Cradoscrupocellaria bertholletii ( Audouin, 1826 ) VIEIRA & JONES & WINSTON, 2013

VIEIRA, LEANDRO M., JONES, MARY E. SPENCER & WINSTON, JUDITH E., 2013, <p class = " HeadingRunIn " align = " left "> <strong> <em> Cradoscrupocellaria </ em>, a new bryozoan genus for <em> Scrupocellaria bertholletii </ em> (Audouin) and related species (Cheilostomata, Candidae): taxonomy, biodiversity and distribution </ strong> </ p>, Zootaxa 3707 (1), pp. 1-63 : 7-12

Cradoscrupocellaria bertholletii ( Audouin, 1826) n. comb.

( Figs 2–3 , Table 1)

Acamarchis Bertholletii Audouin, 1826: 241 ; Savigny 1817: pl. 11, figs 3.1–3.5. [ Egypt]

? Scrupocellaria capreolus Heller, 1867: 11 , pl. 1, fig. 1. [Adriatic]

Scrupocellaria Bertholletii (Audouin) : Hincks 1886: 258, pl. 9, figs 1–2. [Adriatic]

? Scrupocellaria bertholletii (Audouin) : Waters 1909: 133. [Suez Canal]

? Scrupocellaria bertholletii (Audouin) : Waters 1918: 5 (in part). [Mediterranean and Suez Canal]

Scrupocellaria bertholletii (Audouin) : Hastings 1927: 335. [Suez Canal]

Scrupocellaria bertholletii (Audouin) : Calvet 1927: 7. [ Monaco]

Scrupocellaria bertholletii (Audouin) : Marcus 1938: 24 (in part), pl. 5, figs 11A–B. [ Brazil: São Paulo]

? Scrupocellaria bertholletii (Audouin) : Gautier 1962: 85. [Mediterranean]

? Scrupocellaria bertholletii (Audouin) : Prenant & Bobin 1966: 418 (not figs 136–137). [Mediterranean]

? Scrupocellaria bertholletii (Audouin) : Zabala i Limosin 1986: 315 (non fig. 86). [Mediterranean]

? Scrupocellaria bertholletii (Audouin) : Zabala & Maluquer 1988: 97, figs 157–158. [Mediterranean]

Scrupocellaria bertholletii (Audouin) : d’Hondt 1988: 198. [ Israel]

Scrupocellaria bertholletii (Audouin) : Ramalho 2006: 127, fig. 28. [ Brazil: Rio de Janeiro]

Scrupocellaria sp. Gordon et al. 2008: 44, 46, figs 4.1–4.3. [ New Zealand]

Not Scrupocellaria reptans var. bertholletii (Audouin) : Water 1897: 6, pl. 1, figs 18–19. [ Italy]

Not Scrupocellaria bertholletii (Audouin) : Norman 1909: 283, pl. 36, figs 1–2. [Madeira]

Not Scrupocellaria bertholletii (Audouin) : Waters 1918: 5. [ Cape Verde]

Not Scrupocellaria bertholletii (Audouin) : Hastings 1930: 703, pl. 1, figs 1–5. [Galapagos and Gorgona Island]

Not Scrupocellaria bertholletii (Audouin) : Marcus 1938: 24, pl. 5, figs 11A–B. [ Brazil: São Paulo]

Not Scrupocellaria bertholletii (Audouin) : Osburn 1940: 386. [ Bermuda, Puerto Rico and Tortugas]

Not Scrupocellaria bertholletii (Audouin) : Osburn 1947: 20. [N Colombia and Aruba Island]

Not Scrupocellaria bertholetii (sic) (Audouin): Osburn 1950: 133, pl. 18, figs 7–8, pl. 21, fig. 8. [Pacific]

Not Scrupocellaria bertholletii (Audouin) : Banta & Carson 1977: 389, fig. 3 [ Costa Rica]

Neotype (chosen here). NHMUK 1926.9 .6.58 ( Fig. 2A–C ), Scrupocellaria bertholletii, A.B. Hastings det., Suez Canal Expedition, coll. H.M. Fox, 15 December1924, Suez Canal P. 3, 46B.

Additional material examined. NHMUK 1885.8 .12.1, Menipea jeffreysii, S.O. Ridley det., S.O. Ridley Collection, 9 August 1922, Viareggio , Mediterranean . NHMUK 1899.5 .1.264, with Scrupocellaria scruposa, T. Hincks Collection , Adriatic . NHMUK 1899.5.1.341–2, Scrupocellaria bertholletii, T. Hincks det., T. Hincks Collection, Adriatic . NHMUK 1899.5 .1.421, Scrupocellaria bertholletii, T. Hincks det., T. Hincks Collection, Adriatic . NHMUK 1899.7 .1.736 ( Fig. 2D–F ), Scrupocellaria bertholletii , Mediterranean . NHMUK 1911.10 .1.357, Scrupocellaria bertholletii var. aperta, A.M. Norman det., Madeira , 1897 . NHMUK 1911.10 .1.358, Scrupocellaria bertholletii, Prof. Heller det., A.M. Norman Collection, Adriatic . NHMUK 1926.9 .6.60, Scrupocellaria bertholletii, A.B. Hastings det., Suez Canal Expedition, coll. H.M. Fox, 26 October 1924, Suez Canal , barge 690, Port Taufiq . NHMUK 1926.9.6.66–7, Scrupocellaria bertholletii, A.B. Hastings det., Suez Canal Expedition, coll. H.M. Fox, 15 December 1924, Suez Canal P. 3 . NHMUK 1926.9 .6.68, Scrupocellaria bertholletii, A.B. Hastings det., 26 October 1924, Suez Canal , Port Taufig, Barga 690 . NHMUK 1926.9 .6.69, Scrupocellaria bertholletii, A.B. Hastings det., Suez Canal Expedition, coll. H.M. Fox, Suez Canal, Dredger N. 8 . NHMUK 1963.1 .26.2, Scrupocellaria bertholletii, J.S. Ryland det., 30 December 1960, Porto d’Ischia , Italy . NHMUK 1964.3 .12.6, Scrupocellaria reptans , Malta , Mediterranean. NHMUK 2007.11.9.42–3, Scrupocellaria bertholletii, J.S. Ryland det., 26 February 1962, A.P. Austin leg., Malta , Mediterranean. USNM 9083 , Mediterranean . NHMUK 2012.7 .1.11 ( Fig. 3C–F ), Station Cru 82.043, 14 August 1982, Charles H.J.M. Fransen leg., Curaçao , Piscadera Inner Bay , Candelichi, 0–0.5 meters, on Rhizophora roots . MOM 421467 , Scrupocellaria bertholletii, L. Calvet det., Monaco , 652i, slide 84. MOM 42529, Scrupocellaria bertholletii, Calvet det., Port of Monaco , 1903, slide 85. MOM 421681 (part of specimens in NHMUK 2012.7 .1.4, Fig. 3A–B ), L. Calvet det., 30 January 1908, Sur le Coffre du Port Monaco . UFAL 548 , coll. M.D. Correia, November 1987, Station #A, Paranaguá Bay, Paraná, Brazil , colony found on acrylic panels. USNM 9678 , Scrupocellaria bertholletii, A.B. Hastings det., Suez Canal Expedition, coll. H.M. Fox, 26 December 1924, Port Taufig , Barga 690. Cambridge Expedition . USNM 9083 , Scrupocellaria scruposa , Mediterranean .

Diagnosis. Colony with chitinous joints passing across proximal end of opesia in zooids C and D at bifurcation; zooids with 1 inner and 1–2 outer spines, 1 distal spine rarely present; forked scutum sometimes present at inner margin of opesia, overarching to midline of membranous area; small distolateral avicularium present in each zooid; vibracular chamber almost rounded and small; rhizoids with smooth surface.

Redescription. Colony erect, branched, with internodes comprising 5–9 zooids. Internodes almost straight, with acute bifurcating pattern; chitinous joints passing across proximal end of opesia in outer zooids at bifurcation (zooids C and D), and across proximal gymnocyst of inner zooids (F and G). Autozooids elongate, slightly tapering proximally. Oval opesia occupying distal half to three fifths of zooidal length, with two lateral folds where lateral spines are inserted; cryptocyst proximal very narrow, inconspicuous, deep and smooth. Scutum sometimes present, inserted at midline of inner opesial edge, small, forked, angled at 70–90°, overarching to midline of frontal membrane. Distal spines short, unbranched; 1 inner, 1–2 outer spines, rarely 1 distal spine; most proximal outer and inner spines directed forward; axial zooid with 3–4 spines. One distolateral avicularium present in each zooid, small, laterally directed; rostrum triangular, 0.044 –0.062 mm long, with serrated lateral edge and slightly curved tip. Frontal avicularia often present, dimorphic: small frontal avicularium with raised tubular base, 0.040 –0.065 mm long, rostrum slightly serrated, triangular, directed forward; large frontal aquiline avicularium, 0.115–155 mm long, rostrum serrated and slightly curved, directed forward, with triangular curved mandible with hooked tip. A vibracular chamber present on basal surface of each zooid, inconspicuous in frontal view; chamber of vibraculum almost circular, with large rhizoidal foramen on its proximal outer corner; setal groove directed transverse to axis of internode, straight, with smooth setae longer than one autozooid. Single axial vibraculum without rhizoidal foramen. Rhizoids tubular and smooth. Ovicells globular, slightly enlarged distally, with ectooecium perforated by 12–16 medium-sized rounded pores; ovicelled zooids with 1 inner and 2 outer spines.

Remarks. The morphological variation reported for C. bertholletii in the literature (e.g. Prenant & Bobin 1966: 418) led us to examine specimens collected at different localities worldwide. Acamarchis bertholletii was introduced by Audouin (1826), based on colonies from Egypt, but their exact locality is unknown; they could have been collected in either the Red Sea or Mediterranean coast. Audouin’s description was based on specimens illustrated by Savigny (1817), presumably lost (d’Hondt 2006). Two diagnostic characteristics are observed, however, in Savigny’s plates, viz the bifurcated frontal scutum and the shape of the basal vibraculum. These characteristics are also observed in some specimens from the Suez Canal and Mediterranean, as well as in a small specimen from Madeira (NHMUK 1911.10.1.357) identified by Alfred M. Norman (1831–1918) under an unpublished name ( Scrupocellaria bertholletii var. aperta ). Thus, the neotype of C. bertholletii is here selected from the Suez Canal (NHMUK 1926.9.6.58) in order to establish the identity of this species. The majority of specimens deposited at AMNH, NHMUK and USNM assigned to C. bertholletii have scuta with distinct branching patterns and belong to distinct species. Some of those species are reassigned to new species (see below). Unfortunately, we have no access to specimens assigned to C. bertholletii by Gautier (1962), Prenant and Bobin (1966) and Zabala i Limosin (1986), hence their identities remain uncertain.

Cradoscrupocellaria bertholletii has been reported in tropical and subtropical waters worldwide. Hastings (1930) noted distinct differences in morphology among specimens of C. bertholletii from different localities in the Pacific and Egypt, but, notwithstanding, used the same name for three morphologically distinct specimens from Pacific waters. Likewise, morphological differences were described by other authors (e.g. Waters 1897, 1918; Norman 1909; Marcus 1938; Osburn 1940, 1950). Osburn (1950) first described one distinct form of C. bertholletii from California, named Scrupocellaria bertholletii var. tenuirostris , characterized by the elongate shape of the frontal avicularia and the branching pattern of the scutum, now assigned as a distinct species (see below). Later, Fransen (1986) also used morphological characteristics of the scutum and absence of lateral avicularia to distinguish the Caribbean species C. curacaoensis (see below). Among the specimens identified by Marcus (1938) as Scrupocellaria bertholletii , a colony without lateral avicularia was observed; this specimen is assigned to a distinct species (see below), while the specimens with lateral avicularia belong to C. bertholletii .

Six other species with branched scuta previously assigned to Scrupocellaria are known so far— Scrupocellaria bellula , Scrupocellaria ellisi , Scrupocellaria macrorhyncha , Scrupocellaria micheli ( Marcus, 1955) , Scrupocellaria nanshaensis , and Scrupocellaria reptans . Scrupocellaria micheli differs from Cradoscrupocellaria by the presence of large lateral avicularia and a basal vibracular chamber with the setal groove oblique to the internode axis; Scrupocellaria bellula ( Fig. 1 ) is distinguished from Cradoscrupocellaria by its branching pattern, the presence of branched distal spines and its elaborately forked scutum. The other four species are herein assigned to Cradoscrupocellaria ― Cradoscrupocellaria ellisi , C. macrochyncha , C. nanshaensis , and C. reptans .

Cradoscrupocellaria bertholletii is characterized by a forked scutum (often absent, see Figs 2A–B, D–E , 3A, C–E ), the presence of small distolateral avicularia and an almost rounded basal vibraculum chamber. An unidentified specimen from New Zealand (see Gordon et al. 2008, pp. 44, 46, figs. 4.1–4.3) is characterized by the lack of a scutum; this specimen belongs to C. bertholletii . The specimen figured and described by Heller (1867) as Scrupocellaria capreolus from the Adriatic Sea differs from C. bertholletii by the presence of bi- to trifurcate scuta ( Heller 1867, pl. 1, fig. 1); this species, however, requires further examination.

1 NHMUK 1899.7.1.736, Mediterranean. 2 NHMUK 2012.7.1.4, Monaco. 3 NHMUK 2012.7.1.11, Curaçao. 4 NHMUK 2012.7.1.12, Curaçao. 5 USNM 559182, Bermuda.

The specimens reported by Waters (1897) from Italy are distinguished from C. bertholletii by more-branched scuta; their identities, however, remain uncertain. Specimens identified by Waters (1918) from Cape Verde belong to a distinct species (see below), while the identity of his specimens from the Mediterranean and Suez Canal remain uncertain. The figures of C. bertholletii given by Zabala and Maluquer (1986) are characterized by bifurcate scuta, but no specimens were examined by us to confirm their identification. In the MNHN collections, dozens of specimens from the Mediterranean and Red Sea were previously assigned to C. bertholletii ; they also require re-examination.

Distribution. New Zealand; Suez Canal; Mediterranean; Adriatic; Madeira; Curaçao; Brazil; shallow waters.

Figures

FIGURE 1. Scrupocellaria bellula Osburn, 1947. A–D, Uncatalogued specimen (part of AMNH 1518.1), Jamaica. A, Frontal surface of branch with ovicelled zooids. B, Close-up of two ovicelled zooids; note the branched frontal scutum and small frontal avicularium on zooids. C, Frontal surface of branch; note the presence of seven distal spines. D, Close-up of frontal surface of autozooid.

FIGURE 2. Cradoscrupocellaria bertholletii (Audouin, 1826) n. comb. A–C, NHMUK 1926.9.6.58, neotype, Suez Canal; D– F, NHMUK 1899.7.1.736, Mediterranean. A, Frontal surface of branches and two bifurcations. B, Frontal surface of a branch; note zooids with and without scuta. C, Close-up of an abfrontal vibraculum. D, Frontal surface of a branch bifurcation; note the presence of dimorphic frontal avicularia (gigantic on axial zooid) and two zooids with scutum. E, Close-up of an ovicelled zooid with scutum; note the small lateral avicularium. F, Abfrontal surface of branch; note the smooth surface of rhizoids.

FIGURE 3. Cradoscrupocellaria bertholletii (Audouin, 1826) n. comb. A–B, NHMUK 2012.7.1.4, Monaco; C–F, NHMUK 2012.7.1.11, Curaçao. A, Frontal surface of branches and bifurcation. B, Abfrontal surface of branch. C, Frontal surface of branches; note the absence of scuta in the entire colony. D, Frontal surface of bifurcation. E, Close-up of two zooids, one with ovicell; note the absence of scuta in ovicelled and non-ovicelled zooids. F, Abfrontal surface of colony.