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 Author Harmelin, Jean-Georges Author Bishop, John D. D. Author Madurell, Teresa Author Souto, Javier Author Spencer Jones, Mary E. Author Zabala, Mikel text Zoosystema 2019 2019-09-26 41 21 385 418 journal article 10.5252/zoosystema2019v41a21 fbbecb4d-bda5-4215-9147-a47944250471 1638-9387 3725956 urn:lsid:zoobank.org:pub:034A7358-CF99-4908-B047-557074A2E58E Collarina macaronensis Harmelin , n. sp. ( Figs 8B ; 12; 15C; E; 17A, C; 18E; Tables 1-3 ) urn:lsid:zoobank.org:act: 71F62CA0-B1DB-4F12-8646-89F8FFE02477 Cribrilina balzaci – Waters 1899: 9 , figs 31-32. — Norman 1903: 98 , pl. 9, fig. 6; 1909: 292. Collarina balzaci – Harmelin 1978a (in part): 189 , fig. 8 (not pl. I, fig. 7). Cribrilina punctata – Fernández Pulpeiro & Rodriguez Babio 1980: 136 , fig. 2. — Álvarez 1988: 348 , fig. 3B-D. Collarina fayalensis – Arístegui 1984 (in part): 230 , fig. 49b, pl. 11, fig. 6 (not fig. 7). TYPE LOCALITY. — Portugal , Madeira , Porto Santo, Baixo Is . TYPE MATERIAL. — Holotype . Portugal , Madeira , Porto Santo, Baixo Is., R / V Jean-Charcot , ZARCO 1966 St. 23, 33°0’32”N , 16°23’13”W , 5-8 m , dark tunnel, 1 large ovicelled colony (> 1000 zooids) on a pebble, MNHN-IB-2014-1924. Paratypes . Same origin as the holotype: MNHN-IB- 2014-1925: 2 colonies together with the holotype. — MNHN-IB-2014-1926: 4 colonies on 2 pebbles. — MNHN-IB-2014-1927: coated ovicelled colony for SEM examination . OTHER MATERIAL EXAMINED. — Azores , Faial, 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 , coll. by H. Zibrowius, MNHN-IB-2008-7909: 2 colonies on pebble together with holotype and paratypes of C. fayalensis . — Galicia , Sisargas Is., 43°21’22”N , 8°50’17”W , depth unknown, coll. by Fernández-Pulpeiro & Reverter leg., 1 colony ( SEM photos), MHNUSC-Bry-662. — Material from NHMUK collections, SEM photos ( MSJ ): NHMUK 1899.7.1.1344 ( Collarina balzaci ), Madeira , J. Y. Johnson leg., ovicelled colony; NHMUK 1899.7.1.2419 ( Collarina sp.), Madeira , J. Y. Johnson leg., ovicelled colony; NHMUK 1899.7.1.2105 ( Collarina sp.), Madeira , J. Y. Johnson leg., ovicelled colony. 3d; NHMUK 1911.10.1.705 ( Collarina balzaci ), Madeira , ovicelled colony . ETYMOLOGY. — From Macaronesia, biogeographical area including the Canary Islands, Madeira and the Azores where this species was recorded. DIAGNOSIS. — Colonies medium- to large-sized; autozooids with broad marginal gymnocyst, costate shield with a lower central area, predominantly composed of 6-7 costae with smooth surface and long basal part; apertural bar moderately raised, forming an arch; proximal edge of orifice smooth and slightly concave; 3-4 oral spines; paired adventitious avicularia with marked disto-lateral orientation; ooecium formed by distal kenozooid or autozooid; ancestrula with 6 spines. DESCRIPTION Colony encrusting, pluriserial, unilaminar, reaching large size (> 1000 zooids) ( Fig. 18E ). Zooids subpentagonal with distal edge rounded. Gymnocyst visible in frontal view, of variable width laterally, clearly much wider in the proximal part, with some gymnocystal pseudopores ( Fig. 8B ; 12C, H). Costate shield with central part lower than the periphery, composed of 4-9 (6-7:> 60%) short and thick costae, with each ascending basal part forming a clearly distinct lobe, prominent on the gymnocyst and bearing a large pelma, a second, smaller pelma in inner position, before the central, lower part of the shield. Intercostal lacunae 2-3 between adjacent costae, small, irregularly slot-shaped ( Fig. 12 D-H). Apertural bar arched below the orifice, with a moderately raised tip, a pair of medium-sized pseudopores on both sides of the tip and two larger lateral pelmata, near the base of the avicularia ( Figs 8B ; 12F, G; 15C). Adventitious avicularia typically directed laterodistally, almost always paired, rostrum with slightly hooked tip ( Figs 8B ; 12B, D, H; 15C), nested cystids relatively frequent ( Fig. 17A, C ). Orifice wider than long, dimorphic, broader in ovicelled zooids; proximal edge slightly or noticeably concave; condyles triangular, moderately prominent ( Figs 8B ; 12D, F, G). Spines thin, with small conical base, 3 in most cases or 4, 2 arched in ovicelled zooids ( Fig. 12B, D, F ). Ovicell prominent, seemingly acleithral ( Fig. 15C ); ooecium formed by a distal kenozooid at the colony growing edge or by a distal daughter autozooid ( Figs 12 A-C; 15C, E), slightly broader than long, ectooecium with relatively smooth surface, punctured with a dozen medium-sized pseudopores. Ancestrula cribrimorph, with small spinocyst and 6 spines. FIG. 12. — Collarina macaronensis Harmelin , n. sp. : A , B , colony edge,general view and detail with ovicelled and non-ovicelled zooids; C , oblique view: structure of costate shield,apertural bar,paired and apical avicularia; D , G , non-ovicelled zooids with typical traits: concave orifice poster,avicularia directed disto-laterally, costate shield with lower central part and digitate margin, large pelmata also present on the gymnocyst; E , Collarina balzaci ( Audouin, 1826 ) : Harmelin (1978a , fig. 8), Azores, Faial; F , distal part of non-ovicelled zooid with typical orifice, apertural bar with large and small pseudopores, and 3 spines; H , specimen from a continental area. Origin: A, B, NHMUK 1899.7.1.2105, Madeira, J.Y. Johnson leg.; C, NHMUK 1911.10.1.705, Madeira; D, F, G, MNHN-IB-2014-1927, Madeira, Baixo Is.; H, Galicia, Sisargas Is. Scale bars: A, 400 µm; B, C, 200 µm; D, H 100 µm; F, G, 50 µm. REMARKS Collarina macaronensis Harmelin , n. sp. is closely related to C. fayalensis in having orifices with a concave proximal edge, costae with similarly shaped ascending base bearing a large pelma and well-developed gymnocyst. However, in C. macaronensis Harmelin , n. sp. , the paired adventitious avicularia are more distinctly directed disto-laterally and the costate shield involves more costae. Specimens from Faial, Azores recorded and figured as Collarina balzaci by Harmelin (1978a : fig. 8; reproduced here: fig 12E) present these typical characters and belong undoubtedly to C. macaronensis Harmelin , n. sp. This Azorean sample consisted in two colonies on a pebble bearing also the type series of C. fayalensis . The co-occurrence of these two species on the same small substrate confirms they are not ecomorphs of the same species. The discriminant features of C. macaronensis Harmelin , n. sp. are recognizable in part of the figured material from Canary Islands ascribed to C. fayalensis by Arístegui (1984 : fig. 49b; pl. 11, fig. 6) and also in figures of colonies ascribed to Cribrilina punctata by Fernández Pulpeiro & Rodriguez Babio (1980 : Galicia , Ria de Vigo) and Álvarez (1988 : Basque region). SEM examination of specimens from Sisargas Islands, Galicia (coll. by Fernández Pulpeiro, courtesy of O. Reverter) confirmed the occurrence of C. macaronensis Harmelin , n. sp. on the Iberian coast ( Fig. 12H ). SEM examination of three specimens kept at the NHMUK, collected by J. Y. Johnson at Madeira and labelled Collarina balzaci or Collarina sp., has proved that they belong to C. macaronensis Harmelin , n. sp. ( Fig. 12A, B ). As indicated by Norman (1909) in his introduction the Johnson collection was examined by G. Busk, Waters (1899) and himself. Waters (1899) ascribed this material to Cribrilina balzaci and noted thereafter ( Waters 1923 ) that “There are specimens from Madeira , which I took for balzaci , which must perhaps be considered a variety”. The figure of the spinocyst edge of a Madeiran specimen named Cribrilina balzaci by Norman (1903 : pl. 9, fig. 6; same specimen shown here as Fig. 12C ) shows costae with basal parts protruding greatly from the gymnocyst, a typical character of C. macaronensis Harmelin , n. sp. Habitat and geographical distribution In both the Azores and Madeira C. macaronensis Harmelin , n. sp. was found on pebbles and cobbles in shaded conditions ( Fig. 18E ). On the other hand, its absence on rocky walls of dark caves at Madeira despite an extensive sampling survey (JGH, September 2000 ) suggests that this habitat is unsuitable to this Collarina species, in contrast to several Cribrilaria species which were very abundant (JGH, unpublished data). Small substrates in contact with a flat bottom at shallow depth may be an ecological requirement of this species. It is worth noting that this species was noticed by Norman (1909 , as C. balzaci ) on stones between tide-marks at Madeira . At the present state of knowledge, the geographical range of C. macaronensis Harmelin , n. sp. includes the NE Atlantic archipelagoes ( Canaries , Madeira , Azores ) and the northern coast of the Iberian Peninsula.