Cigclisula occlusa ( Busk, 1884 )

Almeida, Ana C. S., Souza, Facelucia B. C., Menegola, Carla M. S., Sanner, Joann & Vieira, Leandro M., 2014, Taxonomic review of the family Colatooeciidae Winston, 2005 (Bryozoa, Cheilostomata), with description of seven new species, Zootaxa 3868 (1), pp. 1-61 : 9-13

Cigclisula occlusa ( Busk, 1884)

( Figs 3–5 , Table 1 )

Escharoides occlusa Busk, 1884: 150 (in part), pl. 21, fig. 8 (part). [Zamboanga, Philippines; not specimen from Cape York, Australia]

Lepralia occlusa: Waters 1888: 26 , pl. 3, figs 32–34. [Zamboanga, Philippines]

Lepralia occlusa: Waters 1909: 152 (in part), pl. 13, fig. 15; pl. 14, figs 1–9, 13 (?in part). [Zamboanga, Philippines]

? Escharoides occlusa: Waters 1913: 519 . [ Zanzibar, Indian Ocean]

? Escharoides occlusa: Robertson 1921: 56 . [Straits of Malacca, Indian Ocean]

? Myriozoum occlusum: Marcus 1923: 435 , pl. 25, figs 8 a, b. [Aru Islands, Indonesia]

Cigclisula occlusa: Canu & Bassler 1929: 291 , pl. 31, figs 3–10. [ Philippines]

Cigclisula occlusa: Harmer 1957: 1057 , pl. 69, figs 16–18. [ Philippines]

Material examined. Lectotype: NHMUK 1944.1 .8.284 [dry specimens; figured by Busk 1884, pl. 21, fig. 8 (part)], Escharoides occlusa, G. Busk coll., Challenger Expedition , Samboanga, Philippines, 10 ftm (18.29 m) . Paralectotypes: NHMUK 1887.12 .9.588 [two dried specimens; figured by Busk , pl. 21, fig. 8] ; NHMUK 1887.12 .9.594 [large dry colony] ; NHMUK 1899.7 .1.2138 [balsam slide] ; NHMUK 1899.7 .1.2138 [balsam slide] ; NHMUK 1899.7 .1.2141 [balsam slide] ; NHMUK 1887.12 .9.589 [balsam slide], Escharoides occlusa, G. Busk coll., Challenger Expedition , Samboanga, Philippines, 10 fths (18.29 m) . NHMUK 1887.12 .9.590 [dry slide] ; NHMUK 1887.12 .9.592 [dry] ; NHMUK 1887.12 .9.592 part [dry slide]; Escharoides occlusa, G. Busk coll., Challenger Expedition, Stn 148, off Possession Island , 24°47’ S, 51°37’ E, South Indian Ocean, 210 ftm (384 m) . Additional specimens: USNM 8039 , Cigclisula occlusa (Busk) , Canu & Bassler 1929, Albatross Stn D 5158, Tinakta Island , Tawi Group, Philippines, 22 m. USNM 8037 , Cigclisula occlusa (Busk) , Albatross Stn D 5137, Jolo Light, Jolo Island , Philippines, 36.6 m .

Description. Colony erect, bilaminar, branching. Zooids hexagonal, longer than wide, limited by slightly raised lateral walls. Frontal shield heavily calcified, smooth-textured, uniformly punctured by 24–40 pseudopores. Primary orifice small relative to frontal shield, somewhat hoof-shaped, as long as wide, sunken, with arcuate anter and concave poster delimited by 2 down-curved condyles at about one-third orifice length. Secondary orifice raised, non-tubercular, almost circular. Suboral avicularium elliptical, laterally placed and distally directed, of 2 sizes, 1 small (0.055 –0.121 mm long), the other double-sized, commonly on zooids in the center of branches. Frontal avicularia small, rare, elliptical, 1–2 per zooid, placed at zooidal margins, sometimes around ooecial margin. Interzooidal avicularia longer than wide, with broadly spatulate mandible; rostrum with rounded proximal edge, median constriction distinct and distal edge concave; calcified palate occupying more than half rostral length, with acutely triangular foramen. Ooecium subglobose, not very prominent, becoming immersed in secondary cacification; ectooecium with longitudinal band of 11–29 irregular pseudopores.

Remarks. Waters (1913) suggested that a new genus was needed to accommodate Escharoides occlusa Busk,

1884 owing the morphology of ooecia and avicularia, but did not name one. Subsequently, Canu & Bassler (1927) designated it as the type species of Cigclisula , characterized by the non-cleithral ooecium with a grating-like surface (longitudinal band of pseudopores). Whereas the grating-like surface of Colatooecia is an entooecium covered by a membranous ectooecial area, in Cigclisula it is a pseudoporous ectooecium.

Busk (1884) described Escharoides occlusa using specimens from Zamboanga ( Philippines), Possession Island (South Indian Ocean) and Cape York ( Australia). Examination of these specimens has revealed differences among colonies from these localities, leading to the recognition of two new species from Australia. Specimens from the Philippines have a smooth frontal shield and ooecia whereas specimens from Australia have a minutely tubercular frontal shield and ooecia ( Cigclisula australis n. sp.), with small protuberances in Cigclisula buski n. sp. Cigclisula occlusa , C. australis n. sp. and C. buski n. sp. have two sizes of interzooidal avicularia that differ in the shape of the opesia—acutely triangular in C. occlusa ; both triangular (larger interzooidal avicularia) and oval (smaller interzooidal avicularia) in C. australis n. sp.; oval in C. buski n. sp. The ectooecium in C. occlusa has 11–29 pseudopores; that in C. australis n. sp. is less porous, with 7–18 pseudopores; in C. buski n. sp. there are 38–50 pseudopores. Additionally, zooids, frontal pseudopores and suboral avicularia are larger in C. occlusa than in C. australis n. sp. and C. buski n. sp. (see remarks under C. australis n. sp. and C. buski n. sp.). It is not possible to recognize diagnostic characters in the original Busk (1884) illustrations and the provenance of the illustrated specimens is unknown, therefore we cannot say if they truly belong to C. occlusa , C. australis n. sp. or C. buski n. sp.

Over time, many specimens assigned to C. occlusa were registered from different localities, including Zanzibar ( Waters 1909, 1913), the Malay Archipelago ( Robertson 1921) and the Indian Ocean and Red Sea ( Waters 1909; Dumont 1981). Specimens from Zanzibar described by Waters (1909, 1913) have two large spatulate avicularia on the margin of the colony and semicircular avicularia scattered over the zooids. Specimens from the Red Sea ( Waters 1909) have a semicircular avicularium placed at the proximal end of the zooid. Busk’s specimens of C. occlusa , C. australis n. sp. and C. buski n. sp. have single large interzooidal avicularia commonly placed at colony margins and elliptical avicularia placed at zooidal margins. Robertson (1921) did not describe specimens from the Malay Archipelago and referred to Waters’s (1909) description. Dumont (1981) also did not describe specimens from the Red Sea and, pending examination of these specimens, as well as specimens from the Indian Ocean, it is not possible to say if they belong to C. occlusa , C. australis n. sp., C. buski n. sp. or other species.

Cigclisula occlusa is characterized by two sizes of suboral avicularia, interzooidal avicularia with a spatulate rostrum and triangular opesia and a subglobose slightly prominent ooecium having an ectooecium with a longitudinal band of 11–29 pseudopores.

Cigclisula occlusa resembles Cigclisula cautium Hastings, 1932 , Cigclisula fissurata ( Ortmann, 1890) and Cigclisula fruticosa Hayward & Ryland, 1995 in having erect bilaminar colonies. Cigclisula occlusa differs from C. cautium in having a shorter zooid size, two sizes of suboral avicularium (one size only in C. cautium ) and more ectooecial pseudopores (11–29 in C. occlusa , 6–7 in C. cautium ). Differences from C. fissurata include the distribution of frontal pseudopores (uniform in C. occlusa , marginal in C. fissurata ), interzooidal and frontal avicularia (present in C. occlusa , absent in C. fissurata ) and number of ectooecial pseudopores (11–29 in C. occlusa , 2–5 in C. cautium ). Differences from C. fruticosa include the elliptical primary orifice and non-tubercular secondary orifice of C. occlusa and the number of ectooecial pseudopores, about 6 in C. fruticosa .

Distribution. Indo-Pacific: Philippines.

Figures

FIGURE 3. Cigclisula occlusa (Busk, 1884). A–F, NHMUK 1944.1.8.284, lectotype, Philippines. A, erect bilaminar colony. B, group of zooids at growing edge. C, close-up of zooid with suboral avicularium. D, close-up of spatulate interzooidal avicularium. E, ovicelled zooids. F, close-up of ovicelled zooid. Scale bars: A = 400 µm; B, E = 200 µm, C, D, F = 100 µm.

FIGURE 4. Cigclisula occlusa (Busk, 1884). A–D, NHMUK 1887.12.9.592 part, paralectotype, Philippines. A, group of zooids. B, close-up of zooids and suboral avicularium. C, primary orifice. D, spatulate interzooidal avicularia. Scale bars: A = 200 µm; B, D = 100 µm, C = 60 µm.

FIGURE 5. Cigclisula occlusa (Busk, 1884). A–D, USNM 8039, Philippines; E–F, USNM 8037, Philippines. A, entire erect colony. B, zooids at bifurcation of erect bilaminar colony. C, primary orifice. D, ovicelled zooids. E, zooids at growing edge of branch and spatulate interzooidal avicularia. F, zooids and small spatulate interzooidal avicularium. Scale bars: A = 1 cm; B = 500 µm; C = 100 µm; D, E, F = 200 µm.