Anaitides Czerniavsky, 1882

Salazar-Vallejo, Sergio I., 2022, New species of hesionid and phyllodocid polychaetes (Annelida, Errantia) from Clipperton Island, Zoosystema 44 (1), pp. 1-26 : 7-11

Genus Anaitides Czerniavsky, 1882 reinstated

Anaitides Czerniavsky, 1882: 148 (key), 158 (diagn.), 159 (species list).

Phyllodoce (Anaitides) – Pleijel 1993b: 298 (list of species).

TYPE SPECIES. — Phyllodoce groenlandica Örsted, 1843 by subsequent designation ( Bergström 1914: 138).

DIAGNOSIS. — Phyllodocids with prostomium with two oblong tapered lateral antennae, median antenna often reduced into a nuchal papilla. Four pairs of oblong tapered tentacular cirri. Dorsal cirrophore without acicula; supracicular lobe usually blunt, as long as subacicular one; ventral cirri usually medially widened. Pharynx with two regions, basal region with large papillae, usually arranged in longitudinal rows, dorsal and ventral areas smooth (without papillae).

ETYMOLOGY. — Anaitis, Anahita or Artemis Anaitis , is the love Godess for Armenians and Persians ( Brosius 1998). Czerniavsky (1882) proposed several new phyllodocid genera, and for some of them followed Théel (1879) who proposed Mystides after Mysta . Anaitides Czerniavsky (1882: 158) was derived from Anaitis Malmgren, 1865 . The latter was preoccupied by Anaitis Duponchel, 1829 (Lepidoptera) and Anaitis Roemer, 1857 (Mollusca) , and was renamed Paranaitis by Southern (1914).

GENDER. — Feminine. After the code ( ICZN 1999, Art. 30.1.4.4), the suffix – ides “is to be treated as masculine unless its author, when establishing the name, stated it had another gender or treated it as such by combining it with an adjectival species-group name in another gender form.” Czerniavsky (1882: 159) listed the species for his new genus, newly transferred from Phyllodoce (feminine) and made no modifications for their corresponding suffixes, thus implying he regarded his new genus-name as having a feminine gender.

DISTRIBUTION. — The species of Anaitides have been described from shallow-water localities in tropical, temperate and polar seas.

KEY TO SPECIES OF ANAITIDES CZERNIAVSKY, 1882

REMARKS

Pleijel (1993b: 295, fig 1G) regarded P. citrina Malmgren, 1865 as incertae sedis especially because it has less than 6 lateral rows of papillae per side on its pharynx basal region. More than six rows were documented for one Anaitides species ( Day 1973: 22), as well as non-regular lateral rows for several other species ( Hartmann- Schröder 1965b: 86; Gathof 1984: 19.33), and differences in the number of papillae per row ( O’Connor 1987: 312). Further, in the new species described below, A. albengai n. sp., variations in the number of lateral rows were noted, and often in the same specimen there was a different number of rows along the left or the right side. However, the presence of smooth areas along dorsal and ventral surfaces are more regular and this explains why the midventral smooth surface is diagnostic, whereas the number of lateral rows is not. Consequently, P. citrina is included in Anaitides as indicated by Uschakov (1972: 130), and after the redescriptions by Pleijel (1988: 143), Pleijel & Dales (1991: 78), and Pleijel (1993a: 35). For the key below, the descriptions or redescriptions were used to key out the species; sometimes, more than one morphological pattern has received the same name, and this explains why the same species name may be reached in two or more alternatives. They are included and might mean problematic records deserving further study. Likewise, the distribution area is incorporated but it does not mean the full distribution of the species.

Anaitides catenula (Verrill, 1873) , A. groenlandica orientalis (Zachs, 1933) , and A. heterocirra Chamberlin, 1919 were not included because of a lack of information in the original descriptions.

1. Dorsal cirri oval ............................................................................................................................................. 2

— Dorsal cirri lanceolate ........................................................................................................................... 26

— Dorsal cirri cordate; ventral cirri foliose, pointed .................................................................................. 41

2(1). Dorsal cirri truncate (upper margin straight) .......................................................................................... 3

— Dorsal cirri non-truncate (upper margin curved) .................................................................................. 13

3(2). Prostomium with eyes ............................................................................................................................. 4

— Prostomium without eyes; ventral cirri pointed, projected far beyond neurochaetal lobes; lateral antennae half as long as prostomial width; pharynx basal papillae globular ............................................................... ............................................................ A. lamella ( de Oliveira, Magalhães & Lana, 2021) n. comb. Brazil.

4(3). Eyes small (1/10-1/15 prostomial width) ................................................................................................... 45

— Eyes large (¼-1/7 prostomial width) ......................................................................................................... 7

5(4). Ventral cirri pointed ............................................................................................................................... 6

— Ventral cirri blunt; prostomium wider than long; lateral antennae as long as 1/6 prostomial width; longest tentacular cirri reach segment 14-15; pharynx basal papillae polygonal ...................................................... .......................................................................................... A. carloensis Kudenov, 1975 Gulf of California.

6(5). Prostomium wider than long; lateral antennae 1/7 as long as prostomial width; longest tentacular cirri reach segment 14-15; pharynx basal papillae conical with a bare middorsal area ................................................. ................................................................................................. A. groenlandica ( Örsted, 1842) Greenland.

— Prostomium as long as wide; lateral antennae 1/5 as long as prostomial width; longest cirri reach segment 15- 18; pharynx basal papillae conical without a middorsal bare area ............................................................... ........................................................................................ A. lineata ( Claparède, 1870) Mediterranean Sea.

7(4). Ventral cirri pointed; lateral antennae ¼ as long as prostomial width ...................................................... 8

— Ventral cirri blunt ................................................................................................................................. 11

8(7). Longest tentacular cirri reach segment 9-10; ventral cirri slightly longer than neurochaetal lobes ............ 9

— Longest tentacular cirri reach segment 12-14 ........................................................................................ 10

9(8). Prostomial posterior notch shallow; longest tentacular cirri reach segment 8-9 .......................................... ................................................................................................. A. australis ( Day, 1975) Western Australia.

— Prostomial posterior notch deep; longest tentacular cirri reach segment 10 ................................................ .......................................................................................................... A. mucosa ( Örsted, 1843) Denmark.

10(8). Ventral cirri slightly longer than neurochaetal lobes .................... A. africana ( Augener, 1918) West Africa.

— Ventral cirri markedly longer than neurochaetal lobes ........ A. mucosa sensu Gathof, 1984 Gulf of Mexico.

11(7). Ventral cirri foliose ............................................................................................................................... 12

— Ventral cirri oval; lateral antennae half as long as prostomial width; longest tentacular cirri reach segment 8 ............................................................................. A. maculata ( Linnaeus, 1767) Northern European seas.

12(11).Eyes ¼ as long as prostomial width; lateral antennae half as long as prostomial width; longest tentacular cirri reach segment 9-13 ................................................................ A. citrina ( Malmgren, 1865) Faeroe Islands.

— Eyes 1/8 as long as prostomial width; lateral antennae 1/3 as long as prostomial width; longest tentacular cirri reach segment 8 ........................................................................... A. williamsi Hartman, 1936 California.

13(2). Ventral cirri foliose ............................................................................................................................... 14

— Ventral cirri cirriform; lateral antennae 2/5 as long as prostomial width; eyes 1/9 as long as prostomial width; longest tentacular cirri reach segment 10, pharynx basal papillae globular and denticulate ......................... ....................................................................................................... A. rosea (M’Intosh, 1877) British Isles.

14(13).Ventral cirri pointed ............................................................................................................................. 15

— Ventral cirri blunt ................................................................................................................................. 22

15(14).Eyes small (1/9-1/12 prostomial width) ..................................................................................................... 16

— Eyes large (1/5-1/7 prostomial width) ....................................................................................................... 24

16(15).Tentacular cirri reach segment 8 ........................................................................................................... 17

— Tentacular cirri reach segment 10-12 .................................................................................................... 18

17(16).Prostomium oval; lateral antennae half as long as prostomial width; eyes central, 1/10 as long as prostomial width .......................................................................... A. pettibonae ( Blake, 1988) Northwestern Atlantic.

— Prostomium cordate; lateral antennae 1/3 as long as prostomial width; eyes posterior, 1/12 as long as prostomial width .............................................................................. A. papillosa ( Uschakov & Wu, 1959) Yellow Sea.

18(16).Dorsal cirri as wide as long, or wider than long ..................................................................................... 19

— Dorsal cirri slightly longer than wide; longest tentacular cirri reach segment 9-10; lateral antennae 1/6 as long as prostomial width ...................................................................... A. koreana Lee & Jae, 1985 Yellow Sea.

19(18).Dorsal cirri barely wider than long; longest tentacular cirri reach segment 11-12; lateral antennae 1/5 as long as prostomial width .............................................................................................................................. 20

— Dorsal cirri 2-3× wider than long .......................................................................................................... 21

20(19).Ventral cirri tips directed downwards; pharynx aperture with 8 papillae ..................................................... ....................................................................... A. longipes ( Kinberg, 1866) Chile (Wesenberg-Lund 1962).

— Ventral cirri tips directed laterally; pharynx aperture with 16 papillae ........................................................ .................................................................. A. longipes sensu Pleijel, 1993 NE Atlantic and Mediterranean.

21(19).Longest tentacular cirri reach segment 12; lateral antennae ¼ as long as prostomial width; dorsal cirri 3× wider than long ............................................................ A. nicoyensis ( Treadwell, 1928) Pacific Costa Rica.

— Longest tentacular cirri reach segment 10; lateral antennae 1/3 as long as prostomial width; dorsal cirri twice wider than long .............................................................. A. panamensis ( Treadwell, 1917) Pacific Panama.

22(14).Eyes large (1/5-1/6 prostomial width) ....................................................................................................... 23

— Eyes small (1/10 prostomial width); prostomium oval, as long as wide; lateral antennae half as long as prostomial width; longest tentacular cirri reach segment 8-9 .............. A. citrina ( Malmgren, 1865) partim NE Atlantic.

23(22).Prostomium oval, as long as wide; lateral antennae half as long as prostomial width; longest tentacular cirri reach segment 7-8 ................................................................................ A. japonica ( Imajima, 1967) Japan

— Prostomium oval, longer than wide; lateral antennae 2/5 as long as prostomial width; longest tentacular cirri reach segment 6 ..................................................................... A. parva Hartmann-Schröder, 1965 Hawaii

24(15).Prostomium as long as wide; longest tentacular cirri reach segment 7; eyes 1/8 as long as prostomial width ..... ............................................................. A. diversiantennata Hartmann-Schröder, 1986 Southern Australia

— Prostomium longer than wide ............................................................................................................... 25

25(24).Longest tentacular cirri reach segment 9; eyes 1/7 as long as prostomial width .. A. longipes ( Kinberg, 1866) Brazil ( de Oliveira et al. 2021)

— Longest tentacular cirri reach segment 12-14; eyes 1/7 as long as prostomial width ...................................... .. A. cf. longipes ( Kinberg, 1866) NE Atlantic ( Pleijel 1988, Fig. 5B ; Pleijel & Dales 1991, Fig. 2B ) Sweden ( Pleijel 1993a).

26(1). Dorsal cirri as long as wide, or slightly longer than wide; prostomium cordate; ventral cirri pointed ..... 27

— Dorsal cirri 2-3× longer than wide ........................................................................................................ 33

27(26).Prostomium wider than long; eyes 1/6 as long as prostomial width ......................................................... 28

— Prostomium as long as wide; eyes 1/6-1/7 as long as prostomial width ...................................................... 31

28(27).Pharynx basal region with about 24 small papillae per row ........................................................................ .................................................................................... A. sanctaevicentis (M’Intosh, 1885) Caribbean Sea.

— Pharynx basal region with 7-13 papillae per row ................................................................................... 29

29(28).Basal pharynx region without middorsal row of papillae ....................................................................... 30

— Basal pharynx region with middorsal row of papillae, with up to 6 papillae; longest tentacular cirri reach segment 11-16 ...................................................................................................................................... 32

30(29).Longest tentacular cirri reach segment 15 ...................... A. erythrophylla ( Schmarda, 1861) Caribbean Sea

— Longest tentacular cirri reach segment 10 ................................ A. medipapillata ( Moore, 1909) California

31(27). Ventral cirri surpassing neurochaetal lobes; pharynx basal region with 8-9 large, foliose papillae per lateral, irregular row ................................................................................................. A. albengai n. sp. Clipperton

— Ventral cirri as long as neurochaetal lobes; pharynx basal region with 11 large, round papillae per lateral row ............................................................................................................................................................ .......................... A. cf medipapillata sensu Villalobos-Guerrero & Tovar-Hernández, 2014 Western Mexico.

32(29).Lateral antennae ¼ as long as prostomial width .................................................................................... 33

— Lateral antennae 1/10-1/16 as long as prostomial width (preserved specimens brownish, sometimes with a middorsal longitudinal band; cirri with darker core) ........................................................................................ A. oculata ( Ehlers, 1887) Florida (incl. A. madeirensis sensu Gathof, 1984 ; Mountford, 1991 Grand Caribbean).

33(32).Longest tentacular cirri reach segment 10 (preserved specimens pale, pharynx papillae and dorsal cirri brownish) ........................................................................................ A. madeirensis ( Langerhans, 1880) Madeira.

— Longest tentacular cirri reach segment 7 (preserved specimens colorful, reddish-violet to brownish) .......... .......................................................................................... A. variabilis Hartmann-Schröder, 1965 Samoa.

33(26).Ventral cirri foliose, pointed .................................................................................................................. 34

— Ventral cirri foliose, blunt ..................................................................................................................... 38

34(33).Eyes visible, large (1/5-1/7 prostomial width) ........................................................................................... 35

— Eyes indistinct; longest tentacular cirri reach segment 7; dorsal cirri twice longer than wide ...................... ................................................................................................. A. dubia Fauchald, 1972 Western Mexico.

35(34).Prostomium as long as wide .................................................................................................................. 36

— Prostomium longer than wide ............................................................................................................... 37

36(35).Eyes 1/5 as long as prostomial width; dorsal cirri twice longer than wide; lateral antennae half as long as prostomial width; dorsal cirri dark brown .......................................... A. fuscacirrata ( Treadwell, 1926) Samoa.

— Eyes 1/7 as long as prostomial width; dorsal cirri 3× longer than wide; lateral antennae 1/3 as long as prostomial width; dorsal cirri pale ........................................... A. salicifolia ( Augener, 1913a) Southwestern Australia.

37(35).Dorsal cirri twice longer than wide; lateral antennae half as long as prostomial width; eyes ¼ as long as prostomial width ................................................................................. A. erythraensis ( Gravier, 1900) Red Sea.

— Dorsal cirri 3× longer than wide; lateral antennae 1/3 as long as prostomial width; eyes 1/5 as long as prostomial width ..................................................................................................... A. elongata Imajima, 1967 Japan.

38(33).Dorsal cirri blunt, without distinct tips ................................................................................................. 39

— Dorsal cirri pointed, with distinct tips; eyes 1/5 as long as prostomial width; longest tentacular cirri reach segment 10 ........................................................................ A. minuta Treadwell, 1937 Gulf of California.

39(38).Prostomial posterior notch deep (reaching level of anterior margin of eyes); tentacular cirri thick, reach segment 7-8; eyes 1/5 as long as prostomial width; lateral antennae 1/3 as long as prostomial width .................. ..................................................................................................... A. sanctijosephi ( Gravier, 1900) Red Sea. — Prostomial posterior notch shallow (reaching level of posterior margin of eyes); tentacular cirri thin; lateral antennae ¼-1/5 prostomial width ........................................................................................................... 40

40(39).Ventral cirri projected far beyond neurochaetal lobe tip; longest tentacular cirri reach segment 9 ............... ................................................................................ A. pseudopatagonica ( Augener, 1922) Juan Fernandez.

— Ventral cirri as long as neurochaetal lobes; longest tentacular cirri reach segment 8 .................................... .............................................................. A. thalia ( de Oliveira, Magalhães & Lana, 2021) n. comb. Brazil.

41(1). Dorsal cirri as long as wide, or slightly longer than wide ....................................................................... 42

— Dorsal cirri twice wider than long ......................................................................................................... 46

42(41).Prostomium cordate ............................................................................................................................. 43

— Prostomium oval, longer than wide; lateral antennae 1/3 as long as prostomial width; eyes 1/6 as long as prostomial width; longest tentacular cirri reach segment 8-9 ..... A. multiseriata ( Rioja, 1941) Western Mexico.

43(42).Prostomium slightly longer than wide; lateral antennae 1/3 as long as prostomial width; eyes 1/6 as long as prostomial width .................................................................................................................................. 44

— Prostomium wider than long ................................................................................................................ 45

44(43).Dorsal cirri with sharp tips; pharynx distal region with series of wider than long tubercles ......................... .................................................................................................... A. adarensis ( Benham, 1927) Antarctica.

— Dorsal cirri with blunt tips; pharynx distal region with large, globular pointed papillae ............................. ........................................................................... A. longipes Kinberg, 1866 sensu Blake, 1997 (California) .

45(43).Eyes 1/7 as long as prostomial width; lateral antennae ¼ as long as prostomial width .................................. .............................................................................................. A. cortezi Kudenov, 1975 Gulf of California.

— Eyes ¼ as long as prostomial width; lateral antennae 1/5 as long as prostomial width .................................. .................................................................................... A. duplex (M’Intosh, 1885) Southeastern Australia.

46(41).Prostomium oval, almost twice wider than long; lateral antennae ¼ as long as prostomial width; eyes minute (1/18 prostomial width) ............................................ A. multipapillata Kravitz & Jones, 1979 Washington.

— Prostomium cordate, slightly wider than long; lateral antennae 1/6 as long as prostomial width; eyes small (1/8 prostomial width) ...................................... A. cuspidata ( McCammon & Montagne, 1979) California.

Figures

FIG. 5. — Pterocirrus bouchardi n. sp., holotype (MNHN-IA-TYPE2046): A, complete, dorsal view; B, anterior region, dorsal view; C, median chaetiger, right parapodium, without dorsal cirrus, posterior view (after Methyl green); D, posterior chaetiger, right parapodium, dorsal cirrus bent posteriorly, posterior view (after Methyl green); E, posterior region. Scale bars: A, 0.6 mm; B, 0.3 mm; C, D: 0.1 mm; E, 0.2 mm.

FIG. 2. — Anaitides albengai n. sp., holotype (MNHN-IA-TYPE2041): A, oblique lateral view of anterior region; B, anterior region, dorsal view (after Shirlastain-A); C, same, oblique lateral view; D, pharynx, anterior end, frontal view; E, chaetiger 50, left parapodium, anterior view (after Methyl green); F, chaetiger 120, left parapodium, anterior view (after Methyl green staining); G, chaetiger 220, right parapodium, anterior view (after Methyl green). Scale bars:A, 0.8 mm; B, 0.5 mm; C, E, 0.3 mm; D, 0.4 mm; F, G, 0.2 mm.