Chloeia Savigny, 1818

Salazar-Vallejo, Sergio I., 2023, Revision of Chloeia Savigny in Lamarck, 1818 (Annelida, Amphinomidae), Zootaxa 5238 (1), pp. 1-134 : 17-20

Chloeia Savigny in Lamarck, 1818

Chloeia Savigny in Lamarck, 1818: 328; Savigny 1822: 14, 58 (diagn.); Audouin & Milne-Edwards 1833: 192, Pl. 9, Figs 11 , 12 ; de Quatrefages 1866: 386 (diagn., syn.); Grube 1878: 9–10; Horst 1910: 170 (key to species); Fauvel 1923: 133–134; Fauvel 1932: 55, 1953: 94; Day 1967: 123; Fauchald 1977: 102 (diagn.); Yáñez-Rivera & Salazar-Vallejo 2022: 508 (diagn.).

Chloenea Kinberg, 1867 (subjective synonym).

Chloochaeta Kinberg, 1867 (subjective synonym).

Strategis Kinberg, 1867 (subjective synonym).

Thesmia Kinberg, 1867: 86 (objective synonym, type species: Aphrodita flava Pallas, 1766 ).

Type species. Amphinome capillata Bruguière, 1789 , by monotypy (junior synonym of Aphrodita flava Pallas, 1766 , see below).

Diagnosis (modif. after Yáñez-Rivera & Salazar-Vallejo 2022: 508). Archinomins with body fusiform. Caruncle trilobate, medial lobe wide, plicate, lateral lobes narrow, plicate, tip tapered or blunt. Dorsal pigmentation patterns variable. Branchiae bipinnate in most chaetigers; a few anterior chaetigers with cirriform branchiae sometimes present. Dorsal cirri with cirrophores. Anus terminal in last chaetigers; posterior end margin with a pair of cirriform lobes.

Remarks. Horst (1910) used, for separating Chloeia species, the pigmentation patterns, start of branchiae, and relative length of ventral cirri of chaetiger 2. The key below follows this method, and some other features have been added such as the size proportions between anterior and posterior eyes, type of branchial ramification (pinnate vs bipinnate), their size trends along body, and the type of chaetae present.

Key to species of Chloeia Savigny in Lamarck, 1818

1 Bipinnate branchiae from chaetiger 3, small; middorsal spots longer than wide, rectangular to slightly widened medially, displaced towards posterior segmental half........................ C. fucata de Quatrefages, 1866 reinstated, Arabian Sea – Bipinnate branchiae from chaetiger 4 ...................................................................... 2 – Bipinnate branchiae from chaetiger 5. .................................................................... 38

2(1) Branchiae progressively smaller towards posterior region..................................................... 3 – Branchiae markedly smaller after a few anterior segments.................................................... 37

3(2) Median segments with middorsal spots.................................................................... 4 – Median segments without pigmentation pattern............................................................ 25

4(3) Middorsal spots oval to circular......................................................................... 5 – Middorsal spots band-shaped............................................................................ 7 5(4) Middorsal spots circular.............................. C. flava (Pallas, 1766) restricted, Bay of Bengal, Indian Ocean – Middorsal spots oval, longer than wide, usually complete, sometimes only posterior half retained..................... 6

6(5) Middorsal spots complete oval, central in each segment; harpoon notochaetae without spurs or minor tines; neurochaetae spurred or furcates with short minor tines.......................... .. C. pulchella Baird, 1868 reinstated, NE Australia

– Middorsal spots half-oval, displaced towards posterior segmental half; harpoon notochaetae with short smooth tines; neurochaetae with long minor tines..................................................................................... C. maculata Potts, 1909 Saint Brandon Rocks, Western Indian Ocean (incl. C. natalensis Day, 1951 South Africa)

7(4) Middorsal bars roughly homogeneously wide along segments.................................................. 8 – Middorsal spots heterogeneous, increasing, decreasing or separate in each segment................................ 13

8(7) Middorsal bands running along each segment, forming a continuous band........................................ 9

– Middorsal bands short, not running along full segment......................... C. slapcinskyi sp. n. Philippine Islands

9(8) Middorsal bands regular, not constricted posteriorly; caruncle pale............................................. 10

– Middorsal bands irregular, truncate anteriorly, constricted posteriorly; caruncle blackish; anterior eyes 3× larger than posterior ones.......................................... C. nuriae Yánez-Rivera & Salazar-Vallejo, 2022 Gulf of California

10(9) Anterior parapodia without harpoon-chaetae.............................................................. 11

– Anterior parapodia with harpoon chaetae and a few furcates; neurochaetae furcates with major tines 3–6 longer than minor ones in anterior chaetigers, 3–8 in median chaetigers................................... C. piotrowskiae sp. n. Philippines

11(10) Anterior notochaetae furcates, major tines 2–3×longer than minor ones......................................... 12

– Anterior notochaetae with tiny spurs; harpoon-chaetae without basal tine............................................................................................. C. gilchristi McIntosh, 1924 reinstated, South Africa (partim)

12(11) Anterior prostomial area pale; neurochaetae furcates with major tines 2× longer than minor ones................................................................................. C. venusta de Quatrefages, 1866 Mediterranean Sea

– Anterior prostomial area blackish; neurochaetae furcates with major tines 5–6× longer than minor ones......................................................................................... C. poupini sp. n. French Polynesia

13(7) Middorsal bands anteriorly widened, or posteriorly widened.................................................. 14

– Middorsal spots medially constricted or separate........................................................... 23

14(13) Middorsal bars expanded anteriorly, decreasing towards posterior segment margin, often T- or Y-shaped............... 15

– Middorsal bars expanding medially or posteriorly, often decreasing towards anterior segment margin................. 21

15(14) Dorsum with middorsal and lateral bands well defined, usually narrower than middorsal band....................... 16

– Dorsum with middorsal band well defined, without lateral bands.............................................. 20

16(15) Additional lateral bands parallel........................................................................ 17

– Additional lateral bands not parallel, oblique, or crossing middorsal spots ........................................ 18

17(16) Median antenna half as long as caruncle; anterior eyes slightly larger than posterior ones; dorsal bands dark purple (orange to reddish in living specimens); neurochaetae acicular or furcates....... C. pseudeuglochis Augener, 1922 Pacific Costa Rica

– Median antenna as long as caruncle, or slightly shorter than it; anterior eyes 2–3× larger than posterior ones; dorsal bands blackish to brownish (unknown in living specimens); neurochaetae acicular or spurred .............. C. mezianei sp. n. Senegal

18(16) Lateral bands oblique................................................................................. 19

– Lateral bands crossing middorsal spots anteriorly, forming a cross in each segment....... C. euglochis Ehlers, 1887 Florida

19(18) Middorsal bands Y-shaped to subtriangular, progressively thinner along each segment.................................................. C. incerta de Quatrefages, 1866 reinstated, restricted, Indonesia, Sulawesi (incl. C. parva Baird, 1868

– Middorsal bands T-shaped, width decreasing rapidly along each segment.............. C. viridis Schmarda, 1861 Jamaica

20(15) Bipinnate branchiae with 11–12 lateral branches; caruncle pale, with black median ridge, with 34 vertical folds. ...................................................................................... C. hutchingsae sp. n. Australia

– Bipinnate branchiae with 6–7 lateral branches; caruncle blackish, with 24 vertical folds.. ..................................................................................... C. zibrowii sp. n. French Polynesia, Marquesas Islands

21(14) Middorsal spots as inverted T.......................................................................... 22

– Middorsal spots expanded medially, never T- or anchor shaped; notochaetae of median chaetigers only harpoon-chaetae without spurs...................................................................... C. amoureuxi sp. n. Madagascar

22(21) Posterior region with middorsal bands narrow, lateral branches short; notochaetae of median chaetigers harpoon-chaetae with tines..................................................................... C. violacea Horst, 1910 Indonesia

– Posterior region with middorsal bands wide, lateral branches well-defined, wide; notochaetae of median chaetigers harpoonchaetae with and without spurs and aciculars............................................. C. gilleti sp. n. Senegal

23(13) Middorsal bands constricted medially or discontinuous, surrounded by a pale area, without lateral bands or barely pigmented.................................................................................................... 24

– Middorsal bands constricted medially, surrounded by anterior expansions of lateral dark bands........................................................................................... C. conspicua Horst, 1910 Indonesia, Java

24(23) Prenotopodial dark band wide; middorsal spots continuous in each segment, slightly wider anteriorly, thinner medially, rounded posteriorly (amphora-like), surrounded by a pale thin band......................... C. amphora Horst, 1910 Indonesia

– Prenotopodial dark band narrow; middorsal bands markedly constricted medially, separated into two spots in median segments..................................................... C. bimaculata Wang, Zhang, Xie & Qiu, 2019 Hong Kong

25(3) Neuropodial furcate chaetae with tines well developed, rarely with spurs........................................ 26

– Neuropodia with most chaetae acicular or spurred (furcates with tiny spurs), furcate chaetae rare..................... 32

26(25) Eyes minute; dorsal cirri pale; second ventral cirri with cirrostyles 3× longer than adjacent ones; first bipinnate branchiae slightly smaller than following ones................... C. paulayi Yáñez-Rivera & Salazar-Vallejo, 2022 Gulf of Mexico

– Eyes large; dorsal cirri with pigmentation, never pale........................................................ 27

27(26) Caruncle tapered; dorsal cirri purple (chaetigers 4–6), or purple dorsal cirrophores (all body), rarely pale.............. 28 – Caruncle blunt, pale; dorsal cirri dark purple... C. entypa Chamberlin, 1919 Western Mexico

28(27) Bipinnate branchiae colorless. .......................................................................... 29

– Bipinnate branchiae purplish, at least basally.............................................................. 30

29(28) Anterior eyes 2× larger than posterior ones; harpoon notochaetae with spurs; neurochaetae spurred...................................................................................... C. pinnata Moore, 1911 Southern California

– Anterior eyes 4× larger than posterior ones; harpoon notochaetae without spurs; neurochaetae furcates with small minor tines........................................................................ C. keablei sp. n. Papua New Guinea

30(28) Branchiae with stems purplish, pigment often extended along branches, at least along anterior chaetigers.............. 31 – Branchiae with stems pale, with reddish to blackish spots basally and distally in median chaetigers, distally in some lateral branches................................................................ C. bemisae sp. n. Philippine Islands

31(30) Chaetiger 3 notochaetae include aciculars, harpoon-chaetae and furcates; anterior eyes 8–10× larger than posterior ones; branchiae massive................................................................ C. fauveli sp. n. Bay of Bengal

– Chaetiger 3 notochaetae include furcates and aciculars, without harpoon-chaetae; anterior eyes 3–4× larger than posterior ones; branchiae delicate, tapered, not massive............................................. C. murrayae sp. n. Australia

32(25) Body fusiform, gradually tapered posteriorly; branchiae with lateral branches tapered.............................. 33

– Body tapered posteriorly (deep-water forms).............................................................. 34

33(32) Body wall brownish; caruncular median ridge with 11 vertical folds; median segments with acicular neurochaetae; anterior neurochaetae spurred.......................................... C. gilchrsti McIntosh, 1924 South Africa, reinstated

– Body wall pale; caruncular median ridge with about 38 vertical folds; all neurochaetae spurred.. .................................................................................................... C. tumida Baird, 1868 India

34(32) Bipinnate branchiae with 8–9 lateral branches............................................................. 35

– Bipinnate branchiae with 11–12 lateral branches, tapered..................................................... 36

35(34) Anterior prostomial area pale; branchiae with lateral branches blunt; body wall pinkish; median segments with notochaetae acicular and harpoon-chaetae without tines.................................... C. gesae sp. n. Northeastern Atlantic

– Anterior prostomial area blackish; branchiae with lateral branches tapered; body wall whitish; median segments with notochaetae acicular and harpoon-chaetae with shorter tines........................................ C. fiegei sp. n. Red Sea

36(34) Anterior prostomial area blackish; bipinnate branchiae with bases pale; notochaetae all furcates.................................................................................................. C. richeri sp. n. New Caledonia

– Anterior prostomial area pale; bipinnate branchiae with bases orange; notochaetae furcates and harpoon-chaetae.. ...................................................................................... C. boucheti sp. n. Indonesia

37(2) Branchiae better developed along chaetigers 4–8; dorsal cirri barely pigmented basally; neurochaetae furcates, major tines smooth............................................................ C. kudenovi Barroso & Paiva, 2011 Brazil

– Branchiae better developed along chaetigers 4–9; dorsal cirri dark purple along their basal haf; neurochaetae furcates, major tines denticulate along inner margin.. .................... C. pocicola Barroso & Kudenov in Barroso et al. 2021 Brazil

38(1) Dorsum pale, with homogeneous pigmentation, or with a middorsal pale band.................................... 39

– Dorsum with two longitudinal bands.................................................................... 43

39(38) Branchiae bipinnate.................................................................................. 40

– Branchiae pinnate; dorsum pink; median segments with harpoon chaetae.............. C. rosea Potts, 1909 Indian Ocean

40(39) Median chaetigers with harpoon chaetae.................................................................. 41

– Median chaetigers without harpoon-chaetae (or rarely present)................................................ 42

41(40) Chaetae of median segments markedly longer than body width; median antenna shorter than caruncle; branchiae short, as long as one successive segment............................................. C. longisetosa Potts, 1909 Maldive Islands

– Chaetae of median segments about as long as body width; median antenna as long as, or longer, than caruncle; branchiae long, as long as two successive segments, without basal spots ........................ C. slapcinskyi sp. n. Philippine Islands

42(40) Notochaetae furcates, major tines 3–4× longer than minor ones; median antenna as long as caruncle; branchiae with a white basal spot...................................................................... C. wangi sp. n. Philippines

– Notochaetae spurred or aciculars; median antenna shorter than caruncle; branchiae without white basal spot................. C. inermis de Quatrefages, 1866 New Zealand (incl. C. spectabilis Baird, 1868 and C. australis Kudenov, 1993 Campbell Plateau (500 km S off New Zealand)

43(38) Anterior eyes 3–4× larger than posterior ones; caruncle with median ridge with 18 vertical folds....................................................................... C. fusca M’Intosh, 1885 restricted, Indonesia ( Moluccas Islands)

– Anterior eyes 6× larger than posterior ones; caruncle with median ridge with 12 vertical folds. ............................................................................................. Chloeia bistriata Grube, 1868 Red Sea

Remarks. The length of ventral cirri of chaetiger 2 has two patterns. In one, these cirri are longer than those present in adjacent segments, whereas in the other, they are of similar length. This feature was used in previous drafts of this key but it is often broken, making this comparison difficult, and this explains why it was removed from the key. Further, the videos available in internet with living specimens clearly show all ventral cirri are directed ventrolaterally; in most preserved specimens the hypertrophied ventral cirri of chaetiger 2 are directed dorsally but this is a preservation artifact.

Figures

FIGURE 11. Chloeia amphora Horst, 1910, non-type specimens (UF 4369). A. Dorsal view. B. Anterior end, dorsal view, after Shirlastain-A pigmentation. C. Chaetigers 10-12, dorsal view (left branchia of chaetiger 10 removed). D. Posterior end, ventral view. E. Another specimen, chaetiger 3, notochaeta (insets: notochaetal tips). F. Same, neurochaetal tips. G. Chaetiger 12, harpoon notochaetae. H. Same, neurochaetal tips. Scale bars: A, 1.5 mm; B, 0.5 mm; C, 0.4 mm; D, 0.7 mm; E, 110 μm; F, 70 μm; G, 160 μm; H, 40 μm.

FIGURE 12. Chloeia bemisae sp. n., holotype (UF 4388).A. Dorsal view. B. Anterior end, dorsal view. C. Left lateral view. D. Paratype (UF 4388b), ventral view. E. Same, chaetiger 4, notochaetal tips. F. Same, neurochaetal tips. G. Chaetiger 11, harpoon notochaetae. H. Same, neurochaetae. Scale bars: A, 1.8 mm; B, 0.5 mm; C, 2.3 mm; D, 1.6 mm; E, 110 μm; F, 25 μm; G, 90 μm; H, 100 μm.