Myrianida Milne Edwards 1845
Myrianida Milne Edwards, 1845 Scolopendra Slabber, 1781: 44 –46, pl 10, fig. 4–5. Nomen oblitum according to Article 23.9. 1 (ICZN 1999). Podonereis Blainville, 1818: 83. Nomen oblitum according to Article 23.9. 1 (ICZN 1999). Myrianida Milne Edwards, 1845: 180, figs 65–68. Nomen protectum according...
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Summary: | Myrianida Milne Edwards, 1845 Scolopendra Slabber, 1781: 44 –46, pl 10, fig. 4–5. Nomen oblitum according to Article 23.9. 1 (ICZN 1999). Podonereis Blainville, 1818: 83. Nomen oblitum according to Article 23.9. 1 (ICZN 1999). Myrianida Milne Edwards, 1845: 180, figs 65–68. Nomen protectum according to Article 23.9. 1 (ICZN 1999). Autolytus Grube, 1850: 310, table facing 281. Diploceraea Grube, 1850: 312, table facing 281. Sacconereis J. Müller, 1853: 31. Crithida Gosse, 1855: 309 –310, pl. 8, fig. 5. Sylline Grube, 1860: 87 –88, pl. 3, fig. 8. Autolytides Malaquin, 1893: 76. Nomenclatural remarks . Scolopendra was described for a female stolon. Podonereis was erected for Nereis punctata (Hesionidae: Nereimyra) and Nereis corniculata , the latter a male stolon. Neither names have been in use since they were introduced, the conditions in Article 23.9. 1.1 (ICZN 1999) are hence fulfilled. The taxon Myrianida has been used by the following authors Day (1960; 1967), Cognetti (1958; 1961), Clark (1961), Hartman (1966 a, b), Rullier (1964), HartmannSchröder (1965 a), Imajima (1966; 1967; 1982), Gidholm (1967), Bellan (1969), Uebelacker (1984), San Martín & Alós (1989), Hartmann Schröder & Rosenfeldt (1990; 1992), Garwood (1991), San Martín (1994; 2003), Parapar, San Martín, Besteiro & Urgorri (1996), López & San Martín (1997), Franke (1999), Nygren & Gidholm (2001), Nygren & Sundberg (2003), thus the conditions in Article 23.9. 1.2 (ICZN 1999) is met with. Linnean name definition . Type species Myrianida fasciata Milne Edwards, 1845, by monotypy. Note that M. fasciata is considered to be a junior synonym to M. pinnigera (Montagu, 1808). Stembased name definition . Myrianida refers to the most inclusive clade comprising M. fasciata Milne Edwards, 1845, M. prolifera (Müller, 1788), and M. irregularis (Imajima & Hartman, 1964), but not Epigamia noroi (Imajima & Hartman, 1964), or Proceraea picta Ehlers, 1864. Apomorphies . Clade supported by two morphological apomorphies (Fig. 4): 1) number of teeth equals 26–30 (character 40), other most parsimonious reconstructions (MPRs) possible, character state change is a parallelism, later reversed and further changed within clade; 2) number of different sizes of teeth in trepan equals 3 (character 41), other MPRs possible, character state change is a parallelism, later reversed and further changed within clade, character state unknown in some taxa. General description . Atokous form . Length 1.2–21 mm for 16–109 chaetigers; width, measured at level of proventricle excluding parapodial lobes, 0.2–1.4 mm. Body shape, excluding parapodial lobes, cylindrical to rounded rectangular in transection, venter flattened; body width fairly constant with tapering end. Ciliation as 1 or 2 trochs per segment (unknown in some taxa). Prostomium rounded rectangular. Four eyes, with lenses, in trapezoid arrangement, anterior pair larger. Eyes confluent or separated; eye spots absent or present. Palps fused at base or completely fused; in dorsal view not projecting in front of prostomium, or projecting 1 / 5 – 2 / 3 of prostomial length. Extension of nuchal epaulettes to between end of tentacular segment and end of chaetiger 12. Prostomium with 3 antennae; median antenna inserted medially on prostomium, lateral antenna on anterior margin. Tentacular cirri 2 pairs. Cirri, cirrophores, cirrostyles may be equal in length or alternating (reported length compared to body width excluding parapodial lobes). Appendages may be cylindrical, thick and swollen, slightly flattened or flattened. Parapodial lobes medium to large in size, rounded, in some taxa with dorsal part prolonged. All chaetae, except bayonet chaetae, compound with bidentate blades; blade serration present. Compounds with small or large distal tooth. Single thin bayonet chaetae, subdistally denticulated. Pharynx with single to multiple sinuations anterior and lateral to proventricle. Trepan with various types of denticulation, arranged in 1 ring. Basal ring absent, or present, variously developed; infradental spines absent or present. Proventricle with varying number of rows of square shaped muscle cells. Pygidium with 2 cirri (reported length compared to body width, excluding parapodial lobes, at level of proventricle, if not otherwise stated), median papilla absent. Epitokes . Male. Body divided into 3 regions, anterior with 2–4 (usually 3) uniramous chaetigers (region a), median with 14–24 biramous chaetigers (region b), and posterior with 0–8 uniramous chaetigers (region c). Body widest in anteromedian half of region b; body width measured in region a, as body width excluding parapodial lobes, and in region b, at the widest part, as body width including parapodial lobes. Ciliation as in stock, 1 or 2 trochs per segment. Prostomium rounded rectangular, wider than long, with straight or concave anterior margin. Eyes large, 2 pairs, with lenses; eyes situated on dorsal and ventral side of prostomium, ventral pair larger. Palps absent. Nuchal epaulettes extending maximally to anterior part of chaetiger 1. Large median antenna inserted on posterior part of prostomium, lateral antennae inserted on anterior margin. Lateral antennae bifid, basal part 1 / 6 – 1 / 2 of total length; outer ventral rami equal to or up to 5 times longer than inner dorsal rami. Basal part and inner dorsal rami segmented with abundant cilia. Pair of small frontal processes inserted anterolaterally to dorsal pair of eyes. Pair of small processes may be present ventral to lateral antennae, proximal to ventral pair of eyes. Tentacular cirri 1 or 2 pairs. First dorsal cirri (corresponding to "tentacular cirri" (e.g. Gidholm 1967) or "lateral horns" (Hamond 1974) in earlier literature, equal in shape and size to median antenna; achaetous knobs absent. Second dorsal cirri situated above first chaetigerous lobes (referred to as cirri on chaetiger 1 to avoid confusion). Cirri length measured as body width excluding parapodial lobes in region a and c; as body width including parapodial lobes in region b. Median antenna with large ceratophore, small cirrophores absent or present on tentacular cirri, mostly too small to be adequately assessed, cirrophores absent or present on dorsal cirri. Appendages may be cylindrical, fusiform, thick, slightly flattened, flattened, or club shaped. Parapodia in region a and c as in stock but generally smaller; in region b large, flattened and equipped with welldeveloped notopodia. Single neuropodial acicula; notopodia in region b supported by 2 anterodorsal aciculae, and 2–3 thick and 3 thin posteroventral aciculae. Neuropodial chaetae as in stock. Notopodial lobes with 15–30 swimming chaetae, in length equal to c. body width in region b including parapodial lobes. Pygidium with 2 cirri (reported length compared to body width in region a, excluding parapodial lobes, if not otherwise stated). Female. Body divided into 3 regions, anterior with 2–6 uniramous chaetigers (region a), median with 14–23 biramous chaetigers (region b), and posterior with 2–15 uniramous chaetigers (region c). Body width fairly constant, slightly wider in region b; body width measured in region a, as body width excluding parapodial lobes, and in region b as body width including parapodial lobes. Ciliation as in stock, 1 or 2 trochs per segment. Prostomium rounded rectangular, wider than long, anterior margin straight or concave. Eyes large, 2 pairs, with lenses; eyes situated on dorsal and ventral side of prostomium, ventral pair larger. Palps absent. Nuchal epaulettes maximally extending to anterior part of chaetiger 1. Median antenna inserted medially on prostomium, lateral antennae inserted on anterior margin. Tentacular cirri 1 or 2 pairs. First dorsal cirri situated above first chaetigerous lobes; achaetous knobs absent. Cirri length measured as body width excluding parapodial lobes in region a, and c; as body width including parapodial lobes in region b. Small cirrophores absent or present on tentacular cirri, mostly too small to be adequately assessed, cirrophores absent or present on dorsal cirri. Appendages may be cylindrical, fusiform, thick, slightly flattened, flattened, or club shaped. Parapodia in region a and c as in stock but generally smaller; in region b with notopodia, not as developed as in male. Single neuropodial acicula; notopodia in region b supported by 2 anterodorsal aciculae, and 2 thick and 2–3 thin posteroventral aciculae. Neuropodial chaetae as in stock. Notopodial lobes with 15–20 swimming chaetae, in length equal to c. 1.5 times body width in region b, including parapodial lobes. Pygidium with 2 cirri (reported length compared to body width in region a, excluding parapodial lobes, if not otherwise stated). Myrianida arborea (Westheide, 1974) comb. n. (Fig. 55 A–C) Autolytus arboreus Westheide, 1974: 319 –320, figure 59 A–F. Material examined . Galápagos : holotype ZMH P 13620 and 1 paratype SMF 10664, Santa Cruz, Bahía Academy, northern side, intertidal, under stones, Jul 1972. Diagnosis . Myrianida with c. 18 equal teeth, equal cirri with equal cirrostyles and cirrophores. Description . Holotype is a rear fragment, description is based on the paratype. Paratype incomplete, length 1.5 mm for 22 chaetigers, width 0.1 mm. Preserved material without colour markings. Ciliation not possible to assess. Eyes separated; eye spots absent. Palps in dorsal view projecting 1 / 5 of prostomial length, fused. Extension of nuchal epaulettes to half of chaetiger 1. Median antenna reaching c. chaetiger 8. Lateral antennae and dorsal tentacular cirri, length 1 / 2 of median antenna. Ventral tentacular cirri 1 / 2 as long as dorsal pair. First dorsal cirri as long as median antenna, second dorsal cirri as long as following dorsal cirri. Alternation in direction of cirri not assessed. Dorsal cirri from chaetiger 3, of equal length, 1 / 3 of body width. Cirrophores present on tentacular cirri, and all dorsal cirri. Cirrophores and cirrostyles equal; cirrophores much shorter than parapodial lobes; cirrophores shorter than cirrostyles. All appendages cylindrical. Parapodial lobes rounded, of medium size. Single acicula in all chaetigers. Chaetal fascicle with 5–6 compounds in all chaetigers. Compound chaetae with small distal tooth; serration present. Single thin bayonet chaetae present in all chaetigers. Pharynx with 1 sinuation anterior to proventricle. Trepan in chaetiger 3–4, with c. 18 equal teeth, arranged in 1 ring (Fig. 55 A). Basal ring moderately to well developed; infradental spines present (Fig. 55 B). Proventricle equal in length to 1.5 segments (Fig. 55 C) in chaetiger 7–8 with c. 20 rows of muscle cells. Pygidium (holotype) has lost its cirri. Reproduction . Unknown Habitat . Intertidal, under stones. Distribution . Central East Pacific. Galápagos. Only known from type locality. Remarks . Myrianida arborea is poorly known, it is similar if not identical to M. brevipes described from Salvador. Myrianida arborea is also very similar to M. edwarsi (Saint Joseph, 1887) but differs in that it lacks the orange colour along the sides in the pharyngeal region found in M. edwarsi the trepan has generally more teeth in M. edwarsi (24–34), compared to c. 18 but variation is not adequately known in M. arborea . Myrianida australiensis (HartmannSchröder, 1982) comb. n. (Fig. 56 A–E) Autolytus prolifer australiensis HartmannSchröder, 1982: 75 –76, figs 73–75; 1983: 136; 1989: 33; 1990: 57; 1991: 43. Autolytus devaneyi HartmannSchröder, 1992 b: 62, figs 26–29. Material examined . Australia : holotype ZMH P 16779, 1 paratype ZMH P 16780, Cape Naturaliste, Eagle bay, tidal flats, algae with sand, 7 Nov 1975; French Polynesia: holotype of Autolytus devaneyi ZMH P 20703, Rangiroa, lagoon, calcareous algae with sand, 7 Sep 1982. Diagnosis . Myrianida with extension of nuchal epaulettes to end of chaetiger 3; 16–18 equal teeth in trepan, thin distinct basal ring; short equal cirrophores and alternating cirrostyles. Description . Length 1.7–2 mm for 31–45 chaetigers, width 0.2 mm. Preserved material whitish, no colour markings. Ciliation not possible to assess. Eyes separated (Fig. 56 B); eye spots present. Palps in dorsal view projecting 1 / 4 – 1 / 3 of prostomial length (Fig. 56 A), fused. Extension of nuchal epaulettes to end of chaetiger 3 (Fig. 56 B). Median antenna lost. Lateral antennae reaching chaetiger 7–9. Tentacular cirri and first dorsal cirri lost. Cirri on chaetiger 2 equal in length to body width. From chaetiger 1– 27 cirri with usual alternation in direction, more posterior difficult to assess. Dorsal cirri from chaetiger 3 alternate in length; short cirri equals 1 / 2 of body width, long cirri equals 3 / 4 of body width. Cirrophores present on tentacular cirri, and all dorsal cirri. Cirrophores equal, cirrostyles unequal; short cirrostyles c. 2 / 3 in length of long cirrostyles; cirrophores shorter than parapodial lobes; cirrophores shorter than cirrostyles. Appendages cylindrical, including lost median antenna (HartmannSchröder 1982). Parapodial lobes rounded, of medium size. Single acicula in all chaetigers. Chaetal fascicle with 5–7 compounds in all chaetigers. Compound chaetae with small distal tooth; serration present (Fig. 56 E). Single thin bayonet chaetae (Fig. 56 D), beginning at chaetiger 4. Pharynx with from one to several sinuations anterior and lateral to anterior half of proventricle (Fig. 56 A). Trepan in chaetiger 1 (Fig. 56 B), with 16–18 equal teeth (Fig. 56 C), arranged in 1 ring. Thin distinct basal ring; infradental spines present (Fig. 56 C). Proventricle equal in length to 3–5 segments in chaetiger 8–13 (Fig. 56 A) with 25–27 rows of muscle cells. Anal cirri lost. Reproduction and morphology of epitokous stage . Schizogamy. The holotype of Autolytus devaneyi has 4 regenerating posterior chaetigers (Fig. 56 A) that possibly represent a developing stolon. HartmannSchröder briefly describes stolons that she refers to Myrianida australiensis . Male stolons with (2–3)+(13–15)+0 chaetigers, for 1.13 mm, female stolons with 25 chaetigers, swimming chaetae in posterior chaetigers only (possibly immature). Habitat . Amongst algae, in intertidal zone. Distribution . East Indian Ocean, South Pacific. West and east Australia, French Polynesia. Remarks . Myrianida australiensis was originally described as a subspecies of Autolytus prolifer , and was later redescribed as A. devaneyi by the same author. The reason for this was probably due to that Myrianida australiensis wrongly was interpreted as having 10 teeth, and nuchal epaulettes reaching end of chaetiger 1. Myrianida australiensis is most similar to M. brevicirrata (Winternitz, 1936) with which it shares the thin basal ring, and combination of equal cirrophores and unequal cirrostyles. However M. australiensis has longer nuchal epaulettes reaching end of chaetiger 3 compared with end of chaetiger 1 in M. brevicirrata , and has only 16–18 teeth compared with 26–30 in M. brevicirrata in addition M. australiensis has shorter cirrophores than M. brevicirrata . Myrianida brachycephala (Marenzeller, 1874) comb. n. (Fig. 57 A–E) Proceraea brachycephala Marenzeller, 1874: 54 –56, pl. 6, fig. 2, 2A–, pl. 7, fig. 2, 2A–B. Proceraea luxurians Marenzeller, 1874: 50 –54, pl. 6, fig. 1, 1A–D, pl. 7, fig. 1. Autolytus mirabilis Verrill, 1882: 367 –368; 1884: 662; Hartman 1944: pl. 13, fig. 8–10. Autolytus punctatus SaintJoseph, 1887: 233 –234, pl. 11 fig. 108–109; Fauvel 1923: 318, fig. 122 L–M. Autolytus ehbiensis in part SaintJoseph, 1887: 228 –233. Autolytus brachycephalus Fauvel 1923 (in part): 316–317, fig 121 G–H; Gidholm 1963; 1965; 1967: 182, 188– 191, figs 7 B, 13 A–B, 14 B, 15, 19A, 20; Hamond 1967: 1 –4, fig. 5 A–B; 1969 a; 1969 c; Hartmann– Schröder 1971: 175 –176; Hamond 1974; Schiedges 1979 a; 1979 b; 1980; San Martín & Alvarado 1981: 223 –224, fig. 2; Kirkegaard 1992: 224 –225, fig 108 A–B; HartmannSchröder 1996: 181; San Martín 2003: 500 –502: fig. 277 A–D; Nygren & Sundberg 2003: GenBank sequences, 16 S rDNA partial sequence AF 474250, and 18 S rDNA partial sequence AF 474296. Autolytus aurantiacus Korringa 1951: 79 –84, figs 8 A(a–d), 8 B(e–f). Autolytus benazzi Cognetti, 1953 b: 89, fig. 1; 1957: 70–71, fig. 14 A–B. Material examined . Italy : 2 syntypes of Proceraea brachycephala on 1 slide, NHMW 2501, Adriatic Sea, 1872; 2 syntypes of P. luxurians on slides, NHMW 2504, Adriatic Sea, 1875. USA : 19 syntypes of A. mirabilis , USNM 10012. Vineyard Sound, 4.75–6 fmns, 18 Aug 1882; 32 female stolons, 1 male stolon YPM 2719, Woods Hole, Massachusetts, surface, 5 Aug 1882. Faroes : 18 spms (13 spms in formalin, 5 spms on slides (2 rear ends in author's collection for DNA analyses)), East of Bordøy, 62 °04.6'N 06° 26.3 'W, triangle dredge, 65–68 m, shellgravel with hydroids, 4 Jul 1997; 6 spms (1 spm in formalin, 4 spms on slide (rear ends in author's collection for DNA analyses), 1 spm in author's collection for DNA analyses), East of Nolsøy, 62 °01.5'N 06° 31.9 'W, triangle dredge, 41 m, bedrock, 27 Jul 1997; 1 spm on slide (rear end preserved for DNA), East of Bordøy, 62 °08.4'N 06° 23.4 'W, triangle dredge, 77 m, shellgravel with hydroids, 4 Jul 1997. Wales : 2 spms on slides (rear ends in author's collection for DNA analyses), Black Point, 53 ° 18.8 'N 04° 2.4 'W, intertidal, algae with epifauna, May 2000. France : 5 syntypes of Autolytus ehbiensis MNHN 1043, 1046, 1050, 1051, 1052, 1054, Dinard, Jul–Aug 1877– 1882; 2 spms on slides (rear ends in author's collection for DNA analyses), Banyulssur Mer, 42 ° 29.6 'N 03° 10.2 'E, epibenthic sledge, 59–62 m, tunicates and shells with epifauna, 26 Apr 2001. Norway : 3 spms (2 spms in formalin, 1 spm on slide (rear ends in author's collection for DNA analyses)), 63 &de : Published as part of Nygren, Arne, 2004, Revision of Autolytinae (Syllidae: Polychaeta)., pp. 1-314 in Zootaxa 680 on pages 115-130, DOI: 10.5281/zenodo.157809 : {"references": ["Milne Edwards, M. (1845) Observations sur le developpement des annelides. Annales des sciences naturelles, Paris, 3, 145 - 182.", "Slabber, M. (1781) Physikalische Belustigungen, oder mikrosckopische Wahrnehmungen von drei und vierzig in- und auslaendischen Wasser- und Landthierchen, Nurnberg, 1 - 99.", "Blainville, H. (1818) Memoire sur la classe des Setipodes, partie des Vers a sang rouge de M. 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