Polydora and Related Genera Associated with Hennit Crabs from the Indo-West Pacific (Polychaeta: Spionidae), with Descriptions ofTwo New Species and a Second Polydorid Egg Predator ofHennit Crabs1

Jason D. Williams 2

Abstract: Polydora and related genera associated with hermit crabs from shallow subtidal coral reef areas of the Indo-West Pacific are described. Over 2000 hermit crabs were collected from localities in the Philippines and Indonesia be­ tween July 1997 and April 1999. In total, 10 species of polychaetes among five genera (Boccardia, Carazziella, Dipolydora, Polydora, and Tripolydora) were identi­ fied and described. Adult morphology of these species was investigated with light microscopy and scanning electron microscopy. The study includes the de­ scription oftwo new Polydora species, including the second known polydorid egg predator of hermit crabs. Six of the species burrow into calcareous substrata, living in burrows within live or dead gastropod shells or coralline algae attached to shells. Two species were found in mud tubes within crevices of gastropod shells inhabited by hermit crabs. The zoogeography and biodiversity of poly­ dorids from the West Pacific are discussed. The diversity of polydorids from the Philippines is comparable with that of other central Pacific and Indo-West Pacific islands, but it is lower than that in areas of the North and Southwest Pacific; lower diversity probably reflects disparity in sampling efforts between these regions. A key to the Philippine polydorids is provided.

POLYCHAETES ARE COMMON associates of Blake 1996, Martin and Britayev 1998, Wil­ hermit crabs, either living on, within, or bur­ liams 2000). The burrowing behavior of these rowing into occupied gastropod shells (e.g., species has been investigated in substantial Jensen and Bender 1973, Stachowitsch 1980, detail because oftheir effects on commercially Hoberg et a1. 1982, Karlson and Shenk 1983, important mollusk species (see Radashevsky Walker 1995). In particular, Polydora and re­ and Williams 1998). Polydorids have been lated genera (termed polydorids, nine genera considered to be facultative or obligate com­ of the family Spionidae that contain a modi­ mensals of hermit crabs, but recent research fied fifth segment) are known to burrow into indicates that the worms can have negative calcareous substrata including gastropod effects on their hosts (Buckley and Ebersole shells occupied by hermit crabs (Blake and 1994, Williams 2000). The purpose of this Evans 1973, Read 1975, Radashevsky 1993, investigation was to examine the systematics and ecology of polydorids associated with hermit crabs from coral reef areas of the 1 This work was supported by the Lerner-Gray Fund for Marine Research (American Museum of Natural Indo-West Pacific. History), a Libbie Hyman Memorial Scholarship (Society The Spionidae is one of the most conspic­ for Integrative and Comparative Biology), and a Gradu­ uous polychaete families ofIndo-West Pacific ate Fellowship from the University of Rhode Island. islands, in many areas second only to the Manuscript accepted 3 February 2001. family Syllidae in number of recorded genera 2 Department of Biological Sciences, 100 Flagg Road, University of Rhode Island, Kingston, Rhode Island (Knox 1957, Bailey-Brock 1995, Paxton and 02881-0816. Current address: Department of Biology, Chou 2000). Polydorids are common in­ Hofstra University, Hempstead, New York 11549-1140. habitants of coral reef areas, and some species are bioeroders found to burrow in live and Pacific Science, vol. 55, no. 4:429-465 dead corals and mollusk shells (Hartman © 2001 by University of Hawai'i Press 1954, Bailey-Brock 1995). In spite of the All rights reserved amount of work completed in the North and

429 430 PACIFIC SCIENCE· October 2001

Southwest Pacific (see Blake and Kudenov 30' N, 120° 56' E; Puerto Galera: Big and 1978, Radashevsky 1993, Sato-Okoshi 1999, Small Lalaguna Beaches, 13° 30' N, 120° 57' Radashevsky and Hsieh 2000a,b), polydorids E; Puerto Galera: Bayanan and Haligi from central Indo-West Pacific areas such as Beaches, 13° 29' N, 120° 53' E), Aldan (Bor­ the Philippines remain largely unknown. Be­ acay: Diniwid Beach, 11° 60' N, 121° 54' E; fore this investigation only two polydorid Boracay: White Beach, 11° 59' N, 121° 55' E; species had been identified from the Philip­ Boracay: Rocky Beach, 11° 57' N, 121° 56' pines, although a number offaunistic surveys E), Palawan (El Nido: Magbautoc Island, 10° of the polychaetes from the Philippines and 59' N, 119° 36' E; El Nido: Imbaladan Island, surrounding areas have been completed 10° 58' N, 119° 34' E; El Nido: Apulit Island, (Pillai 1965, Williams 2000). Part of the suc­ too 58' N, 119° 37' E), and Cebu (Mactan cess of polydorids is due to the possession of Island, too 18' N, 123° 58' E; Olango Island, long prehensile palps allowing the worms to too 16' N, 124° 03' E) Provinces of the Phil­ suspension and deposit feed or a combina­ ippines (Figure 1) and in Bali (Sanur, 8° 41' S, tion ofboth (Williams and McDermott 1997). 115° IS' E), Indonesia, from June to August Other taxonomically important features of 1997 and January to April 1999. All Philip­ these worms include major spines of the fifth pine specimens were collected by the author. segment, branchial distribution, neuropodial Worms were removed from burrows after hooded hook morphology, and pygidium cracking the gastropod shells with a mortar morphology. and pestle constructed of galvanized steel This paper represents part of a series of pipe (60 mm in diameter). Specimens were studies on the systematics, ecology, and feed­ relaxed in 3% magnesium chloride in sea­ ing biology of polydorids associated with water, fixed in 4% formalin-seawater solution hermit crabs from the Indo-West Pacific. (1 part 39% formalin and 9 parts seawater), The first paper provided the description of rinsed in tap water, and stored in 70% ethyl Polydora robi, an obligate commensal of her­ alcohol. Sketches of live and preserved speci­ mit crabs from the Philippines and Indonesia mens were completed using a compound mi­ (Williams 2000). Polydora robi was docu­ croscope with a drawing tube attachment. mented to ingest the embryos of host hermit These sketches were scanned into a Macin­ crabs, and the reproduction and larval devel­ tosh computer and images were prepared opment of the species have been investigated using the programs Adobe Photoshop and (Williams 2001). The study reported here Adobe Illustrator. provides records of 10 species of polydorids Adult morphology was examined with from the Philippines and Indonesia, nine of SEM. Specimens stored in 70% ethanol were which are previously unrecorded from the dehydrated in an ascending ethanol series Philippines and two are new to science. followed by four changes of 100% ethanol. Morphology of representative species was in­ Material was dehydrated with Peldri II by vestigated with scanning electron microscopy placing the specimens into a 1: 1 mixture of (SEM). Data on the ecology of these species 100% ethanol and Peldri II for 1 hr at 34°C. and feeding biology of the second known egg The specimens were transferred to 100% predator ofhermit crabs are provided. Peldri II for 3 hr and then placed in a cool water bath and allowed to sublime overnight. MATERIALS AND METHODS Dried specimens were mounted on a copper Hermit crabs inhabiting gastropod shells stub, coated with gold-palladium mixture, and were collected intertidally and shallow sub­ viewed in a SEM (lEaL 1200EX). tidally «5 m) in Bataan (Mabayo, 14° 44' N, Specimens were borrowed from the Na­ 120° 16' E; Morong, 14° 41' N, 120° 16' E), tional Museum of Natural History (USNM) Batangas (Anilao, 13° 46' N, 120° 56' E; and the National Museum of Victoria (NMV). Sombrero Island, 13° 42' N, 120° 50' E; Type and representative materials from this Sepoc Point, 13° 41' N, 120° 50' E), Orien­ study are deposited in the American Museum tal Mindoro (Puerto Galera: Coco Beach, 13 ° of Natural History (AMNH) and USNM. 118' 0122' 126'

o(J 0 200km , I

0 ':J 0 0

18' 18' Pacific Ocean

South China Sea

14' 14'

10° 10°

SuIu Sea Mindanao

6° Celebes Sea 118° 122° 126'

FIGURE 1. Map of the Philippines with six provinces indicated in the text: 1, Bataan; 2, Batangas; 3, Oriental Mindoro; 4, Aldan; 5, Palawan; 6, Cebu. 432 PACIFIC SCIENCE· October 2001

KEY TO THE POLYDORID SPECIES FROM THE PHILIPPINES (Figure numbers in parentheses) 1. Segment 5 with one type of major spines 2 1. Segment 5 with two types of major spines 10 2. Branchiae begin on segment 2; tridentate hooded hooks (15J); segment 4 with modi- fied notosetae (15G); segment 5 weakly modified Tripolydora spinosa 2. Branchiae begin posterior to segment 5; bidentate hooded hooks; segment 5 highly modified 3 3. Notosetae present on segment 1 (SA; 7A; 9A); hooded hooks without constriction on shaft (7E; 9C) 4 3. Notosetae absent on segment 1 (llA; 12A; 14A); hooded hooks with constriction on shaft (11 C; 12E) 6 4. Posterior notopodial spines present; spines thick, acicular (5B,C; 6C,G,E) ...... Dipolydora armata 4. Posterior notopodial spines absent 5 5. Major spines ofsegment 5 with small protuberance (8C,D); pygidium with large ventral lobe and 2 smaller dorsal lobes (7B,D; 8B) Dipolydora socialis 5. Major spines of segment 5 with two lateral teeth (9B; lOB,D); small cuff-shaped pygi- dium (9D; lOC) Dipolydora tridenticulata 6. Bundles of needlelike notosetae protruding through cuticle in posterior segments; pygidium with anal cirri Polydora robi 6. Bundles of posterior notosetae absent; cup-shaped pygidium 7 7. Major spines of segment 5 simple, falcate (14D,F) Polydora mabinii 7. Major spines of segment 5 with lateral tooth or flange 8 8. Occipital tentacle present Polydora cavitensis* 8. Occipital tentacle absent ...... 9 9. Palps crossed by bars of black pigmentation (1IA); caruncle to end of segment 3 (1IA) Polydora sp. A 9. Palp pigmentation absent; caruncle to middle of segment 2 (12A,D; BA,C) ...... Polydora umangivora 10. Branchiae begin on segment 2 (2A; 3A,B) Boccardia berkeleyorum 10. Branchiae begin on segment 7 (4A) Carazziella reishi *Species not encountered during this study.

SYSTEMATIC ACCOUNT Paraboccardia Rainer, 1973:550. Type species: Paraboccardia syrtis Rainer, 1973, by origi­ Family SPIONIDAE Grube, 1850 nal designation. Genus Boccardia Carazzi, 1893 Neoboccardia Buzhinskaja, 1985:129. Type Perialla Kinberg, 1866:353. Type species: species: Polydora perata Khlebovitsch, 1959, Perialla claparedei Kinberg, 1866, by mono­ by monotypy. typy. (See Remarks section.) Boccardia Carazzi, 1893:15. Type species: REMARKS: The genus Boccardia has been Polydora polybranchia Haswell, 1885, by reviewed by Blake and Woodwick (1971) and monotypy. emended by Blake and Kudenov (1978). To Polydora and Related Genera Associated with Hermit Crabs . Williams 433 date 20 species of Boccardia have been de­ Boccardia berkeleyorum Blake & W oodwick, scribed (see Blake and Kudenov 1981, 1971:34-36, fig. 2a-k; Sato-Okoshi and Hutchings and Turvey 1984, Blake 1986, Okoshi, 1997:486. Guerin 1990). As indicated by Blake (1983), the genus name Perialla Kinberg, 1866, has MATERIAL EXAMINED: Philippines: Ba­ priority over Boccardia Carazzi, 1893. How­ tangas: 5 spec. (USNM 187516: 4 spec.; USNM ever, since the description of Perialla clapar­ 187517: 1 spec., on 2 SEM stubs), from edei by Kinberg (1866) the genus name has Drupa rubusidaeus (Roding) inhabited by un­ not been utilized by any other author, al­ identified hermit crabs, Sombrero Island, 13 though Kinberg's description of this species July 1997; 4 spec. (AMNH 4246) from dead was reprinted (Kinberg 1910). Article 23.9.1 bivalve shells and Drupella coruus (Roding) of the International Code of Zoological No­ inhabited by Dardanus lagopodes, Sombrero menclature (ICZN 1999) indicates that pre­ Island, 30 January 1999; Oriental Mindoro: vailing usage must be maintained when a 1 spec. (USNM 187518), from Drupa sp. in­ senior synonym has not been used as a valid habited by Calcinus gaimardii (Milne­ name after 1899 (23.9.1.1) and when the ju­ Edwards), Puerto Galera: Big Lalaguna nior synonym has been used in at least 25 Beach, 31 July 1997. United States: Califor­ publications by at least 10 authors in the last nia: Cayucos: 2 paratypes (USNM 48761), from 50 yr (23.9.1.2). The second condition of Tegula brunnea (Philippi) occupied by Pagurus article 23.9.1 is fulfilled, but the first con­ granosimanus (Stimpson), 28 August 1961, leg. dition is not strictly met, although in a prac­ K. H. Woodwick. tical sense it is because no author other than DESCRIPTION: Up to 12.1 mm long, Kinberg has utilized the name Perialla after 0.7 mm wide at segment 7; 80 segments. 1899. For these reasons and to serve taxo­ Prostomium with slight indentation; caruncle nomic stability, I conclude that Perialla extending to end of segment 3; occipital ten­ should be considered a nomen oblitum and tacle and eyes absent (Figures 2A, 3A). Palps that Boccardia be considered a nomen pro­ short, with bars of white pigmentation. Black tectum to preserve the current usage of Boc­ pigmentation on sides of prostomium and cardia. peristomium, and dusky black pigmentation DIAGNOSIS: Prostomium with anterior in­ on dorsal side of all segments, most promi­ cision or rounded; caruncle extending poste­ nent on posterior segments (Figure 2A). riorly, surrounded by cilia of nuchal organ. Slight dusky black pigmentation on ventral Segment 1with orwithoutnotosetae. Segment side of prostomium and anterior segments; 5 modified with two types of heavy spines pigment nearly absent after preservation; in arranged in a double row: dorsal row simple, alcohol body opaque white to light tan. falcate; ventral row with expanded tip and ac­ Segment 1 with neurosetae, weakly devel­ cessory structures bearing bristles. Posterior oped notopodiallobes, lobes more prominent notosetae in form of capillaries or specialized in life, notosetae absent (Figures 2A, 3A,B). spines. Bidentate hooded hooks begin on Winged capillary notosetae of segments 2-4, segments 7-11; main fang at approximately 6, and subsequent segments in two rows, no­ right angle to shaft, with wide angle between tosetae ofposterior row longer (Figure 3A,B); main fang and apical tooth, no constriction middle segments with two acicular spines on shaft. Branchiae on segments 2-4, 6, and accompanied by two capillary setae; in some subsequent segments. Pygidium lobed, cuff­ specimens reduced to one acicular spine in shaped or disk-shaped with or without sepa­ posterior segments and two capillary setae rate lobes. (Figures 2B, 3C). Winged capillary neuro­ setae of segments 2-4, 6, and subsequent Boccardia berkeleyorum Blake & W oodwick, segments in two rows (Figure 3B); four bi­ 1971 dentate hooded hooks begin on segment 7, Figures 2, 3 up to five in series at segment 15, accom- 434 PACIFIC SCIENCE· October 2001

A

B

C,D

FIGURE 2. Boccardia berkeleyorum Blake & Woodwick (AMNH 4246). A, Anterior end, dorsal view; B, posterior end, dorsal view; C, bidentate hooded hook from anterior segment; D, falcate and bristle-topped spines and companion setae offifrh segment, lateral view. Scale: A, B, 150 flm; C, D, 50 flm. panied by one or two winged capillaries; dorsal to five to seven bristle-topped spines hooks with wide angle between main fang and (Figures 2D, 3B,E). apical tooth, approximately right angle be­ Branchiae on segments 2-4, 6, and sub­ tween main fang and shaft, without constric­ sequent segments (Figures 2A, 3A,B); attain­ tion on shaft, with fine bristles on hood ing full size at segment 8, absent from (Figures 2C, 3D). posterior one-fourth of body; nototrochs Segment 5 with ventral fascicle of four from segment 2. winged capillary neurosetae; notosetae ab­ Pygidium with three to four irregularly sent. Major spines of two types in curved, shaped lobes surrounding anus, lobes with double row; four to six simple falcate spines nonmotile cilia (Figures 2B, 3C). Polydora and Related Genera Associated with Hermit Crabs . Williams 435

FIGURE 3. Boccardia berkeleyorum Blake & Woodwick (USNM 187517), SEM micrographs. A, Anterior end, dorsal view; B, anterior end, lateral view of right side; C, posterior end, dorsal view; D, bidentate hooded hooks from anterior segment; E, falcate and bristle-topped spines and companion setae offifth segment, lateral view. Scale: A-C, 100 Ilm (B, C, see scale bar in A); D, E, 10 Ilm.

REMARKS: The Philippine specimens ECOLOGY: Boccardia berkeleyorum has been agree well with the original description by found in V-shaped burrows within gastropod Blake and Woodwick (1971). However, these shells occupied by Calcinus gaimardii, Darda­ specimens possess weakly developed notopo­ nus lagopodes, and an unidentified hermit crab, dial lobes on segment 1 not observed in the in addition to dead bivalve shells. In Califor­ holotype, possibly due to contraction after nia B. berkeleyorum is found within Tegula preservation. Black pigmentation on the new brunnea Philippi occupied by Pagurus gran­ specimens is observed in life, but is absent osimanus (Stimpson) (Blake and Woodwick from specimens preserved in alcohol for more 1971). Boccardia berkeleyorum is one of eight than 6 months; white bars of pigment on the members of the genus known to bore into palps are also absent after preservation, but calcareous substrata: B. acus Rainer, B. anop­ have been observed in Boccardia berkeleyorum thalma (Rioja), B. chilensis, B. galapagense specimens from Vancouver Island (Sato­ Blake, B. lamellata Rainer, B. otakouica Rainer, Okoshi and Okoshi 1997) and Boccardia chil­ and B. tricuspa (Hartman). ensis Blake & W oodwick from New Zealand DISTRIBUTION:Vnited States: California (Read 1975). (Blake and Woodwick 1971); Canada: Van- 436 PACIFIC SCIENCE· October 2001 couver Island (Sato-Okoshi and Okoshi Segment 1 with noto- and neurosetae. 1997); Philippines: Batangas, Mindoro (first Unilimbate capillary notosetae of segments record: this paper). 2-4, 6, and subsequent segments in two rows; reduced to three to four simple capillary no­ Genus Carazziella Blake & Kudenov, 1978 tosetae in posterior segments; no specialized posterior spines. Unilimbate capillary neuro­ Carazziella Blake & Kudenov, 1978:240. setae of segments 2-4, 6, and subsequent Type species: Polydora citrona Hartman, segments in two rows; two bidentate hooded 1941, by original designation. hooks begin on segment 8, up to three in se­ ries at segment 15, accompanied by one cap­ DIAGNOSIS: Prostomium with anterior incision or rounded; caruncle short, some­ illary seta; hooks with wide angle between times divided with second ridge. Segment 1 main fang and apical tooth, approximately with or without notosetae. Segment 5 modi­ right angle between main fang and shaft, no fied with two types of heavy spines arranged constriction on shaft, hood serrated (apparent in a double row: dorsal row falcate with or via SEM only) (Figure 4H). without bristles; ventral row with expanded Segment 5 with posterioventral fascicle of tip and accessory structures bearing bristles. four winged neurosetae. Notosetae absent in Bidentate hooded hooks begin on segments largest specimen; anteriodorsal fascicle oftwo 7-14; main fang at approximately right angle capillary notosetae present in smaller speci­ to shaft, wide angle between main fang and men. Major spines of segment 5 of two types, apical tooth, no constriction on shaft. Bran­ ventral row offour bristle-topped spines with chiae begin on segments 7-10. Pygidium a cone lacking bristles, cone oriented apically with two to four lobes or digitiform cirri. (Figure 4D) or projecting laterally (Figure 4E-G), and a dorsal row of three simple fal­ Carazziella reishi (Woodwick, 1964) cate spines (Figure 4B, C, G). Figure 4 Branchiae begin on segment 7, attaining full size at segment 8, broad and overlapping Pseudopolydora reishi Woodwick, 1964: 152­ at midline, continuing for four to nine 154, fig. 3(1-3); Kahn and Lloyd, 1973a: segments. 381; non Reish (1968:84). Pygidium with four glandular lobes (Fig­ Carazziella reishi (Woodwick): Blake, 1979a: ure 41,J), dorsal and ventral notches promi­ 477-479, fig. 7; Sato-Okoshi, 1999:838, nent, lateral divisions obscured (Figure 4J). appendix 1, appendix 2. Gizzardlike structure in the digestive tract MATERIAL EXAMINED: Philippines: Ori­ absent. ental Mindoro: 3 spec. (USNM 187520: 2 spec.; REMARKS: One specimen differed from USNM 187519: 1 spec., on SEM stub), from the halotype in presence of four eyes, Conus sp. inhabited by Ciliopagurus strigatus rounded prostomium, and absence of notose­ (Herbst), Puerto Galera: Big Lalaguna Beach, tae on segment 5. However, these characters 31 July 1997. Marshall Islands: Eniwetok are variable in other species ofthe genus (e.g., Atoll: halotype (USNM 32611), from sand Carazziella carrascoi Blake) and therefore are among coral rock, Engebi, 7 September 1956, insufficient for the erection of a new species. leg. D. J. Reish. ECOLOGY: The specimens were found in DESCRIPTION: Largest Philippine speci­ detrital burrows beneath encrusting coralline men 2.4 mm long, 0.3 mm wide at segment 7; algae. The burrows of the worms extended 47 segments. Prostomium rounded; triangu­ through the algae, but not into the gastropod lar caruncle extending to middle of segment shells (Conus sp.) inhabited by Ciliopagurus 2; occipital tentacle absent; four eyes present strigatus. In the Marshall Islands and Indone­ (Figure 4A). Palps extending back to segment sia, Carazziella reishi has been found in coral 18. In alcohol slight black pigmentation on rocks (Woodwick 1964, Blake 1979a) and is prostomium (Figure 4A), body opaque white the only member of the genus known to bur­ to light tan. row into calcareous substrata. , D J

H

A,I,J

B-H

FIGURE 4. Carazziella reishi (Woodwick) (USNM 187520). A, Anterior end, dorsal view; B, simple and bristle-topped spines and companion setae offifth segment, lateral view; C, falcate spines offifth segment. D-F, bristle-topped spines offifth segment; G, falcate and bristle-topped spines of fifth segment; H, bidentate hooded hook from anterior seg­ ment; I, posterior end, dorsal view; J, posterior end, lateral view. Scale: A, I, J, 100 J.lm; B-H, 30 J.lffi. 438 PACIFIC SCIENCE· October 2001

DISTRIBUTION: Johnston Atoll (Ward teriorly, surrounded by cilia of nuchal organ. 1981); Marshall Islands: Eniwetok Atoll Segment 1 with notosetae. Segment 5 modi­ (Woodwick 1964); Japan: Chikura, Shingu fied with one type of major spines in a single (Sato-Okoshi 1999); Philippines: Mindoro curved row, with or without companion setae. (first record: this paper); Indonesia: Bali, Pula Posterior notopodial spines present or absent. Boonda (Blake 1979a). Bidentate hooded hooks begin on segments 7-17; obtuse angle between main fang and Genus Dipolydora Verrill, 1879, sensu Blake, shaft, narrow angle between main fang and 1996 apical tooth; no constriction on shaft. Bran­ chiae begin posterior to segment 5. Pygidium Dipolydora Verrill, 1879:174. Type species: variable; cuffiike, cup-shaped, or lobed. With Polydora concharum Verrill, 1879, des­ or without gizzardlike structure in anterior ignated by Verrill (1881). part of digestive tract.

DIAGNOSIS: Prostomium with anterior Dipolydora armata (Langerhans, 1880) incision or rounded; caruncle extending pos- Figures 5, 6

FIGURE 5. Dipolydora armata (Langerhans) (USNM 187530). A, Anterior end, dorsal view; B, posterior end, dorsal view; C, posterior end, lateral view. Scale; A-C, 100 /UIl. Polydora and Related Genera Associated with Hennit Crabs . Williams 439

FIGURE 6. Dipo/ydora armata (Langerhans) (USNM 187525), SEM micrographs. A, Anterior end, dorsal view; B, seg­ ments 3 to 7, lateral view; C, posterior end, dorsal view; D, modified spines and noto- and neurosetae ofsegment 5, lateral view; E, posterior end, dorsal view; F, modified spines and noto- and neurosetae of segment 5, lateral view; G, posterior end, apical view. Scale: A, C, 50 ~; B, E, G, 20 ~; D, 5 ~; F, 10 ~.

Polydora armata Langerhans, 1880:93-94, Woodwick, 1964:147, fig. 2; Day, 1967: pI. 4, fig. 5a-c; Fauvel, 1927:55-56, fig. 466-468, fig. 18.2i-j; Kahn and Lloyd, 19a-e; Okuda, 1937:230-231, fig. 10; 1973a:381; Rainer, 1973:558, fig. 7; Read, Hartman, 1941:306; 1969:127, 5 figs.; 1975:412-413; Blake and Kudenov, 1978:

------440 PACIFIC SCIENCE· October 2001

255-256, fig. 43a-g; Hartmann-Schroder, Island, 9 July 1997. Indonesia: Bali: 10+ spec. 1979:134, figs. 299-302; Kojima and Ima­ (USNM 187531), from Conus sp. inhabited by jima, 1982:31-35; Blake, 1983:258-260; Dardanus fagopodes, Sanur, 5 August 1997; Ward, 1987:352-353, fig. 3.II.129. 30+ spec. (USNM 187532), from Bursa sp., Polydora monifaris Ehlers, 1905:43-44, pI. 6, Drupe/fa comus, and Turbo sp. inhabited by figs. 5-14; fide Day (1954). Calcinus gaimardii, C. fatens, and Cfibanarius Dipolydora armata (Langerhans): Blake, 1996: englaucus (Ball & Haig), and live Drupeffa cor­ 196-198 [synonomy], fig. 4.36; Lewis, nus, Sanur, 6 August 1997; Banjak Island: 10 1998:651-662, figs. 4-8; Sato-Okoshi, spec. (USNM 45325), from coral reef limestone 1999:838, appendix 1, appendix 2. platforms, Pulo Melila (ZO 15' N, 9r 25' E), November 1963, leg. A. J. Kohn. Australia: MATERIAL EXAMINED: Philippines: Ba­ Point Gellibrand (37° 52' S, 144° 54' E): 70+ tangas: 8 spec. (USNM 187521), from Latirus spec. (NMV F42881), in shell of Hafiotis ruber turritus (Gmelin) inhabited by an unidentified Leach, 5 April 1997, leg. J. A. Blake and J. D. hermit crab, Sombrero Island, 5 July 1997; Kudenov; Tiparra (34° 5' S, 13r 29' E): 1 50+ spec. (AMNH 4247), from Drupa rubusi­ spec. (NMV F43056), in shell of Hafiotis roei daeus, Drupe/fa comus, and Latirus turritus Gray, 19 May 1971, leg. S. Shepherd. Carib­ inhabited by Calcinus gaimardii, Dardanus fa­ bean Sea, Barbados, 94 spec., from the hy­ gopodes, and unidentified hermit crabs, Som­ drocoral Miffepora compfanata Lamarck, brero Island, 13 July 1997; 10+ spec. (USNM Tropicana reef on the west coast, 11 Sep­ 187522), from Drupa rubusidaeus inhabited by tember 1994, leg. J. Lewis. Calcinus fatens (Randall), Sepoc Point, 5 July DESCRIPTION: Maximum length 3.3 mm 1997; Oriental Mindoro: 10+ spec. (USNM long, 0.2 mm wide at segment 7; up to 33 187523), from Drupeffa comus inhabited by segments. Prostomium bifid; caruncle trian­ Calcinus fatens, Puerto Galera: Coco Beach, gular extending from middle to end of seg­ 12 January 1999; 300+ specimens (USNM ment 2 (Figures 5A, 6A); occipital tentacle 187524: 275+ spec.; USNM 187525: 11 spec., and eyes absent. Palps extend posteriorly for on 5 SEM stubs), from Drupella comus and 10-15 segments (Figures 5A, 6A). In alcohol Cymatium rubecufum (Linnaeus) inhabited by body opaque white to light tan; no pigmen­ unidentified hermit crabs, Puerto Galera: Big tation present. Lalaguna Beach, 19 July 1997; 55+ spec. Segment 1 with notosetae and longer neu­ (AMNH 4248), from Drupeffa comus inhabited rosetae (Figure 6A). Unilimbate capillary no­ by Cafcinus minutus (Buitendijk) and un­ tosetae of segments 2-4, 6, and subsequent identified hermit crabs, Puerto Galera: Big segments in two rows (Figure 6B); four to Lalaguna Beach, 21 July 1997; 40+ spec. six posteriormost segments containing 9-17 (USNM 187526), from Drupella comus and thick acicular spines with straight or curved Chicoreus pafmarosae (Lamarck) inhabited by tip, often in curved arrangement accom­ Calcinus gaimardii and C. fatens, Puerto Ga­ panied by two capillary setae (Figures 5B,C, lera: Big Lalaguna Beach, 31 July 1997; Pala­ 6C,E,G); following segment containing thin­ wan: 7 spec. (USNM 187527); Aldan: 10+ spec. ner needlelike spines (Figures 5B,C, 6G). (USNM 187528), from Drupeffa comus in­ Unilimbate capillary neurosetae of segments habited by Calcinus gaimardii, Boracay: Rocky 2-4, 6, and subsequent segments in two rows Beach, 12 April 1999; 45+ spec. (USNM (Figure 6B); three bidentate hooded hooks 187529), from Drupeffa comus inhabited by begin on segment 7, up to six in series at Calcinus gaimardii, Boracay: Rocky Beach, 15 segments 8-10; hooks with obtuse angle April 1999; Cebu: 100+ spec. (USNM 187530), between main fang and shaft, narrow angle from Astralium sp., Bursa sp., Cantharus sp., between main fang and apical tooth, no con­ Conus sp., Gyrineum gyrinum (Linnaeus), and striction on shaft, with fine bristles covering unidentified gastropod shells inhabited by hood; with capillary companion setae (Figure Calcinus gaimardii, C. fatens, Dardanus fago­ lOP). podes, and unidentified hermit crabs, Olango Segment 5 almost twice as large as seg- Polydora and Related Genera Associated with Hermit Crabs . Williams 441

ments 4 and 6, with two to three spines; covered by bristles, and larger cup-shaped posterioventral fascicle of three to four uni­ pygidium. limbate capillary neurosetae and anteriodorsal Lewis (1998) reported on reproduction fascicle of three to four geniculate notosetae of Dipolydora armata associated with the hy­ (Figure 6B,D,F). Major spines bidentate with drozoan Millepora complanata from Barbados. lateral hood connecting teeth; hood with fine Examination of the Barbados specimens bristles; companion setae absent (Figure showed that he misidentified peritrichous 6D,F). ciliates (Ciliophora: Oligohymenophorea) at­ Branchiae begin on segment 7, attaining tached to the notosetae as spermatophores of full size on segments 8-9, terminating on Dipolydora armata. The peritrichous ciliates segment 11; nototrochs from segment 2 (Cothumia-like sp.) attached to the notosetae (Figures SA, 6A). of Dipolydora armata are approximately 50 11m Gizzardlike structure in the digestive tract in length and have a 3D-11m smooth stalk. absent. The ciliates are found attached to 16-35% of Pygidium cuff-shaped with dorsal gap and Dipolydora armata specimens from Barbados, ventral notch giving appearance of two lobes and worms have 2-12 ciliates attached to the (Figures 5B,C, 6C,E,G). notosetae ofthe middle body segments (Lewis REMARKS: The morphology of the Philip­ 1998). In comparison, spermatophores of pine and Indonesian specimens agrees with Polydora comuta Bosc are about 220 11m in previous descriptions of this broadly defined length and have a tail approximately 1100 11m species. Blake (1983) indicated that two spe­ long covered with short, hairlike projections cies might be contained within Dipolydora (Rice 1978). Upon contact with the palps of a armata. Specifically, he suggested that a form female worm the spermatophores break apart with thinner posterior spines found in colder and release enclosed sperm (Rice 1978). latitudes might represent an undescribed Spermatophores are probably present in the species. Although the specimens reported genus Dipolydora but their production and here exhibit thin needlelike spines in one morphology remain unknown (Rice 1978, segment following the acicular spines, in all Blake and Arnofsky 1999). other respects they exhibit the typical mor­ ECOLOGY: Dipolydora armata is a common phology of Dipolydora armata. More recently, associate ofhermit crabs from the Philippines examination of specimens from Australia, and Indonesia. The worms were found in 1­ Barbados, and Belize has indicated the exis­ 24% of the hermit crabs examined from the tence of several undescribed sibling species six provinces sampled in the Philippines (J. A. Blake, pers. comm.; Blake, unpubl. data; (6.2 ± 6.3; total percentage ± SD, n = 1578). Radashevsky, unpubl. data). The specimens The species is found in burrows penetrating reported here may represent an additional the coralline algal crusts covering live Dru­ undescribed species within the Dipolydora pella comus or gastropod shells inhabited by armata complex, but a comprehensive anal­ hermit crabs. Typically the worms burrow ysis of type material and sibling species is through the coralline algae and construct de­ required. trital tubes in the space between the shell Martin (1996) indicated that Dipolydora surface and algal covering. When present, the armata and D. blakei (Maciolek) may be syn­ worms are often abundant, with approxi­ onyms, but examination of type material of mately 35-40 worms per square centimeter, D. blakei (USNM 81928) showed that the spe­ and hundreds of worms are found in a single cies are distinct. Dipolydora armata differs shell. The worms have been found in 18 gas­ from D. blakei based on the short triangular tropods: Astralium sp., Bursa sp., Cantharus caruncle, bidentate major spines with cowl­ sp., Cerithium tenellum, Chicoreus palmarosae, ing, and cuff-shaped pygidium rather than Conus sp., Coralliophila neritoidea, Cymatium triangular caruncle extending laterally to the rubeculum, Drupa grossularia, D. rubusidaeus, base of the notopodiallamellae on segment 1, Drupella sp., D. comus, D. rugosa, Gyrineum major spines falcate with accessory flange gyrinum, Latirolagena smaragdula (Linnaeus), 442 PACIFIC SCIENCE· October 2001

Latirus turritus, Peristernia nassatula, and Turbo Dipolydora socialis (Schmarda): Blake, 1996: sp. inhabited by eight species ofhermit crabs: 189-192, fig. 4.34 [synonomy]. Calcinus gaimardii, C. latens, C. minutus, C. pulcher (Forest), Ciliopagurus strigatus, Cliba­ MATERIAL EXAMINED: Philippines: Ba­ narius englaucus, Dardanus lagopodes, and Pa­ taan: 1 spec. (USNM 187533), from Cantharus guristes runyanae. undosus (Linnaeus) inhabited by Calcinus gai­ DISTRIBUTION: United States: California mardii, Mabayo, 21 February 1999; Batangas: (Blake 1996); New Zealand (Read 1975, 3 spec. (AMNH 4249), from Drupa rubusidaeus Blake 1983); Australia (Blake and Kudenov inhabited by Dardanus lagopodes, Anilao, 13 1978); Japan: northern limit to Chikura February 1999; 13 spec. (USNM 187534: 10 (Sato-Okoshi 1999); Philippines: Batangas, spec., USNM 187535: 3 spec., on 3 SEM stubs), Oriental Mindoro, Palawan, Aldan, Cebu from Drupa rubusidaeus, Drupella cornus, and (first record: this paper); Indonesia: Bali (this an unidentified gastropod shell inhabited by paper); Europe (Carazzi 1893); South Amer­ Calcinus minutus and unidentified hermit ica: Argentina, Chile, Ecuador (Blake 1983); crabs, Sombrero Island, 13 July 1997; 1 spec. Caribbean Sea: Barbados (Lewis 1998). (USNM 187536), from Drupella cornus in­ habited by Calcinus minutus, Sombrero Island, 30 January 1999; Palawan: 1 spec. (USNM Dipolydora socialis (Schmarda, 1861) 187537) from Drupella rugosa inhabited by Figures 7, 8 Paguristes runyanae, Magbautoc Island, 16 March 1999. Australia: Port Phillip Bay: 5 Leucodore socialis Schmarda, 1861:64, figs. a-c, spec. (NMV F43159), from silt/sand, Geelong pI. 26, fig. 209. Arm (38 0 2.3' S, 1440 34.5' E), 18 November Polydora caeca var. magna Berkeley, 192 7:419; 1971, leg. Marine Pollution Studies Group. Pettibone, 1967: II. DESCRIPTION: Largest Philippine speci­ Polydora magna Berkeley & Berkeley, 1936: men measured 7.0 mm long, 0.4 mm wide 473; 1952:21. at segment 7; 56 segments. ProstOInium Polydora socialis plena Berkeley & Berkeley, strongly incised on anterior margin; caruncle 1936:468-469; 1952:22; Pettibone, 1967: extending to segments 3-5; occipital tentacle II. and eyes absent (Figures 7A, 8A). Palps ex­ Polydora caeca Oersted, sensu Berkeley & tending posteriorly to segments 11-15; with Berkeley, 1936:469; 1952:20-21; non dusky black pigmentation on ventral side on Oersted, 1843;fide Blake (1979b). some specimens (Figure 7A). Color in alco­ Polydora socialis (Schmarda): Hartman, 1941: hol opaque white; no body pigmentation. 310-311, pI. 48, figs. 41-42; 1969:147; Segment 1 with noto- and neurosetae and Hartmann-Schroder, 1962:137-139, figs. large notopodial lobes (Figures 7A, 8A). 167-168; 1965:209-211, figs. 200-203; Winged capillary notosetae of segments 2-4, Blake, 1971:20-23, figs. 13-14 [synon­ 6, and subsequent segments arranged in three omy]; 1975:215, figs. 237-238; 1979b:607­ rows; no specialized posterior spines. Winged 609 [synonomy]; 1981:950; 1983:264; capillary neurosetae of segments 2-4, 6, and Carrasco, 1974:194-196, figs. 27- 32; Light, subsequent segments in two rows; two or 1977:71; 1978:179-181, fig. 180; Blake and three bidentate hooded hooks from segment Kudenov, 1978:248-250, fig. 38d-e; John­ 7, up to five in series at segment 11, accom­ son, 1984:6-28 to 6-30, figs. 6-19 and 6­ panied by one or two capillary setae on seg­ 20; Sato-Okoshi and Okoshi, 1997:486. ments 7-10; hooks curved with obtuse angle Polydora plena Foster, 1971:24-25, figs. 22­ between main fang and shaft, narrow angle 29. between main fang and apical tooth, without Polydora neocardalia Hartman, 1961:96-98, pI. constriction on shaft (Figure 7E). 14, figs. 1-4; 1969:141, 2 figs.; Lissner et Segment 5 with posterioventral fascicle of aI., 1986: appendix D; Steinhauer and four geniculate neurosetae and anteriodorsal Imamura, 1990: F-l;fide Blake (1996). fascicle of two or three geniculate notosetae; FIGURE 7. Dipolydora socia/is (Schmarda) (USNM 187533). A, Anterior end, dorsal view; B, posterior end, dorsolateral view; C, falcate spines and companion setae offifth segment, lateral view; D, posterior end, lateral view; E, bidentate hooded hook from anterior segment. Scale: A, B, D, 100 jl!Il; C, E, 30 )lm. 444 PACIFIC SCIENCE· October 2001

FIGURE 8. Dipolydora socia/is (Schmarda) (USNM 187535), SEM micrographs. A, Anterior end, dorsal view; B, posterior end, dorsolateral view; C, falcate spines and companion setae offifth segment, dorsal view; D, falcate spines and com­ panion setae of fifth segment, dorsal view. Scale: A, 50 ~m; B, 20 ~m; C, D, 5 ~m.

slightly curved row of five exposed major REMARKS: The Philippine specimens agree spines alternating with pennoned companion well with recent descriptions of Dipolydora setae (Figures 7C, 8C,D). Major spines fal­ socialis (Blake 1996). However, the specimens cate, with subterminal protuberance (Figures are mostly smaller than those previously re­ 7C,8C,D). corded, although Blake and Kudenov (1978) Branchiae begin on segments 8-10; reach­ also found smaller specimens in collections ing posterior one-third of body. from Australia. Considerable debate exists Gizzardlike structure apparent in some over the burrowing activity of Dipolydora specimens at segments 14-18. socialis. The species has been documented to Pygidium with large ventral lobe and two construct burrows in soft bottom sediments smaller dorsal lobes; dorsal lobes incon­ and calcareous substrata, including gastropod spicuous in some specimens (Figures 7B,D, shells inhabited by hermit crabs (Blake 1971, 8B). 1981; ].D.W., unpubl. data). However, Rada- Polydora and Related Genera Associated with Hermit Crabs . Williams 445 shevsky (1993) and Sato-Okoshi and Okoshi Dipolydora tridenticulata (Woodwick, 1964) (1997) considered the species to be a non­ Figures 9, 10 burrower restricted to soft bottom areas. The gross morphology of the gizzardlike Polydora tridenticulata W oodwick, 1964: 153­ structure of the anterior portion of the di­ 157, fig. 4(1-5); Ward, 1987:354-355, fig. gestive tract has been described in consider­ 3.11.132. able detail for Dipolydora socialis and other Dipolydora tridenticulata (Woodwick): Blake, related species (Blake 1971), but the function 1996:186. of the structure has not been investigated. In the description of Pygospio muscularis, Ward MATERIAL EXAMINED: Philippines: Ba­ (1981) proposed that the structure aids in the tangas: 3 spec. (USNM 187538: 1 spec.; USNM removal of bacterial and algal films attached 187539: 2 spec., on 3 SEM stubs), from Bursa to sand grains. Histological and behavioral granularis (Roding), inhabited by Calcinus feeding studies of species possessing the giz­ minutus and Dardanus lagopodes, Anilao, 13 zardlike structure are required to determine February 1999; 8 spec. (USNM 187540), from its functional significance. Drupella cornus inhabited by Dardanus lago­ ECOLOGY: In this study Dipolydora socialis podes and dead bivalve shells, Sombrero Is­ was found in sediment tubes constructed land, 30 January 1999; 6 spec. (AMNH 4250: 5 within the crevices of damaged gastropod spec.; USNM 187541: 1 spec., on SEM stub), shells (Cantharus undosus, Drupa rubusidaeus, from Drupa rubusidaeus inhabited by Calcinus Drupella cornus, and D. rugosa) inhabited by gaimardii and Dardanus lagopodes, Sombrero hermit crabs (Calcinus gaimardii, C. minutus, Island, 13 July 1997; Oriental Mindoro: 2 Dardanus lagopodes, and Paguristes runyanae). spec. (USNM 187542: 1 spec.; USNM 187543: 1 Three specimens were found with perit­ spec., on SEM stub), from Drupella cornus richous ciliates (Ciliophora: Oligohymeno­ inhabited by Calcinus minutus, Puerto Galera: phorea) attached to the notosetae of approxi­ Big Lalaguna Beach, 21 July 1997; 20 spec. mately four to six segments, with one ciliate (USNM 187544: 15 spec.; USNM 187545: 5 per fascicle. Additional reports of peritri­ spec., on 7 SEM stubs), from Drupa sp., chous ciliates associated with spionid poly­ Drupella cornus, Terebra sp., and an unidenti­ chaetes include Boccardiella magniovata fied gastropod shell inhabited by Calcinus (Read), Polydora neocaeca Williams & Rada­ gaimardii, C. minutus, and C. latens, Puerto shevsky (Read 1975, Williams and Rada­ Galera: Big Lalaguna Beach, 31 July 1997. shevsky 1999), and Dipolydora armata (see Marshall Islands: Eniwetok Atoll: holotype previous Remarks section). Williams and (USNM 32612), burrowing in coral rock, Rigili, Radashevsky (1999) used SEM to examine the 29 August 1956, leg. D.]. Reish. stalks of peritrichs attached to the hooded DESCRIPTION: Largest Philippine speci­ hooks of Polydora neocaeca. Little is known men 3.1 mm long, 0.3 mm wide at segment 7; about the association between peritrichs and 69 segments. Prostomium slightly incised; polychaete hosts, but the peritrichs may ben­ broad caruncle extending to segments 3-4; efit by extending between the branchiae of occipital tentacle absent; eyes deeply em­ the worms and feeding within the flow of bedded and irregularly shaped or eyes absent water drawn through the burrow. (Figures 9A, lOA). Palps extending posteri­ DISTRIBUTION: Mexico (Blake 1981); orly to segment 8. In alcohol body opaque United States: east and west coasts (Blake white; no pigmentation. 1971, 1996); Canada: Vancouver Island (Sato­ Segment 1 with noto- and neurosetae Okoshi and Okoshi 1997); New Zealand (Figures 9A, lOA). Winged capillary notose­ (Read 1975); Australia (Blake and Kudenov tae of segments 2-4, 6, and subsequent seg­ 1978); Philippines: Bataan, Batangas, Palawan ments in three rows; no specialized posterior (first record: this paper); Falkland Islands spines. Winged capillary neurosetae of seg­ (Blake 1983); South America: Argentina, ments 2-4, 6, and subsequent segments in Chile, Ecuador (Blake 1983). two rows; one to three bidentate hooded c

B,C

FIGURE 9. Dipolydora tridenticulata (Woodwick) (USNM 187540). A, Anterior end, dorsal view; B, spines and companion setae of fifth segment, dorsal view; C, bidentate hooded hook from anterior segment; D, posterior end, dorsal view. Scale: A, D, 100 Jilll; B, C, 30 Jilll. FIGURE 10. Dipolydora tridenticulata (Woodwick) (USNM 187539) (A-D), (USNM 187541) (B), (USNM 187545) (C, E) and Dipolydora armata (Langerhans) (USNM 187525) (F), SEM micrographs. A, Anterior end, dorsal view; B, spines and companion setae offifth segment, dorsolateral view; C, posterior end, dorsal view; D, spines and companion setae of fifth segment, dorsal view; E, peritrichous ciliate attached to notosetae; F, bidentate hooded hooks of anterior segment. Scale: A, 100 )lID; B, D, 5 )lffi; C, E, 10 )lffi; F, 1 )lffi. 448 PACIFIC SCIENCE· .October 2001

hooks from segment 7, up to five in series at Only one ciliate was found per fascicle of segment 15, accompanied by one capillary notosetae. See discussion on ecology of Di­ seta on segments 7-9; hooks with obtuse polydora socialis for additional references to angle between main fang and shaft, narrow peritrichlspionid associations. angle between main fang and apical tooth, no DISTRIBUTION: Hawai'i (Ward 1987); constriction on shaft (Figure 9C). Marshall Islands (Woodwick 1964); Philip­ Segment 5 with posterioventral fascicle of pines: Batangas: Oriental Mindoro (first rec­ four winged neurosetae and anteriodorsal ord: this paper). fascicle of two geniculate notosetae; slightly curved row of three to five exposed major Genus Polydora Bosc, 1802, sensu Blake, 1996 spines alternating with pennoned companion setae (Figures 9B, IOB,D). Major spines Polydora Bosc, 1802:151. Type species: Poly­ falcate with two lateral teeth (Figures 9B, dora cornuta Bosc, 1802, by monotypy. 10B,D). Teeth of unworn posterior spines Leucodore Johnston, 1838:66. Type species: curved with a sharp point; one tooth extends Leucodore ciliatus Johnston, 1838, by mono­ beyond main fang of spine, other tooth below typy; fide Blake and Maciolek (1987). main fang; worn anterior spines with rounded Leipoceras Mobius, 1874:254. Type species: teeth below main fang (Figures 9B, IOB,D). Leipoceras uviferum Mobius, 1874, by Branchiae begin on segments 8-10; at­ monotypy;fide Blake and Maciolek (1987). taining full size at segment 11; terminating Protopolydora Czerniavsky, 1881:360. Type on segments 15-20, nototrochs between bran­ species: Polydora hamata Langerhans, 1880, chiae. by original designation. Homonym of Gizzardlike structure absent in digestive Polydora hamata Webster, 1879, renamed tract. Polydora posthamata Jones, 1962. Pygidium small, glandular cuff-shaped with dorsal notch (Figures 9C, IOE). DIAGNOSIS: Prostomium with anterior REMARKS: Dipolydora tridenticulata belongs incision or rounded; caruncle extending pos­ to the D. giardi group of six species known teriorly, surrounded by cilia of nuchal organ. to burrow into calcareous substrata (Blake Segment 1 without notosetae. Segment 5 1996). The lateral teeth of the major spines modified with one type of major spine in a are worn on the anterior spines (Figures 9B, single curved row, with or without compan­ IOB,D) , and, as indicated by Woodwick ion setae; juveniles may possess large, falcate (1964), the tridentate morphology of the major spines that are replaced by subsequent spines is not apparent if care is not taken to spines. Posterior notopodial spines present view them from various angles. However, or absent. Bidentate hooded hooks begin on SEM shows that the two lateral teeth are segments 7-14, main fang at approximately observable even in worn spines and are best right angle to shaft, wide angle between main seen in lateral view (Figure lOB). fang and apical tooth, with constriction on ECOLOGY: Dipolydora tridenticulata is a shaft. Branchiae begin posterior to segment 5. burrower found in gastropod shells (Bursa Pygidium variable: cuff-shaped, cup-shaped granularis, Drupa sp., D. rubusidaeus, Drupella with dorsal gap, lobed, scoop-shaped, or with cornus, and Terebra sp.) inhabited by hermit digitiform cirri. Anterior portion of digestive crabs (Calcinus gaimardii, C. latens, C. minutus, tract lacks gizzardlike structure. and Dardanus lagopodes). The species has also been found in dead bivalve shells. Pe­ Polydora sp. A ritrichous ciliates were found attached by a Figure 11 short stalk to the notosetae of anterior seg­ ments in three specimens. The anterior end MATERIAL EXAMINED: Philippines: Ori­ of the ciliates extends to a position between ental Mindoro: 1 spec. (USNM 187546), from the branchiae of the worms (Figure 10E). Coralliophila sp. inhabited by Calcinus gai- Polydora and Related Genera Associated with Hennit Crabs . Williams 449

11A). In alcohol body opaque white to light tan, palps crossed with distinct bars of black pigmentation (Figure 11A). Segment 1 with neurosetae, without noto­ setae, with weakly developed notopodial lobes. Winged capillary notosetae of seg­ ments 2-4, 6, and subsequent segments arranged in three successive rows; no special­ ized posterior notosetae. Winged capillary neurosetae of segments 2-4, 6, and subse­ quent segments arranged in two vertical rows; five bidentate hooded hooks begin on seg­ ment 7, not accompanied by capillaries, up to eight in series at segment 23; hooks with approximately right angle between main fang and shaft, wide angle between main fang and apical tooth, with constriction on shaft (Figure 11 C). Segment 5 almost twice as large as seg­ c ments 4 and 6, with slightly curved row offive exposed major spines alternating with pen­ noned companion setae; with posterioventral fascicle of four winged capillary neurosetae; anteriodorsal fascicle of two geniculate noto­ setae. Major spines falcate, with rounded, lat­ eral tooth (Figure lIB). Branchiae begin on segment 7; attaining full size at 12. Pygidium unknown. REMARKS: Polydora sp. A belongs to the Polydora ciliata/websteri group. The species -A most closely resembles Polydora sp. (Blake 1996), Polydora sp. (Caceres-Martinez et al. B,C 1999), Polydora sp. (Sato-Okoshi 1999), P. agassizi Claparede, P. cf. agassizi (Radashevsky and Hsieh 2000a), and P. limicola Annenkova in having palps crossed by black bars, car­ uncle length, and major spine morphology. FIGURE 11. Polydora sp. A (USNM 187545). A, Anterior Lack of a posterior end and pygidium pre­ end, dorsal view; B, spines and companion setae of fifth cludes determination of the species. Polydora segment, dorsal view; C, bidentate hooded hook from anterior segment. Scale: A, 200 J.lIIl; B, C, 30 ~m. possessing palps with distinct bars or blotches of black pigment have recently been reviewed (Williams and Radashevsky 1999, Rada­ mardii, Puerto Galera: Big Lalaguna Beach, shevsky and Hsieh 2000a). 31 July 1997. ECOLOGY: The species was found asso­ DESCRIPTION: Specimen broken, lacking ciated with the gastropod shell Coralliophila posterior end; 7.9 mm long, 0.3 mm wide at sp. inhabited by the hermit crab Calcinus gai­ segment 7; 48 segments. Prostomium slightly mardii. It is unknown whether Polydora sp. A bifid; caruncle terminating at end of segment is a shell burrower or occupied a mud tube in 3; eyes and occipital tentacle absent (Figure the crevices of the gastropod shell. 450 PACIFIC SCIENCE· October 2001

Polydora umangivora Williams, n. sp. (USNM 187556), from Coralliophila neritoidea, Figures 12, 13 Drupella cornus, and Latirolagena smaragdula inhabited by Calcinus gaimardii, Puerto Ga­ HOLOTYPE: Philippines: Batangas (USNM lera: Coco Beach, 12 January 1999; 3 para­ 187553), from Thais mancinella inhabited by types (USNM 187557), from Drupella cornus Dardanus lagopodes, Sombrero Island, 30 Jan­ and D. rugosa inhabited by Calcinus gaimardii uary 1999. and C. latens, Puerto Galera: Coco Beach, 14 OTHER MATERIAL EXAMINED: Philip­ January 1999; 2 paratypes (USNM 187558), pines: Bataan: 31 paratypes (AMNH 4251), from Drupella cornus inhabited by Calcinus from Cantharus undosus, Coralliophila nerit­ gaimardii, Puerto Galera: Coco Beach, 15 oidea (Lamarck), Cymatium sp., Cypraea sp., January 1999; 1 paratype (USNM 187559), Latirolagena smaragdula, Latirus polygonus from Drupella cornus inhabited by Calcinus (Gmelin), L. turritus, and an unidentified gaimardii, Puerto Galera: Big Lalaguna gastropod shell inhabited by Calcinus gai­ Beach, 19 July 1997; 2 paratypes (USNM mardii, C. minutus, and Dardanus lagopodes, 187560), from Cymatium rubeculum inhabited Mabayo, 21 February 1999; 1 paratype (USNM by Calcinus gaimardii and Clibanarius sp., Pu­ 187547), from an unidentified gastropod shell erto Galera: Big Lalaguna Beach, 21 July inhabited by Diogenes sp., Morong, 6 Febru­ 1997; 11 paratypes (USNM 187561), from ary 1999; 7 paratypes (AMNH 4252), from Drupella cornus, Latirolagena smaragdula, Tur­ Cantharus undosus, Coralliophila neritoidea, bo sp., and unidentified gastropod shells in­ Latirolagena smaragdula, and an unidentified habited by Calcinus gaimardii and Clibanarius gastropod shell inhabited by Calcinus gai­ englaucus, Puerto Galera: Big Lalaguna Beach, mardii and C. minutus, Morong, 28 February 31 July 1997; 6 paratypes (USNM 187562), 1999; 1 paratype (USNM 187548), from Can­ from Coralliophila neritoidea, Drupella cornus, tharus undosus inhabited by Calcinus gaimardii, Strombus labiatus labiatus Roding, and an un­ Morong, 25 April 1999; Batangas: 6 paratypes identified gastropod shell inhabited by Calci­ (USNM 187549), from Terebra sp. inhabited by nus gaimardii, C. latens, C. minutus, and an an unidentified hermit crab, Anilao, 20 June unidentified hermit crab, Puerto Galera: Big 1997; 31 paratypes (USNM 187550), from Lalaguna Beach, 28 March 1999; 2 paratypes Latirus turritus, Pisania fasciculata (Reeve), and (USNM 187563), from Drupella cornus and an an unidentified gastropod shell inhabited by unidentified gastropod shell inhabited by Clibanarius sp. and Dardanus sp., Anilao, 28 Calcinus gaimardii, Bayanan Beach, 13 January June 1997; 14 paratypes (USNM 187551), from 1999; Aklan: 3 paratypes (USNM 187564), from Bursa granularis, Drupa rubusidaeus, Drupella Drupella cornus and an unidentified gastropod cornus, D. rugosa, Drupella sp., Gyrineum sp., shell inhabited by Calcinus gaimardii and C. Latirus turritus, Thais mancinella (Linnaeus), latens, Boracay: Diniwid Beach, 13 April 1999; and an unidentified gastropod shell inhabited 2 paratypes (USNM 187565), from Cantharus by Calcinus gaimardii, C. minutus, C. pulcher, undosus and Drupella rugosa inhabited by Cal­ and Dardanus lagopodes, Anilao, 13 February cinus latens, Boracay: Rocky Beach, 12 April 1999; 3 paratypes (USNM 187552), from Dru­ 1999; 5 paratypes (USNM 187566), from Cor­ pella cornus and Thais mancinella inhabited by alliophila neritoidea, Drupa sp., Drupella cornus, Calcinus minutus and Dardanus sp., Sombrero D. rugosa, and an unidentified gastropod shell Island, 13 July 1997; 4 paratypes (USNM inhabited by Calcinus gaimardii, C. latens, and 187554), from Chicoreus palmarosae, Drupella C. minutus, Boracay: Rocky Beach, 15 April cornus, and Thais mancinella inhabited by Cal­ 1999; Palawan: 5 paratypes (USNM 187567), cinus latens and Dardanus lagopodes, Sombrero from Drupella rugosa and an unidentified Island, 30 January 1999; 3 paratypes (USNM gastropod shell inhabited by Paguristes runya­ 187555), from Conus sp., Drupella cornus, and nac, Magbautoc Island, 16 March 1999; 3 Latirus turritus inhabited by Calcinus latens paratypes (USNM 187568), Conus sp. and Dru­ and an unidentified hermit crab, Sepoc Point, pella rugosa inhabited by Calcinus latens and 5 July 1997; Oriental Mindoro: 7 paratypes D. lagopodes, Apulit Island, 15 March 1999; A,B D

FIGURE 12. Polydora umangivora, n. sp.: holotype (USNM 187553) (B, D) and paratype (USNM 187570) (A, C, E). A, Anterior end, dorsal view; B, posterior end, dorsal view; C, spines and companion setae offifth segment, dorsal view; D, anterior end, dorsal view; E, bidentate hooded hook from anterior segment. Scale: A, B, 200 !lm; D, 300 !lm; C, E, 30 !lm. 452 PACIFIC SCIENCE· October 2001

1 paratype (USNM 187569), from Cerithium Branchiae begin on segment 7, continuing tel/enum Sowerby inhabited by Paguristes to terminal segments of body; attaining full runyanae, Apulit Island, 17 March 1999; size at segment 12. Dorsal ciliary bands on Cebu: 38 paratypes (USNM 187570: 35 para­ segments 1-4 and 7 to terminal segments types; USNM 187571: 3 paratypes, on 6 SEM (Figures 12D, l3A,C). stubs), from Astralium rhodostoma (Lamarck), Pygidium broad cup-shaped with non­ Bursa sp., Cantharus sp., Conus sp., Drupella motile cilia on rim of pygidium (Figures 12B, cornus, Strombus sp., Turbo sp., and an un­ 13B). identified gastropod shell inhabited by Calci­ ETYMOLOGY: The species is named after nus gaimardii, C. latens, Clibanarius englaucus, its behavior of ingesting (vora, to eat in Dardanus lagopodes, and an unidentified her­ Greek) the embryos of host hermit crabs mit crab, Olango Island, 9 July 1997. Indo­ (umang, hermit crab in the Ilokano language nesia: Bali: 2 paratypes (USNM 187572), from of the Philippines). Bursa sp. and Drupella cornus inhabited by REMARKS: Polydora wnangivora belongs to Calcinus gaimardii, Sanur, 6 August 1997. the Polydora ciliata/websteri group and most DESCRIPTION: 31.7 mm long, 0.8 mm closely resembles P. woodwicki Blake & Ku­ wide at segment 7; 186 segments. Ovigerous denov and P. pacifica Takahashi. Polydora female with ova in segments 27-107. Prosto­ umangivora differs from P. woodwicki in lack of mium rounded to slightly indented when eyes, lack of notosetae on segment 5, major viewed ventrally; caruncle short, terminating spines with distinct accessory tooth, and on segments 1-2; occipital tentacle and eyes larger cup-shaped pygidium rather than four absent (Figures 12A,D, l3A,C). Palps extend eyes, notosetae present on segment 5, spines posteriorly for 10-12 segments. In life, palps with accessory flange, and cuff-shaped pygi­ and body are pale yellow to orange; no pig­ dium. mentation present. After preservation color Polydora umangivora differs from P. pacifica usually dark yellow/orange on dorsal and in rounded prostomium, caruncle length, ventral sides of prostomium, peristomium, number and morphology ofmajor spines, and and all segments (Figure 12A). habitat. Polydora pacifica is a commensal of Segment 1 with neurosetae, without noto­ pearl oysters, Pinctada margaritifera (Lin­ setae, with weakly developed notopodiallobes naeus), from the Palau Islands. Polydora (Figure l3A,C). Winged capillary notosetae umangivora reaches maximal lengths of ap­ of segments 2-4, 6, and subsequent segments proximately 32 mm and 190 segments and has in three rows (Figure l3D); no specialized only four to seven exposed major spines; P. posterior notosetae. Winged capillary neuro­ pacifica reaches lengths of 85 mm and 390 setae of segments 2-4, 6, and subsequent segments and has up to 17 major spines. Ta­ segments in two rows (Figure l3D); five bi­ kahashi (1937:159-160, fig. 8) indicated that dentate hooded hooks begin on segment 7, the hooded hooks of this species have a pro­ up to 11 in series at segment 23, not accom­ cess below the main fang; his figure of the panied by capillaries; hooks with approxi­ hooded hook shows a spur extending from mately right angle between main fang and the shaft ofthe hooded hook toward the main shaft, wide angle between main fang and api­ fang. Such a process has not been noted for cal tooth, with constriction on shaft, with fine any other polydorid species. Unfortunately, bristles on hood (Figures 12E, l3P). the presence of this structure could not be Segment 5 almost twice as large as seg­ confirmed because attempts to locate type ments 4 and 6, with slightly curved row offive specimens were unsuccessful. exposed major spines and one developing VARIABILITY: All specimens exhibited spine alternating with pennoned companion rounded prostomium and short caruncle ex­ setae; posterioventral fascicle of winged neu­ tending to the middle of segment 1 through rosetae; notosetae absent (Figure l3D,E). the end of segment 2 (n = 45). In some Major spines falcate, with lateral tooth (Fig­ specimens the caruncle was in the form of a ures 12C, l3D,E). raised ridge or keel. Approximately 67% of FIGURE 13. Polydora umangivora, n. sp.: paratypes (USNM 187571), SEM micrographs. A, Anterior end, dorsal view, arrowhead indicates segment 2 with two parapodia on right side; B, posterior end, dorsal view; C, anterior end, dorsal view; D, segments 4 and 5 with modified spines and accompanying setae; E, spines and companion setae of fifth seg­ ment, dorsal view; F, hooded hooks, apical view. Scale: A-C, 100 !lm; D, E, 10 l!ID; F, 1 !lm. 454 PACIFIC SCIENCE· October 2001 those specimens examined (n = 46) possessed Winged capillary notosetae of segments 2-4, distinct bright yellow/orange coloring that 6, and subsequent segments in two rows; in developed after preservation and covered the segments 2-4 anterior and posterior rows dorsal and ventral sides of anterior and pos­ subequal, with anterior row slightly shorter; terior body segments. One specimen pre­ on segment 6 anterior row geniculate and pared for SEM was unique in possessing two approximately half the length of the posterior parapodia on the right side of segment 2 row; reduced to three to five bilimbate noto­ (Figure 13A). setae in posterior segments; no specialized ECOLOGY: Polydora umangivora constructs posterior notosetae. Winged capillary neuro­ V-shaped burrows within gastropod shells setae of segments 2-4, 6, and subsequent inhabited by hermit crabs. The species has segments in two rows, anterior row shorter been found to ingest the embryos attached to then posterior; two to three bidentate hooded the pleopods of three host hermit crabs: Cal­ hooks begin on segment 7, up to six in series cinus gaimardii, Dardanus lagopodes, and Pa­ at segment 14, not accompanied by capil­ guristes runyanae. This report represents the laries; hooks with approximately right angle second polydorid documented to prey on between main fang and shaft, wide angle be­ host hermit crab embryos. The feeding biol­ tween main fang and apical tooth, with slight ogy and ecology of the species will be treated constriction on shaft (Figure 14E). in forthcoming papers (unpubl. data). Segment 5 slightly larger than preceding DISTRIBUTION: Philippines: Bataan, Ba­ segments; row of two to four exposed major tangas, Oriental Mindoro, Palawan, Aldan, spines alternating with pennoned companion Cebu; Indonesia: Bali. setae; with posterioventral fascicle of three to four winged neurosetae; anteriodorsal fascicle Polydora mabinii Williams, n. sp. of two to four large geniculate notosetae with Figure 14 fine bristles (Figure 14D,F). Major spines simple, falcate (Figure 14D,F). HOLOTYPE: Philippines: Batangas (USNM Branchiae begin on segments 8-9, con­ 187575), from Drupella cornus inhabited by tinuing to middle portion of body; attaining Calcinus latens, Sombrero Island, 13 July 1997. full size at segment 11. OTHER MATERIAL EXAMINED: Philip­ Pygidium cup-shaped with dorsal gap; pines: Batangas: 1 paratype (USNM 187573), covered with small papillae (Figure 14C). from Latirus turritus inhabited by an un­ ETYMOLOGY: The species is named after identified hermit crab, Anilao, 28 June 1997; Mabini, a small port on Batangas Bay. The 1 paratype (USNM 187574), from Drupella cor­ species is dedicated to the boatmen who work nus inhabited by Calcinus latens, Sombrero at the port and made travel to surrounding Island, 13 July 1997. islands possible. DESCRIPTION: Holotype broken, consist­ REMARKS: Polydora mabinii belongs to the ing of 33 anterior segments. Largest complete Polydora ciliata/websteri group and most specimen measures 15.7 mm long, OJ mm closely resembles P. aggregata Blake and P. wide at segment 7; 97 segments. Ovigerous limicola. Polydora mabinii differs from P. ag­ female with ova in segments 20-58. Prosto­ gregata by possessing simple, falcate major mium bifid; caruncle terminating on seg­ spines and anteriodorsal group oflarge genic­ ments 3-4; occipital tentacle absent; two or ulate setae, caruncle extending to segments four large eyes arranged in trapezoid (Figure 3-4, and number of total segments (up to 97 14A) or eyes absent. Palps extend posteriorly segments and 16 mm long) instead of seg­ for 14-15 segments. In alcohol small spots of ment 5 with major spines with small elevation black pigmentation sparsely distributed along and anteriodorsal group of winged capillary dorsal side and pygidium in some specimens; notosetae, caruncle terminating bluntly on body opaque white. segment 2, and small number of segments Segment 1 with neurosetae, without noto­ (35-40 segments and 6-8 mm long) (Blake setae, with well-developed notopodial lobes. 1971). Polydora mabinii differs from P. limicola A

A,B

c

D,F

E

FIGURE 14. Polydora mabinii, n. sp.: holotype (USNM 187575) (A, B, D-F) and paratype (USNM 187573) (C). A, Anterior end, dorsal view; B, anterior end, lateral view; C, posterior end, dorsal view; D, fifth segment, lateral view ofleft side; E, bidentate hooded hook from anterior segment; F, fifth segment, dorsal view ofright side. Scale: A, B, C, 100 1llJl; D, E, F,30J.lm. 456 PACIFIC SCIENCE· October 2001 in lack of palp pigmentation, segment 5 with Oriental Mindoro: 12 spec. (USNM 187581), simple, falcate major spines and anteriodorsal from Astralium rhodostoma, Conus sp., Cypraea group of large geniculate setae instead of sp., Drupa rubusidaeus, Drupella cornus, D. ru­ palps with black pigment bands, and segment gosa, Latirolagena smaragdula, Strombus labi­ 5 with major spines with small tooth and an­ atus labiatus, and an unidentified gastropod teriodorsal group of winged capillary notose­ shell inhabited by Calcinus gaimardii, C. latens, tae (Hartman 1961, Blake 1996). In addition, and C. minutus, Puerto Galera: Coco Beach, mature Polydora mabinii possess only four to 12 January 1999; 4 spec. (USNM 187582), from five major spines in the fifth segment, but P. Drupella cornus inhabited by Calcinus gaimar­ limicola contains 8-15 major spines. Diffi­ dii, C. latens, and C. minutus, Puerto Galera: culties in the taxonomy ofP. limicola exist, and Coco Beach, 14 January 1999; 2 spec. (USNM reports of this species may refer to more than 187583), from Drupella cornus and an uniden­ one species (Williams and Radashevsky 1999). tified gastropod shell inhabited by Calcinus ECOLOGY: Polydora mabinii has been found gaimardii, Puerto Galera: Coco Beach, 15 Jan­ associated with gastropod shells (Drupella uary 1999; 3 spec. (USNM 187584), from Dru­ cornus and Latirus turritus) inhabited by her­ pella cornus inhabited by Calcinus gaimardii, mit crabs (Calcinus latens). It is unknown if the C. latem, and an unidentified hermit crab, species burrows into occupied shells or con­ Bayanan Beach, 13 January 1999; Aklan: 1 structs tubes within the crevices of the shells. spec. (USNM 187585), from Strombus labiatus The presence of three gastropod veliger lar­ labiatus inhabited by Calcinus latem, Boracay: vae within the gut of one of the specimens Rocky Beach, 12 April 1999; 3 spec. (USNM indicates that the species is a suspension 187586), from Cantharus undosus, Conus sp., feeder, capable of feeding on motile zoo­ and an unidentified gastropod shell inhabited plankton. by Calcinus gaimardii and C. latens, Boracay: DISTRIBUTION: Philippines: Batangas. Rocky Beach, 15 April 1999; Palawan: 2 spec. (USNM 187587), from Drupella rugosa in­ Polydora robi Williams, 2000 habited by Paguristes runyanae, Magbautoc Island, 16 March 1999; 2 spec. (USNM Polydora robi Williams, 2000:539, figs. 1-4. 187588), from Tectus sp. and an unidentified gastropod shell inhabited by Calcinus gai­ MATERIAL EXAMINED: Philippines: Ba­ mardii and Paguristes runyanae, Apulit Island, taan: 12 spec. (USNM 187576), from Cantharus 17 March 1999; 1 spec. (USNM 187589), from undosus, Cypraea sp., Latirolagena smaragdula, Peristernia nassatula (Lamarck) uninhabited and an unidentified gastropod shell inhabited by hermit crab, Apulit Island, 18 March 1999. by Calcinus gaimardii and Dardanus lagopodes, DESCRIPTION: Up to 30.4 mm long, 1.1 Mabayo, 21 February 1999; 5 spec. (USNM mm wide at segment 7; 171 segments. Pro­ 187577), from Conus sp., Latirolagena sma­ stomium rounded; caruncle extending to ragdula, and an unidentified gastropod shell middle of segment 2; short, triangular occipi­ inhabited by Calcinus gaimardii, Mabayo, 28 tal tentacle at anterior margin of segment 1; February 1999; 1 spec. (USNM 187578), from eyes usually absent but two or four may be an unidentified gastropod shell inhabited by present in a row in juveniles. Palps extend Diogenes sp., Morong, 6 February 1999; 3 posteriorly for 10-15 segments. In life, palps spec. (USNM 187579), from Cantharus undosus, and body pale yellow, without pigmentation; Cypraea sp., and an unidentified gastropod in alcohol body opaque white. shell inhabited by Calcinus gaimardii and Cal­ Segment 1 with neurosetae, without noto­ cinus sp., Morong, 25 April 1999; Batangas: 9 setae. Winged capillary notosetae ofsegments spec. (USNM 187580), from Conus sp., Cor­ 2-4,6, and subsequent segments in two rows alliophila neritoidea, Cypraea sp., Drupella cor­ and a superior group of longer and thinner nus, D. rugosa, and Thais armigera (Link) setae; notosetae in posterior one-third of inhabited by Calcinus gaimardii, C. latens, C. body in flared or cone-shaped bundles offine minutus, C. pulcher, Ciliopagurus strigatus, and needlelike spines protruding through cuticle; Dardanus lagopodes, Anilao, 13 February 1999; approximately 85-95 spines in posterior Polydora and Related Genera Associated with Hermit Crabs . Williams 457 segments with two to four longer anterior Cebu (Williams 2000; this paper); Indonesia: notosetae. Winged capillary neurosetae of Bali (Williams 2000). segments 2-4, 6, and subsequent segments in two rows and an inferior tuft of capillaries. Genus Tripolydora Woodwick, 1964 Bidentate hooded hooks begin on segment 7, 15-27 hooks in middle body segments, not Tripolydora Woodwick, 1964:155. Type spe­ accompanied by capillaries; hooks with ap­ cies: Tripolydora spinosa Woodwick, 1964, proximately right angle between main fang by monotypy. and apical tooth, with constriction on shaft, accompanying setae absent. DIAGNOSIS: Prostomium rounded, ex­ Segment 5 almost twice as large as seg­ tending posteriorly as a broad caruncle to ments 4 and 6, with slightly curved row of segment 3. Segment 1 without notosetae, four to eight exposed major spines alternating notopodial lobes greatly reduced or absent. with pennoned companion setae; with poste­ Segment 4 with anterior row of modified rioventral fascicle of four to nine winged notosetae with inflated sheaths around a pro­ neurosetae; notosetae absent. Major spines truding shaft. Modified spines of segment 5 falcate, with lateral obliquely curved flange. with terminal tooth and short knobs. Seg­ Tear-shaped gland on each side of the fifth ments 7-10 with anterior row of unilimbate segment, ventral to the major spines with a capillary notosetae containing riblike sheaths; duct leading to the sides of segment 5; gland posterior notosetae in bundles of long capil­ in juveniles only. laries. Unilimbate capillary neurosetae re­ Branchiae begin on segment 7, continuing placed by tridentate hooded hooks on to terminal segments of body, small at first, segment 9. Tridentate hooded hooks lack attaining full size at segment 10, diminishing constriction on shaft, accompanied by capil­ in size in posterior segments. Dorsal ciliary lary setae. Branchiae on segments 2-4, 6, and bands begin on segment 7. Posterior seg­ continuing to posterior segments. Pygidium ments with well-developed digitiform noto­ composed of two ventral lappets and two podia and enlarged neuropodia. smaller dorsal lappets. Pygidium small, with digitiform composite cirri surrounding anus; anal cirri longer on Tripolydora spinosa W oodwick, 1964 ventral side of pygidium, with irregular Figure 15 knoblike projections and nonmotile cilia. REMARKS: Polydora robi was recently de­ Tripolydora spinosa Woodwick, 1964:155-157, scribed by Williams (2000), who provided fig. 4(6-9); Kohn and Lloyd, 1973b:700; data on the variation, ecology, and feeding Blake and Woodwick, 1981:352-362, figs. biology of the species. Previously the largest 1-5; Ward, 1987:368-369, fig. 3.II.I48. recorded specimen was 25.0 mm long for 117 segments. The original description was based MATERIAL EXAMINED: Philippines: Ba­ on specimens collected from three provinces tangas: 1 spec. (USNM 187590), from Drupella of the Philippines (Batangas, Cebu, and Min­ comus inhabited by Calcinus latens, Anilao, 13 doro) and Indonesia. Polydora robi has now February 1999. Chile: Easter Island: 1 spec. been found in six provinces of the Philippines (USNM 49537), in tide pool, between Hanga and is widely distributed in the country. Poly­ Roa and Hanga Pika, 15 February 1969, leg. dora robi ranges from 24 to 171 segments J. Randall. (75 ± 25, n = 111) and 3-41 mm (16.4 ± 9.5, DESCRIPTION: 6.6 mm long, 0.4 mm wide n = 40) in size. Polydora robi was the first at segment 7; 39 segments. Ovigerous female polydorid documented to feed on host hermit with ova in segments 12 to 26. Prostomium crab eggs. The ecology, feeding biology, re­ rounded; broad caruncle reaching end of production, and larval development are segment 2; occipital tentacle and eyes absent treated in forthcoming papers (unpubl. data). (Figure 15A). Palps extend posteriorly for 15 DISTRIBUTION: Philippines: Bataan, Ba­ segments. In life, black pigmentation bands tangas, Oriental Mindoro, Palawan, Aldan, on palps, prostomium, and dorsal side of an- 458 PACIFIC SCIENCE· October 2001

A

-A -B C·F,I,J -G,H E F

FIGURE 15. Tripolydora spinosa Woodwick (USNM 187590). A, Anterior end, dorsal view; B, posterior end, dorsal view; C, major spines offifth segment, dorsal view; D, major spines offifth segment, ventral view; E, fifth segment notosetae, anterior row; F, fifth segment notosetae, posterior row; G, modified setae of the fourth segment; H, notosetae of the fourth segment, middle row; I, unilimbate capillary notosetae of segment 7; J, tridentate hooded hook from anterior segment. Scale: A, 100 1JllI; B, 50 J.lm; C-F, I, J, 30 J.lm; G, H, 15 1JllI.

terior and posterior segments (Figure 15A). tae of segments 2-10 arranged in three rows; After preservation, black pigmentation bands no specialized posterior notosetae. Notosetae absent on palps; color in alcohol opaque of segments 2-3 and 6 are unilimbate capil­ white. laries. Segment 4 bears four notosetae with Segment 1 with neurosetae; greatly re­ inflated sheaths containing bristles in anterior duced notopodia, without notosetae. Notose- row; shaft of notosetae protruding from Polydora and Related Genera Associated with Hennit Crabs Williams 459 sheath (Figure 15G); middle row with short DISCUSSION bilimbate notosetae with bristles, posterior row with long bilimbate notosetae (Figure Table 1 summarizes data on the diversity of ISH). Segments 7-10 possess unilimbate polydorids in seven of the best-studied areas capillary notosetae with riblike sheaths (Fig­ of the Pacific. The central Pacific and Indo­ ure 151); approximately eight unilimbate West Pacific areas (Philippines, Hawai'i, and notosetae at segment 30 increasing to 24 at Marshall Islands) contain approximately segment 36 and subsequent segments. Neu­ equal numbers of polydorids (7-14 species rosetae of segments 2-8 with an anterior row among four to six genera), whereas North of four to five pennoned setae and seven to and Southwest Pacific areas contain approxi­ eight longer unilimbate setae; four tridentate mately two to three times as many species hooded hooks begin on segment 9 (Figure (26-30 species among six to seven genera). 15J), initially accompanied by five to seven The coral reef areas of the Indo-West Pacific unilimbate capillaries, up to six hooded hooks harbor some ofthe most highly diverse faunal in series at segment 11; unilimbate capillaries assemblages in the world. In particular, a replaced by two to three simple capillaries small triangle (termed the East Indies Trian­ and sabre setae on segment 27. gle) formed by the Philippines, the Malay Segment 5 approximately same size as Peninsula, and New Guinea is home to the preceding segments, with three major spines greatest concentration of species within many and a ventral row of bilimbate neurosetae; tropical marine groups (Briggs 1999) includ­ two dorsal rows of bilimbate notosetae, pos­ ing polychaetes (Knox 1957, Bailey-Brock terior row longer than anterior row (Figure 1995). The lower abundance of polydorids in 15E,F). Major spines with extended tooth and the Indo-West Pacific is most likely due to two short knobs; worn spines with reduced lack of sampling rather than a true represen­ tooth (Figure 15C,D). tation of the number of species in these areas. Branchiae begin on segment 2, continuing In fact, sampling efforts for spionids (in to segment 24; attaining full size at segment terms of numbers of researchers sampling 7. as well as regions and habitats covered) in Pygidium composed of two ventral lappets Japan and Australia have been considerably and two dorsal lappets; lappets possessing greater than in Indo-West Pacific areas such nonmotile cilia (Figure 15B). as the Philippines (see Blake and Kudenov REMARKS: The single Philippine specimen 1978, Radashevsky 1993, Radashevsky and agrees well with the published accounts ofthe Hsieh 2000a,b). My study was restricted to species (Woodwick 1964, Blake and Wood­ polydorids associated with hermit crabs, a wick 1981, Ward 1987). The pigmentation relatively small group of species that are observed has not been previously noted, pre­ predominately burrowers of calcareous sub­ sumably because it disappears after preserva­ strata. When compared with the numbers of tion. burrowers found in other areas, the diversity ECOLOGY: The specimen was in a mud in the Philippines is closer to that in the tube within the gastropod shell Drupella cor­ North and Southwest Pacific (Table 1). nus, inhabited by Cakinus latens. The species Of the 10 polydorid species recorded from elsewhere is known to inhabit mud tubes the Philippines and Indonesia, four species constructed among coral rock, sand, algae, (Dipolydora tridenticulata, Polydora umangivora, and sponges in addition to burrowing into Polydora mabinii, and Polydora robi) are re­ dead coral (Woodwick 1964, Blake and stricted to the Indo-West Pacific. Carazziella Woodwick 1981, Ward 1987). reishi is found throughout the Indo-West Pa­ DISTRIBUTION: Chile: Easter Island cific, central Pacific islands, and Japan. Simi­ (Blake and Woodwick 1981); Hawai'i (Ward larly, Tripolydora spinosa is found throughout 1987); Marshall Islands: Eniwetok Atoll the Indo-West Pacific, central Pacific islands, (Woodwick 1964); Philippines: Batangas and Easter Island. Polydora cavitensis Pillai was (first record: this paper). previously recorded from among oysters in 460 PACIFIC SCIENCE· October 2001

TABLE 1 Biodiversity of Nine Polydorid Genera within Seven Areas of the Pacific

Marshall New Northwest Genus Hawai'ia Islands Zealand Australia Pacific Philippines Taiwan

Amphipolydorab Boccardia 1 (1) 8 (5) 4 (1) 2 (1) Boccardiella 1 (1) 1 2 (1) 1 (1) Carazziella 1 (1) 1 (1) 4 2 (1) 1 (1) Dipolydora 2 (2) 2 (2) 3 (3) 8 (6) 12 (9) 3 (3) Polydora 5 (2) 2 (2) 7 (5) 7 (6) 5 (3) 5 (1) Polydorella 2 2' Pseudopolydora 3 (1) 3 (2) 3 2 8 Tripolydora 1 (1) 1 (1) 1 (1) Total 14 (9) 7 (6) 15 (10) 30 (13) 26 (17) 13 (9) 13 (1)

Note: Numbers indicate total species recorded within each genus; numbers in parentheses indicate species known to burrow in calcareous substrates. References: Hawai'i: Abbott 1946, Ward 1981, 1987, Bailey-Brock 1990, 2000; Marshall Islands: Woodwick 1964; New Zealand: Rainer 1973, Read 1975; Australia: Blake and Kudenov 1978, Hutchings and Turvey 1984; Northwest Pacific: Imajima and Hartman 1964, Radashevsky 1993, 1994a,b, Sato-Okoshi 1998, 1999; Philippines: Pillai 1965, Williams 2000; Taiwan: Radashevsky and Hsieh 2000a,b.

fl Hawai'i contains two polydorids (Polydora nuchalis and Boccardia proboscidea) introduced via aquacultural products (Bailey-Brock 1990, 2000). b Amphipolydora is a monotypic genus known only from Argentina (Blake 1983).

C Two Polydorella species have recendy been found associated with sponges from the Philippines (unpubl. data).

the northern Philippines (Pillai 1965). The ACKNOWLEDGMENTS species has not been subsequently reported by any authors and was not encountered during I thank Filipina B. Sotto for access to space at this study; the species appears to be endemic the Marine Laboratory of the University of to the Philippines. As currently defined, Di­ San Carlos and Jason G. Young for his field polydora armata and D. socialis are widely dis­ assistance. I appreciate the help of Paul Cas­ tributed species in Atlantic and Pacific waters. sidy (Western Washington University) and Previously, Boccardia berkeleyorum was Alan W. Harvey (Georgia Southern Univer­ known only from the eastern Pacific (Cali­ sity) in identification of hermit crabs, John fornia and Vancouver Island). This report Clamp (North Carolina Central University) represents the first record of the species in in identification of ciliates, and Christopher the Indo-West Pacific and represents a large B. Boyko (University of Rhode Island and the range extension. Introductions of spionid American Museum of Natural History) in polychaetes are well documented from nu­ identification of gastropod species. Linda merous areas around the world via ballast Ward (National Museum ofNatural History) water and aquacultural products (Bailey­ kindly tracked down older literature refer­ Brock 1990, 2000, Carlton and Geller 1993, ences. I thank my committee members, Blake 1996, Rohner et al. 1996). For example, Robert Bullock, Emily Carrington, Barbara Polydora nuchalis W oodwick was presumably Sullivan, and James Blake, for their reviews of introduced to Hawai'i through shipments of early versions of the manuscript. shrimp from Mexico. Similarly, Boccardia pro­ Literature Cited boscidea was recently introduced to Hawai'i by shipments of Ostrea edulis Linnaeus from Abbott, D. P., Jr. 1946. Some polychaetous Maine (Bailey-Brock 1990, 2000). It is possi­ annelids from a Hawaiian fish pond. Univ. ble that Boccardia berkeleyorum was introduced Hawaii Res. Publ. 23:5-24. to the Philippines through such human influ­ Bailey-Brock, J. H. 1990. Polydora nuchalis ence. The species is a burrower of calcareous (Polychaeta: Spionidae), a new Hawaiian substrates and could have impacts on mari­ record from aquaculture ponds. Pac. Sci. culture in the Philippines. 44:81-87. Polydora and Related Genera Associated with Hermit Crabs . Williams 461

---. 1995. Polychaetes of western Pacific (Polychaeta: Spionidae) from the Gala­ Islands: A review of their systematics and pagos Islands and redescription of Boc­ ecology. Pages 1-424 in M. N. A Pe­ cardia basi/aria Hartman from southern terson, J. E. Maragos, L. G. Eldredge, J. E. California. Bull. South. Calif. Acad. Sci. Bardach, and H. F. Takeuchi, eds. Marine 85:16-21. and coastal biodiversity in the tropical --.1996. Family Spionidae Grube, 1850. island Pacific Region. East-West Center, Including a review of the genera and spe­ Honolulu. cies from California and a revision of the ---. 2000. A new record ofthe polychaete genus Polydora Bose, 1802. Pages 81-223 Boccardia proboscidea (Family Spionidae), in J. A Blake, B. Hilbig, and P. H. Scott, imported to Hawai'i with oysters. Pac. Sci. eds. Taxonomic atlas of the benthic fauna 54:27-30. of the Santa Maria Basin and western Berkeley, E. 1927. Polychaetous annelids Santa Barbara Channel. Vol. 6. Santa Bar­ from the Nanaimo District. 3. Leodicidae bara Museum of Natural History, Santa to Spionidae. Contrib. Can. BioI. Fish., Barbara, California. n. s., 3:405-422. Blake, J. A, and P. L. Arnofsky. 1999. Re­ Berkeley, E., and C. Berkeley. 1936. Notes production and larval development of the on Polychaeta from the coast of western spioniform Polychaeta with application to Canada. 1. Spionidae. Ann. Mag. Nat. systematics and phylogeny. Hydrobiologia Hist., ser. 10, 18:468-476. 402:57-106. --. 1952. Annelida. Polychaeta Seden­ Blake, J. A, and J. W. Evans. 1973. Polydora taria. Can. Pac. Fauna 9b. Fisheries Re­ and related genera as borers in mollusk search Board of Canada Part 2:1-139. shells and other calcareous substrates. Blake, J. A 1971. Revision of the genus Poly­ Veliger 15:235-249. dora from the east coast of North America Blake, J. A, and J. D. Kudenov. 1978. The (Polychaeta: Spionidae). Smithson. Con­ Spionidae (Polychaeta) from southeastern trib. Zool. 75:1-32. Australia and adjacent areas with a revision --. 1975. Phylum Annelida: Class Poly­ of the genera. Mem. Natl. Mus. Vic. chaeta. Pages 151-243 in R. I. Smith andJ. 39:171-280. T. Carlton, eds. Light's Manual, Intertidal ---. 1981. Larval development, larval nu­ invertebrates of the central California trition, and growth for two Boccardia spe­ coast. University of California Press, cies (Polychaeta: Spionidae) from Victoria, Berkeley. Australia. Mar. Ecol. Prog. Ser. 6:175­ ---. 1979a. Four new species of Caraz­ 182. ziella (Polychaeta: Spionidae) from North Blake, J. A., and N. J. Maciolek. 1987. A re­ and South America, with a redescription of description of Polydora cornuta Bose (Poly­ two previously described forms. Proc. BioI. chaeta: Spionidae) and designation of a Soc. Wash. 92:466-481. neotype. Proc. BioI. Soc. Wash. 7:11-15. ---. 1979b. Revision of some polydorids Blake, J. A., and K. H. Woodwick. 1971. A (Polychaeta: Spionidae) described and review of the genus Boccardia Carazzi recorded from British Columbia by Edith (Polychaeta: Spionidae) with descriptions and Cyril Berkeley. Proc. BioI. Soc. Wash. of two new species. Bull. South. Calif. 92:606-617. Acad. Sci. 70:31-42. ---. 1981. Polydora and Boccardia species ---. 1981. The morphology of Tripolydora (Polychaeta: Spionidae) from western spinosa Woodwick (Polychaeta: Spionidae): Mexico, chiefly from calcareous habitats. An application of the scanning electron Proc. BioI. Soc. Wash. 93:947-962. microscope to polychaete systematics. --. 1983. Polychaetes of the family Proc. BioI. Soc. Wash. 94:352-362. Spionidae from South America, Antarctica Bose, L. A G. 1802. Histoire naturelle des and adjacent seas and islands. Antarct. Res. vers, contenant leur description et leurs Ser. 39:205-288. moeurs, avec figures dessinees d'apres na­ ---. 1986. A new species of Boccardia ture. Vols. 1-3: 1-324. Deterville, Paris. 462 PACIFIC SCIENCE· October 2001

Briggs, J. C. 1999. Coincident biogeographic velle espece de spionide (annelides, poly­ patterns: Indo-West Pacific Ocean. Evo­ chetes) Boccardia semibranchiata. Ann. Inst. lution 53:326-335. Oceanogr.66:37-45. Buckley, W. J., and J. P. Ebersole. 1994. Hartman, O. 1941. Some contributions to the Symbiotic organisms increase the vulnera­ biology and life history of Spionidae from bility of a hermit crab to predation. J. Exp. California. Allan Hancock Found. Pac. Mar. BioI. Eco1. 182:49-64. Exped.7:289-323. Buzhinskaja, G. N. 1985. Polychaeta of the ---. 1954. Marine annelids from the shelf off South Sakhalin and their ecology. northern Marshall Islands. Prof. Pap. U.S. Issled. Fauny Morei SSSR 30:72-224. Geo1. Surv. (no. 260): 615-644. Caceres-Martinez, J., G. D. Tinoco, M. L. U. ---. 1961. Polychaetous annelids from Bustamante, and I. M. Gomez-Humaran. California. Allan Hancock Pac. Exped. 1999. Relationship between the burrowing 25:1-226. worm Polydora sp. and the black clam ---. 1969. Atlas of the sedentariate Chione fluctifraga Sowerby. J. Shellfish Res. polychaetous annelids from California. 18:85-89. Allan Hancock Foundation, University of Carazzi, D. 1893. Revisione del genero Poly­ Southern California, Los Angeles. dora Bose, e cenni su due specie che vivono Hartmann-Schroder, G. 1962. Die Poly­ sulle ostriche. Mitt. Zoo1. Stat. Neapel chaeten des Eulitorals. Pages 57-167 in G. 11:4-45. Hartmann-Schroder and G. Hartmann, Carlton, J. T., and J. B. Geller. 1993. Eco­ eds. Zur Kenntnis des Eulitorals der chi­ logical roulette: The global transport of lenischen Pazifikkiiste und argentinis­ nonindigenous marine organisms. Science chen Kiiste. Mitt. Hamb. ZOO1. Mus. Inst. (Washington, D.C.) 261:78-82. 60. Carrasco, F. D. 1974. Spionidae (Polychaeta) ---. 1965. Die Polychaeten des Sub­ provenientes de la Bahia de Concepcion y litorals. Pages 59-305 in G. Hartmann­ lugares adyacentes. Bo1. Soc. BioI. Con­ Schroder and G. Hartmann, eds. Zur cepcion 48:185-201. Kenntnis des Sublitorals der chilenischen Czerniavsky, V. 1881. Materialia ad zoo­ Kiiste under besonderer Berucksichtigung graphiam Ponticam comparatum. 3 der Polychaeten und Ostracoden. Mitt. Vermes. Bull. Soc. Imp. Nat. Moscou Hamb. Zoo1. Mus. Inst. 60. 56:338-420. ---. 1979. Die Polychaeten der tropi­ Day, J. H. 1954. The Polychaeta of Tristan schen Nordwestkiiste Australiens (zwi­ da Cunha. Results of the Norwegian Sci­ schen Derby im Norden und Port entific Expedition to Tristan da Cunha Hedland im Siiden). Pages 75-218 in G. 1937-1938. Nor. Vidensk. Akad. Oslo Hartmann-Schroder and G. Hartmann, Arbok. 29:1-35. eds. Zur Kenntnis der Eulitorals der ---. 1967. A monograph on the Poly­ australischen Kiisten unter besonderer chaeta of southern Africa. Br. Mus. (Nat. Berucksichtigung der Polychaeten und Rist.) Pub1. 656:1-878. Ostracoden (Teil 2 und Teil 3). Mitt. EWers, E. 1905. Neuseelandische Anneliden. Hamb. Zoo1. Mus. Inst. 76. Abh. Koniglichen Ges. Wiss. GOttingen, Haswell, W. F. 1885. On a destructive para­ Mathematisch-Physikalische Klasse 3: 1­ site of the rock oyster. Proc. Linn. Soc. 80. N.S.W.1O:273-275. Fauvel, P. 1927. Polychetes sedentaires. Ad­ Hoberg, M. K., S. G. McGee, and H. M. denda aux Errantes, Archiannelides, My­ Feder. 1982. Polychaetes and amphipods zostomaires. Faune de France 16:1-494. as commensals with pagurids from the Foster, N. M. 1971. Spionidae (Polychaeta) Alaska shelf. Ophelia 21:167-179. of the Gulf of Mexico and the Caribbean Hutchings, P. A., and S. P. Turvey. 1984. Sea. Stud. Fauna Curacao Other Caribb. The Spionidae of South Australia (Annel­ lsI. 36:1-183. ida: Polychaeta). Trans. R. Soc. S. Aust. Guerin, J. P. 1990. Description d'une nou- 108:1-20. Polydora and Related Genera Associated with Hermit Crabs . Williams 463

(ICZN) International Commission on Zoo­ Haliotis diversicolor aquatilis chiefly on spe­ logical Nomenclature. 1999. International cies Polydora. Bull. Jpn. Soc. Sci. Fish. Code of Zoological Nomenclature, 4th ed. 48:31-35. ICZN, London. Langerhans, P. 1880. Die Wurmfauna von Imajima, M., and O. Hartman. 1964. The Madeira. Z. Wiss. Zool. 34:87-143. polychaetous annelids of Japan. Part II. Lewis, J. B. 1998. Reproduction, larval de­ Allan Hancock Found. Occas. Pap. 26: velopment and functional relationships of 239-452. the burrowing, spionid polychaete Dipoly­ Jensen, K, and K Bender. 1973. Inverte­ dora armata with the calcareous hydrozoan brates associated with snail shells inhabited Millepora complanata. Mar. BioI. (Berl.) by Pagurus bernhardus (L.) (Decapoda). 130:651-662. Ophelia 10:185-192. Light, W. J. 1977. Spionidae (Annelida: Johnson, P. G. 1984. Family Spionidae Polychaeta) from San Francisco Bay, Cali­ Grube, 1850. Pages 6-1 to 6-99 in J. M. fornia: A revised list of nomenclatural Uebelacker and P. G. Johnson, eds. Taxo­ changes, new records, and comments on nomic guide to the polychaetes of the related species from the northeastern northern Gulf of Mexico. Barry A. Vittor Pacific. Proc. BioI. Soc. Wash. 90:66-88. and Associates, Inc., Mobile. ---. 1978. Invertebrates of the San Fran­ Johnston, G. 1838. Miscellanea Zoologica. cisco Bay estuary system. Family Spionidae Aricidae. Mag. Zool. Bot. 2:63-72. (Annelida, Polychaeta). The Boxwood Jones, M. L. 1962. On some polychaetous Press, Pacific Grove, California. annelids from Jamaica. Bull. Am. Mus. Lissner, A., C. Phillips, D. Cadien, R. Smith, Nat. Hist. 124:173-212. B. Bernstein, R. Cimberg, T. Kauwling, Karlson, R. H., and M. A. Shenk. 1983. Epi­ and W. Anikouchine. 1986. Assessment of faunal abundance, association, and over­ long-term changes in biological commun­ growth patterns on large hermit crab ities in the Santa Maria Basin and western shells. J. Exp. Mar. BioI. Ecol. 70:55-64. Santa Barbara Channel. Phase I. Final re­ Khlebovitsch, V. V. 1959. [Some species of port submitted for the Minerals Manage­ polychaete worms from the Kurile Islands, ment Service: U.S. Department of the which are new or recorded for the first Interior, Pacific OCS Office. Contract No. time in the U.S.S.R.] Zool. Zh. 28:167­ 14-12-0001-30032. NMS OCS Study, 181. MMS 86-0012. Kinberg, J. 1866. Annulata nova. Ofversight Martin, D. 1996. A new species of Polydora af Kungliga Vetenskaps-Akademiens For­ (Polychaeta, Spionidae) associated with the handlingar, Stockholm 22:239-258. excavating sponge Cliona viridis (Porifera, ---. 1910. Kongliga Svenska Fregatten Hadromerida) in the northeastern Medi­ Eugenies Resa omkring jorden under beW terranean Sea. Ophelia 45:159-174. af C.A. Virgin aren 1851-1853. Zoologi. Martin, D., and T. A. Britayev. 1998. Symbi­ 3. Annulater., pp. 1-78. otic polychaetes: Review of known species. Knox, G. A. 1957. The distribution of poly­ Oceanogr. Mar. BioI. Annu. Rev. 36:217­ chaetes within the Indo-Pacific. Proc. 8th 340. Pac. Sci. Congr. 3:403-411. Mobius, K 1874. Mollusken, Wiirmer, Echi­ Kohn, A. J., and M. C. Lloyd. 1973a. Poly­ nodermen und Coelenteraten. Die zweite chaetes of truncated reef limestone sub­ deutsche Nordpolarfahrt in den Jahren strata on eastern Indian Ocean coral reefs: 1869 und 1870 unter Fiihrung des Kiipi­ Diversity, abundance, and taxonomy. Int. tans Karl Koldewey. Herausgegeben von Rev. Gesamten Hydrobiol. 58:369-399. den Verein fuer di deutsche Norpo­ ---. 1973b. Marine polychaete annelids of larfahrtin Bremen. Wiss. Ergeb. 2:246­ Easter Island. Int. Rev. Gesamten Hydro­ 261. bioI. 58:691-712. Oersted, A. S. 1843. Annulatorum danicorum Kojima, H., and M. Imajima. 1982. Burrow­ conspectus. Naturhistorisk I Maricolae. ing polychaetes in the shells of abalone Tidsskrift Kebenhavn 4: 1-51. 464 PACIFIC SCIENCE· October 2001

Okuda, S. 1937. Spioniform polychaetes from Trans. San Diego Soc. Nat. Hist. 15:67­ Japan. J. Fac. Sci. Hokkaido Univ. 5:217­ 106. 254. Rice, S. A. 1978. Spermatophores and sperm Paxton, H., and L. M. Chou. 2000. Poly­ transfer in spionid polychaetes. Trans. Am. chaetous annelids from the South China Microsc. Soc. 97:160-170. Sea. Raffles Bull. Zool. Suppl. 8:209-232. Rohner, M., Bastrop, R, and K. Jiirss. 1996. Pettibone, M. H. 1967. Type-specimens of Colonization of Europe by two American polychaetes described by Edith and Cyril genetic types or species of the genus Mar­ Berkeley (1923-1964). Proc. U.S. Nat!. enzelleria (Polychaeta: Spionidae). An elec­ Mus. 119:1-23. trophoretic analysis of allozymes. Mar. Pillai, T. G. 1965. Annelida Polychaeta from BioI. (Berl.) 127:277-287. the Philippines and Indonesia. Ceylon J. Sato-Okoshi, W. 1998. Three new species of Sci. BioI. Sci. 5: 11 0-177. polydorids (Polychaeta, Spionidae) from Radashevsky, V. I. 1993. Revision of the ge­ Japan. Species Diversity 3:277-288. nus Polydora and related genera from the ---. 1999. Polydorid species (Polychaeta, North West Pacific (Polychaeta: Spioni­ Spionidae) in Japan, with descriptions of dae). Publ. Seto Mar. BioI. Lab. 36:1-60. morphology, ecology and burrow struc­ ---. 1994a. Species of the genus Polydora ture. 1. Boring species. J. Mar. BioI. Assoc. (Polychaeta, Spionidae) from the middle U.K. 79:831-848. Kurile Islands. Bull. Nat!. Sci. Mus. Ser. A Sato-Okoshi, W., and K. Okoshi. 1997. Sur­ (Zool.) 20:67-76. vey of the genera Polydora, Boccardiella and ---. 1994b. Life history of a new Polydora Boccardia (Polychaeta, Spionidae). Bull. species from the Kurile Islands and evolu­ Mar. Sci. 60:482-493. tion of lecithotrophy in polydorid genera Schmarda, L. K. 1861. Neue wirbellose (Polychaeta: Spionidae). Ophelia 39:121­ Thiere beobachtet und gesammelt auf ei­ 136. ner Reise urn die Erde 1853 bis 1857. Vol. Radashevsky, V. I., and H. L. Hsieh. 2000a. 1. Turbellarien, Rotatorien und Anneli­ Polydora (Polychaeta: Spionidae) species den. Pt. 2:1-164. from Taiwan. Zool. Stud. 39:203-217. Stachowitsch, M. 1980. The epibenthic and --. 2000b. Pseudopolydora (Polychaeta: endolithic species associated with the gas­ Spionidae) species from Taiwan. Zool. tropod shells inhabited by the hermit crabs Stud. 39:218-235. Paguristes oculatus and Pagurus cuanensis. Radashevsky, V. I., and J. D. Williams. 1998. Mar. Ecol. 1:73-101. Polydora websteri Hartman in Loosanoff & Steinhauer, M., and E. Imamura. 1990. Cali­ Engle, 1943 (Annelida, Polychaeta): Pro­ fornia OCS Phase II Monitoring Program posed conservation of the specific name by three year annual report. Vol. I. Submitted a ruling that it is not to be treated as a to the U.S. Department of the Interior, replacement name for P. caeca Webster, Minerals Management Service, Pacific 1879, and designation of a lectotype for OCS Region, under Contract No. 14-12­ P. websteri. Bull. Zool. Nomencl. 55:212­ 0001-30262. 216. Takahashi, K. 1937. Notes on the poly­ Rainer, S. 1973. Polydora and related genera chaetous annelid, Polydora pacifica n. sp., (Polychaeta: Spionidae) from Otago waters. which bores holes in Pinctada margaritifera J. R Soc. N.z. 3:545-564. (Linne). Stud. Palao Trop. BioI. Stn. Read, E. 1975. Systematics and ecology of 2:155-167. polydorid species (Polychaeta: Spionidae) Verrill, A. E. 1879. Notice ofrecent additions from Wellington Harbour. J. R Soc. N.Z. to the marine invertebrata, of the north­ 5:395-419. eastern coast ofAmerica, with descriptions Reish, D. J. 1968. A biological survey of the of new genera and species and critical Bahia de Los Angeles, Gulf of California, remarks on others. Part 1, Annelida, Ge­ Mexico. II. Benthic polychaetous annelids. phyrea, Nemertina, Nematoda, Polyzoa, Polydora and Related Genera Associated with Hermit Crabs . Williams 465

Tunicata, Mollusca, Anthozoa, Echino­ Indo-West Pacific and first record of host dermata, Porifera. Proc. U.S. Nat!. Mus. hermit crab egg predation by a commensal 2:165-205. polydorid worm. ZooI. J. Linn. Soc. --.1881. New England Annelida. Part 1. 129:537-548. Historical sketch, with annotated list of ---. 2001. Reproduction and larval de­ species hitherto recorded. Trans. Conn. velopment of Polydora robi (Polychaeta: Acad. Arts Sci. 4:285-324. Spionidae), an obligate commensal of her­ Walker, S. E. 1995. Taphonomy of modern mit crabs from the Philippines. Invertebr. and fossil intertidal gastropod associations BioI. 120:237-247. from Isla Santa Cruz and Isla Santa Fe, Williams, J. D., and J. J. McDermott. 1997. Galapagos Islands. Lethaia 28:371-382. Feeding behavior of Dipolydora commensalis Ward, L. A. 1981. Spionidae (Polychaeta: (Polychaeta: Spionidae): Particle capture, Annelida) from Hawaii, with descriptions transport, and selection. Invertebr. BioI. offive new species. Proc. BioI. Soc. Wash. 116:115-123. 94:713-730. Williams, J. D., and V. I. Radashevsky. 1999. --. 1987. Family Spionidae. Pages 340­ Morphology, ecology, and reproduction of 369 in D. M. Devaney and L. G. Eldredge, a new Polydora species (Polychaeta: Spio­ eds. Reef and shore fauna of Hawaii, 64. nidae) from the east coast of North Amer­ Bishop Museum Press, Honolulu. ica. Ophelia 51:115-127. Webster, H. E. 1879. Annelida Chaetopoda Woodwick, K. H. 1964. Polydora and related of the Virginia coast. Trans. Albany Inst. genera (Annelida, Polychaeta) from Eni­ N.Y. 9:202-269. wetok, Majuro, and Bikini Atolls, Marshall Williams, J. D. 2000. A new species of Poly­ Islands. Pac. Sci. 18:146-159. dora (Polychaeta: Spionidae) from the