Pectinaria antipoda Schmarda, 1861

Zhang, Jinghuai & Hutchings, Pat, 2019, A revision of Australian Pectinariidae (Polychaeta), with new species and new records, Zootaxa 4611 (1), pp. 1-70 : 38-47

Pectinaria antipoda Schmarda, 1861

Figs 22–23 , Table 3

Pectinaria antipoda Schmarda, 1861 , neotype (BMNH 1886.8.20.1) designated and described by Hutchings & Peart, 2002): 99 – 127, figs 9 – 10 & 12

Material examined. (Body length greater than 10 mm). Australia, Queensland: AM W.25446, 1 spec., Lindeman Island, Lindeman Group , 20°27’S 149°02’E, sand, coll. G.P. Whitley, 1935 ; AM W.33201, 1 spec., Heron Island , 23°26’53”S 151°54’50”E, 15 m, on holothurian in coral garden, coll. P.A. Hutchings, 24 Jul 2003 ; AM W. 200622 , 1 spec., Heron Reef, Heron Island, Great Barrier Reef , 23°27’S 151°55’E, 10 m, in Gorgonian hole in medium sediment, coll. G.W. Rouse, 29 Aug 1984 . New South Wales : AM W.50685, 2 specs, Solitary Islands Marine Park , 29°56’S 153°23’E, 55 m, 20 May 1998 ; AM W.1760, 1 spec., Port Jackson , 33°51’S 151°16’E, Nov 192 ; AM W.1761, 1 spec., Port Hacking, Gunnamatta Bay , tidal flats, 34°04’30”S 151°08’54”E, coll. Museum Party, Nov 1925 ; AM W.2578, 1 spec., Port Jackson, between Sow and Pigs Shoal and Shark Island , 33°51’S 151°16’E, 4 m, coll. M. Ward, 01 Apr 1928 ; AM W.6461, 1 spec., SE of Malabar , 33°58’08”S 151°16’52”E, 51 m, coll. Australian Museum Shelf Benthic Survey, 02 Jan 1973 ; AM W.6462, 2 specs, E of Malabar , 33°58’16”S 151°20’E, coll. Australian Museum Shelf Benthic Survey, 29 May 1973 ; AM W.28592, 1 spec., E of Malabar, Sydney, 33°58’41”S 151°17’51”E, 80.5 m, coll. EPA - Malabar Deep Ocean Outfall Study ( DOOM), 24 Feb 1998 ; AM W.6460, 1 spec., 1 km SE of Little Bay , 33°59’09”S 151°15’40”E, 25 m, coarse gravel, sand and shell fragments, coll. Australian Museum Shelf Benthic Survey, 12 May 1972 ; AM W.6966, 1 spec., 2.3 km SE of Malabar , 33°59’27”S 151°16’48”E, 66 m, coll. Australian Museum Shelf Benthic Survey, Oct 1973 ; AM W.3672, 5 specs, Port Hacking, Gunnamatta Bay , 34°04’30”S 151°08’54”E, muddy sand, coll. E.C. Pope & party, Oct 1957 , at low water mark of spring tides; AM W.23077, 1 spec., Wattamolla, off Providential Head , 34°08’S 151°08’30”E, 45 m, coll. The Ecology Lab for RMI / Pioneer Project , 01 Feb 1990 ; AM W.5617, 1 spec., Jervis Bay , 35°03’S 150°44’E, 15 m, sandy, coll. N. Coleman, 23 Apr 1973 ; AM W.21203, 3 specs, Jervis Bay, Honeymoon Bay , 35°03’48”S 150°45’24”E, 20 m, unvegetated sediment, coll. P.A. Hutchings & party, 05 Jun 1989 ; AM W.25420, 1 spec., Jervis Bay, Honeymoon Bay , 35°03’48”S 150°45’24”E, 20 m, unvegetated sediment, coll. P.A. Hutchings & party, 21 Aug 1989 ; AM W.25421, 1 spec., Jervis Bay, Honeymoon Bay , 35°03’48”S 150°45’24”E, 20 m, coll. P.A. Hutchings & party, 27 Feb 1990 ; AM W.25423, 2 specs, Jervis Bay, Honeymoon Bay , 35°03’48”S 150°45’24”E, 20 m, coll. P.A. Hutchings & party, 21 Feb 1991 ; AM W.194491, 1 spec., Jervis Bay, Murrays Basin sandbank, 35°07’30”S 150°45’30”E, Halophila , sand, coll. NSW State Fisheries, 17 Oct 1972 ; AM W.10700, 1 spec., Wagonga River, 400 m SW of Narooma Bridge , 36°12’55”S 150°07’13”E, coll. Australian Museum Eurobodalla Shire Estuary Survey, 17 Sep 1974 ; AM W.10498, 1 spec., Wagonga River, 900 m SW of Narooma Bridge , 36°13’10”S 150°07’30”E, coll. Australian Museum Eurobodalla Shire Estuary Survey, 17 Sep 1974 ; AM W.17111, 1 spec., Merimbula Channel, Lake Merimbula , 36°53’30”S 149°54’30”E, intertidal, sand, among Posidonia , coll. J.H. Day and party, 04 Dec 1975 ; AM W.19254, 1 spec., Lake Merimbula , 36°55’S 149°54’E, 6 m, mud, coll. NSW State Fisheries, 25 Nov 1977 . South Australia : AM W.25410, 1 spec., Spencer Gulf, Port Pirie , 10 km NW of Port Davis Creek, 33°16’S 137°51’E, 12.1 m, unvegetated sand, coll. T.J. Ward et al., Aug 1979 ; AM W.25409, 1 spec., Bay of Shoals, Kangaroo Island , 35°50’S 137°15’E, 1 m, coll. D. Hoese, 11 Mar 1978 . Tasmania : AM W.26160, 1 spec., Tinderbox, D’ Entrecasteaux Channel , 43°04’S 147°20’E, 8 m, coll. G. Edgar, 18 Mar 1999 ; AM W.26162, 2 specs, Dennes Point , 43°05’S 147°21’E, 5 m, coll. G. Edgar, 28 Mar 1999 ; AM W.26163, 3 specs, Creeses Mistake , 43°07’S 147°47’E, 15.5 m, coll. R.B. Mawbey, 30 Apr 1999 ; AM W.26164, 2 specs, Simmons Point , 43°12’S 147°17’E, 17.9 m, coll. R.B. Mawbey, 29 Jan 1999 . Victoria : AM W.16204, 1 spec., Port Phillip Bay, off Patterson River , 37°58’S 144°54’E, 4 m, fine sand, coll. Marine Studies Group, 10 Mar 1971 ; AM W.16203, 1 spec., Port Phillip Bay, off Patterson River , 38°16’18”S 144°41’30”E, 9 m, sand, coll. Marine Pollution Studies Group, Ministry for Conservation, Victoria, 09 Dec 1971 . Western Australia : AM W.25406, 4 specs (1 spec. mounted for SEM), Rottnest Island , 32°00’S 115°30’E, Pocillopora reef, coll. P.A. Hutchings, 14 Jan 1991 ; AM W.30720, 1 spec., Woodman Point , 32°08’S 115°44’E, 10 Jul 1994 .

Small material examined (Body length shorter than 10 mm). Australia, Queensland: AM W.45037, 1 spec., Lizard Island, 500 m off S Watsons Bay , 14°39’41”S 145°26’27”E, 15 m, sandy mud with filamentous algae, coll. M.T. Aguado, M. Capa, P.A. Hutchings & A. Murray, 23 Aug 2013 ; AM W.43945, 2 specs, Lizard Island, front of reef between Bird and South Islands , 14°41’43”S 145°27’56”E, 21 m, sand, coll. A. Zhadan, 14 Aug 2013 ; AM W.44029, 1 spec., Lizard Island, front of reef between Bird and South Islands , 14°41’52”S 145°27’50”E, 12 m, sand, coll. N. Budaeva, 14 Aug 2013 ; AM W.45851, 1 spec., SE of Lizard Island, reef on NW of North Direction Island , 14°44’36”S 145°30’20”E, 16 m, sand, coll. A. Zhadan, 15 Aug 2013 . New South Wales : AM W.29146, 1 spec., S of Smoky Cape, Black Rock , 30°56’58”S 153°04’32”E, 13 m, algal turf on top of rock, coll. R.T. Springthorpe, 13 Feb 2002 . AM W.24315, 1 spec., E of Marley , 34°08’05”S 151°09’39”E, 60 m, sand, coll. Fisheries Research Institute ( NSW), 02 May 1991 ; Tasmania : AM W.26161, 1 spec., North East River , 39°45’S 147°56’E, coll. G. Edgar, 08 Apr 1997 .

Material examined (but not measured). Australia, New South Wales: AM W.14894, 1 spec., E Botany Bay , 33°59’S 151°10’E, 14 m, sand, coll. NSW State Pollution Control Commission, 26 Jan 1977 ; AM W.25413, 1 spec., SE of Bate Bay , 34°06’48”S 151°11’E, 45 m, coll. The Ecology Lab for RMI / Pioneer Project , 25 Jun 1990 ; AM W.25414, 1 spec., SE of Bate Bay , 34°06’48”S 151°11’E, 45 m, coll. The Ecology Lab for RMI / Pioneer Project , 25 Jun 1990 ; AM W.25415, 1 spec., SE of Bate Bay , 34°06’48”S 151°11’E, 45 m, coll. The Ecology Lab for RMI / Pioneer Project , 25 Jun 1990 ; AM W.25416, 1 spec., SE of Bate Bay , 34°06’48”S 151°11’E, 45 m, coll. The Ecology Lab for RMI / Pioneer Project , 25 Jun 1990 ; AM W.25417, 1 spec., SE of Bate Bay , 34°06’48”S 151°11’E, 35 m, coll. The Ecology Lab for RMI / Pioneer Project , 25 Jun 1990 ; AM W.25418, 1 spec., Bass Point , 34°36’S 150°54’E, 65 m, coll. The Ecology Lab for RMI / Pioneer Project , 29 Oct 1990 ; AM W.25419, 1 spec., Bass Point , 34°36’S 150°54’E, 35 m, coll. The Ecology Lab for RMI / Pioneer Project , 25 Jun 1990 ; AM W.25424, 3 specs, Jervis Bay, Green Point , 35°01’S 150°45’12”E, 12 m, unvegetated sediment, coll. P.A. Hutchings & party, 18 Jun 1991 ; AM W.25428, 2 specs, Jervis Bay, Montagu Roadstead , 35°02’12”S 150°46’E, 12 m, coll. P.A. Hutchings & party, 05 Jun 1990 ; AM W.25422, 1 spec., Jervis Bay, Honeymoon Bay , 35°03’48”S 150°45’24”E, 20 m, coll. P.A. Hutchings & party, 17 Aug 1989 ; AM W.25614, 1 spec., Jervis Bay, Honeymoon Bay , 35°03’48”S 150°45’24”E, 20 m, muddy sediments, coll. P.A. Hutchings, 05 Jun 1989 ; AM W.27528, 1 spec., Jervis Bay, Honeymoon Bay , 35°03’48”S 150°45’24”E, 20 m, unvegetated sediment, coll. P.A. Hutchings & party, 05 Jun 1989 ; AM W.25425, 1 spec., Jervis Bay , S of Plantation Point, 35°04’48”S 150°41’48”E, 20 m, unvegetated sediment, coll. P.A. Hutchings & party, 18 Jun 1991 ; AM W.194145, 1 spec., Jervis Bay, Murrays Beach , 35°07’30”S 150°45’30”E, coll. NSW State Fisheries, 25 Apr 1972 ; AM W.194259, 1 spec., Jervis Bay, Murrays Beach , 35°07’30”S 150°45’30”E, coll. NSW State Fisheries, 25 Apr 1972 ; AM W.194358, 2 specs, Jervis Bay, Murrays Beach , 35°07’30”S 150°45’30”E, coll. NSW State Fisheries, 25 Apr 1972 ; AM W.194372, 1 spec., Jervis Bay, Murrays Basin sandbank, 35°07’30”S 150°45’30”E, sand, coll. NSW State Fisheries, 17 Oct 1972 ; AM W.194398, 1 spec., Jervis Bay, Murrays Beach , 35°07’30”S 150°45’30”E, coll. NSW State Fisheries, 25 Apr 1972 ; AM W.194469, 2 specs, Jervis Bay, Murrays Basin sandbank, 35°07’30”S 150°45’30”E, Zostera & sand, coll. NSW State Fisheries, 17 Oct 1972 ; AM W.194521, 1 spec., Jervis Bay, Murrays Beach , 35°07’30”S 150°45’30”E, sand, coll. NSW State Fisheries, 25 Apr 1972 ; AM W.194584, 2 specs, Jervis Bay, Murrays Basin sandbank, 35°07’30”S 150°45’30”E, sand, coll. NSW State Fisheries, 17 Oct 1972 ; AM W.20835, 1 spec., Jervis Bay, Hole in the Wall , 35°07’36”S 150°44’48”E, 12 m, unvegetated sediment, coll. P.A. Hutchings & party, 20 Feb 1989 ; AM W.25426, 1 spec., Jervis Bay, Hole in the Wall , 35°07’36”S 150°44’48”E, 12 m, unvegetated sediment, coll. P.A. Hutchings & party, 18 Jun 1991 ; AM W.25427, 2 specs, Jervis Bay, Hole in the Wall , 35°07’36”S 150°44’48”E, 12 m, unvegetated sediment, coll. P.A. Hutchings & party, 18 Jun 1991 ; AM W.10698, 1 spec., Wagonga River , 400 m SW of Narooma Bridge, 36°12’55”S 150°07’13”E, coll. Australian Museum Eurobodalla Shire Estuary Survey, 17 Sep 1974 ; AM W.10699, 1 spec., Wagonga River , 400 m SW of Narooma Bridge, 36°12’55”S 150°07’13”E, coll. Australian Museum Eurobodalla Shire Estuary Survey, 17 Sep 1974 ; South Australia: AM W.25412, 1 spec., Spencer Gulf, Port Pirie , 3 km NW of 1st Creek , 33°12’S 138°00’E, 4.1 m, coll. T.J. Ward et al., Mar 1980 ; AM W.25411, 1 spec., Spencer Gulf, Port Pirie , 10 km NW of Port Davis Creek, 33°16’S 137°51’E, 12.1 m, unvegetated sand, coll. T.J. Ward et al., Aug 1979 . Western Australia: AM W.25407, 2 specs, Rottnest Island, North Point , 31°59’18”S 115°30’30”E, underneath boulders, coll. P.A. Hutchings, 22 Jan 1991 ; AM W.25408, 1 spec., Rottnest Island, Thompsons Bay , 32°00’S 115°30’E, 12 m, coll. P.A. Hutchings, 17 Jan 1991 .

Description. Based on all specimens measured. Preserved specimen pale or reddish in colour, conical in shape ( Fig. 22A, B ). Body length 10.0 – 46.8 mm including paleae and scaphe, width 1.0 – 7.5 mm at cephalic regions.

Cephalic veil oval, free, with 16 – 29 smooth cirri on anterior margin and 2/3 of lateral margins ( Figs 22C – D ; 23A – B ). Pair of ear-shaped lobes adjacent to both sides of dorsal base of cephalic veil. Buccal tentacles with wide longitudinal grooves, arising posterior to cephalic veil ( Figs 22C – D ; 23B ).

Operculum semicircular; dorsal and lateral margin well developed, smooth; ventral margin (opercular ridge) with 8 – 13 pairs of stout and golden paleae, curved dorsally, with blunt tips ( Figs 22A – C ; 23A – B ).

First pair of tentacular cirri not extending beyond tips of paleae, with annuli, arising from connection of opercular margin and paleal ridge ( Figs 22B, F ; 23A ). Pair of small ventral lappets present behind tentacular cirri near cephalic veil on segment 1 ( Figs 22C – D ; 23A ). Ventral region of segment 1 covered by buccal tentacles and ventral lobes of segment 2 ( Figs 22C – D ; 23B ).

Second pair of tentacular cirri extending beyond opercular margin, with annuli, on latero-median connecting ridge on segment 2, inserted more dorsally than 1 st pair of tentacular cirri ( Figs 22B, F ; 23A ). Segment 2 with ventral lobes as pair of narrow ventro-lateral lobes separated by grooves from base of 2 nd pair of tentacular cirri, and pair of broad mid-ventral lobes about twice as wide as ventro-lateral lobes. Segment 2 with pair of short dorsolateral lobes, not extending to dorsum, connected to base of 2 nd pair of tentacular cirri.

Comb-like branchiae on segments 3 – 4, consisting of series of dense, flat lamellae ( Figs 22C, F ; 23A ). Branchiae on segment 3 larger and inserted more ventrally than those of segment 4 ( Figs 22E ; 23A ). Pair of small dorso-lateral glandular pads present on dorsal side of branchiae on segments 3 and 4 ( Figs 22F ; 23A ).

Distinct ventral glandular lobes present on segments 3 – 6, becoming progressively more lateral and broader on segments 3 – 5 ( Figs 22A, C ; 23A – B ). Hump present near the branchiae on segment 4. Segment 3 with broad ventral lobe, curved anteriorwards at mid-line. Segment 4 with pair of broad ventro-lateral lobes and mid-ventral lobe about 1/2 width of ventro-lateral lobes, separated from those by shallow grooves. Segment 5 with pair of broad ventro-lateral lobes and narrow mid-ventral lobe about 1/4 width of lateral lobes, separated from those by shallow grooves. Segment 6 with pair of broad ventral lobes, separated from each other by deep notch, with continuous papillae ( Figs 22A, C ; 23A – B ).

Notopodia other than those of segment 1 which bear paleae, on segments 5 – 21 (17 pairs), each bearing two kinds of notochaetae; one winged, bordered with serrations along distal portion, recurved dorsally, covered progressively with more spines from about middle to front of wing, on anterior surface; other stout, straight, tapering to pointed tip, covered progressively with more spines from mid-anterior portion to tip, on anterior surface ( Figs 22B, G ; 23C – D ). Neuropodia, 13 pairs on segments 8 – 20, each with slightly raised torus with transverse row of uncini; each uncinus with U-shaped peg embedded into torus, several rows of minor teeth, and two longitudinal rows of major teeth, each with 7 – 8 teeth ( Figs 22B, G ; 23E ). Segment 21 with ventro-lateral lobes near notopodia.

Scaphe ovoidal, flattened dorsally, divided into three anterior narrow lobes and three posterior broadly triangular lobes on each lateral margin,4 th lobe largest with pointed tip ( Figs 22A – B, F ; 23F ). Anal flap triangular tongue with short anal cirrus, with smooth margin. Scaphal hooks 6 – 12 pairs (more frequently 6 – 8), amber, with blunt tips, slightly curved dorsally ( Figs 22G ; 23F – G ).

Tube with wide opening, slightly curved, fragile, made of large sand grains ( Fig. 22E ).

Methyl Green stained body distinctly green on cirri of cephalic veil, ventral lobes of segments 2–6, dorsolateral lobes of segment 2, dorso-lateral pads of segments 3–4, ventro-lateral regions adjacent to neuropodia, dorsal and ventro-lateral regions of segment 21, and dorsum of anal flap except for anal cirrus and margin ( Fig. 22 ).

Small individuals. Based on all specimens (body length less than 10 mm) measured. Body length 2.2 – 9.2 mm including paleae and scaphe, width 0.3 – 2.6 mm at cephalic regions. Cephalic veil oval, free from operculum, with 14 – 20 smooth cirri on anterior and lateral margins. Opercular with 7 – 11 pairs of golden paleae, curved dorsally, with blunt tips. Scaphal hooks 2 – 6 pairs, amber, blunt bent tips. All other morphological characters similar to those of larger individuals.

Distribution. SE Queensland, New South Wales, Australian Capital Territory, Victoria, Tasmania, SW Western Australia ( Fig. 1 ).

Habitat. Found in mud or muddy sand sediments and in amongst algae, from intertidal to 80.5 m.

Remarks. Pectinaria antipoda has been widely reported from Australia since it was described by Schmarda, 1861 (Augener 1921; Knox & Cameron 1971; Poore et al. 1975; Hutchings & Peart 2002; Wong & Hutchings 2015). As the type of P. antipoda is lost, Hutchings & Peart (2002) designated a neotype from the type locality Port Jackson, from material collected several decades after Schmarda collected his material, and deposited in the Natural History Museum in London. Hutchings & Peart (2002) suggest that in those few decades after Schmarda collected his type material little development had occurred in Port Jackson and there is, therefore, confidence of the validity of the neotype. Wong & Hutchings (2015) also described several specimens from Lizard Island, but all are less than five mm in length with fewer scaphal hooks (2 – 5 pairs) and cirri (14–17) on cephalic veil. Larger specimens of P. antipoda have more cirri on the cephalic veil, paleae and scaphal hooks ( Fig. 24 ). Hutchings & Peart (2002) suggest that the ventral lobes with continuous papillae on segment 6 are important taxonomic characters. All the specimens of P. dodeka Hutchings & Peart, 2002 and P. kanabinos Hutchings & Peart, 2002 deposited in the Australian Museum were examined and the continuous papillae on segment 6 are also present in ventral lobes of both species. Pectinaria antipoda can be easily distinguished from the other Australian species by the paleae which have compact blunt tips, relatively more (usually 6 – 8 pairs) scaphal hooks and dorso-lateral lobes on segment 2 ( Table 3 ). Pectinaria antipoda also differs from P. kanabinos which has elongated anal flap, cirri on latero-posterior margins of anal flap, more longitudinal rows (four rows) of major uncinial teeth and fewer pairs of scaphal hooks which are distinctly curved; Pectinaria carnosus lacks continuous papillae on the ventral lobes on segment 6 and an anal cirrus; and P. dodeka has a long and narrow compact anal flap.

Figures

FIGURE 1. Distribution of the species of Amphictene, Lagis and Pectinaria in Australia.

FIGURE 22. Pectinaria antipoda Schmarda, 1861 Methyl Green stained body and tube (AM W.3672): A. Ventral view of entire body; B. Lateral view of entire body; C. Ventral view of anterior end; D. Close up of anterior end, in ventral view; E. Tube; F. Lateral view of anterior end; G. Lateral view of posterior end. Abbreviations: ac, anal cirrus; af, anal flap; an, anus; br, branchiae; bt, buccal tentacles; cv, cephalic veil; dll, dorso-lateral lobe; dlp, dorso-lateral pads; op, operculum; or, opercular rim; p, paleae; s, segment; sc, scaphe; tc, tentacular cirri; vl, ventral lappets.

FIGURE 23. Pectinaria antipoda Schmarda, 1861 SEM (AM W.25406): A. Laterior view of anterior end; B. Ventral view of anterior end; C–D. Close up of notochaetae, segment 9; E. Uncini, segment 9; F. Dorsal view of posterior end; G. Scaphal hooks. Abbreviations: af, anal flap; br, branchiae; bt, buccal tentacles; cv, cephalic veil; dll, dorso-lateral lobe; dlp, dorso-lateral pads; nec, neurochaetae; noc, notochaetae; op, operculum; or, opercular rim; p, paleae; s, segment; sc, scaphe; sh, scaphal hooks; tc, tentacular cirri; vl, ventral lappets.

FIGURE 24. Relationship between body width and body length, number of cirri on cephalic veil, number of paleae, number of scaphal hooks for Pectinaria antipoda Schmarda, 1861.

Tables

TABLE 3. (Continued) Major teeth of uncini Papillae Anal flap Scaphal hooks on Species Nos. of Nos. of Large ventral Reference Anal Nos. of longitudinal teeth medial surface Shape Margin Shape of tips cirrus pairs rows per row tooth of scaphe broad pointed, slightly Pallas 1766; Malmgren 1866; Hessle 1917; P. belgica 2‒4 7‒8 absent absent smooth present 6‒12 tongue-like curved Fauvel 1927; Nilsson 1928; Holthe 1986b; broad P. regalis 2‒3 6‒10 absent absent smooth present 0‒4 pointed, straight Verrill 1901; Hartman 1941; Long 1973 triangular P. gouldii 2‒4 6‒8 absent absent semicircular crenulated present 8‒22 pointed, straight Verrill 1874; Nilsson 1928; Long 1973 pointed, strongly P. californiensis 2 5 present absent tongue-like crenulated present 13 Hartman 1941 curved P. hartmanae 2 7 absent absent n.r. crenulated present 8‒10 blunt Reish 1968 pointed, slightly P. meredithi 2‒3 6‒8 absent absent semicircular 4‒5 papillae present 7‒9 Savigny 1822 curved pointed, slightly P. chilensis 2 7‒8 absent absent tongue-like crenulated present 13‒15 Hartman 1941 curved pointed, strongly P. aegyptia 2 7‒8 absent absent short broad smooth present 4‒5 Nilsson 1928 curved long pointed, strongly P. nana n.r. n.r. absent absent 3 pappillae present 3 Wesenberg-Lund 1949 tongue-like curved P. panava n.r. 8 absent absent semilunar crenulated pesent 7 n.r. Willey 1905 P. dimai 3‒4 6‒7 n.r. absent n.r. n.r. n.r. n.r. n.r. Zachs 1933 broad pointed, strongly P. hiuchiensis 2 8 absent absent smooth absent 8‒9 Kitamori 1965; Nishi et al. 2014 tongue-like curved blunt, strongly Imajima & Hartman 1964; Nishi et al. P. okudai 3‒4 7‒9 absent absent tongue-like crenulated present 12‒13 curved 2014 blunt, strongly P. torquata 2 7‒8 absent absent tongue-like crenulated absent 4‒5 Zhang & Qiu 2017 curved ……continued on the next page

	Major teeth of uncini			Papillae on		Anal flap		Scaphal hooks
Species	Nos. of longitudinal rows	Nos. of teeth per row	Large medial tooth	ventral surface of scaphe	Shape	Margin	Anal cirrus	Nos. of pairs	Shape of tips	Reference
P. belgica	2 ‒ 4	7 ‒ 8	absent	absent	broad tongue-like	smooth	present	6 ‒ 12	pointed, slightly curved	Pallas 1766; Malmgren 1866; Hessle 1917; Fauvel 1927; Nilsson 1928; Holthe 1986b;
P. regalis	2 ‒ 3	6 ‒ 10	absent	absent	broad triangular	smooth	present	0 ‒ 4	pointed, straight	Verrill 1901; Hartman 1941; Long 1973
P. gouldii	2 ‒ 4	6 ‒ 8	absent	absent	semicircular	crenulated	present	8 ‒ 22	pointed, straight	Verrill 1874; Nilsson 1928; Long 1973
P. californiensis	2	5	present	absent	tongue-like	crenulated	present	13	pointed, strongly curved	Hartman 1941
P. hartmanae	2	7	absent	absent	n.r.	crenulated	present	8 ‒ 10	blunt	Reish 1968
P. meredithi	2 ‒ 3	6 ‒ 8	absent	absent	semicircular	4 ‒ 5 papillae	present	7 ‒ 9	pointed, slightly curved	Savigny 1822
P. chilensis	2	7 ‒ 8	absent	absent	tongue-like	crenulated	present	13 ‒ 15	pointed, slightly curved	Hartman 1941
P. aegyptia	2	7 ‒ 8	absent	absent	short broad	smooth	present	4 ‒ 5	pointed, strongly curved	Nilsson 1928
P. nana	n.r.	n.r.	absent	absent	long tongue-like	3 pappillae	present	3	pointed, strongly curved	Wesenberg-Lund 1949
P. panava	n.r.	8	absent	absent	semilunar	crenulated	pesent	7	n.r.	Willey 1905
P. dimai	3 ‒ 4	6 ‒ 7	n.r.	absent	n.r.	n.r.	n.r.	n.r.	n.r.	Zachs 1933
P. hiuchiensis	2	8	absent	absent	broad tongue-like	smooth	absent	8 ‒ 9	pointed, strongly curved	Kitamori 1965; Nishi et al. 2014
P. okudai	3 ‒ 4	7 ‒ 9	absent	absent	tongue-like	crenulated	present	12 ‒ 13	blunt, strongly curved	Imajima & Hartman 1964; Nishi et al. 2014
P. torquata	2	7 ‒ 8	absent	absent	tongue-like	crenulated	absent	4 ‒ 5	blunt, strongly curved	Zhang & Qiu 2017

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