Collarina fayalensis Harmelin, 1978a: 190 Unexpected diversity of the genus Collarina Jullien, 1886 (Bryozoa, Cheilostomatida) in the NE Atlantic-Mediterranean region: new species and reappraisal of C. balzaci (Audouin, 1826) and C. fayalensis Harmelin, 1978 Harmelin, Jean-Georges Bishop, John D. D. Madurell, Teresa Souto, Javier Spencer Jones, Mary E. Zabala, Mikel Zoosystema 2019 2019-09-26 41 21 385 418 X2VF Harmelin, 1978 Harmelin 1978 [256,651,1811,1838] Gymnolaemata Cribrilinidae Collarina Animalia Cheilostomatida 10 393 Bryozoa species fayalensis  ( Figs 5; 7B; 15B; 17D; 18F; Tables 1-3)      Collarina fayalensis Harmelin, 1978a: 190, fig. 9. — Arístegui 1984(in part): 230, pl. 11, fig. 7 (not fig. 6).   TYPEMATERIAL. —   Holotypeand paratypes. Azores, Faial Is., Castelo Branco, R/ V Jean-Charcot, BIAÇORES 1971 St. P.11, 38°31’12”N,  28°42’7”W,  5-7 m,  12.X.1971, H. Zibrowius leg., 12 small colonies on pebble together with colonies of  C. macaronensisHarmelin, n. sp.(recorded as  C. balzaci: Harmelin 1978a), MNHN-IB-2008-7909. OTHER MATERIAL EXAMINED. —   Azoresarchipelago. Formigas Is., R/V  Jean-Charcot, BIAÇORES 1971 St. P.43, 37°16’14”N, 24°46’54”W,  15 m,  7.XI.1971, H. Zibrowiusleg., 1 small colony on pebble, MNHN. —  São Miguel Is., Ilhéu de Vila Francado Campo, Azores Bio-Oceanographic Survey1979, ABOS St.AC- 79.6, 37°42’20”N, 25°26’33”W,  15 m,  24.VII.1979, coll. by JGH, c.30 colonies on 7 pebbles, 5 colonies on broken  Pinnashell, MNHN, NHMUK. —  São Miguel Is., Ponta da Galera, ABOS St.AC-79.7, 37°42’25”N, 25°30’32”W,  14 m, 24.VII.1079, coll. by JGH, 3 colonies on 2 small pebbles in submarine cave; 3 colonies on  Pinnashell, MNHN, NHMUK. —  São Miguel Is., Capelas, ABOS St.AC-79.9, 37°50’37”N, 35°41’19”W,  15 m,  25.VII.1979, coll. by JGH, 6 colonies on 1 pebble, MNHN. —  São Jorge Is., Ponta da Rosais, ABOS St.AC-79. 29, 38°45’06”N, 28°18’59”W, 20 -22 m,  6.VIII.1979, coll. by H. Zibrowius, 1 colony on pebble, MNHN. —  Graciosa Is., Ilhéu da Praia, ABOS St.AC-33, 39°03’36”N, 27°57’11”W,  20 m,  8.VIII.1979, coll. by JGH, 1 ovicelled colony on pebble from a tunnel, MNHN.  DIAGNOSIS. — Colonies small to large; autozooids longer than broad, with very extensive gymnocyst and minute costate shield, formed by 2-5 very short costae with smooth surface and a large marginal pelma; presence of large pseudopores on the gymnocyst; apertural bar poorly prominent and pierced by large pelmata; orifice longer than broad with proximal edge slightly concave, 2-5 thin oral spines; adventitious avicularia paired, directed disto-laterally; ooecium kenozooidal bearing a distal avicularium; ancestrula with 5 or 6 spines.  DESCRIPTION Colony encrusting, pluriserial, unilaminar, small- to large-sized (> 1000 autozooids). Autozooids longer than broad, roughly pentagonal with shorter distal side bearing the oral spines. Gymnocyst occupying the largest part of the frontal surface, moderately convex, punctured by several large gymnocystal pseudopores at the zooid margin or occasionally nearer to the costate shield ( Fig. 5B, D, E). Costate shield very small, with only 3-4 costae in most cases (range 2-5), with smooth surface and poorly demarcated basal, ascending part, the latter bearing a large pelma and generally with a second smaller pore near the centre of the shield; intercostal lacunae generally limited to a single, very small one per pair, centrally located and a much larger, semi-rectangular “basal intercostal space” with a tiny median slot ( Fig. 5G). Apertural bar low, with a slightly prominent centre, pierced by four pseudopores, two medium-sized on either side of the midline and two larger lateral pores, close to and below the base of the avicularia ( Figs 5B, C, F; 7B). Adventitious avicularia paired in most cases (> 95%), directed laterally with a slight distal angle ( Figs 5A, B, D, F; 14C), nested cystids occasionally present ( Fig. 17D). Orifice longer than broad ( Table 1), proximal edge slightly concave and often irregularly indented, with a small knob at each corner; condyles small ( Figs 5B, F; 7B). Oral spines with relatively narrow base, 2-5 innon-ovicelled zooids, 3 inmajority (75%), 2 relatively thin in ovicelled zooids ( Fig. 5C). Ovicell present in large colonies, less frequently in medium-sized ones, bearing a distal avicularium in most cases (> 80%); ooecium kenozooidal, formed at the colony edge, with surface pierced by about 12-15 medium- sized pseudopores, orifice about 40% larger on average than that of a non-ovicelled zooid, a distal avicularium in most cases ( Figs 5C; 15B). Ancestrula similar to older zooids but smaller, 3 very small costae, 6 spines, orifice longer than broad.  FIG. 5. —  Collarina fayalensisHarmelin, 1978: A -C, non-ovicelled and ovicelled autozooids showing the structure and limited extent of the costate shield, orifice dimorphism, small spines and relatively narrow avicularia; D, non-ovicelled zooid, for marginal pores; E, drawing from Harmelin (1978: fig. 9); F, distal part of a non-ovicelled zooid from a colony edge; G, proximal half of costate shield, note the distribution of pelmata and the shape of the basal intercostal spaces. Origin: Azores - Saô Miguel, Vila Franca Is., 15 m (A-C, F, G); Formigas Is., 15 m, ‘Jean Charcot’ Biaçores 1971, P.43 (D); Faial, ‘Jean Charcot’ Biaçores 1971, P.11 (E). Scale bars: A, 200 µm; B-D, 100 µm, F, 50 µm; G, 25 µm. DISTINCTIVE FEATURES   Collarina fayalensisis easily identified by the minute size of the costate shield, formed by 2-4 very short costae, which contrasts with the huge development of the gymnocyst. The greater number of costae (4-6) noted by Harmelin (1978a)included the pair forming the apertural bar. The latter is typically poorly prominent and pierced by large pelmata. The presence of large pseudopores on the gymnocyst is also a peculiar feature, shared with  C. macaronensisHarmelin, n. sp., in which these marginal pseudopores are less numerous. The Azorean material examined here, much more abundant than the one (small colonies on a single pebble) of the type-series ( Harmelin 1978a, herewith Fig. 5E), showed that this species can develop large colonies and provided information on the ovicell and the habitat. Ovicells were not frequent, present only in relatively large colonies. This feature may indicate that  C. fayalensisis not able to develop fertile colonies on ephemeral substrates, in contrast to  C. balzaci.  HABITAT AND GEOGRAPHICAL DISTRIBUTION Most sampled specimens, including the types, encrusted pebbles and cobbles at shallow depth ( Fig. 18F). On the other hand, few colonies were found on empty shells of  Pinna rudisLinnaeus, 1758from the same sites which, otherwise, were colonized by many other bryozoans. Therefore, pebbles can be considered the typical habitat of  C. fayalensis. These pebbles and cobbles were of volcanic origin, with a smooth surface, small- to medium-sized (up to 15 cm, but sample constraints may have limited this maximum size), and densely colonized on all faces by various species of bryozoans and secondarily by serpulids. This typeof colonization, equally distributed all around the subspherical substrates, indicates that the sampling sites were washed by active water circulation limiting sedimentation. The most frequent bryozoans in the assemblage including  C. fayalensiswere  Microporellasp.,  Schizoporellasp.,  Chorizopora brongniartii( Audouin, 1826)and  Disporellasp. All examined specimens of  C. fayalensiscame from various localities of five Azoresisles: Faial, São Miguel, Formigas, São Jorge, Graciosa. This species has also been recorded in the CanariesIslands by Arístegui (1984), but his figures indicate that  C. fayalensisco-occurs with  C. macaronensisHarmelin, n. sp.in the Canaries(see below). 5891636302 1971-10-12 MNHN-IB- Faial Is. & V Jean-Charcot & St. P. 6 38.52 Azores 19 -28.701944 10 393 2 Azores holotype 2580041167 1971-11-07 MNHN St. P. & H. Zibrowius Jean-Charcot 15 37.270554 Azores 19 -24.781666 Formigas Is. 10 393 1 2580041185 1979-07-24 MNHN, NHMUK Sao Miguel Is. France St. 15 37.705555 Ilheu de Vila 19 -25.4425 Azores Bio-Oceanographic Survey 10 393 1 2580041163 MNHN, NHMUK Sao Miguel Is. France 14 37.706944 Ponta da Galera 19 -25.508888 St. 10 393 1 2580041141 1979-07-25 MNHN Sao Miguel Is. France 15 37.84361 Capelas 19 -35.688614 St. 10 393 1 2580041143 Sao Jorge Is. France 38.751667 Ponta da Rosais 19 -28.316387 St. 10 393 1 2580041105 1979-08-06 MNHN Zibrowius France 10 393 1 2580041178 1979-08-08 MNHN Graciosa Is. France 20 39.059998 Ilheu da Praia 19 -27.953056 St. 10 393 1