Proceraea fasciata ( Bosc, 1802) (Fig. 10 A –F)

Nereis fasciata Bosc, 1802: 144, pl. 5, fig. 6.

Autolytus (?), banded species, Verrill 1873: 398.

Proceraea ornata Verrill, 1873: 746; Mensch 1900 a; 1900 c.

Proceraea tardigrada Webster, 1879: 227 –230; Andrews 1891: 282 –283; 1892: 188, pl. 11, fig. 41–43, pl. 12, fig. 57 –58, 62–65, 69– 70.

Proceraea? coerulea Webster, 1879: 230.

Autolytus longisetosus Hartman 1944: 338, pl. 13, fig. 1.

Autolytus alexandri Hartman 1944: 338, pl. 13, fig. 2.

Autolytus ornatus Hartman 1945: 16; Allen 1957: 49 –50

Autolytus fasciatus Pettibone 1963: 141 –143, figs 37 A –B, 38–39; Allen 1964.

Proceraea fasciata Gardiner 1976: 127 –129, fig. 10 E –H.

Material examined. USA: holotype of Proceraea ornata Verrill, 1873, YPM 24226, Connecticut, New Haven Co. Branford, Thimble Islands, shallow depths, among hydroids and bryozoans, 2 Oct 1873; 7 spms USNM 52909 ( 15), North Carolina, Bogue Sound, Morehead City, shallow depths, shell gravel, 6 Apr 1974.

Diagnosis. Proceraea with broad red­brown bands on nearly every segment, or on every second to fourth segment.

Description. Length 12–15 mm for 83–90 chaetigers; width 0.5 mm. Live specimens white with transverse intersegmental red­brown bands on dorsal side on nearly every segment, or on every second to forth segment (Fig. 10 A); nuchal epualettes brownish; parapodial bases with single red spot; eyes red. Segmental bands sometimes preserve. Ciliation present on prostomium, nuchal epaulettes, and a few segments ventrally.

Eyes confluent (Fig. 10 A); eye spots present. Palps in dorsal view projecting 1 / 4 of prostomial length (Fig. 10 A), fused. Extension of nuchal epaulettes to end of chaetiger 1 (Fig. 10 A).

Median antenna reaching chaetiger 8–12 (n= 7) in preserved specimens. Lateral antennae and dorsal tentacular cirri, length 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, 1 / 4 – 1 / 3 of body width (Fig. 10 A). Cirrophores on tentacular segment and first dorsal cirri present; cirrophores otherwise absent. All appendages cylindrical.

Parapodial lobes rounded conical, small (Fig. 10 B). Anterior chaetigers with 2 aciculae, 1 in median and posterior. Chaetal fascicle with 10–15 compounds in anterior chaetigers, 3–10 in median and posterior. Compound chaetae with small distal tooth in anterior 10–15 chaetigers (Fig. 10 C, D), more posterior with large distal tooth (Fig. 10 E); serration present. Single thick bayonet chaetae (Fig. 10 F), beginning between chaetiger 14–15.

Pharynx with sinuation anterior to proventricle (Fig. 10 A). Trepan in chaetiger 2 (Fig. 10 A), 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 –2,5 segments in chaetiger 7–9 (Fig. 10 A) with 35–38 rows of muscle cells (n= 5). Anal cirri equal in length to 1 / 3 of body width.

Reproduction and morphology of reproductive stages. Schizogamy by anterior scissiparity. 3 specimens with developing stolons behind chaetiger 13. Pettibone ( 1963) gives descriptions on stolon morphology. Stolons with same colour pattern as in stock. As judged from figures ( Pettibone 1963: figs 37 A, 39 A, B) all appendages cylindrical, both in male and female. Male stolons with 6 +( 21–29)+( 13–53) chaetigers. Nuchal epaulettes triangular, reaching half of first segment ( Pettibone 1963: fig. 37 A). Tentacular cirri 2 pairs, first dorsal cirri situated above achaetous knobs ( Pettibone 1963: fig. 37 B). Female stolons with 6 +( 19–24)+( 38–48) chaetigers. Nuchal epaulettes reaching half or end of chaetiger 1 ( Pettibone 1963: fig. 39 B). Tentacular cirri 2 pairs, first dorsal cirri situated above achaetous knobs ( Pettibone 1963: fig. 39 C). Eggs carried in a 3 ­lobed egg sac; eggs whitish, blue or purplish. Breeding occurs almost all year round ( Allen 1957).

Habitat. Intertidal to 40 m, among algae, hydrozoans and bryozoans.

Distribution. North West Atlantic. Massachusetts to South Carolina, Puerto Rico.

Remarks. Proceraea fasciata is unique among Proceraea in its colour pattern. The colour pattern is not always retained in preserved specimens, and can thus be confused with preserved specimens of P. p i c t a, P. scapularis ( Claparède, 1864), and P. pleijeli sp. n., which all have about the same length in their nuchal epaulettes and about the same type of compound chaetae. The proventricle is generally longer in these three species measuring 3–4 segments with> 43 rows of muscle cells compared to 2–2.5 segments and 33–38 rows. The synonymy of Proceraea tardigrada, Proceraea? coerulea, and Proceraea ornata is concluded from their descriptions.

Proceraea filiformis Hartmann­Schröder, 1989 (Fig. 11 A –F)

Proceraea filiformis Hartmann­Schröder, 1989: 33, figs 47–51; 1991: 43.

Material examined. Australia: holotype ZMH P­ 19669, Lake Macquarie, south of sea entrance, algae, 15 Jan 1976.

Diagnosis. Proceraea with large palps, large confluent eyes, and with an 8 ­sinuated pharynx.

Description. Length 2.2 mm for 18 chaetigers, incomplete, width 0.2 mm. Preserved material whitish, no colour markings; eyes reddish orange. Ciliation not possible to assess.

Eyes large, confluent; eye spots absent. Palps in dorsal view projecting 1 / 2 of prostomial length, fused (Fig. 11 B). Extension of nuchal epaulettes to end of tentacular segment (Fig. 11 A, B).

Median antenna reaching chaetiger 6. Lateral antennae and dorsal tentacular cirri, length 2 / 3 of median antenna. Ventral tentacular cirri 1 / 3 as long as dorsal pair. First dorsal cirri lost, second dorsal cirri as long as ventral tentacular cirri. Dorsal cirri from chaetiger 3, of equal length, 1 / 3 of body width (Fig. 11 A). Cirrophores on tentacular segment and first dorsal cirri present; cirrophores otherwise absent. All appendages cylindrical, including lost first dorsal cirri ( Hartmann­Schröder 1989).

Parapodial lobes rounded conical, small (Fig. 11 A). Anterior chaetigers with 2 aciculae, 1 in posterior. Chaetal fascicle with 5–6 compounds. Compound chaetae with small distal tooth in anterior 3 chaetigers (Fig. 11 E), more posterior with large distal tooth (Fig. 11 F); serration present. Single thick bayonet chaetae (Fig. 11 F), beginning at chaetiger 15.

Pharynx with twisted sinuation anterior to proventricle (Fig. 11 A). Trepan in chaetiger 1, with 18 unequal teeth; 9 large and 9 smaller; 1 large alternating with 1 small (Fig. 11 C, D), arranged in 2 rings. Basal ring present; infradental spines absent. Proventricle equal in length to 2.5 segments in chaetiger 8–10 with 35 rows of muscle cells. Pygidium lost.

Reproduction. Unknown

Habitat. Algae.

Distribution. South West Pacific. East coast of Australia.

Remarks. Proceraea filiformis is most similar to P. aurantiaca which also may have a twisted pharynx. However, the nuchal epaulettes are shorter in P. filiformis than in P. aurantiaca. See also remarks for P. misakiensis.

Proceraea gigantea Nygren & Gidholm, 2001 (Fig. 12 A –F) Proceraea gigantea Nygren & Gidholm, 2001: 187 –190, figs 8–10.

Material examined. USA: holotype (LACM­POLY 1966), 1 paratype (LACM­AHF POLY 1967) and additional 6 spms ( 2 mounted for SEM), Santa Catalina Island, Wrighley Marine Science center, the vicinity, 33 ° 24 ’N, 118 ° 30 ’W, dredge, 10 m, Haliotis with algae, sponges, vermetids, 15 Apr 1971; 1 spm, Santa Catalina Island, Wrighley Marine Science center, the harbor, 33 ° 27 ’N, 118 ° 29 ’W, 0.5 m, eel­grass, stones and green algae, 14 Apr 1971; 5 spms (rear ends in author's collection for DNA analyses), California, Santa Catalina Island, 33 ° 26.70 'N 118 ° 29.05 'W, 1 m, floating dock, outside Wrighley marine science center, amongst hydroids, sponges, bryozoans, 14–17 Jan 2001.

Diagnosis. Large Proceraea, up to 50 mm, with large teeth in trepan, orange in colour with single red spot in parapodial lobes.

Description. Length up to 48 mm for 118 chaetigers in live specimens. Length in preserved specimens 4.8–36.8 mm for 43–118 chaetigers, width 0.15–0.42 mm. Live specimens without colour markings, yellowish to orange (Fig. 12 A), sometimes with scattered reddish­brown pigments anteriorly; parapodial bases with single red spot (Fig. 12 B); intestine yellowish­brown without light reflecting granular accumulations; proventricle pinkish­orange; eyes red. Preserved specimens without colours. Ciliation present on prostomium, nuchal epaulettes, and a few segments ventrally.

Eyes confluent or almost confluent; eye spots present. Palps in dorsal view projecting c. 1 / 2 of prostomial length (Fig. 12 A), fused. Nuchal epaulettes extending over anterior part of chaetiger 1 (Fig. 12 A).

Median antenna reaching chaetiger 15–20 in live specimens (n= 6). Lateral antennae and dorsal tentacular cirri, length 1 / 3 – 2 / 3 of median antenna. Ventral tentacular cirri 1 / 3 – 1 / 2 as long as dorsal pair. First dorsal cirri 2 / 3 of median antenna, second dorsal cirri 1–1.5 times ventral tentacular cirri. Alternation in direction of cirri not assessed. Dorsal cirri from chaetiger 3, of equal length, 1 / 3 – 1 / 2 of body width (Fig. 12 A). Cirrophores on tentacular segment and first dorsal cirri present; cirrophores otherwise absent. All appendages cylindrical.

Parapodial lobes rounded conical, small. Anterior chaetigers with 3–4 aciculae, 1–2 in median and posterior. Chaetal fascicle with 8–16 compounds in anterior chaetigers, 4–7 in median and posterior. Compound chaetae with small distal tooth (Fig. 12 D) in anterior 5– 10 chaetigers, more posterior with large distal tooth (Fig. 12 E); serration present. Single thick bayonet chaetae beginning between chaetiger 5–20 (Fig. 12 F).

Pharynx with 1 sinuation anterior and lateral to anterior half of proventricle (Fig.

12 A). Trepan in chaetiger 2–3 (Fig. 12 A), with 18 unequal teeth (Fig. 12 C), 9 large and 9 smaller; 1 large alternating with 1 small in 2 rings. Basal ring present; infradental spines absent. Proventricle equal in length to 3–5 segments in chaetiger 6–10 (Fig. 12 A) with 50– 60 rows of muscle cells (n= 6). Anal cirri equal in length to 1 / 2 – 2 / 3 of body width.

Reproduction. Schizogamy by anterior scissiparity. A few specimens are in different stages of developing heads behind chaetiger 13 (collected in January). Specimens with developed swimming chaetae not observed.

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

Distribution. North East Pacific. California. Only known from type locality.

Remarks. Proceraea gigantea was described as the largest known Proceraea, and with extraordinary large teeth in its trepan. Size is not a very good character for separating taxa as the size­range for other Proceraea is not fully understood. Large trepan teeth is also found in P. monoceros Ehlers, 1907, and P. longilappeta Imajima, 1966; these taxa have longer nuchal epaulettes reaching end of chaetiger 1 and beginning of chaetiger 2 respectively; further these taxa have small distal tooth in all chaetigers. As size of teeth in the trepan of P. misakiensis is not known, synonomy with this taxon cannot be excluded.