Proceraea madeirensis

Nygren, Arne, 2004, Revision of Autolytinae (Syllidae: Polychaeta)., Zootaxa 680, pp. 1-314 : 56-64

Proceraea madeirensis nom. n. (Fig. 15A–B)

Proceraea fasciata Langerhans, 1879: 581 , fig. 33A–C. Junior homonym of Nereis fasciata Bosc, 1802 .

? Autolytus (Proceraea) fasciata Augener 1913: 264 –265.

? Proceraea fasciata Westheide 1974: 323 –325, figs 61–62; Hartmann­Schröder 1987: 44 –45, figs 20–22.

Material examined. Madeira: holotype NHMW 2512.

Diagnosis. Proceraea with brown intrasegmental bands; antennae, dorsal tentacular cirri, and first dorsal cirri brown.

Description. Holotype on slide, in poor condition, dried, length 4.2 mm for 35 chaetigers, width 0.7 mm (compressed specimen). Preserved material brown, no colour markings. Live specimens with brown intrasegmental bands, as well as brown antennae, dorsal tentacular cirri and first dorsal cirri ( Langerhans 1879). Ciliation not possible to assess.

Eyes faded, not seen. Palps in dorsal view projecting 1/3 of prostomial length (Fig. 15A), fused. Nuchal epaulettes extending to beginning of first chaetiger.

Anterior appendages curled, not evaluated. Dorsal cirri from chaetiger 3, of equal length, 1/5–1/4 of body width. Cirrophores on tentacular segment and first dorsal cirri not possible to assess, cirrophores on other cirri absent. All appendages cylindrical.

Parapodial lobes small. Aciculae not seen. Chaetal fascicle with 6–7 compounds in anterior chaetigers, 4–5 in median and posterior. Compound chaetae in median chaetigers with large distal tooth (Fig. 15B), anterior chaetigers not possible to examine; serration present. Single thick bayonet chaetae, (Fig. 15B), beginning at chaetiger 20.

Pharynx with sinuation anterior to proventricle (Fig. 15A). Trepan in chaetiger 4 (Fig. 15A) with 18 unequal teeth, 9 large and 9 smaller; 1 large alternating with 1 small, arranged in 2 rings. Basal ring present; infradental spines absent. Proventricle equal in length to 3–4 segments in chaetigers 7–10, with 55–60 rows of muscle cells. Pygidium with 2 anal cirri.

Reproduction. Unknown.

Habitat. Intertidal, amongst algae.

Distribution. North East Atlantic. Madeira.

Remarks. The identity of Augener's, Westheide's and Hartmann­Schröder's specimens to Langerhans' taxon is doubtful. Both Westheide and Hartmann­Schröder describe unidentate chaetae on their specimens, the latter author also examined specimens collected by Augener and found unidentate chaetae in these; this type of chaetae are not present in the holotype and are not described by Langerhans. In addition Augener and Westheide describe the brown transverse bands to be intersegmental rather than intrasegmental. New material from Madeira or from the vicinity is needed to get a more accurate description and opinion of the taxon.

Etymology. Langerhans' species has long been a junior homonym to P. fasciata ( Bosc, 1802) , and the new name madeirensis is introduced, meaning "from Madeira".

Proceraea micropedata ( Hartmann­Schröder, 1962) (Fig. 16A–C) Odontosyllis micropedata Hartmann­Schröder, 1962: 100 –103, figs 87–92. Proceraea micropedata Orensanz 1974: 28 .

Material examined. Chile: holotype ZMH P­ 14762, mouth of Rio Andalien, algae and barnacles, 10 Mar 1960.

Diagnosis. Proceraea with egg shaped dorsal cirri (see remarks).

Description. Holotype incomplete anterior fragment, length 1.9 mm for 13 chaetigers, width 0.5 mm. Preserved material brown, without colour markings. Ciliation not possible to assess.

Eyes separated; eye spots absent. Palps in dorsal view projecting 1/3 of prostomial length (Fig. 16A), fused. Extension of nuchal epaulettes to end of tentacular segment.

Median antenna reaching chaetiger 5–6. Lateral antennae and dorsal tentacular cirri, length 2/3 of median antenna. Ventral tentacular cirri 2/3 as long as dorsal pair. First dorsal cirri as long as median antenna, second dorsal cirri 2/3 as long as ventral tentacular cirri. Alternation in direction of cirri not assessed. Cirri from chaetiger 3, of equal length, c. 1/5 of body width. Cirrophores on tentacular segment and first dorsal cirri present; cirrophores otherwise absent. Anterior appendages cylindrical, dorsal cirri from chaetiger 3 egg shaped (Fig. 16A).

Parapodial lobes rounded, small. All chaetigers with 1–2 aciculae. Chaetal fascicle with 8–9 compounds. Compound chaetae with small distal tooth (Fig. 16B, C); serration present. Single thick bayonet chaetae (Fig. 16B), beginning at chaetiger 1.

Pharynx with 1 sinuation anterior to proventricle. Trepan in chaetiger 1 with 18 unequal teeth, 9 large and 9 smaller; 1 large alternating with 1 small, arranged in 2 rings. Basal ring present; infradental spines absent. Proventricle equal in length to 2.5 segments in chaetiger 3–5, with c. 30 rows of muscle cells. Pygidium lost.

Reproduction. Schizogamy by anterior scissiparity behind chaetiger 13 (Hartmann­ Schröder 1962).

Habitat. Amongst algae, barnacles, Mytilus .

Distribution. South East Pacific. From central Chile to Magellan Strait.

Remarks. Only the holotype could be located in the Museum of Hamburg, so the material of this species is restricted to one anterior fragment. The egg shaped dorsal cirri could be a distinguishing character but in the description the paratype is drawn with cylindrical dorsal cirri. In other respects the taxon is very similar to other Proceraea with short nuchal epaulettes and similar compounds, foremost P. cornuta , P. okadai and P. nigropunctata . At present it can not be determined if any of these taxa are the same as P. micropedata . Proceraea micropedata was described as having ventral cirri and, perhaps for this reason, referred to Odontosyllis , but these structures ( Hartmann­Schröder 1962: fig. 88) could not be detected.

Proceraea misakiensis ( Imajima, 1966) comb. n. (Fig. 17A–C) Autolytus (Regulatus) misakiensis Imajima, 1966: 61 –63, fig. 18A–H.

Material examined. Japan: 1 paratype NSMT­Pol P­ 16, Misaki, Onagawa, intertidal, Apr 1964.

Diagnosis. Proceraea without colour markings, and with large distal tooth in the compound chaetae of median, and posterior chaetigers.

Description. Paratype complete, length 9.2 mm for 94 chaetigers, width 0.4 mm. Live specimens orange ( Imajima 1966). Preserved material whitish without colour markings; eyes brownish red. Ciliation not possible to assess.

Eyes confluent; eye spots absent. Palps in dorsal view projecting 1/4–1/3 of prostomial length, fused. Nuchal epaulettes extending over anterior part of chaetiger 1.

Antennae and anterior appendages curled, not possible to assess (Fig. 17A). Alternation in direction of cirri not assessed. Dorsal cirri from chaetiger 3, of equal length, 1/3–1/4 of body width (Fig. 17A). Cirrophores on tentacular segment and first dorsal cirri present; cirrophores otherwise absent. All appendages cylindrical (Fig. 17A).

Parapodial lobes rounded conical, small. Anterior chaetigers with 2 aciculae, 1 in median and posterior. Chaetal fascicle with 10 compounds in anterior chaetigers, 6–10 in median and posterior. Compound chaetae with small distal tooth in anterior 5 chaetigers, more posterior with large distal tooth (Fig. 17B); serration present. Single thick bayonet chaetae (Fig. 17C), beginning at chaetiger 7.

Pharynx dissected. Proventricle equal in length to 3 segments (Fig. 17A) in chaetiger 7–9 with c. 40 rows of muscle cells. Anal cirri equal in length to dorsal cirri.

Reproduction. Unknown.

Additional information. Imajima (1966) described the trepan as having 18 unequal teeth, 9 large and 9 smaller; 1 large alternating with 1 small, arranged in 2 rings.

Habitat. Intertidal.

Distribution. North West Pacific. Central Japan, Misaki, Onagawa.

Remarks. Imajima described the colour as orange in P. misakiensis , hence lacking colour markings. Similar species without distinct colour markings, with nuchal epaulettes reaching over anterior part of chaetiger 1, and similar chaetae with large distal tooth, includes P. gigantea , P. aurantiaca and P. paraurantiaca . Proceraea gigantea has large trepan teeth but the size of these is not known for P. misakiensis . Proceraea paraurantiaca has a distinctive ciliated ridge on their palps, not found in P. misakiensis . Proceraea aurantiaca may also be a close relative, and with present material it is not possible to separate P. misakiensis from either of P. aurantiaca or P. gigantea .

Proceraea monoceros ( Ehlers, 1907) comb. n. (Fig. 18A–B) Pterautolytus monoceros Ehlers, 1907: 8 –10, figs 1–3.

Autolytus monoceros Augener 1924a: 60 –63; 1924b: 396–399; Benham 1927: 61.? Autolytus monoceros Fauvel 1934: 313 –314. (= Proceraea vulgaria according to Imajima 1966).

Material examined. New Zealand: holotype ZMB 6747, Port Chalmers.

Diagnosis. Proceraea with nuchal epaulettes reaching end of chaetiger 1 and with a conical tubercle on dorsum of second segment.

Description. Holotype almost complete, lacking posterior­most end; length c. 20 mm for 87 chaetigers, width 0.8 mm. Preserved material brown; eyes reddish brown. Ciliation not possible to assess.

Eyes separated; eye spots absent. Palps in dorsal view projecting 1/3 of prostomial length, fused. Extension of nuchal epaulettes to end of chaetiger 1 (Fig. 18A), distinctly curved, with a conical tubercle on dorsum of second segment.

Median antenna curled, reaching chaetiger 6–7. Lateral antennae and first dorsal cirri lost. Dorsal tentacular cirri equal in length to median antenna. Ventral tentacular cirri, length 1/4 as long as dorsal pair. Second dorsal cirri 3/ 4 in length of dorsal tentacular cirri. Alternation in direction of cirri not assessed. Dorsal cirri from chaetiger 3, of equal length, 1/4–1/3 of body width (Fig. 18A). Cirrophores on tentacular segment and first dorsal cirri present; cirrophores otherwise absent. All appendages cylindrical (Fig. 18A) including lost antennae and first dorsal cirri ( Ehlers 1907).

Parapodial lobes rounded, small to medium in size. Aciculae numbering 2 in chaetiger 46. Most chaetae lost, c. 10 compounds in anterior chaetigers, 5–10 in median and posterior. Compound chaetae with small distal tooth; serration present. Single thick bayonet chaeta, beginning at c. chaetiger 50.

Pharynx with sinuation anterior to proventricle. Trepan in chaetiger 2, with 18 unequal teeth, 9 large and 9 smaller; 1 large alternating with 1 small (Fig. 18B), arranged in 2 rings; trepan teeth large. Basal ring present; infradental spines absent. Proventricle equal in length to 4 segments in chaetiger 8–11 with 45–55 rows of muscle cells. Pygidium lost.

Reproduction. Unknown.

Habitat. Unknown.

Distribution. South West Pacific. New Zealand.

Remarks. Pterautolytus was erected for a single specimen with unpaired conical tubercle between the nuchal epaulettes and with ventral cirri. However, the structure that Ehlers refers to as ventral cirri is part of the parapodial lobes, and later authors have not been able to find the conical tubercle in newly collected specimens. As a consequence Pterautolytus has been synonomized with Autolytus ( Augener 1924a) . The trepan structure has notoriously been difficult to assess. Ehlers and Augener described the trepan to have c. 8 teeth. I found the holotype to have a typical Proceraea ­trepan as described above. The bayonet chaeta is also of the thick type normally found in Proceraea . Proceraea monoceros shares the long nuchal epaulettes with P. p i c t a, P. scapularis , P. fasciata and P. longilappeta . Proceraea monoceros has a smaller distal tooth in compound chaetae than does P. picta , and P. scapularis ; P. monoceros has a shorter proventricle than P. longilappeta , equal in length to 4 segments with 45–55 rows of muscle cells compared to 5–6 segments with 65–71 rows. As only one specimen of each has been compared, P. monoceros and P. longilappeta needs to be reassessed.

Proceraea mukaishima ( Imajima, 1966) comb. n. (Fig. 19A–C) Autolytus (Regulatus) mukaishimus Imajima, 1966: 73 –75, fig. 23A–E.

Material examined. Japan: 2 paratypes NSMT P­ 54, Mukaishima , intertidal, among seaweed, 6 Jan 1964.

Diagnosis. Proceraea with unique colour pattern consisting of broad brown bands on every segment, medially narrowed; trepan with 34–44 teeth, 1 large alternating with 2–4 smaller, in 2 rings.

Description. 2 complete paratypes; length 5.8–8.5 mm for 39–45 chaetigers, width 0.45 mm. Preserved material yellowish with colour markings consisting of a brown band covering chaetiger 1, following chaetigers with faint brown bands, medially narrowed (Fig. 19A); eyes reddish brown. Ciliation not possible to assess.

Eyes separated; eye spots absent. Palps in dorsal view projecting 1/4 of prostomial length (Fig. 19A), fused. Nuchal epaulettes extending over tentacular segment (Fig. 19A).

Median antenna reaching chaetiger 7–8. Lateral antennae and dorsal tentacular cirri, length 2/3 of median antenna. Ventral pair of tentacular cirri 1/3–1/2 as long as dorsal pair. First dorsal cirri as long as median antenna, second dorsal cirri as long as ventral tentacular cirri. Alternation in direction of cirri not assessed. Dorsal cirri from chaetiger 3, of equal length, 1/5 of body width (Fig. 19A). Cirrophores on tentacular cirri and first dorsal cirri present; cirrophores otherwise absent. All appendages cylindrical.

Parapodial lobes rounded, small. Anterior chaetigers with 2 aciculae, 1 in median and posterior. Chaetal fascicle with 6–8 compounds in anterior chaetigers, 4–5 in median and posterior. Compound chaetae with small distal tooth (Fig. 19B); serration present. Single thick bayonet chaetae (Fig. 19C), beginning at chaetiger 1.

Pharynx with 1 sinuation anterior and lateral to anterior half of proventricle. Trepan in chaetiger 2, with 34–44 unequal teeth, 9 large and 25–35 smaller; 1 large alternating with 2–4 smaller, in 2 rings. Basal ring present; infradental spines absent. Proventricle equal in length to 2–2.5 segments in chaetiger 4–6, with 35 rows of muscle cells (n=2). Anal cirri half as long as dorsal cirri.

Reproduction. Schizogamy by anterior scissiparity behind chaetiger 13. One of the paratypes with developing head, with small rudimentary antennae behind chaetiger 13 (Fig. 19A).

Habitat. Amongst algae, intertidal.

Distribution. North West Pacific. Northern Japan, only known from the type locality. Remarks. Proceraea mukaishima is easily identified from its colour pattern and trepan structure, both unique in Proceraea . The reproductive mode has not been reported earlier.

Proceraea nigropunctata Nygren & Gidholm, 2001 (Fig. 20A–H)

Proceraea nigropunctata Nygren & Gidholm, 2001: 181 –184, figs 1A–E, 2A–C, 3A–C, 4; Nygren & Sundberg 2003: GenBank sequences, 16S rDNA partial sequence AF474274 , and 18S rDNA partial sequence AF474320 .

Material examined. USA: holotype (LACM­POLY 1962), 3 paratypes (LACM­AHF POLY 1963) and additional 15 spms, Santa Catalina Island, Wrighley Marine Science center, the vicinity, 33°24’N, 118°30’W, ca 10 m, dredge, red algae mainly Gelidium purpurescens , 15 Apr 1971; 2 spms, Santa Catalina Island, Wrighley Marine Science center, the vicinity, 33°24’N, 118°30’W, ca 10 m, dredge, Haliotis with algae, sponges, vermetids, 15 Apr 1971; 2 spms, Santa Catalina Island, Wrighley Marine Science center, the harbour, 33°27’N, 118°29’W, 0.5 m, eel­grass, stones and green algae, 14 Apr 1971; 10 spms (3 spms mounted for SEM), Friday Harbor, Peavine pass between Orcas and Blakely Islands, 48°35.5’N, 122°48.0’W, 18 m, dredge, dead Balanus nobilis , hydroids, shells, 6 May 1971; 10 spms, Friday Harbor, Dot rock, SE side of Decatur Island, 48°29.5’N, 122°37.5’W, 18 m, dredge, gravel and shells, 7 May 1971; 40 spms (several rear ends in author's collection for DNA analyses), Santa Catalina Island, Wrighley Marine Science center, floating dock outside laboratory, 33°26.7’N, 118°29.05’W, 0.5 m, algae, hydroids, Jan 2001; 10 spms (5 spms in formalin, 5 spms on slides (rear ends in author's collection for DNA analyses), Washington, San Juan Island, 48°32.7' N 123°00.8'W N, epifauna on floating dock outside Friday Harbor laboratory, sponges, hydroids, barnacles, 22 Jan 2001.

Diagnosis. Proceraea with seven dorsal dark brown spots arranged in 2 transverse rows across each segment.

Description. Length in preserved specimens 2.9–12.0 mm for 30–64 chaetigers; width c. 0.3 mm. Live specimens with dark brown colour pattern (Fig. 20A–C); 7 dorsal dark brown spots arranged in 2 transverse rows across each segment, 2 weaker spots in parapodial lobes, and a midventral longitudinal band (Fig. 20B); anterior appendages yellowish; intestine yellowish, intestinal granular accumulations forming light reflecting narrow middorsal band interrupted in segment borders (Fig. 20C); eyes brown. Brown colour pattern often retained in preserved specimens. Ciliation present on prostomium, nuchal epaulettes, and a few segments ventrally.

Eyes confluent or almost confluent (Fig. 20A); eye spots present. Palps in dorsal view projecting 1/4–1/3 of prostomial length (Fig. 20A), fused. Extension of nuchal epaulettes to end of tentacular segment (Fig. 20A).

Median antenna reaching chaetiger 7–9 in preserved specimens (n=15). Lateral antennae and dorsal tentacular cirri, length 1/2–2/3 of median antenna. Ventral tentacular cirri 1/ 3–1/2 as long as dorsal pair. First dorsal cirri as long as median antenna, second dorsal cirri as long as ventral tentacular cirri. Alternation in direction of cirri not assessed. Dorsal cirri from chaetiger 3, of equal length, c. 1/2 of body width. Cirrophores on tentacular segment and first dorsal cirri present; cirrophores otherwise absent. All appendages cylindrical.

Parapodial lobes rounded conical, small. Single acicula in all chaetigers. Chaetal fascicle with 6–8 compounds in anterior chaetigers, 3–5 in median and posterior. Compound chaetae with small distal tooth (Fig. 20D, E); serration present. Single thick bayonet chaetae (Fig. 20F), beginning between chaetiger 2–9.

Pharynx with 1 sinuation anterior to proventricle. Trepan in chaetiger 1–2, with 18 unequal teeth, 9 large and 9 smaller; 1 large alternating with 1 small, arranged in 2 rings. Basal ring present; infradental spines absent. Proventricle equal in length to 1.5 segments in chaetiger 5–7 with 34–36 rows of muscle cells (n=4). Anal cirri equal in length to 1/2 body width.

Reproduction. Schizogamy by anterior scissiparity behind chaetiger 13. Specimens with stolons found at all collecting times, i.e. January, April, and May.

Morphology of epitokous stages.

Male. Length in preserved specimens 4–5.1 mm for 6+(20–21)+(4–10) chaetigers (n=5), width in region a 0.25–0.3 mm, width in region b 0.7–0.8 mm. Same colour pattern as in stock (Fig. 20H), but black midventral band sometimes absent. Ciliation absent except for nuchal epaulettes.

Prostomium with concave anterior margin. Nuchal epaulettes rounded, prominent, reaching beginning of chaetiger 1.

Median antenna reaching posterior part of region b (c. chaetiger 20). Lateral bifid antennae, 2–3 times as long as prostomial width; basal part 1/4 of total length, outer ventral rami slightly shorter than inner dorsal rami. Frontal processes, equal to 1/3 of prostomial width. Tentacular cirri 2 pairs, dorsal pair as long as 1.5 times prostomial width, ventral pair 1/ 4 in length of dorsal. First dorsal cirri, equal in length to median antenna; achaetous knobs present. Cirri in region a equal to 2/3 of body width, cirri in region b and c successively shorter, in region b equal to 1/3 of body width, in region c 1/4 of body width. Ceratophore on median antenna, small cirrophores on tentacular cirri, and large cirrophores on first dorsal cirri, present; cirrophores otherwise absent. Frontal processes club shaped, cirri on chaetiger 3–6 fusiform, other appendages cylindrical.

Single neuropodial acicula in all chaetigers; 2 anterodorsal, and 4 thick and 3 thin posteroventral notopodial aciculae in region b. Neuropodial chaetal fascicle with 4–10 compounds, single bayonet chaetae beginning at chaetiger 1. Notopodial chaetal fascicle with 20–25 swimming chaetae. Anal cirri equal in length to cirri in region c.

Female. Length in preserved specimens 4–5.6 mm for 6+(13–15)+(15–18) chaetigers (n=5), width in region a 0.2–0.3 mm, width in region b 0.4–0.7 mm. Same colour pattern as in stock (Fig. 20G). Ciliation absent except for nuchal epaulettes.

Prostomium with straight anterior margin. Nuchal epaulettes rounded, prominent, reaching beginning of chaetiger 1.

Median antenna reaching chaetiger 2. Lateral antennae, slightly shorter than median antenna. Tentacular cirri 2 pairs, dorsal pair 1/2 as long as prostomial width, ventral pair 1/ 3–1/2 as long as dorsal pair. First dorsal cirri, about as long as lateral antennae; achaetous knobs present. Cirri in region a equal, as long as or slightly longer than body width; cirri in region b slightly longer than in a, equal to 1/2–2/3 of body width; cirri in region c successively shorter, c. 1/4 of body width. Cirrophores absent or very small on tentacular segment, cirrophores on first dorsal cirri, and on cirri in region a present; cirrophores otherwise absent. All appendages cylindrical.

Single neuropodial acicula in all chaetigers; 2 anterodorsal, and 2 thick and 3 thin posteroventral notopodial aciculae in region b. Neuropodial chaetal fascicle with 4–8 compounds; single bayonet chaetae beginning at chaetiger 1. Notopodial chaetal fascicle with 15–20 swimming chaetae. Anal cirri equal in length to posterior cirri in region c.

Habitat. Among algae, sponges, hydroids on stones and shells. Shallow waters to about 18 m depths.

Distribution. North East Pacific. California to Washington.

Remarks. Proceraea nigropunctata is easily identified on the characteristic colour pattern. However, these colours are not always retained in formalin­fixed material, and thus it could be impossible to separate P. nigropunctata from similar taxa in preserved condition. Similar species with nuchal epaulettes reaching over only tentacular segment, and the same type of compound chaetae, includes P. okadai , P. c o r n u t a, and P. micropedata . The two former are distinct from P. nigropunctata in their coloration; the colour pattern of P. micropedata is not known, but the anterior fragment that exist of this species have egg shaped dorsal cirri not found in either of the three other species.