taxonID	type	description	language	source
537A7F7AFF94FD2B564E9B71707DFBE9.taxon	description	(figures 1 – 5)	en	Williams, Jason (2004): Reproduction and morphology of Polydorella (Polychaeta: Spionidae), including the description of a new species from the Philippines. Journal of Natural History 38 (11): 1339-1358, DOI: 10.1080/0022293031000155395, URL: http://www.informaworld.com/openurl?genre=article&doi=10.1080/0022293031000155395&magic=crossref||D404A21C5BB053405B1A640AFFD44AE3
537A7F7AFF94FD2B564E9B71707DFBE9.taxon	materials_examined	HOLOTYPE: Philippines, Puerto Galera, Agus Beach, y 3 m, 1 July 2000, from Clathria (Thalysias) cervicornis (Thiele, 1903) (USNM 1008710). PARATYPES: same data as for holotype (USNM 1008711: 270 z paratypes; USNM 1008712: 10 paratypes, on nine SEM stubs). Philippines, Puerto Galera, Small La Laguna Beach, y 3 m, 24 July 2000, from Clathria (Thalysias) cervicornis (AMNH 4347: 180 z paratypes). Etymology. The specific name kamakamai is derived from the Filipino word for supernatural beings that are small, human-like creatures possessing long beards and nails used to pinch offending children (Jocano, 1969), reminiscent of the acicular neurosetae of anterior segments in this species. Description Holotype. 1.3 mm long, 0.3 mm wide at segment 7; 14 segments. Prostomium bifid; caruncle short, terminating between posterior margin of segment 1 and middle of segment 2; occipital tentacle and eyes absent (figures 1 – 3). Palps extend posteriorly for 10 – 12 segments. Black pigmentation in irregular patches on palps, sides of peristomium, prostomium, along caruncle, dorsal and ventral sides of all segments and pygidium (figures 1, 3). Segment 1 without notopodial lobes or notosetae; neuropodia lobes present and neurosetae absent. Capillary notosetae of segments 2 – 4, 6 and subsequent segments in two rows; no specialized posterior notosetae. Neurosetae of segments 2 – 5 of two types, arranged in two rows; dorsal row composed of up to three acicular spines on segment four, ventral row composed of up to four capillaries on segment four (figure 1 C, E); segments 6 and 7 with reduced number of acicular spines and capillaries; three bidentate hooded hooks begin on segment 8, not accompanied by capillaries; hooks with approximately right angle between main fang and shaft, narrow angle between main fang and apical tooth, with constriction on shaft (figure 1 F). Segment 5 slightly modified, about the same size as segments 4 and 6, with acicular spines and capillary neurosetae in same arrangement as on segments 2 – 4; notosetae absent (figure 2 A). Two types of major spines on segment 5 in approximately straight, horizontal rows; ventral row of four acicular spines with small digitiform bosses (y 0.3 m m) closely applied to the apical ends of the spines, lacking on tips of the spines (figures 1 D, 2 A, B), and a dorsal row of four falcate spines (figures 1 D, 2 A, B). Branchiae absent. Dorsal ciliary bands apparent from segment 3 continuing to terminal segments of stock individual (figure 2 C – E). Narrow pharynx to segment 7 leading to broad intestine, narrowing in posterior segments; without gizzard-like structure in digestive tract. Pygidium reduced, knob-shaped (figure 1 B); with large cells (y 30 m m) within pygidium and terminal segments (figure 3). Variability. All specimens examined exhibited black pigmentation on the palps, peristomium, prostomium, surrounding the caruncle, sides of anterior and posterior segments, and on the pygidium. Pigmentation was most prominent on the dorsal side but was also present on the ventral side of segments. A bifid prostomium and caruncle extending to middle of segment two was observed in all specimens; no specimens possessed branchiae. Acicular neurosetae were found in approximately vertical rows reaching four in number on segment 5, accompanied by a dorsal row of typically four to six capillary neurosetae. The number of acicular neurosetae was reduced to two or three in segments 6 and 7, accompanied by one or two capillary neurosetae. The fifth segment contained four to seven major spines in both the dorsal and ventral rows; 20 % of the specimens examined (N ~ 50) possessed one more additional spine in the ventral row than the dorsal row; all other specimens possessed equivalent numbers of spines in both rows. Hooded hooks numbered three or four on segment 8, reaching a maximum of three to five hooded hooks on segments 8 – 11. The pygidium was variable, from a reduced, knob-shape (figures 1 B, 2 C, D) to a more elongate conical shape (figure 3); small bumps were often observed surrounding the anus. Distribution Philippines: Oriental Mindoro. Remarks Polydorella kamakamai is unique within the genus in possession of acicular neurosetae in segments 2 – 7 and the fifth segment containing ventral row of major spines with digitiform bosses. The species is further differentiated from P. dawydoffi, P. prolifera and P. stolonifera by the lack of branchiae. Ecology Polydorella kamakamai was found to construct small mud tubes (up to 5 mm) on the surface of the sponge, Clathria (Thalysias) cervicornis. The worms were found in fairly high abundance, with approximately 280 worms on a sponge consisting of eight main branches approximately 100 mm long and 7 mm in diameter (y 1892.0 cm 2). Some specimens contained material in the gut the same colour as the preserved sponge; additional behavioural and histological studies are required to determine if the worms feed on the tissue of sponges as has been documented in some other polychaetes (see Martín and Britayev, 1998). Many tubes contained two individuals with dorsal sides apposed and anterior ends facing in opposite directions; tubes containing greater than two individuals were not observed. When disturbed, worms were able to move quickly out of their tubes and efficiently crawl across the surface of the sponge. Asexual reproduction Most specimens examined (N ~ 290) exhibited some stage of asexual reproduction via paratomy (figure 4). Single parental or stock individuals consist of 13 – 15 segments (figure 5 A). Stock individuals then exhibit a growth zone which follows segment 10 (figures 1 B, 2 C, 5 B). The growth zone is distinct from anterior and posterior segments in its opaque colouring and smooth form. The opaque colouring of the growth zone may come from the breakdown of muscular material as the zone prepares for the formation of a new stolon body (Schroeder and Hermans, 1975). The growth zone exhibits two sections: an anterior section that will become the new segment 11 (denoted as 11 ’, following Radashevsky (1996) where ’ indicates newly developed stock segments) and the posterior portion which produces the developing stolon 1. After further development of the growth zone, palp buds are formed and the new stock segment anterior to the head of the developing stolon 1 exhibits setae. New segments of the stolon body then appear to form simultaneously (overt distinction of 10 segments displayed at once from subdivision of the elongated growth zone). Ultimately, stolon 1 consists of 10 new anterior segments and three to four posterior segments derived from the stock individual (figure 5 D). Stolon 2 is formed following 11 ’. Prior to complete development of the first 10 segments of stolon 1 (i. e. lacking externally observable setae), a secondary growth zone is formed following segment 10. Formation of stolon 2 proceeds as described above and the secondary growth zone of stolon 1 then exhibits palp buds (figures 2 D, E, 5 D). A third stolon (stolon 3) is observed in some individuals behind 11 ’. In this way, a chain of five developing individuals is formed (stock, stolon 3, stolon 2, stolon 1 and the secondary stolon) (figures 3 A, B, 5 E). The process of asexual reproduction is shown in figure 5. It appears that in most cases paratomic division takes place when four or five individuals are present in a chain. Stolon 1 (with secondary stolon) breaks from the stock individual (stolon 3 and stolon 2) (figure 5 E). Prior to division, the posterior segments of stolon 2 are found in a wedge-shape under the head of stolon 1 (figures 2 E, 3 B). At this time, palps of stolon 1 are approximately one-half to threequarters the length of stock individuals. Following division, the pygidium is fully formed on the posterior end of stolon 2. Stolon 1 (with secondary stolon) and stock (with stolon 3 and stolon 2) individuals then repeat the course of asexual reproduction described above. Following division, the worms apparently do not immediately leave their original tubes and construct new ones, as evidenced by tubes containing two individuals. The division of paratomic chains with stock individuals lacking a stolon 3 or stolon 2 leads to the presence of single individuals within the population. In only seven (2.4 %) cases were the stock individuals within paratomic chains observed to possess 13 – 15 stock segments prior to division; this low number explains the relative rarity of single individuals lacking growth zones (figure 4). The colonies were dominated by specimens composed of chains of two individuals (sometimes with developing growth zones); chains of three and four individuals were approximately half as abundant as chains of two individuals (figure 4). Only rarely were single individuals and chains of five individuals or greater observed (figure 4). Sexual reproduction In addition to asexual reproduction, one specimen composed of 15 segments was found with eggs (figure 1 A). Segments 10 – 12 of the specimen were narrower than preceding segments (no growth zone was exhibited), while segments 13 – 15 were expanded and contained yolky eggs. Eggs were oval in shape; 43.3 m m (¡ 5.8) by 62.2 m m (¡ 6.8) (N ~ 11). Based on the volume of the eggs within this specimen, segments 13 – 15 are estimated to contain 36 – 42, 26 – 34 and 15 – 20 eggs, respectively.	en	Williams, Jason (2004): Reproduction and morphology of Polydorella (Polychaeta: Spionidae), including the description of a new species from the Philippines. Journal of Natural History 38 (11): 1339-1358, DOI: 10.1080/0022293031000155395, URL: http://www.informaworld.com/openurl?genre=article&doi=10.1080/0022293031000155395&magic=crossref||D404A21C5BB053405B1A640AFFD44AE3
537A7F7AFF9AFD2F55C69B3D7535FBDC.taxon	description	(figures 6, 7)	en	Williams, Jason (2004): Reproduction and morphology of Polydorella (Polychaeta: Spionidae), including the description of a new species from the Philippines. Journal of Natural History 38 (11): 1339-1358, DOI: 10.1080/0022293031000155395, URL: http://www.informaworld.com/openurl?genre=article&doi=10.1080/0022293031000155395&magic=crossref||D404A21C5BB053405B1A640AFFD44AE3
537A7F7AFF9AFD2F55C69B3D7535FBDC.taxon	materials_examined	Material examined. Polydorella dawydoffi: 10 paratypes from Thailand, Gulf of Thailand, Thom Island of Anthoi Archipelago, 7 m, 2 February 1986, from Xestospongia testudinaria (Lamarck, 1815) (USNM 148722; one prepared for SEM); 60 z specimens from Philippines, Puerto Galera, Small La Laguna Beach, y 3 m, 30 June 2000, from Chalinula sp., (USNM 1008713); 50 z specimens from same locality and sponge as above, 1 July 2000, (AMNH 4348); 18 specimens from same locality and date as above, from Niphates sp. (USNM 1008714); 60 z specimens from Philippines, Puerto Galera, Agus Beach, y 3 m, 1 July 2000, from Niphates sp. (AMNH 4349); 30 specimens from same locality and sponge as above, 25 July 2000 (USNM 1008715: 24 specimens; USNM 1008716: six specimens, on three SEM stubs); seven specimens from Philippines, Puerto Galera, Big La Laguna Beach, y 3 m, 24 July 2000, from Petrosia sp. (USNM 1008717); 37 specimens from Red Sea: Egypt, Hurghada, 0.1 m, November 2001, from Xestospongia sp. (USNM 1008718: 32 specimens; USNM 1008719: five specimens, on two SEM stubs), coll. Pliascheva. Description Single individuals with 14 – 15 segments; approximately 1.5 mm in length and 0.4 mm in width at segment 5. Prostomium bifid; caruncle short, reaching anterior margin or middle of segment 2; occipital tentacle absent; two round, white eyes observable in life only. Palps extend posteriorly for 8 – 12 segments. Pigmentation variable, present on dorsal and ventral sides of anterior and posterior segments or absent (figure 6 A, B). Segment 1 without notopodial lobes or notosetae; neuropodial lobe present without neurosetae (figure 6 A). Unilimbate capillary notosetae of segments 2 – 4, 6 and subsequent segments in two rows. Unilimbate capillary neurosetae of segments 2 – 4, 6 and 7 arranged in two rows; up to six bidentate hooded hooks begin on segment 8, not accompanied by capillaries; hooks with approximately right angle between main fang and shaft, narrow angle between main fang and apical tooth, with constriction on shaft (figure 6 F). Segment 5 slightly modified, about the same size as segments 4 and 6, with posteroventral fascicle of unilimbate capillary neurosetae in two rows; with anterodorsal fascicle of unilimbate notosetae. Two types of major spines of segment 5 in approximately horizontal curved row; ventral row of up to four to six spines with distally enlarged ends, with small digitiform bosses on sides and main shaft, approaching apical edge with eight to nine rounded or pointed teeth, posterior end of spines with capillary extension (figures 6 E, 7 A – D) and dorsal row of up to three to four acicular spines with an apical shelf (figures 6 E, 7 B). Single pair of branchiae present on segment 7 (figure 6 A, B). Dorsal ciliary bands from segment 2 continuing to terminal segments. Posterior segments with large cells (y 20 – 30 m m); pygidium cylindrical, slightly tapering to distal end (figure 6 A, B). Asexual reproduction The sequence of stolon body formation closely follows that described by Radashevsky (1996), although the growth zone position may vary between segments 10 and 11. Eighteen specimens examined from the Philippines and the Red Sea exhibited a growth zone that appeared to follow segment 10 (these individuals possessed a recently formed growth zone corresponding to stolon 1). Radashevsky (1996) noted that the growth zone may be incorrectly interpreted to form after segment 10 during secondary stolon formation. Yet even during formation of stolon 1 specimens exhibited a growth zone following segment 10. Thus, it appears that the growth zone can form following segment 10, soon after which the anlage of segment 11 is formed. Individuals later in development (with stolon 1 possessing new thoracic segments and newly formed palps) possess a fully developed segment 11 and often a developing segment 12. Records of specimens possessing a growth zone following segment 11 may represent individuals later in development rather than initial position of the growth zone. SEM examination confirmed that 10 segments simultaneously arise in developing stolons (figure 6 C); early in this stage the setae of the developing segments are not visible externally and develop later in segment formation. Feeding Examination of specimens collected from the Red Sea showed that nearly all individuals contained sand grains within the digestive tract. The sand grains were of considerable size (164.5 ¡ 100.3 m m, mean maximal length ¡ SD; N ~ 20) compared to the digestive tract and distributed throughout the digestive tract of paratomic chains (figure 6 D). These sand grains indicate the worms have the ability to ingest large food particles; their mode of feeding (deposit, suspension, or deposit and suspension) remains unknown but it is likely that they exhibit a combination of deposit and suspension feeding as documented in other polydorids (e. g. Dauer et al., 1981; Williams and McDermott, 1997). If the worms remove sand grains and other deposited material on the surface of sponges they could be benefiting sponges by keeping their surfaces free of debris, as documented in other polychaetes associated with sponges (Martín et al., 1992). Distribution South China Sea: south-eastern coast of Vietnam, Philippines; Red Sea. Remarks SEM examination has allowed more detailed analysis of the fifth segment spine morphology, leading to the observation of the capillary extension at the apical end of the spines and the digitiform bosses (both difficult to observe with light microscopy). The spine morphology of P. dawydoffi specimens from the Philippines differs slightly from those collected in the Red Sea and Vietnam. The Philippine specimens exhibit spines which are more squat on the denticulate edge and possess a thicker capillary extension (figure 7 A, B); Vietnam and Red Sea specimens exhibit spines which are more elongate along the denticulate edge and have a rounded distal end with a thinner distal capillary (figure 7 C, D). No other morphological differences were noted between specimens from the three localities and without further morphological or molecular evidence, the distinctions in the spines do not appear to warrant the erection of a new species. The Philippine specimens exhibited slight pigmentation on anterior and posterior segments while those of the Red Sea were heavily pigmented (figure 6 A, B); Radashevsky (1996) found no pigmentation on specimens collected from Vietnam. One of the paratypes examined was composed of 18 segments and contained an endoparasitic copepod within segments 11 – 15; additional parasitic copepods were found in specimens from Vietnam (Radashevsky, 1996). This is the first record of the species from the Red Sea and the Philippines.	en	Williams, Jason (2004): Reproduction and morphology of Polydorella (Polychaeta: Spionidae), including the description of a new species from the Philippines. Journal of Natural History 38 (11): 1339-1358, DOI: 10.1080/0022293031000155395, URL: http://www.informaworld.com/openurl?genre=article&doi=10.1080/0022293031000155395&magic=crossref||D404A21C5BB053405B1A640AFFD44AE3
537A7F7AFF9EFD31551B9B49706EFB2A.taxon	description	(figures 6, 8)	en	Williams, Jason (2004): Reproduction and morphology of Polydorella (Polychaeta: Spionidae), including the description of a new species from the Philippines. Journal of Natural History 38 (11): 1339-1358, DOI: 10.1080/0022293031000155395, URL: http://www.informaworld.com/openurl?genre=article&doi=10.1080/0022293031000155395&magic=crossref||D404A21C5BB053405B1A640AFFD44AE3
537A7F7AFF9EFD31551B9B49706EFB2A.taxon	materials_examined	Material examined. Twelve paratypes from Australia, Victoria, Westernport Bay, Crawfish Rock, 5 m, 3 January 1972, from Ancorina cordicata (Carter, 1879) (NMV F 42898, previously registered as G 2898; two prepared for SEM). Description Largest single individual 3.8 mm long and 0.6 mm wide at segment 7; 16 segments, with endoparasitic copepod (see Remarks); single individuals with developing growth zone of stolon 1 and 16 segments 2.5 mm long and 0.4 mm wide at segment 7 (figure 8 A). Prostomium bifid; caruncle short, terminating at posterior margin of segment 1; occipital tentacle and eyes absent (figure 8 A, B). Palps extend posteriorly for 8 – 12 segments. Pigmentation absent. Segment 1 with small notopodial lobe without notosetae; neuropodial lobe with two or three winged capillary neurosetae (figure 8 A, B). Unilimbate capillary notosetae of segments 2 – 4, 6 and subsequent segments in two rows. Unilimbate capillary neurosetae of segments 2 – 4, 6 and 7 arranged in two rows; up to eight bidentate hooded hooks begin on segment 8, not accompanied by capillaries (figure 8 A, B); hooks with approximately right angle between main fang and shaft, narrow angle between main fang and apical tooth, with constriction on shaft. Segment 5 slightly modified, about the same size as segments 4 and 6, with posteroventral fascicle of bilimbate capillary neurosetae in two rows; notosetae absent. Two types of major spines of segment 5 in approximately horizontal curved row; ventral row of three to six spines with expanded tips covered by fine bristles and a dorsal row of up to five simple acicular spines (figure 6 G). Branchiae present on segments 7 and 8 (figure 8 A, B). Dorsal ciliary bands from segment 2 continuing to terminal segments (figure 8 B). Pygidium with small cirri surrounding terminal anus (figure 8 A). Asexual reproduction and parasitism Of 12 individuals examined, 10 exhibited asexual reproduction with two individuals joined in a chain (figure 8 A). The two single individuals consisted of up to 16 segments; both possessed endoparasitic copepods within segments 8 – 12. Individuals undergoing asexual reproduction contain a growth zone that developed after segment 10. Formation of new posterior stock segments is apparent in side and ventral views (figure 8 A). The sequence of stolon body formation appears to be similar to that described for Polydorella kamakamai, although no specimens were composed of more than two individuals. Distribution South China Sea: south-eastern coast of Vietnam, Philippines; Red Sea. Remarks Blake and Kudenov (1978) indicated that brown pigmentation was present on the branchiae, peristomium and sides of anterior segments; this pigmentation has been lost from the specimens stored in alcohol for 29 years. This is the first report of parasitic copepods within Polydorella stolonifera, although Radashevsky (1996) noted such parasites in P. dawydoffi. The copepod parasites apparently interfere with the process of asexual reproduction in Polydorella. These parasites may induce larger numbers of segments and longer length in infected individuals; 16 segments in P. stolonifera that contained parasites versus 15 segments in those lacking parasites and 17 – 18 segments in parasitized P. dawydoffi versus 15 segments in those lacking parasites. The present findings are in contrast to those of McCurdy (2001), who found earlier fragmentation in asexual reproduction via architomy by Pygospio elegans Claparède, 1863 when parasitized by trematode metacercariae. Such differences are not unexpected because host responses to parasites could be influenced by a variety of factors including mode of asexual reproduction (i. e. paratomy versus architomy) and type of parasite (crustacean versus trematode).	en	Williams, Jason (2004): Reproduction and morphology of Polydorella (Polychaeta: Spionidae), including the description of a new species from the Philippines. Journal of Natural History 38 (11): 1339-1358, DOI: 10.1080/0022293031000155395, URL: http://www.informaworld.com/openurl?genre=article&doi=10.1080/0022293031000155395&magic=crossref||D404A21C5BB053405B1A640AFFD44AE3
