Polycirrus Grube 1850

Polycirrus Grube, 1850

Aphlebine Quatrefages in Milne Edwards, 1844: 18.

Torquea Leidy, 1855: 146.

Apneumea Quatrefages, 1865: 380.

Ereutho Malmgren, 1866: 391.

Leucariste Malmgren, 1866: 390.

Cyaxares Kinberg, 1867: 348.

Dejoces Kinberg, 1867: 348.

Anisocirrus Gravier, 1905: 437.

Pseudoampharete Hartmann-Schröder, 1960: 39–41.

Litancyra Hutchings, 1977: 9.

Type species: Polycirrus medusa Grube, 1850 by monotypy. Type considered lost and neotype designated herein.

Type locality. France, Villa Franca, Mediterranean Sea, 43˚47ʹ11.24ʺ N, 7˚36ʹ26.11ʺ E. (see treatment of P. medusa for neotype locality).

Diagnosis. Polycirridae with an expanded, often trilobed, upper lip bearing tentacles of two or three kinds. Outer lower lip enlarged and resembling a ventral pad. Notopodia from segment 3 continuing for 10–70 segments; neuropodia from segment 7–18, continuing to pygidium. Notochaetae capillaries of various types including smooth, hirsute, and pinnate (= spinose or corn-eared), present in anterior body only. Neurochaetae avicular uncini, present in anterior and/or posterior body. Five or more pairs of nephridia (segmental organs) in anterior segments opening through pre-gular nephridial papillae (segments 3 and 4) and post-gular nephridial papillae (segment 5 onward).

Remarks. The diagnosis is modified from Holthe (1986b) and Hutchings & Glasby (1986). The genus was shown not to be monophyletic in Nogueira et al. (2013).

Key to the valid world species of Polycirrus

(Region of type locality indicated after species name; P. mexicanus appears twice in the key because of the uncertainty over the starting position of its neuropodia.)

1(0). Neurochaetae beginning well before last notochaetigerous segment.............................................. 2 Neurochaetae beginning on or about (within one or two segments of) last notochaetigerous segment................... 29 Neurochaetae beginning well after last notochaetigerous segment.............................................. 47

2(1). 18 or fewer notochaetigerous segments.................................................................... 3 19–21 notochaetigerous segments........................................................................ 13 22 or more notochaetigerous segments.................................................................... 14

3(2). Uncini of posterior body with long neck and concave base (Type 2; Fig. 3g)....................................... 4 Uncini of posterior body with short neck and straight to convex base (Type 1; Fig. 3 e,f)............................. 9

4(3). Neurochaetae beginning on segment 12 or earlier; mid-ventral groove from segment 2 or 3........................... 5 Neurochaetae beginning on segment 13–16; mid-ventral groove from segment 4 or 5................... P. medius (Japan) 5(4). Notochaetae hirsute (Fig. 3c); ventral pads incised........................................ P. porcatus (S Australia) Notochaetae pinnate (Fig. 3d); ventral pads more-or-less smooth................................................ 6

6(5). Outer lower lip subconical lobe protruding above venter....................................................... 7 Outer lower lip flat, shield-like........................................................................... 8

7(6). Ventro-lateral pads well defined, segmentally discrete (e.g. Fig. 1a, b)......................... P. abrolhensis (N Brazil) Ventro-lateral pads poorly defined (e.g. Fig. 1c, d)............................................. P. clavatus (Brazil)

8(6). Anterior body uncini atypical Type 2 (almost flat base and smaller in size than those of posterior body)..................................................................................................... P. holthei (Caribbean) All uncini typical Type 2.............................................................. P. pumilis (E Australia)

9(3). Neurochaetae beginning on segment 12 or earlier; outer lower lip flat, shield-like.................................. 10 Neurochaetae beginning on segment 13–16; outer lower lip subconical lobe protruding above venter..................................................................................................... P. nephrosus (E Australia)

10(9). Notopodia with lobes distally flared; notochaetae hirsute.............................. P. kerguelensis (Kerguelen Is.) Notopodia with prechaetal lobe low, postchaetal lobe digitiform; notochaetae smooth or hirsute...................... 11 Notopodia with lobes triangular; notochaetae smooth (narrowly winged)...................... P. denticulatus (N France)

11(10). 10 notochaetigerous segments............................................................ P. rosea (E Australia) 14–20 notochaetigerous segments....................................................................... 12

12(11). Lower lip longer than wide; uncini with 1–4 teeth above main fang............................ P. norvegicus (Norway) Lower lip wider than long; uncini with 11–15 teeth above main fang...................... P. elisabethae (Scotland, UK)

13(2). Notochaetae smooth (may be narrowly winged).......................................... P. fedorovi (Arctic Ocean) Notochaetae pinnate................................................................ P. eous (Sea of Okhotsk)

14(2). Neurochaetae beginning on segment 12 or earlier........................................................... 15 Neurochaetae beginning on segment 13–16................................................................ 26

15(14). Uncini with long neck and concave base (Type 2); notochaetae pinnate.......................... P. twisti (Suez Canal) Uncini with short neck and straight to convex base (Type 1); notochaetae smooth, narrowly winged, or hirsute........... 16

16(15). Notopodial prechaetal lobe low, postchaetal lobe digitiform; uncini with subrostral process present or absent............ 17 Notopodial lobes both similar, low and rounded; uncini with subrostral process present as a low protuberance........... 24 Notopodial lobes triangular; uncini with subrostral process absent.............................................. 25

17(16). Ventro-lateral pads more-or-less smooth (e.g. Fig. 1a)........................................................ 18 Ventro-lateral pads incised, forming corrugations or grooves (e.g. Fig. 1b, c)...................................... 22

18(17). Dorsum anteriorly smooth............................................................................. 19 Dorsum anteriorly papillose, ridged, or tessellated........................................................... 20

19(18). Notopodia digitiform; uncini lacking subrostral process.................................. P. coibensis n.sp. (Panama) Notopodia more-or-less rectangular; uncini with subrostral process as low protuberance.......... P. swakopianus (Namibia)

20(18). Upper lip with well-developed medial lobe, no lateral lobes; mid-ventral groove from segment 4–5..................................................................................................... P. californicus (California) Upper lip trefoiled (well-developed medial and lateral lobes); mid-ventral groove from segment 2–3.................. 21

21(20). Neuropodial tori ridge-like; uncini with subrostral process present as low protuberance................ P. chilensis (Chile) Neuropodial tori erect pinnules; uncini lacking subrostral process...................... P. hamiltoni (Macquarie Island)

22(17). Outer lower lip oblong, longer than wide............................................... P. arenivorus (N France) Outer lower lip inverted T-shaped..................................................... P. perplexus (California) Outer lower lip subtriangular and pointing toward mouth..................................................... 23

23(22). Upper lip trefoiled (well-developed medial and lateral lobes); neuropodial tori erect pinnules........ P. dubius (N Carolina) Upper lip with well-developed medial lobe, no lateral lobes; neuropodial tori ridge-like............ P. luminosus (Bermuda) 24(16). Notochaetae one type only, faintly hirsute............................................. P. phosphoreus (E Canada) Notochaetae of two types, hirsute and pinnate.......................................... P. carolinensis (N Carolina)

25(16). Outer lower lip grooved, oblong, wider than long; dorsum anteriorly smooth.................... P. tentaculatus (N Peru) Outer lower lip tessellated, subtriangular, and pointing toward mouth; dorsum anteriorly tessellated..................................................................................................... P. multisetigerous (Chile)

26(14). Notochaetae smooth or hirsute; notopodia more-or-less rectangular; neuropodial tori ridge-like....................... 27 Notochaetae pinnate; notopodia digitiform; neuropodial tori erect pinnules...................... P. multus (Hong Kong)

27(26). Uncini lacking subrostral process....................................................... P.purpureus (Jamaica) Uncini with subrostral process present as a low rounded protruberance.......................................... 28

28(27). Uncini with enlarged median tooth plus smaller teeth above main fang......................... P. aurantiacus (France) Uncini with 2 rows of smaller teeth above main fang.......................................... P. nervosus (Japan)

29(1). Neurochaetae begin on segment 12 or earlier................................................. P. paivai (N Brazil) Neurochaetae begin on segment 13 to 16.................................................................. 30 Neurochaetae begin on segment 17 or later................................................................ 42

30(29). Uncini with long neck and concave base (Type 2)........................................................... 31 Uncini with short neck and straight to convex base (Type 1)................................................... 34

31(30). Notochaetae smooth, narrowly winged, or hirsute; outer lower lip flat, shield-like; inner lower lip divided into two equal lobes................................................................................ P. quadratus (Hong Kong) Notochaetae pinnate; outer lower lip subconical lobe protruding above venter; inner lower lip otherwise................ 32

32(31). Notopodial prechaetal lobe low, postchaetal lobe digitiform; teeth above main fang arranged in double transverse series... 33 Notopodial pre- and post-chaetal lobes both similar, low and rounded; teeth above main fang arranged in single transverse series........................................................................... P. disjunctus (E Australia)

33(32). Oral surface of upper lip without grooves leading to mouth; poorly defined ventro-lateral pads........................................................................................................ P. boholensis (Philippines) Oral surface of upper lip glandular, ciliated, with grooves leading to mouth; discrete ventro-lateral pads present............................................................................................ P. glaucus (W Australia)

34(30). Outer lower lip oblong, wider than long.................................................................. 35 Outer lower lip subtriangular and pointing toward mouth..................................................... 39 Outer lower lip inverted T-shaped...................................................... P. hesslei (Subantarctic)

35(34). Notochaetae include both smooth, narrowly winged and pinnate types in each notopodium...... P. decipiens (Gulf of Aden) Notochaeta all of one type, of various degrees of hirsuteness................................................. 36

36(35). Oral surface of upper lip without grooves leading to mouth; notochaetae within a chaetiger of two distinct lengths........ 37 Oral surface of upper lip with grooves leading to mouth; notochaetae within a chaetiger graded long from dorsal to ventral..................................................................................................... 38

37(36). Upper lip trefoiled (well-developed medial and lateral lobes); discrete ventro-lateral pads present....................................................................................................... P.antarcticus (Antarctic) Upper lip with medial lobe only, well developed; poorly defined ventro-lateral pads.............. P. dodeka (Hong Kong)

38(36). Dorsum anteriorly smooth; neuropodial tori ridge like.................... P. macintoshi replacement name (Kerguelen Is) Dorsum anteriorly tessellated; neuropodial tori erect pinnules........................... P. latidens (Skagerrak, Europe)

39(34). Upper lip with medial lobe only, well developed; notopodia digitiform.......................................... 40 Upper lip trefoiled (well-developed medial and lateral lobes); notopodia more-or-less rectangular.................... 41

40(39). 17–18 notochaetigerous segments; pre-gular nephridial papillae present (on segments 3, 4)....... P. arcticus (Arctic Ocean) 11–14 notochaetigerous segments; pre-gular nephridial papillae absent............................. P. nonatoi (Brazil)

41(39). Ventro-lateral pads corrugated (Fig. 1c); subrostral process of uncini as sharp protuberance.... P. broomensis (NW Australia) Ventro-lateral pads smooth; subrostral process of uncini low and rounded..................... P. breviuncinatus (Brazil)

42(29). Uncini with long neck and concave base (Type 2); notochaetae comprise only pinnate types......................... 43 Uncini with short neck and straight to convex base (Type 1); notochaetae comprise only smooth (winged) types, or both smooth and pinnate types.............................................................................. 44

43(42). Up to 18 notochaetigerous segments; lower lip (outer region) subconical lobe protruding above venter............................................................................................... P. aquila (Malay Archipelago) Notochaetigerous segments 19 to 21; lower lip (outer region) flat, shield-like..................... P. coccineus (Red Sea)

44(42). Outer lower lip subconical lobe protruding above venter...................................... P. plumosus (Norway) Outer lower lip flat, shield-like.......................................................................... 45

45(44). Uncini with subrostral process present as low protuberance.................................................. 46 Uncini lacking subrostral process............................................... P. variabilis (Great Barrier Reef)

46(45). Uncini with a single tooth above main fang....................................... P. mexicanus, in part (W Mexico) Uncini with two rows of teeth above main fang, terminal row comprising 7–9 teeth....... P. bicrinalis (Great Barrier Reef)

47(1). Uncini with long neck and concave base (Type 2); subrostral process absent...................................... 48 Uncini with short neck and straight to convex base (Type 1); subrostral process present as low protuberance............. 49

48(47). Neurochaetae begin on segment 13–16; teeth above main fang arranged in single transverse series...................................................................................................... P. paucidens (S Australia) Neurochaetae begin on segment 17 or later; teeth above main fang arranged in multiple series...... P. octosetus (E Australia)

49(47). Notochaetae include both smooth and pinnate types......................................................... 50 Notochaetae are all more or less smooth (includes minutely hirsute and narrowly winged types)...................... 51

50(49). Outer lower lip oblong, wider than long; uncini with a single tooth above the main fang.... P. mexicanus, in part (W Mexico) Outer lower lip subtriangular and pointing toward mouth; uncini with a transverse row of teeth above main fang......... 52

51(49). Ventro-lateral pads more or less smooth; midventral groove well defined..................... P. medusa (Mediterranean) Ventro-lateral pads papillated; midventral groove narrow, poorly defined due to encroachment of papillae.................................................................................................... P. papillosus (Brazil)

52(50). Ventro-lateral pads more or less smooth, on segments 3–9................................. P. parvus (NW Australia) Ventro-lateral pads tessellated, on segments 3–15........................................ P. tesselatus (S Australia)