Porella compressa Revised 1805

Porella compressa (J. Sowerby, 1805) (Figs 3, 29–44; Table 4) Millepora compressa J. Sowerby, 1805: 83, pl. 41. Smittia grimaldii Jullien in Jullien & Calvet 1903: 99, pl. 11, fig. 1. Smittia decipiens Jullien in Jullien & Calvet, 1903: 101, 149, pl. 12, fig. 5. Smittia fallax Jullien in Jullien & Calvet, 1903: 102, pl. 12, fig. 6. Smittia immersa Jullien in Jullien & Calvet, 1903: 105, pl. 13, fig. 6. Smittina immersa: Reverter-Gil & Fernández-Pulpeiro, 2001: 110. Porella compressa: Reverter-Gil & Fernández-Pulpeiro, 1998: 46, fig. 2; Hayward & Ryland, 1999: 158, figs 55, 57C, D. Material examined. Lectotype of Smittia decipiens (designated here): MNHN 3924, Hirondelle st. 53, 43.74722’N, 05.86278°W, 135 m depth, Bay of Biscay, Calvet Coll. Additional material examined: MNHN 3934, Hirondelle st. 60, 43.95000°N, 07.11278°W, 300 m depth, Bay of Biscay, Calvet Coll. as Smittia decipiens. Lectotype of Smittia fallax (designated here): MOM 420222: Hirondelle st. 58, 43.66667°N, 06.57944°W, 134 m depth, Bay of Biscay, Calvet Coll. Paralectotypes of Smittia fallax (designated here): MOM 420124: Hirondelle st. 53, 43.74722°N, 05.86278°W, 135 m depth, Bay of Biscay, Calvet Coll. MOM 420149: Hirondelle st. 56, 43.64167°N, 06.13778°W, 90 m depth, Bay of Biscay, Calvet Coll. MOM 420240: Hirondelle st. 59, 43.88333°N, 06.67944°W, 248 m depth, Bay of Biscay, Calvet Coll. MOM 420255: Hirondelle st. 60, 43.95000°N, 07.11278°W, 300 m depth, Bay of Biscay, Calvet Coll. Holotype (by monotypy) of Smittia grimaldii: MNHN 5966, Hirondelle st. 85, 46.51667°N, 04.52944°W, 180 m depth, Bay of Biscay, Calvet Coll. Lectotype of Smittia immersa (designated here): MNHN 3907, Hirondelle st. 60, 43.95000°N, 07.11278°W, 300 m depth, Bay of Biscay, Calvet Coll. Paralectotype of Smittia immersa (designated here): MOM 420256, Hirondelle st. 60, 43.95000°N, 07.11278°W, 300 m depth, Bay of Biscay, Calvet Coll. Additional material: MHNUSC-Bry 317: Doniños, 43.50105°N, 08.35002°W, 44 m depth. MHNUSC-Bry 539: Malpica, 43.42528°N, 08.82361°W, 119 m depth, 16/11/2001. MHNUSC-Bry 545: Malpica, 43.39667°N, 09.18139°W, 186 m depth, 02/11/2003. MHNUSC-Bry 540: Corme, 43.31083°N, 09.03083°W, 100 m depth, 31/ 12/2002. MHNUSC-Bry 541, 542: Laxe, 43.29139°N, 09.20667°W, 143 m depth, 06/11/2004. MHNUSC-Bry 543: Camariñas, 43.20139°N, 09.32167°W, 148 m depth, 18/02/2002. MHNUSC-Bry 547: Ría Muros, Porto do Son, 42.72194°N, 09.03500°W, 16 m depth, 08/03/2002. MHNUSC-Bry 546: Ría Muros, 42.68528°N, 09.10306°W, 44 m depth, 22/03/2002. MHNUSC-Bry 548: A Guarda, 41.98472°N, 08.95056°W, 69 m depth, 21/03/2005. SD, standard deviation; N, number of measurements. Additional localities studied (specimens not stored): O Barqueiro, 43.69806°N, 07.39083°W, 68 m depth, 13/ 12/2002. A Coruña, 43.54167°N, 08.43722°W, 94 m depth, 10/01/03. A Coruña, 43.43139°N, 08.40750°W, 63 m depth, 02/03/2002. A Coruña, 43.41056°N, 08.51528°W, 85 m depth, 15/05/2002. Malpica, 43.35222°N, 08.83778°W, 23 m depth, 02/06/2002. Laxe, 43.36639°N, 09.32778°W, 223 m depth, 29/01/2002. Laxe, 43.34389°N, 09.09611°W, 127 m depth, 30/01/2005. Laxe, 43.39944°N, 09.18833°W, 190 m depth, 18/09/2005. Camelle, fragments, 43.28194°N, 09.14333°W, 95 m depth, 14/01/2003. Camelle, several dead fragments, 43.21417°N, 09.11500°W, 35 m depth, 05/05/2004. Fisterra, one small colony, 42.84778°N, 09.39750°W, 152 m depth, 08/12/2002. A Guarda, 41.90833°N, 08.97667°W, 85 m depth, 04/02/2005. Description. Colony erect, about 5–10 cm high, orange, with a thickly calcified encrusting base, irregularly branching, becoming thinner toward the growing margin (Figs 29, 42). Branches bilaminar, initially flat, but becoming irregularly rounded in later ontogeny. Autozooids rectangular to hexagonal, initially in regular series, but then irregularly arranged when frontal budding occurs in older parts of the colony (Fig. 44). Frontal shield slightly convex, with few frontal pores and large marginal pores; rapid thickening from secondary calcification does not substantially change its aspect. Primary orifice approximately as long as wide, with distal margin semicircular and proximal margin straight with a broad, low lyrula, occupying up to three-quarters of the proximal margin; condyles rounded, not well developed (Figs 31–35). Avicularium suboral, immediately proximal to the lyrula, oriented slightly sloping or almost perpendicular to the plane of the orifice, proximally directed (Figs 31–35). Rostrum semicircular, sometimes with a very small denticulation (Figs 32, 35, 39); crossbar complete, leaving a small oval proximal opesia. The secondary calcification soon conceals avicularium and the primary orifice with a deep peristome. Secondary orifice circular with a thin raised rim bordered by a narrow groove inside the peristome (Fig. 30). Whenever new zooids bud frontally and secondary calcification increases, different layers of zooids are formed, adapting the longitude of the peristome to the position of the zooid in the different layers, been more elongated in zooids with more internal position (Figs 40, 41). Vicarious spatulate avicularium sometimes present, positioned mainly in branching points; rostrum rounded and lightly cupped distally, crossbar complete with a small but robust columella (Figs 36–38). Ovicell globular, wider that long, imperforate and visible only in early ontogeny, soon completely immersed (Figs 42, 43). Remarks. Porella compressa is regarded as a Boreal-Arctic species (Hayward & Ryland 1999), although distributed from the Faroe Islands to southwest of the Iberian Peninsula (Reverter-Gil et al. 2014). We have revised material of this species collected NW of the Iberian Peninsula between 16 and 300 m depth. On the other hand, Smittia decipiens, S. fallax and S. immersa were described by Jullien (in Jullien & Calvet 1903) from material collected by l’Hirondelle in the southern area of the Bay of Biscay, and S. grimaldii from the northern area. Smittia decipiens was collected at st. 53 (two colonies at 135 m depth) and st. 60 (one sample at 300 m depth) (Figs 35, 38, 39). Several fragments of S. fallax were collected at sts 53, 56, 57, 58, 59, 60, between 90 and 300 m depth (Figs 30, 31, 36, 40, 41). Two broken colonies of S. immersa were collected only at st. 60 (300 m depth) (Figs 32, 37). Finally, S. grimaldii was collected only at st. 85 (several fragments at 180 m depth) (Fig. 33). None of the species have been collected again, but S. decipiens was already considered a junior synonym of P. compressa by Reverter-Gil & Fernández-Pulpeiro (1998), and S. immersa was included, as Smittina immersa, in a Galician checklist (Reverter-Gil & Fernández-Pulpeiro 2001). Samples of the four species are now stored at the MNHN and MOM, and lectotypes and paralectotypes are here designated. The study of all this material proves that these species of Smittia are indistinguishable from P. compressa because they show all the diagnostic features of the species. Moreover, three of these species (S. decipiens, S. fallax, S. immersa) were collected at the same station (st. 60), and it is incomprehensible why Jullien & Calvet (1903) described the material as three different species without discussing the perceived differences. On the other hand, the original description of S. grimaldii (Jullien in Jullien & Calvet 1903, p. 99) is very similar to that of P. compressa (see description above). Nonetheless, the authors merely stated that this species is quite similar to S. fallax, but without further comments. The primary orifices of all the colonies are similar (Figs 31–35, 39; Hayward & Ryland 1999, fig. 55C), as is the suboral avicularium (Figs 31–35, 39; Hayward & Ryland 1999, fig. 55A, B). The secondary orifice shows the same outline (Figs 30, 36, 41; Hayward & Ryland 1999, figs 55D, 57C). All the colonies are erect, with branches initially flat-sectioned, but becoming cylindrical in later ontogeny. There are, however, some differences. In the type material of S. decipiens and S. immersa the proximal border of the rostrum of the avicularia is finely dentate (Figs 32, 35, 39), a character not seen in the material of the other two species (S. fallax and S. grimaldii) or mentioned in any of the available descriptions of P. compressa. In any case, this feature is not visible using optical microscopy and is rapidly obscured by secondary calcification. In addition, the edge of the avicularia in S. fallax and S. grimaldii is eroded in the material examined, so denticles may be missing. Another difference is the presence of large, spatulate avicularia in material of S. decipiens, S. fallax and S. immersa (Figs 36–38), mainly at the margin of the branches. Such avicularia were not found in material of S. grimaldii, and were not reported in the literature on P. compressa. Their presence, however, seems inconstant and they are very difficult to see owing to the general disorder of the zooids in many parts of the colony. In fact, these avicularia were originally reported by Jullien & Calvet (1903) only in S. immersa, but we have found them in the type material of S. decipiens and S. fallax, as well as in colonies collected in Laxe (NW Spain, MHNUSC-Bry 541). Perhaps these variations are due to environmental or geographical factors or intraspecific variations which, however, are insufficient to define separated species. In summary, there is no single character, or set of characters, that makes it possible to distinguish between the four species described by Jullien & Calvet (1903), or between them and P. compressa. Although new, freshly collected material may allow undertaking future genetic studies, we currently assume that S. decipiens, S. fallax, S. grimaldii and S. immersa are junior subjective synonyms of P. compressa, as was already stated by Reverter-Gil & Fernández-Pulpeiro (1998) for S. decipiens. The variability in colonial morphology is remarkable and mainly related to secondary calcification, which results in flat to circular-section branches that develop very long peristomes (Figs 39–41). We were unable to observe polypide activity in these zooids from the innermost layers, but nothing indicates that they are nonfunctional. These long tubular structures are therefore probably related to morphological adaptations of the polypide, as for instance a larger development of the introvert, which enables expanding the tentacle crown outside of the peristome. The proximal channel along the peristome no doubt helps with water exchange during the process of polypide extension and retraction.Published as part of Souto, J. & Reverter-Gil, O., 2019, Identity of bryozoan species described by Jullien & Calvet from the Bay of Biscay historically attributed to Smittia, pp. 105-123 in Zootaxa 4545 (1) on pages 115-120, DOI: 10.11646/zootaxa.4545.1.6, http://zenodo.org/record/261870

Raymondcia gemmata Souto & Reverter-Gil 2019, n. comb.

Raymondcia gemmata (Jullien in Jullien & Calvet, 1903) n. comb. (Figs 2, 23–28; Table 3) Smittia gemmata Jullien in Jullien & Calvet, 1903: 105, pl. 13, fig. 4. Material examined. Lectotype (designated here): MOM 420125: Hirondelle st. 53, 43.74722°N, 05.86278°W, 135 m depth, Bay of Biscay, Calvet Coll., dry fragment on slide. Paralectotype (designated here): MOM 420125: Hirondelle st. 53, 43.74722°N, 05.86278°W, 135 m depth, Bay of Biscay, Calvet Coll., fragment on ethanol. Description. Colony encrusting, unilaminar, beige in preserved material, colour unknown when alive. Zooids hexagonal to oval, longer than wide, separated by fine sutures with a lateral laminar wall visible in frontal view. Frontal wall slightly convex, finely granulated, evenly pierced by large round pores (Fig. 23). Primary orifice wider than long, with a convex proximal border, but lacking a lyrula. Two small quadrangular condyles in the proximolateral corners (Figs 24, 27, 28). Oral spines not observed. Secondary orifice oval, wider than long, formed by the extension of secondary calcification of the adjacent zooids distally by the proximal margin of succeeding zooids, and in some cases by the disto-lateral zooids (Figs 24, 27). Avicularia suboral, placed within the secondary orifice and almost perpendicular to the primary orifice. Apparently oval in outline, not well preserved in the studied material, with a short, rounded mandible proximally directed, and a crossbar formed by two lateral condyles (Figs 27, 28). Ovicell subimmersed, formed by the distal zooid, spherical, completely calcified with only one central pore; secondary calcification finely granular, covering the ovicell from the distal and lateral zooids, with the sutures well visible between the implicated zooids (Figs 25, 26). Ancestrula unknown. Remarks. Jullien (in Jullien & Calvet 1903) described Smittia gemmata from two fragments collected at st. 53 of l’Hirondelle (Bay of Biscay, 135 m depth) (Fig. 2), now stored at the MOM. These specimens are here designated as the lectotype and paralectotype. The original description of the species was incomplete; among other characters, the suboral avicularium was not reported. The species does not seem to have been found again since its original description. Soule et al. (1995) established the genus Raymondcia for two smittinid species with apertural rim and ovicells composed of distal, lateral and frontal (in orifice) segments, and ovicell sometimes with one or two small central pores. Currently, five species with a mainly circumpolar distribution are included in this genus (Bock & Gordon 2018): R. klugei (Gontar, 1982), R. majuscula (Smitt, 1867), R. mcginitiei Soule, Soule & Chaney, 1995, R. osburni Soule, Soule & Chaney, 1995 and R. rigida (Lorenz, 1886). However, Soule et al. (1995) already indicated in the original description of the genus that good examples of the ovicell of Raymondcia were present in species such as Smittina beringia Kluge, 1952 or the description of Prenantia bella (Busk, 1860) by Hayward & Ryland (1979). It remains unclear whether the authors transferred these species to Raymondcia or not. In fact, a few pages later they discussed Lepralia bella with the new species Dengordonia uniporosa Soule, Soule & Chaney, 1995, but without referring the former to Raymondcia. Years later, Grischenko et al. (2007) reported the species as ‘ R. bella (Busk, 1860) ’ (sic.). Accordingly, the genus Raymondcia is already known from European Atlantic waters, as R. bella was reported from Aberdeen, Shetland and west Norway (Hayward & Ryland 1999). We consider that S. gemmata can fit the characters of the genus Raymondcia, and we therefore propose here, though provisionally, to transfer the species to this genus. The original material of the species is scarce and not well preserved, so fresh material is needed to undertake new studies on its morphology.Published as part of Souto, J. & Reverter-Gil, O., 2019, Identity of bryozoan species described by Jullien & Calvet from the Bay of Biscay historically attributed to Smittia, pp. 105-123 in Zootaxa 4545 (1) on page 114, DOI: 10.11646/zootaxa.4545.1.6, http://zenodo.org/record/261870

Identity of bryozoan species described by Jullien & Calvet from the Bay of Biscay historically attributed to Smittia

Souto, J., Reverter-Gil, O. (2019): Identity of bryozoan species described by Jullien & Calvet from the Bay of Biscay historically attributed to Smittia. Zootaxa 4545 (1): 105-123, DOI: https://doi.org/10.11646/zootaxa.4545.1.

Smittina cervicornis Cook 1968

Smittina cervicornis (Pallas, 1766) (Figs 1, 4–22; Table 2) Millepora cervicornis Pallas, 1766: 252. Smittia colletti Jullien in Jullien & Calvet, 1903: 100, 149, pl. 12, fig. 3. Smittia cervicornis (Ellis & Solander): Jullien & Calvet, 1903: 151, pl. 17, fig. 6a–d. Smittia cervicornis (Pallas): Barroso, 1912: 42, fig. 8. Porella cervicornis (Pallas): Calvet, 1931: 93; Gautier, 1962: 204. Smittina cervicornis (Pallas): Cook, 1968: 210; Zabala & Maluquer, 1988: 121, fig. 266, pl. 12C; Hayward & McKinney, 2002: 49, fig. 22A–C. Smittina colletti (Jullien & Calvet): Reverter-Gil & Fernández-Pulpeiro, 2001: 109. ?Not Porella colletti (Jullien & Calvet): Gautier, 1962: 206; Harmelin, 1969: 297, fig. 5:4. Material examined. Holotype (by monotypy) of Smittia colletti: MNHN 3913, Hirondelle st. 57, 43.74167°N, 06.20444°W, 240 m depth, Bay of Biscay, Calvet Coll. Additional material: MOM 420122, Hirondelle st. 53, 43.74722°N, 05.86278°W, 135 m depth, Bay of Biscay, Calvet Coll. as Smittia colletti. MOM 420180: Hirondelle st. 57, 43.74167°N, 06.20444°W, 240 m depth, Bay of Biscay, Calvet Coll. as Smittia colletti. MOM 420183: Hirondelle st. 57, 43.74167°N, 06.20444°W, 240 m depth, Bay of Biscay, Calvet Coll. as Smittia cervicornis. MHNUSC-Bry 422: Point Etxandarri, Bay of Biscay, 43.44225°N, 02.92000°W, 25 m depth. MHNUSC-Bry 423: Avilés Canyon, Bay of Biscay, 43.6709°N, 06.07703°W, 86 m depth. MHNUSC-Bry 534: one colony, A Coruña, 43.56056°N, 08.63361°W, 147 m depth, 12/ 04/2003. MHNUSC-Bry 536: fragments, Malpica, 43.42528°N, 08.82361°W, 119 m depth, 16/11/2001. MHNUSC-Bry 535: two small colonies, Malpica, 43.39667°N, 09.18139°W, 186 m depth, 02/11/2003. MHNUSC- Bry 537: fragments of colonies, Corme, 43.31083°N, 09.03083°W, 100 m depth, 31/12/2002. MHNUSC-Bry 538: a small colony, Fisterra, 42.84778°N, 09.39750°W, 152 m depth, 08/12/2002. MHNUSC-Bry 426: Portimão, 37.13611°N, 08.52722°W, 27/03/2004, many specimens in fishing nets. MNCN 25.03 /4142: Portimão, 37.13611°N, 08.52722°W, 27/03/2004, many specimens in fishing nets. MHNUSC-Bry 424: Menorca Channel, st. P52, 39.85008°N, 03.48209°E, 69 m depth, 12/09/2011. Western Mediterranean; MHNUSC-Bry 449: Imperial de Terre, Riou Island, 43.1729°N, 5.39327°E, 20–22 m depth, 28/09/2018, coll. A. Ostrovsky. MHNUSC-Bry 450: Imperial de Terre, Riou Island, 43.1729°N, 5.39327°E, 20–22 m depth, 28/09/2018 coll. A. Ostrovsky. Additional localities studied (specimens not stored): O Barqueiro, 43.69806°N, 07.39083°W, 68 m depth, 13/ 12/2002. Malpica, 43.35222°N, 08.83778°W, 23 m depth, 02/06/2002. Menorca Channel, st. P5, 39.80500°N, 03.49200°E, 56 m depth, 02/09/2011. Menorca Channel, st. P6, 39.80620°N, 03.49070°E, 55 m depth, 02/09/2011. Menorca Channel, st. P14, 39.7942°N, 03.60070°E, 72 m depth, 04/09/2011. Menorca Channel, st. P50, 39.8507°N, 03.47450°E, 68 m depth, 12/09/2011. Menorca Channel, st. P51, 39.8507°N, 03.48000°E, 69 m depth, 12/09/2011. Menorca Channel, st. P54, 39.83275°N, 03.47425°E, 60 m depth, 13/09/2011. Description. Colony erect, rigid, up to 10–15 cm in height, orange (Figs 4–5). Colonies with abundant branches, branching pattern by bifurcation. Branches bilaminar, flat-sectioned, with up to 12 lineal series of zooids by layer, widest just before bifurcations (Figs 9, 22). In old parts of the colony frontal budding occurs after secondary calcification of the first layer. Frontal budding gave successive layers with the zooids in irregular disposition (Figs 21, 22). Autozooids quadrangular, hexagonal or irregular in shape in successive frontal layers; separated by sutures between slightly raised thin ridges; frontal shield umbonuloid, flat or lightly convex (Figs 6– 16). Large frontal pores along the frontal surface, being larger in the central area (Figs 9–17). Primary orifice semicircular with a short, broad lyrula in the proximal border (Figs 17–20). No spines. Secondary orifice very variable, depending on the degree of secondary calcification and frontal budding. Initially the orifice is keyholeshaped, with a larger distal section corresponding to the primary orifice, and a smaller proximal area, where the suboral avicularium is supported (Figs 10, 11, 18, 20). As the calcification increases, the secondary orifice grows as a peristome, initially with a proximal channel toward the avicularium but quickly forming a thickly calcified peristome, cylindrical, projecting from colony surface. In heavily calcified colonies, only a perfectly round orifice persists, with circular growing marks (Figs 8, 13, 21). Avicularium suboral, prominent, with rounded rostrum and a semicircular mandible, directed proximally. Distal part of avicularium with a raised cup. Crossbar complete, with a robust columella which is frequently bifurcate (Figs 18–20). Ovicell not seen in studied specimens. Remarks. Jullien (in Jullien & Calvet 1903) described Smittia colletti based on one colony from st. 53 of the l’Hirondelle (Bay of Biscay, 135 m depth). This sample is the holotype of the species and is now stored at the MNHN. Later, Calvet (in Jullien & Calvet 1903) reported two isolated fragments from st. 57 (240 m depth). Smittia colletti was never recorded again in the Atlantic, but the original records were compiled by Reverter-Gil & Fernández-Pulpeiro (2001) as Smittina colletti. In the Mediterranean, one specimen was reported by Gautier (1962) at 162 m depth in the Banc du Esquerquis (Tunisia), and Harmelin (1968, 1969) recorded specimens from two other Mediterranean localities: Strait of Sacarpanto, 29–33 m depth and 60–80 m depth, and in Santorini at 100– 110 m depth. Other records were included as nominal references by Harmelin (1979), Zabala & Maluquer (1988), Rosso (2003) and Rosso & Di Martino (2016), but apparently no more material was studied. Calvet (in Jullien & Calvet 1903) also recorded four samples of S. cervicornis at the locality where he recorded S. colletti (st. 57). Smittina cervicornis is a common and abundant species in the Mediterranean, where it was described by Pallas (1766) and later recorded by several authors (e.g. Calvet 1907, 1927, 1931; Gautier 1962; Zabala & Maluquer 1988; Álvarez 1994; Harmelin et al. 1994; Hayward & McKinney 2002; Rosso 2003; Sanfilippo et al. 2013; Rosso & Di Martino 2016), but the type material of the species, described from Riou Island (Marseille, France) appears to be lost, as is the rest of the Pallas collection (Kukliński & Taylor 2008). Outside the Mediterranean, S. cervicornis was also recorded from different localities: Morocco (Canu & Bassler 1925, 1928; d’Hondt 1978), Gulf of Cádiz (López de la Cuadra 1991; Álvarez 1994), south of Portugal (Reverter-Gil et al. 2014), Canary Islands (Arístegui Ruíz & Fernández Gil 2003) and tropical west Africa (Cook 1968), and also from the Bay of Biscay (Jullien & Calvet 1903; Calvet 1907; Barroso 1912; Aguirrezabalaga et al. 1986; Álvarez 1991). Although S. cervicornis and S. colletti, collected at the same locality, seemed very similar, Jullien & Calvet (1903) did not discuss the differences between the two species. It is a mystery why, of all the material collected at st. 57, Calvet (in Jullien & Calvet 1903) decided to report four of the samples as S. cervicornis, whereas he preferred to identify two fragments as a different species, S. colletti. Perhaps he was simply retaining an earlier, unpublished identification of the sample made by Jullien, who was actually the person who described the species in the first part of the work. The first author to provide diagnostic characters for S. colletti was Gautier (1962), who stated that the species comprises white colonies with fewer and narrower branches, and with smaller zooids, and that it seems to live in deeper waters than S. cervicornis. However, these statements were based on material collected in the Mediterranean and were not reported in the original description of S. colletti. More importantly, these features are not visible in the original material of the species. We have studied the original specimens of S. colletti (Figs 6, 7, 9–11, 18, 20), together with other newly collected material from several localities in the Bay of Biscay between 25 and 86 m depth, off the NW Iberian Peninsula between 23 and 186 m depth, in the Balearics between 55 and 72 m depth and specimens collected at 20– 22 m depth from Riou Island, Marseille (type locality of S. cervicornis); and also several colonies collected in the Algarve (south Portugal) previously reported by Souto et al. (2010). Our observations were compared with existing descriptions of S. cervicornis (e.g. Gautier 1962; Zabala 1986; Zabala & Maluquer 1988; Hayward & McKinney 2002). All the material shows wide inter- and intra-colony variability. This pertains especially to frontal-shield morphology, mainly correlated with secondary calcification. It also involves the peristomial area, which is initially keyhole-shaped with a variable coarse calcification that changes to form a circular rim in older zooids. The suboral avicularium seems to be more stable morphologically, but varies in the size of the proximal opesia. There is a welldeveloped central columella, which is typically bifurcate. None of the variations shows a consistent pattern, however, so it is not possible to separate the studied material into different groups. Even the biometrics of all the characters in the colonies studied are similar (see Table 2). Recently, a new species of Smittina was described from Mauritania, Smittina imragueni Matsuyama et al., 2015, which closely resembles S. cervicornis. According to Matsuyama et al. (2015) this species differs from S. cervicornis mainly in the position and size of the avicularia, which are smaller and almost perpendicular to the growing plane. Moreover, the frontal shield of the zooids figured by Matsuyama et al. (2015), seem to be more arcuate than in the specimens studied during this work, although this character could be variable and dependent on secondary calcification. Similar differences were highlighted by Matsuyama et al. (2015) to differentiate S. colletti from S. imragueni, but the authors also indicated the presence of one medial denticle on top of the lyrula of S. colletti as a difference. We were unable to observe this denticle in type specimens of S. colletti or in any other specimen studied here, so in our opinion this difference is incorrect. 9909090909090SD, standard deviation; N, number of measurements. In the absence of significant differences we conclude that the type specimens of S. colletti and the other specimens studied here are actually referable to S. cervicornis, which becomes a senior subjective synonym. Mediterranean material reported by Gautier (1962) and Harmelin (1968, 1969) as Porella colletti probably corresponds to a different species, but confirmation is beyond the scope of the present paper. Smittina cervicornis seems abundant locally in the Bay of Biscay, forming groups of colonies that were observed between 20 and 30 m depth off Point Etxandarri. It is also very abundant in the Mediterranean, where the species has been described as an important component of coralligenous biocoenoses (e.g. Chimenz & Faraglia 1995; Ballesteros 2006; Harmelin 2017).Published as part of Souto, J. & Reverter-Gil, O., 2019, Identity of bryozoan species described by Jullien & Calvet from the Bay of Biscay historically attributed to Smittia, pp. 105-123 in Zootaxa 4545 (1) on pages 108-114, DOI: 10.11646/zootaxa.4545.1.6, http://zenodo.org/record/261870

Bowerbankia citrina

<i>Bowerbankia citrina</i> (Hincks, 1877) <p>(Figures 5, 6, 7B)</p> <p> <i>Valkeria citrina</i> Hincks 1877, p 215.</p> <p> <i>Bowerbankia citrina</i> (Hincks): Hincks 1880, p. 524, plate 76, figures 6–8; Prenant and Bobin 1956, p. 300, figures 122 (VII), 135, 136 (I–III); d’Hondt 1983, p. 61, figure 32E; Hayward 1985, p. 142, figure 48; De Blauwe 2009, p. 86, figures 66–68.</p> <p> <i>Material examined</i></p> <p> MNCN-25.03 <i>/</i> 3830: Ría de Ferrol, 43 <i> ◦</i> 28 <i> ′</i> 01 <i> ′′</i> N, 08 <i> ◦</i> 16 <i> ′</i> 34 <i> ′′</i> W, 8–10 m, 15 November 2009: one colony on a stone. Personal collection: Ría de Ferrol, 43 <i> ◦</i> 28 <i> ′</i> 01 <i> ′′</i> N, 08 <i> ◦</i> 16 <i> ′</i> 34 <i> ′′</i> W, 8–10 m, 15 November 2009: several colonies on a stone. Portugal: Baleal Beach, Peniche, intertidal. June 2010: two colonies.</p> <p> <i>Description</i></p> <p> Colony arborescent and branching, forming dense tangled tufts, less chitinized and less stiff than <i>B. pustulosa</i>; up to 5 cm high in the material examined. Bright yellow when alive, becoming light beige after fixation.</p> <p>Erect branches formed by cylindrical kenozooids separated by transverse septa. Almost all the kenozooids branching; occasionally, some internodes may be formed by 2–3 in-line kenozooids. Branching lateral, frequent; branches developing in different planes around the main axis, making the colony three-dimensional.</p> <p> Rhizoids produced near proximal end of the kenozooids, especially from the basal kenozooids; less numerous than in <i>B. pustulosa</i>, they descend to the base of the colony and are joined to its axis. Structure and possible functions similar to those described for <i>B. pustulosa</i>.</p> <p>Each kenozooid bears in its distal half a dense group of 20–30 closely spaced, free autozooids. Groups form a rather short helix that undergoes slightly more than a half spiral turn around the stolon, causing a slight torsion of the kenozooid bearing them.</p> <p>Autozooids elliptical, budded at the growing tips of the colony as small vesicles clearly arranged in two spiral series; arrangement becoming less clear later in ontogeny. Autozooids deciduous, lacking in extensive areas of the colony, especially older parts, their previous presence being revealed by basal uniporous septula. New autozooids are budded within groups so that the original biserial arrangement is lost. Polypide with eight tentacles, each tentacle with a bright yellow line continuous with a basal line around the tentacular base. Yellow colouration disappears after fixation. Gizzard large, level with the base of the tentacle sheath when the polypide is retracted. Setose collar conspicuous.</p> <p>Zooids with brown bodies have the same shape as feeding autozooids, but are clearly shorter and slightly wider.</p> <p> <i>Measurements</i></p> <p> ZL: 0.518 <i>±</i> 0.0654 (0.423 –0.615; 13); ZW: 0.150 <i>±</i> 0.0164 (0.125 –0.173; 13); ZbL: 0.305 <i>±</i> 0.0204 (0.279 –0.326; 8); ZbW: 0.174 <i>±</i> 0.0256 (0.140 –0.221; 9); SL: 2.080 <i>±</i> 0.1826 (1.741 –2.389; 9); SW: 0.181 <i>±</i> 0.0356 (0.148 –0.259; 9); GL: 1.051 <i>±</i> 0.1319 (0.852 –1.296; 9); G <i>/</i> S: 50.5 <i>±</i> 4.10 (45.0–59.3; 9).</p> <p> <i>Remarks</i></p> <p>Prenant and Bobin (1956) mentioned that the entire autozooid, particularly the polypides, are a lemon-yellow colour in this species. However, Hayward (1985) found the cystid wall to be transparent but the polypide to be coloured. We have observed that the yellow colour is actually due to a narrow, bright lateral line present in each tentacle, and a narrow circular line that surrounds the basis of the tentacle crown (Figure 6A). This can also be seen in the figure of De Blauwe (2009).</p> <p> Both the geographical distribution and number of references to <i>B. citrina</i> are more limited than for <i>B. pustulosa</i>. The species seems to be common in the south of Great Britain and in Brittany, although known with certainty from only a few locations (Hayward 1985). It has also been recently recorded in the Netherlands (De Blauwe 2009). In the Iberian peninsula, this species has been reported only in Aveiro (N Portugal), by Marchini et al. (2007) in a paper that included no description or figures. In the present study we include material from the north-western Iberian peninsula, where the species was hitherto unknown (Reverter-Gil and Fernández- Pulpeiro 2001), and also the Portuguese coast (Peniche), representing the southernmost record for the species.</p>Published as part of <i>Souto, J., Fernández-Pulpeiro, E. & Reverter-Gil, O., 2011, Presence of rhizoids in two species of the genus Bowerbankia (Bryozoa: Ctenostomata) and their systematic relevance, pp. 2543-2557 in Journal of Natural History 45 (41 - 42)</i> on pages 2549-2550, DOI: 10.1080/00222933.2011.597003, <a href="http://zenodo.org/record/5204605">http://zenodo.org/record/5204605</a&gt

Molecular phylogenetic analysis confirms the species status of <i>Electra verticillata</i> (Ellis and Solander, 1786)

Electra verticillata was original described by Ellis and Solander ((1786) The natural history of many curious and uncommon zoophytes collected from various parts of the globe by the late John Ellis, systematically arranged and described by the late D. Solander, London), and since then the species status of this bryozoan has been in dispute. Many bryozoologists considered E. verticillata as one variety of colony morphology of Electra pilosa (Linnaeus 1767). To test the species status of E. verticillata, we analysed DNA sequences from material from the Bay of Douarnenez (near Morgat, France), together with sequences from E. pilosa, E. posidoniae, E. scuticifera, E. indica, and Electra omanensis. Phylogenetic analyses based on fragments of the 18S, 16S and 12S ribosomal RNA genes confirmed the status of E. verticillata as a separate species. We also examined the morphology of specimens of E. pilosa and E. verticillata in various institutions as well as in own collections. This study revealed morphological and ecological differences between these two species and clarified the geographical distribution of E. verticillata

New and little known species of <i>Celleporina</i> Gray, 1848 (Bryozoa, Cheilostomata) from the Atlantic-Mediterranean region

Celleporina parvula, a species present in shallow waters along the NW African coast, is re-described from its original material. Celleporina fragilis, only known from the Canaries, and Celleporina derungsi, from south Portugal, are newly reported from the Gorringe Bank and NW Iberian Peninsula respectively. Two new species are described: Celleporina algarvensis n. sp., from south Portugal, and Celleporina mediterranea n. sp., from NW Mediterranean. The structure of the ooecium and its tabula, and the structure of the frontal wall in the genus Celleporina are discussed in the light of new observations