Islands

MV65-IV-58 Dec. 14 33 038.2 ' 78°56.6' Otter trawl Juan Fernandez Islands

MV65-IV-63 Dec. 15 33 041.2 ' 78°57' Otter trawl Juan Fernandez 33 040.7' 78°51.8 ' Islands

Drab 126 Dec. 11 33°38' 78°49' 0-1 Hand Juan Fernandez Islands

Drab 130 Dec. 12 33°38 78°49' 46 Snapper Juan Fernandez Islands

Drab 134 Dec. 13 SCUBA Juan" Fernandez Islands, Chile Bay, behind Punta Seroeste

Drab 135 Dec. 14 33°34.3' 78°54.9' 160-180 Otter trawl Juan Fernandez 33°35.5' 78°55.5' Islands

6669 May 8 02°12.20' 80°52.10' 8-9 SCUBA Ecuador, Bah{a de Santa Elena, 200 m offshore

6670 May 8 02°11.28 ' 80°56.31' 8-9 SCUBA Ecuador, Bah!a de Santa Elena

6671 May 8 02°13 .09' 80°54.38' 0-3 SCUBA Ecuador, Bah!a de Santa Elena

66122 May 21 00°44.36' 90°13.51' 17-18 SCUBA Galapagos Islands

*S. Earle, collector.

TABLE A3. List of stations taken by the USNS Eltanin (1962-1972) from the Southern Oceans

Latitude Depth Gear Cruise Station Date S Longitude m Type Area

3 72 June 24, 1962 31°06.30' 71°48.50'W 805-970 MT off western Chile 5 203 Aug. 31, 1962 35 039' 73008 'w 436 PG off western Chile 5 208 Sept. 11, 1962 37 029 ' 73 055'W 957 CG off Western Chile 5 222 Sept. 27, 1962 53 015' 66051'W 79-80 OT Tierra del Fuego 5 265 Oct. 19, 1962 62059' 67051'W 3691-3693 MT Drake Passage

63 001 ' 67039 'w 5 272 Oct. 21, 1962 64054 ' 68021'W 412 BT off Antarctic

68018'W Peninsula 5 303 Oct. 30, 1962 62 001' 70059W 4077 MT Drake Passage

62006' 70051 'w 6 338 Dec. 3, 1962 53 009' 59037'W 587-595 RD Falkland Islands

53 008' 59035'W 6 344 Dec. 4, 1962 54004' 58046 'w 119 MT south of Falkland

58°45'W Islands Burwood Bank

274

Cruise Station Date

6

6

6 6 6

6 6

6

6

6

6 7

7

7

9

9

9 9

9

9

9

11 11

11

12

12

12 12 12

12

12

14

363 Dec. 7, 1962

408 Dec. 31, 1962

410 Dec. 31, 1962 412 Jan •. 1, 1963 418 Jan. 2, 1963

419 Jan. 3, 1963 428 Jan. 5,.1963

432 Jan. 7, 1963

437 Jan. 9, 1963

439 Jan. 9, 1963

453 Jan. 21, 1963 463 Feb. 11, 1963

490 Feb. 20, 1963

500 Feb. 21, 1963

671 Aug. 23, 1963

748 Sept. 26, 1963

749 Sept. 26, 1963 750 Sept. 26, 1963

752 Sept. 26, 1963

753 Sept. 26, 1963

758 Sept. 27, 1963

958 Feb. 5, 1964 960 Feb. 6, 1964

962 Feb. 6, 1964

993 March 13, 1964

997 March 14, 1964

1001 March 15, 1964 1003 March 15, 1964 1079 April 13, 1964

1082 April 14, 1964

1089 April 17, 1964

1209 Aug. 11, 1964

BIOLOGY OF THE ANTARCTIC SEAS XIV


Latitude S

57 0 09' 580 00 ' 61016'

61 016 ' 62 006' 62 0 39' 62 0 40' 62 0 14' 62041 ' 62 0 39' 62 052' 62 0 55 ' 62 0 50' 62 0 51' 63 0 51' 63050 ' 54 0 27' 54 0 10' 54 0 13' 62 006'

62 006' 62 0 07' 54041'

54 0 38' 33 0 00'

33 0 00' 33 0 01'

33 0 14 '

33 0 16'

33 0 15 '

52 056' 52 0 40 ' 52036' 53056' 53 055 ' 61025 '

61 0 44 ' 61 046' 62039 ' 62 0 41 61 026 ' 61 0 24' 60 050 ' 60 0 52' 60 0 47' 60 0 48 ' 58018 ' 58 0 19'

Longitude

58058 'w 58050 'w 56 0 11'W

56009 'w 56 000'W 56010'W 56008'W 58017'W 57051'W 57046'W 59027'W 59015 'w 60040'W 60035'W 62038'W 62035 'w 66012'W 44049'W 44 046'W 45 008'W 45 010'W 45 012'W 45 0 13 'w 38038'W

38031 'w 71 0 51 'w 71 0 52 'w 71052'W 71 0 53 'w

71 047'W

71 0 47 'w

71 049'W

75 000'W 74°58'W 74 055'W 71015'W 71012' 56030'W

55 056'W 55054'W 54046'W 54 043 'w 41 055'W

42 055'W 42 056'W 53 030'W 53 031'W 160003'W

Depth ID

Gear Type

PC

223-225 MT

220-240 BT 1180 MT 311-426 BT

509-549 RD 622-1120 BT

935-884 BT

267-311 BT

128-165 BT

31 PG 3403-3484 PG

485 RD

489-490 PG

220-320 BT

1025 PG

1007 PG 624 PG

209 PG

192 PG

156 PG

92-101 64

256-320

300

769

238 210-230 593-598

298-302

641

3697-3784

BT BT

BT

BT

BT

CG BT BT

BT

BT

MT

Area

Falkland Islands

South Shetland Islands

Bransfield Strait Bransfield Strait South Shetland

Islands Bransfield Strait Bransfield Strait

Bransfield Strait


Antarctic Peninsula

Tierra del Fuego shag rocks, near

South Georgia south of South

Orkney Islands south of South

Orkney Islands South Georgia

off western Chile

off western Chile off Valparaiso,

Chile

off Valparaiso, Chile



southwestern Chile southwestern Chile

Strait of Magellan



Bransfield Strait Bransfield Strait SE of South Orkney

Islands E of South Orkney

Islands NE of South

Shetland Islands South Pacific

Ocean, Albatross Cordillera



Cruise Station Date Latitude

S

14 1248 Aug. 25, 1964 59 057'

16 1418 Feb. 10, 1965 54 0 32 ' 45 0 37' 45 035 33 003 '

16 1431 Feb. 23, 1965

21 194 Nov. 23, 1965

21 204 Nov. 26, 1965 33°44'

21 205 Nov. 26, 1965 33 043'

22

22

25 25

27

27

27

27

32

32

32

32

32

32 32

32

32 32

32

32

32

32

1536 March, 1966 54 031' 54 0 29'

1596 March 14, 1966 54039'

300 Sept. 22, 1966 362 Nov. 11, 1966 56°10'

56°13' 1870 Jan. 14, 1967 71°17'

71 °16' 1885 Jan. 16, 1967 74°30'

74°32' 1930 Jan. 28, 1967 74°19'

74 0 20' 1944 Feb. 2, 1967 67°23'

1989

1996

2005

2012

2016

2020 2029

2031

2035 2050

2065

2073

2126

2143

Jan. 1, 1968

Jan. 10, 1968

Jan. 12, 1968

Jan. 13, 1968

Jan. 14, 1968

Jan. 15, 1968 Jan. 17, 1968

Jan. 17, 1968

Jan. 18, 1968 Jan. 22, 1968

Jan. 26, 1968

Jan 29, 1968

Feb. 13, 1968

Feb. 26, 1968

67° 24'

73°59' 73°58' 73°58' 73° 5 9' 74°06' 75°00'

74°39' 74°59' 74°32' n O Ol' n003' 78°23'

Longitude

136°57'W 136040'W

159 002'W 170058'E 170059'E 71 047'W

80 043'W

39°19 'w 39022'W 57009'W

156°09'W 156012'W 171 033'E 171029'E 170°10 'E 170012'E 176°39'W 176°34'W 180000'W 189058'W

169 048'E 169045'E

172 008'E 172°09'E

176°54 'E 176°50'E 170051'E 170058'E 176°11'E 176016'E 179°11' W 176°42'E 176°40 'E 172018'E 172°12 'E 168017'E 168038'E 168023'E 173°06'W 173002'W 163026'W

178°35 'E 178°34'E

Depth m

3386-3495

86-101 51

137-141

183-283

128-123

659-686

124

o 200

659-714

311-328

831-836

516-595

589-594

348-352

864-870

589-608

581-586

256 335-338

535

876 909-923

473-475

503

143

2010-2100

Gear Type Area

MT South Pacific Ocean, Albatross

Cordillera BT MacQuarie Island OT New Zealand, east

of South Island PG off Valparaiso,

Chile BT W of Juan

Fernandez Islands BT W of Juan

Fernandez Islands W of South BT

BT Georgia

S of Falkland Islands, Burwood Bank

hand Montemar, Chile IKMT South Pacific

Ocean BT Ross Sea, off

Cape Adare BT Ross Sea

BT Ross Sea

BT South Pacific Ocean, near Scott Island

BT

BT

BT

BT

BT

CG BT

BT

CG BT

BT

CG

CG

BT

South Pacific Ocean, SW Campbell Island

Ross Sea, off Moubray Bay, Victoria Land

Ross Sea

Ross Sea

Ross Sea

Ross Sea Ross Sea

Ross Sea

Ross Sea Ross Sea, N of

Ross Island Ross Sea, off Ross

Ice Shelf Ross Sea,

Scott Coast Ross Sea,

Robertson Bay South Pacific

Ocean, near Antipodes Island

275



La titude Cruise Station Date S Longitude

Depth m

38 7GR7 March 31, 1969 64°13.05' 150006.67'E

51 5761 Feb. 8, 1972 76°01.5' 76°01.6'

179049.9'E 179053.5'E

3543

388-399

Gear Type Area

CG South Pacific Ocean, off George V Coast, Antarctica

MT Ross Sea, Pennell Bank

*Code for gear types: BT, Blake trawl; CG, Campbell (Camera) grab; midwater trawl; MT, Menzies trawl; OT, Otter trawl; PC, Phleger corer; RD, rock dredge.

IKMT, Isaacs-Kidd PG, Petersen grab;

TABLE A4. List of Stations Taken by the R/V Hero (1971-1973) From South America and the Antarctic Peninsula-(Cruises 712, 721, 731)

Station

656

658

663

664

674 71-2-2

71-2-6

71-2-8

71-2-14

71-2-16 71-2-18 71-2-19 71-2-21 71-2-30 71-2-31 71-2-32 71-2-33 71-2-37 71-2-39 71-2-40 71-2-41

726

767

Date

April 26, 1971

April 28, 1971

May 9, 1971

May 10, 1971

May 20, 1971 April 17, 1971

April 21, 1971

April 23, 1971

April 25, 1971

April 27, 1971 May 2, 1971 May 3, 1971 May 4, 1971 May 10, 1971 May 12, 1971 May 13, 1971 May 13, 1971 May 17, 1971 May 20, 1971 May 21, 1971 May 22, 1971

Dec. 26, 1971

Jan. 4, 1972

Latitude S

56 0 48'

56046.7' 54 047.5' 54°46.8 '

54°46.1'

54045.4' 52 025'

53°17'

54 047.8'

54 048.2'

54046.2 ' 54050 ' 54049.2 ' 54°47.3' 54046.4' 54045 ' 54°44.25 ' 54043.9' 54039.25 ' 54045.45' 54045.45 ' 54 043.05'

62 019.3' 62 019.2' 64045'

Longitude W

64042'

64°42.7' 64041.5 ' 64004'

63°57.9'

64°09.8' 71°00 '

65 016 '

65014.7'

64042.7' 64°26.8' 64°27.8' 64°18.7' 63°57.7' 63°53.3 63°51.25' 63°52' 64°02.3' 64°10.1' 64°09.55' 64°13.6'

59011.8' 39°11.7' 64005'

Depth, m

18

13-34

31

29

30 o

0-1

o

o

o 0-1 o o o o o o o 0-1 o 0-1

64-82

Type Gear*

PG

MT

PG

PG

trawl hand

hand

hand

hand

hand hand hand hand hand hand hand hand hand dip net hand Poison

Sta. BT

SCUBA

Area

Tierra del Fuego

Tierra del Fuego

Tierra del Fuego

Tierra del Fuego

Staten Island Straits of

Magellan Straits of

Magellan Tierra del

Fuego Tierra del

Fuego Staten Island Staten Island Staten Island Staten Island Staten Island Staten Island Staten Island Staten Island Staten Island Staten Island Staten Island Sta ten Island


Antarctic Peninsula, off Anvers Island

BLAKE: SOUTH AMERICAN AND ANTARCTIC SPION IDAE 277


Latitude Longitude Depth, Type Station Date S W m Gear* Area

819 Jan. 17, 1972 62058.4 ' 60041. 9' 30 PG South Shetland Islands, Deception Island

843 Jan. 26, 1972 64047.5' 64007.2' 107 BT Antarctic Peninsula, S of Arthur Harbor

848 Jan. 26, 1972 64 047.4' 64 006.9' 94-165 BT Antarctic Peninsula, S of Arthur Harbor

970 Dec. 7, 1971 64049.04' 63032.8' 102 PG Antarctic Peninsula, Bismarck Straits

983 Dec. 9, 1971 65014.4' 64 015.5' 38 PG Antarctic Peninsula, Grandidier Channel

1060 Dec. 19, 1971 62019.0 ' 59011.4 ' 44 PG South Shetland Islands

1063 Dec. 19, 1971 62019.0' 59 011.4' 44 BT South Shetland Islands

1066 Jan. 26, 1972 64047.4' 64006.8 PG Antarctic Peninsula, outside of Arthur Harbor

1075 Feb. 23, 1972 64047.4' 64 007.2' 91-110 BT Antarctic Peninsula, S of Arthur Harbor

5444 March 29, 1972 64046.45' 64004.90' 34-40 grab Antarctic Peninsula, Arthur Island

1897 March 6, 1973 64046.41' 64004.04' 22 OPG Antarctic Peninsula, Arthur Harbor

1937 March 9, 1973 64 052.20' 63 032.40' 96-133 BT Antarctic 64 052.36' 63 033.22' Peninsula,

Bismarck Strai ts

*Code for gear types: BT, Blake trawl; MT, Menzies trawl; OPG, Orange Peel grab; PG, Peters en grab.

TABLE A5. List of Stations Taken by the Deep Freeze I-IV Expeditions (1956-1959) and International Wedde11 Sea Expedition (1968-1969)

Ship Station Area Latitude

USS Ec;!isto 3 Ross Sea, 71032'S Robertson Bay

4 Ross Sea, SW 71°30'S Robertson Bay, Relay Bay

6 Ross Sea, 73 019' S Robertson Bay

8 Ross Sea, 77 026'S McMurdo Sound

USCGC Glacier 1 McMurdo Sound 77030'S between Cape Royds and Cape Evans

USS Staten 19 Wedde11 Sea 77°37'S Island

USS Staten 20- Wedde11 Sea 77°08'S Island OP-5

USCGC Westwind 7 Wedde11 Sea, 77039' S off Filchener shelf ice, Vahse1 Bay

USCGC Westwind 9 Bransfie1d Strait, 62°24'S South Shetland Islands near Deception Island

USS Atka 24 near Wi1kes 66°15.4' S Station, off Vincennes Bay, between Budd and Knox Coasts, Antarctica

Longitude Date

Deep Freeze I 170018'E Feb. 6, 1956

169032 'E Feb. 7, 1956

169015'E Feb. 12, 1956

169030'E Feb. 18, 1956

Deep Freeze 11 166°04'E Oct. 28, 1946

43015'W Jan. 11, 1957

45 010'W Jan. 21, 1957

Deep Freeze III 44°50'W Jan. 16, 1958

59044'W Jan. 26, 1958

1l0028.40'E Jan. 23, 1958

Method

Orange Peel grab

Orange Peel grab

dredged

dredged

Depth, m

27

400

100

321

400

430

270

225

170

Collector

J. Q. Tierney

W. L. Tress1er

W. H. Litt1ewood

W. H. Litt1ewood

J. Q. Tierney

J. Q. Tierney

L. Wi1son

Deep Freeze IV

USCGC Northwind 8 Ross Sea, 72 016.4'S 170018'E Jan. 12, 1959 135 1. W. Wilson Moubray Bay, off Cape Ha11ett

USS Staten 10 Victoria Land, 71021.30 's 17000S'E Jan. 24, 1959 volcanic 128 R. B. Starr Island Ross Sea area, gravel and

Robertson Bay, hard mud, W of Cape Adare Challenger-

type trawl USS Edisto 20, Weddel1 Sea, 77 040' S 3s030'W Jan. 28, 1959 384

trawl off Vahsel Bay 5

International Weddell Sea EXpedition, 1968-1969 USCGC Glacier 68-1 70 007'S 39038'W Feb. 6, 1968 epibenthic 650 J. S. Rankin

sled USCGC Glacier 68-5 76 000'S SsoOO'W Feb. 9, 1968 Van Veen 400 J. S. Rankin

grab

USCGC Glacier 68-9 73 031' S 60 003'W Feb. 11, 1968 Van Veen 526 J. S. Rankin grab

USCGC Glacier 68-18 72 046'S 42 04S'W Feb. 18, 1968 epibenthic 1664 J. S. Rankin sled

USCGC Glacier 68- Arthur Harbor March 17, 1968 40 J. S. Rankin Pa1mer anchorage,

II Anvers Island, Antarctic Peninsula

USCGC Glacier 69-1 74°28.1'S 30031.7 'w Feb. 24, 1969 anchor 513 J. S. Rankin dredge

USCGC Glacier 69-2 7S031'S 30008'W Feb. 25, 1969 anchor 412 J. S. Rankin dredge

USCGC Glacier 69-4 77005.5 's 3S004'W Feb. 26, 1969 anchor 743 J. S. Rankin dredge

USCGC Glacier 69-6 760S0'S 400SS'W March 1, 1969 anchor 513 J. S. Rankin dredge

USCGC Glacier 69-7 77016' S 42038'W March 1, 1969 anchor 512 J. S. Rankin dredge

USCGC Glacier 69-8 77 036.2 's 42 030'W March 2, 1969 anchor 585 J. S. Rankin dredge

USCGC Glacier 69-10 77 O S0's 4200S'W March 4, 1969 anchor 659 J. S. Rankin dredge

USCGC Glacier 69-22 730 28.4'S 30026.9'W March 13, 1969 anchor 3111 J. S. Rankin dredge

USCGC Glacier 69-23 72°49.6'S 30029.7'W March 14, 1969 anchor 3697 J. S. Rankin dredge

TABLE A6. List of Some Stations From the Lund University Chile Expedition (1948-1949)

Station

M-56

M-60

Area

Peninsula Laqui, Punta Corona, northeastern point

Isla Tenglo, the bay on the south side

Latitude

40047'S

41030'15"S

Longitude

73 053'W

72°58'50"W

Bottom Type

intertidal, flat rocks with small holes and very shallow rock pools

intertidal in sand

Date

Feb. 26 and 28, 1949

March 25 and 29, 1949

M-113 Estrecho de Magallanes, Punta Santa 53 022'S Maria, near Agua Fresca

70057'W intertidal, sand, gravel, and muddy May 2, 1969 clay covered with boulders

M-115 Estrecho de Magallanes, near the 53011'S 70055'W intertidal, gravel and clay mixed May 3, 1949 estuary of Rio los Ciervos, S of

M-ll6 M-121

M-123

M-131

M-133

Punta Arenas Seno Almirantazgo, Caleta Mar!a Bah!a San Vicente, Punta Lilies

W of San Vicente Montemar, north of Valparaiso,

"Estaci6n de Biologia Marina" Iquique, southern part of town

Iquique, the harbor

*Data rro~m 1IraTt:stromandDahC(T951).

> <

...... ro ...... ...... ...... "''1 '" '" '" .......... w w W OP '" N N

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with mud and covered with boulders

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pools intertidal, rocks with oysters

exposed intertidal red rocks with rock pools

intertidal, sheltered rocks and boulders

May 7, 1949 June 9, 1949

Oct. 17-21, 1949

July 1,4, and 6,1949

July 2, 1949

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>: H <:


1972

Banse. 1963

Banse. 1968

Banse. 1968

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1979b Revision of some polydorids (Polychaeta: Spionidae) described and recorded from British Columbia by Edith and Cyril Berkeleley. Proc. bioI. Soc. Wash •• 92(3): 606-617. 3 figs.

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1939

1940

1941

1943

1945

The polychaetous annelids collected by the Presidential Cruise of 1938. Smithson. misc. collns, ~: 1-22, 3 figs. Boccardici proboscidia, a new species of spionid worm from California. J. Wash. Acad. Sci., 30: 382-387. Some contributions to the biology and life history of Spionidae from California. Allan Hancock Pacif. Exped., ~(4): 289-324, pIs. 45-48. Description of Polydora websteri, n. sp. In V. L. Loosanoff and J. B. Engl~ Polydora in oysters suspended in the water. BioI. Bull. Woods Hole, Mass., ~: 69-78, fig. 1. The marine annelids of North Caro-


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Documents

Blake, 1983_Spionidae S America and Antarctica