Celleporina canariensis Aristegui, 1989: 147

HAYWARD, PETER J. & McKINNEY, FRANK K., 2002, Northern Adriatic Bryozoa From The Vicinity Of Rovinj, Croatia, Bulletin of the American Museum of Natural History 2002 (270), pp. 1-139 : 86-96

Celleporina canariensis Aristegui, 1989: 147 .

DESCRIPTION (AMNH 988; CMRR 2260): Colonies cream­colored or light yellow, encrusting, multilaminar, forming small pisiform nodules, to 3 mm diameter in present material. Autozooids small, with smoothly calcified frontal shield and large, conspicuous marginal pores. Primary orifice longer than wide, with a short, U­shaped sinus midproximally and rounded, prominent condyles. No oral spines. Peristome deep, tubular, obscuring orifice, its rim shallowly notched midproximally; peristomial avicularia paired, lateral, rostrum terminal, elliptical, ca. 0.09 mm long, with complete crossbar, bluntly triangular, directed distolaterally. Distal to each avicularium, peristome rim is produced as a short, blunt, medially directed process. Vicarious avicularia sparse, dimorphic: elliptical, 0.2 mm long, with complete crossbar, thickened columella, no palate; or larger, to ca. 0.4 mm, with broadly spatulate rostrum, lacking palate, angled at 45° to minute proximal opesia and directed basally. Ovicell about as wide as long, frontal surface level with peristome rim; entooecial tabula occupying most of the frontal surface, bordered by a single row of pores. Aperture edged by a broad band of smooth ectooecium, often produced medially into a short umbo.

Tentacles light yellow, 12; lophophores flaring conical, radially symmetrical.

REMARKS: This species is most similar to C. caminata , a common Mediterranean spe­ cies which develops robust nodular colonies. It is distinguished from that species principally by its smaller orifice and proportionately shorter sinus. Further, the peristome rim in C. caminata typically bears three columnar avicularia with elliptical rostra. Aristegui (1989) described only spatulate vicarious avicularia in his type suite but both these and the smaller elliptical type occur in single colonies of the material described here.

DISTRIBUTION: Celleporina canariensis is presently known from several localities around the Canary Isles, and from caves along the Catalan coast.

MEASUREMENTS (SKELETAL): AAL 98 ± 10 µm, 77–113 (3, 28), AAW 55 ± 7, 47–72 (3, 8), DO 354 ± 53, 259–452 (3, 30), OL 130 ± 6, 119–139 (5, 22), OW 103 ± 5, 89– 112 (5, 22), OvL 145 ± 18, 116–174 (3, 12), OvW 218 ± 18, 189–224 (3, 12), SL 23 ± 5, 15–36 (5, 22), SW 23 ± 4, 15–31 (5, 22), VAL 267 ± 49, 213–308 (2, 3), VAW 166 ± 41, 142–213 (2, 3). (POLYPIDE): IH 0 µm (1, 2), LD 880 (1, 10), TL 560–680 (1, 2).

Celleporina siphuncula , new species Figure 40A–D

DIAGNOSIS: Colonies pisiform, small; orifice widely sinuate; peristome long, tubular; peristomial avicularium single, proximo­medial.

HOLOTYPE: AMNH 989.

PARATYPES: AMNH 990 –994; CMRR 2261.

ETYMOLOGY: The species name is from the Latin sipho (bent tube) and – culum (suffix denoting diminutive) because of the shape of the small peristomial avicularium.

DESCRIPTION: Colonies cream colored, pisiform, to 4.5 mm diameter in present material. Autozooids closely packed, oval in outline with a smooth, convex frontal shield, with a few indistinct marginal pores; commonly 0.3 mm in length, with a cylindrical peristome up to 0.375 mm long. Autozooid boundaries distinct only in the smallest colonies, of fewer than 20 autozooids; subsequent astogeny proceeds through fronto­lateral budding, with peristomes projecting perpendicularly to colony surface. Primary orifice slightly wider than long; a shallow, u­shaped sinus occupies two­thirds of the proximal border, between short, broad condyles, its depth equivalent to one­fifth total orifice length. No oral spines. Peristome encircles whole of primary orifice, developing as a tall cylinder, its rim slightly flared and produced into a variable number of short lobes. A small avicularium present midproximally on peristome rim, originating from a septulum proximolateral to orifice, either right or left, traversing peristome as a slender tube and expanding into an ovate cystid on its distal edge; rostrum acute to orifice plane, proximally directed, oval, with a finely denticulate distal rim, crossbar complete, slen­ der, mandible semielliptical. Vicarious avicularia frequent, 0.25–0.3 mm long; rostrum broadened distally and deeply cupped, but not spatulate, crossbar complete, slender, without a columella; palate simply a narrow shelf bordering a foramen occupying most of length and breadth of rostrum. Ovicell hemispherical; a broad tabula occupying most of visible frontal surface, with a thickened ectooecial rim and a single marginal series of large pores; aperture wide and arched, opening into proximal portion of peristome. Above the tabula lateral lobes of peristome curve medially and abut, forming a suture as peristome lengthens into a tube.

Tentacles cream­colored, 14–16; lophophores bell­ to funnel­shaped, radially symmetrical between chimneys, obliquely truncate adjacent to chimneys.

Embryo color pale orange.

REMARKS: This tiny species is distinguished from other Celleporina species in the northeast Atlantic­Mediterranean region by its widely sinuate orifice, long tubular peristome and single, proximo­median peristomial avicularium.

OCCURRENCE: It was abundant on Cellaria fistulosa in the 35 m­deep meadow west of Banjole Island, together with Celleporina caminata .

MEASUREMENTS (SKELETAL): AAL 63 ± 9 µm, 51–82 (3, 11), AAW 45 ± 8, 34–65 (3, 10), DO 295 ± 47, 206–389 (3, 30), OL 89 ± 9, 79–101 (3, 5)m OW 93 ± 5, 86–101 (3, 6), OvL 136 ± 15, 116–167 (3, 18), OvW 215 ± 15, 194–242 (3, 18), SL 19 ± 2, 16– 22 (3, 7), SW 54 ± 3, 50–59 (3, 7), VAL 244 ± 24, 216–272 (2, 4), VAW 159 ± 14, 142–176 (2, 4). (POLYPIDE): IH 202 ± 27 µm, 160–240 (1, 10), LD 634 ± 74, 500–740 (1, 10), MD 30 (1, 2), TL 610 ± 50, 540–680 (1, 10).

Celleporina tubulosa ( Hincks, 1880) Figure 40E–G

Cellepora costazii var. tubulosa Hincks, 1880: 412 .

Celleporina hassallii var. tubulosa: Zabala and Maluquer, 1988: 159 .

Celleporina tubulosa: Hayward and Ryland, 1999: 330 .

DESCRIPTION (AMNH 995): Colony pisiform. Primary autozooidal orifice transversely oval with broad, shallow sinus occupying entire proximal width of anter. Tall, tubular peristome slightly flared apically, narrowly transversely oval at aperture; avicularia paired, situated laterally on peristome rim, with small oval rostra acute to orifice plane, directed laterally and with mandibles facing each other. Ovicell with a peripheral row of large pores, opening into peristome at about halfway along its length. Vicarious avicularia broadly spatulate, with an extensive palatal foramen and a very slender crossbar lacking a columella.

REMARKS: A single specimen of this species was found, but it is sufficiently distinct from other European species of Celleporina to be readily recognized. It is especially distinguished by its primary orifice.

DISTRIBUTION: This rare species has been reported reliably on just three previous occasions. The type locality is unknown and the type specimen lost, but Hayward and Ryland (1999) described and figured a specimen from the Scilly Isles, southwest England, which closely matched Hincks’ (1880) original description and figure. Zabala and Maluquer (1988) figured what is clearly the same species from the Catalan coast. The specimen listed by Gautier (1962) from Sicily was said to be identical in morphology and measurements to C. hassallii (Johnston) except for an elongate peristome; no mention was made of the distinctive primary orifice and the positioning of the peristomial avicularia, and it is thus not certain that the specimen belonged to C. tubulosa .

MEASUREMENTS (SKELETAL): AAL 78 ± 9 µm, 69–93 (1, 10), AAW 37 ± 3, 33–44 (1, 10), DO 354 ± 37, 301–405 (1, 10), OL 130 (1, 2), OW 103–109 (1, 2), AL 314–318 (1, 2), AW 238–258 (1, 2).

GENUS TURBICELLEPORA RYLAND, 1963

Turbicellepora avicularis ( Hincks, 1860) Figure 41A–D

Cellepora avicularis Hincks, 1860: 278, 1880:

406.

Turbicellepora avicularis: Hayward and Ryland,

1999: 336.

DESCRIPTION (AMNH 996–998; CMRR 2262): Colonies orange, with variable form, from low mounds and nodules to stout, irregularly lobed or erect forms with tapered branches; to 15 mm high in present material. Autozooids broad, convex, with boundaries visible only in few, frontally budded individuals; frontal shield smooth, with a single series of few, conspicuous marginal pores. Primary orifice slightly longer than wide, a deep, V­shaped proximal sinus comprising about one­quarter total length, flanked by thick, rounded condyles; no spines. Peristome low, absent midproximally, adjacent to lateral suboral avicularium with laterally directed, elongate triangular rostrum, hooked distally, oblique to orifice plane; crossbar stout, with distinct columella. Vicarious avicularia characteristic, to 0.6 mm long, broadly spatulate to more nearly parallel­sid­ ed, rounded distally; always with an extensive palate, and a thick crossbar with a stout columella often bifid distally. A second, less frequently occurring vicarious avicularium has a broadly triangular, distally hooked rostrum and thick crossbar with equally thick columella. Ovicell broader than long, smooth surfaced, with 10–15 small round pores.

Tentacles light orange, 16–18; lophophores bell­shaped, radially symmetrical midway between chimneys to obliquely truncate, supported by long introverts.

Embryo color bright, medium orange.

DISTRIBUTION: This is the most commonly occurring species of Turbicellepora in the northeast Atlantic region, distributed from northern Norway and southern Iceland southwards to the Mediterranean, where it is widespread in shallow coastal waters.

MEASUREMENTS (SKELETAL): AAL 140 ± 19 µm, 113–172 (3, 19), AAW 96 ± 13, 62– 120 (3, 19), DO 441 ± 102, 262–624 (3, 30), OL 132 ± 16, 110–167 (3, 26), OW 128 ± 10, 108–145 (3, 26), SL 43 ± 7, 26–54 (3, 26), SW 69 ± 16, 47–103 (3, 26), VAL 448 ± 40, 412–537 (3, 10), VAW 276 ± 26, 248–338 (3, 10). (POLYPIDE): IH 383 ± 51 µm, 320–460 (1, 10), LDMn 655 ± 74, 540–800 (1, 14), LDMx 695 ± 59, 600–800 (1, 17), MD 30 ± 3, 25–35 (1, 8), TLMn 554 ± 38, 500–620 (1, 10), TLMx 707 ± 140. 600–960 (1, 10).

Turbicellepora camera Hayward, 1978 Figure 41E–H

Turbicellepora camera Hayward 1978: 570 , figs. 4N–P, 5T, 10.

DESCRIPTION (AMNH 999; CMRR 2263): Colonies orange, initially small, domed nodules, some of the present material developing the slender, cylindrical, branching growth characteristic of the species. Autozooids oval, convex, with boundaries visible only in newly budded individuals; smoothly calcified frontal shield bordered by few, small marginal pores. Primary orifice slightly longer than broad, with a deep, V­shaped proximal sinus comprising almost half total length; condyles broad and thick, no spines. A low, thickened peristome developing in early ontogeny; lowest midproximally, adjacent to proximolateral suboral avicularium, with elliptical, round­ended rostrum, oblique to orifice plane and laterally directed. Vicarious avicularia characteristic: rostrum broadly spoon­shaped, 0.25–0.3 mm long, cupped distally; crossbar slender, with an indistinct median columella; completely lacking a palate and in cleaned material appearing as an open chamber. Additional adventitious avicularia sporadic, abundant in some areas of colony, identical to suboral type. Ovicell hemispherical, smooth, with few (<10) large, irregular pores.

REMARKS: The present material does not differ from the Aegean type specimen ( Hayward, 1978); the shape of the orifice, the distinctive vicarious avicularia and the proliferating adventitious avicularia together distinguish T. camera from all other Mediterranean species of Turbicellepora .

DISTRIBUTION: The species is apparently endemic to the Mediterranean and the adja­ cent Atlantic coast, described originally from the Aegean Sea ( Hayward, 1978) and subsequently reported from the Bay of Cadiz and the Strait of Gibraltar ( Harmelin and d’Hondt, 1992).

MEASUREMENTS (SKELETAL): AAL 155 ± 24 µm, 124–186 (2, 9), AAW 133 ± 22, 106–169 (2, 9), DO 389 ± 70, 252–503 (3, 29), OL 112 ± 8, 96–128 (3, 25), OW 119 ± 11, 101–136 (3, 25), SL 42 ± 5, 35–53 (2, 12), SW 88 ± 9, 74–105 (2, 12), VAL 456 ± 47, 394–518 (3, 7), VAW 320 ± 28, 267–394 (3, 7), ZL 483 ± 28, 446–512 (1, 5), ZW333 ± 28, 301–360 (1, 5).

FAMILY PHIDOLOPORIDAE GABB AND HORN, 1862

GENUS RETEPORELLA BUSK, 1884

Reteporella grimaldii ( Jullien, 1903) Figure 42A–F

Retepora grimaldii Jullien , in Jullien and Calvet, 1903: 62.

Sertella septentrionalis Harmer, 1933: 620 . Gautier, 1962: 233. Geraci, 1975: 244. Zabala, 1986: 544. Zabala and Maluquer, 1988: 154.

Reteporella septentrionalis: Hayward and Ryland, 1996: 109, 1999: 374 .

Reteporella grimaldii: Reverter­Gil and Fernández­Pulpeiro, 1999: 1416 .

DESCRIPTION (AMNH 1000–1002; CMRR 2264): Colonies orange, erect, brittle, consisting of bilaminar fenestrate plates, complexly infolded and fused into rigid three­dimensional structure; inwards­facing surfaces bearing autozooids, outer faces consisting of sheets of kenozooids. Colony size and architecture variable, probably dependent upon microhabitat; a single colony from locality 12 measuring 18 X 14 X 13 cm, but larger in situ specimens reported up to about 70 cm diameter and 35 cm thick perhaps consisted of several closely intergrown colonies. Fenestrulae elongate oval, commonly 1.5 X 0.5 mm; trabeculae of three to five alternating longitudinal autozooid series. Autozooids at growing edge elongate, rectangular, convex, with indistinct boundaries. Frontal shield smooth, with few, large marginal pores around proximal margins. Primary orifice broader than long; distal border finely denticulate, proximal border shallowly concave between indistinct condyles. Single pair of oral spines present, one on each side of orifice, persisting in late ontogeny. Peristome developing in early ontogeny, obscuring primary orifice; deep, slightly flared frontally, with conspicuous medio­proximal pore, linked to peristome rim by clear groove. Inner face of peristome coarsely denticulate, thickened spine bases prominent in each proximal corner. Avicularia adventitious. Frontal avicularia dimorphic: short, elliptical, 0.06 mm long, normal to frontal plane, with varying orientation; or larger, with narrowly triangular, distally hooked rostrum, ca. 0.3 mm long, orientated perpendicular to frontal plane and very conspicuous. Single large, transversely orientated avicularia in proximal, abfrontal axils of fenestrulae while short elliptical type are common across abfrontal surfaces. Ovicell slightly longer than wide, flattened frontally, with narrow fissure extending for about half its length; aperture overhung by short, straight­edged labellum.

Tentacles light orange, 12; lophophores distally flaring conical, radially symmetrical to slightly obliquely truncate, commonly overlapped with adjacent tentacles up to 60 µm.

Embryo color deep, bright reddish­orange.

REMARKS: Colonies are small and delicate in quiet environments such as within the Cellaria turf west of Banjole Island (fig. 42A), but overall colony size and branch width are much more robust in more exposed, less confined environments (fig. 42B).

DISTRIBUTION: Offshore, typically on hard substrata. This species is common in the northeast Atlantic region, from the White Sea, the Kara Sea, Iceland, and western Greenland south to the Mediterranean, where it seems to occur in shallower habitats than to the north. Hayward and Ryland (1996) could not distinguish Mediterranean from North Atlantic specimens.

MEASUREMENTS (SKELETAL): AAL(large) 324 ± 77 µm, 192–494 (3, 30), AAW(large) 202 ± 45, 133–297 (3, 26), AL(fenestrulae) 248 ± 23, 331–386 (1, 10), DO 273 ± 41, 199–359 (4, 40), OW 97 ± 11, 79–122 (3, 30), OvL 193 ± 25, 149–248 (3, 24), OvW 173 ± 16, 143–208 (3, 24), ZL 353 ± 17, 331–386 (1, 10), ZW 187 ± 13, 160–202 (1, 10). (POLYPIDE): IH ~0 µm, LD 450 ± 35, 380–500 (1, 23), TLMn 324 ± 22, 300–360 (1, 5), TLMx 447 ± 25, 400–520 (1, 23).

GENUS RHYNCHOZOON HINCKS, 1895

Rhynchozoon neapolitanum Gautier, 1962

Figure 43A–E

Rhynchozoon neapolitanum Gautier, 1962: 243 .

Zabala, 1986: 531. Zabala and Maluquer, 1988:

149.

DESCRIPTION (AMNH 968, 1003; CMRR 2265): Colonies cream­colored to tan, encrusting, multilaminar; developing frontally budded, massive nodular forms, frequently exceeding 3 X 3 X 3 cm. Autozooids with vitreous, nodular frontal shield bordered by a single series of conspicuous marginal pores. Primary orifice as wide as long; distal and lateral rim with large, rounded, closely spaced denticulations; proximal border with short, narrow U­shaped sinus accentuated by stout, faceted condyles. No oral spines. Peristome thick, deepening through ontogeny; distal and lateral border developing short knobs; proximal border characteristically with a pair of stout columnar processes flanking a deep median embayment; at base of inner surface of one process, a thick angular uncinate lateral process projects into the lumen of the peristome. Median suboral avicularium present in some autozooids with uncinate process at its proximal end; rostrum broadly triangular, hooked distally, acute to plane of orifice and facing laterally. Frontal avicularia often frequent in later ontogeny; characteristic: with voluminous cystid occupying more than half frontal surface of bearing autozooid, rostrum elongate triangular, acute to frontal plane, directed proximally or proximolaterally. Ovicell short and broad; flat frontally, with a transversely oval area of membranous ectooecium; aperture straight edged, with a short notch in each corner.

Tentacles cream­colored, 13–15; lophophores distally flared conical, radially symmetrical to obliquely truncate.

Embryo color pale orange.

REMARKS: The largest colonies often incorporate tubular channels opening over the surface, suggesting they had grown in association with hydroids (as in Rhynchozoon larreyi Auct. ; see Ristedt and Schuhmacher, 1985).

DISTRIBUTION: This common and distinctive species is widely distributed throughout the Mediterranean.

MEASUREMENTS (SKELETAL): DO 447 ± 79 µm, 301–635 (2, 20), OL 106 ± 6, 93–117 (2, 20), OW 125 ± 4, 117–131 (2, 20), OvL 196 ± 16, 173–213 (2, 20), OvW 287 ± 20, 258–311 (2, 20), SL 14 ± 3, 11–20 (2, 20), SW 25 ± 5, 19–36 (2, 20), ZL 614 ± 54, 513–733 (2, 20), ZW 388 ± 50, 292–462 (2, 20). (POLYPIDE) IH 69 ± 42 µm, 1–120 (1, 10), LD 709 ± 61, 600–800 (1, 10), MD 30 ± 4, 25–35 (1, 4), TL 663 ± 61, 560–780 (1, 10).

Rhynchozoon pseudodigitatum Zabala and Maluquer, 1988 Figure 43F–H

Figures

Fig. 40. A–D. Celleporina siphuncula. A. General aspect (paratype, AMNH 990; 0.5 mm). B. Autozooids and maternal zooids with ovicells (holotype, AMNH 989; 0.1 mm). C. Vicarious avicularia (holotype, AMNH 989; 0.1 mm). D. Autozooidal orifice (holotype, AMNH 989). E–G. Celleporina tubulosa (AMNH 995). E. General aspect (0.5 mm). F. Maternal zooids with ovicells and (top right) vicarious avicularium (0.1 mm). G. Autozooidal orifice (0.05 mm).

Fig. 41. A–D. Turbicellepora avicularis. A. general aspect (AMNH 996; 0.5 mm). B. autozooid (top left), maternal zooid with ovicell (top center) and (bottom center) vicarious avicularium (AMNH 997; 0.1 mm). C. reparative budding of autozooid within vicarious avicularium (AMNH 998; 0.1 mm). D. autozooidal orifice (AMNH 996; 0.05 mm). E–H. Turbicellepora camera (AMNH 999). E. general aspect (0.5 mm). F. autozooids and (top center) vicarious avicularium (0.1 mm). G. ontogenetically old autozooid with multiple vicarious avicularia (0.1 mm). H. autozooidal aperture (0.05 mm).

Fig. 42. A–F. Reteporella grimaldii. A. Delicate colony from within Cellaria meadow (AMNH 1000; 1 mm). B. Robust colony from more exposed locality (AMNH 1001; 1 mm). C. Maternal zooids with partially immersed ovicells (AMNH 1000; 0.1 mm). D. Oblique view of colony surface with autozooids and dimorphic adventitious avicularia (AMNH 1002; 0.1 mm). E. Autozooids (right) and maternal zooids with incompletely formed ovicells (AMNH 1002; 0.1 mm). F. Reverse surface with kenozooids and dimorphic avicularia (AMNH 1001; 0.2 mm).

Fig. 43. A–E. Rhynchozoon neapolitanum. A. General aspect of frontally budded region (AMNH 1003; 0.5 mm). B. Colony margin with laterally budded zooids (AMNH 968; 0.5 mm). C. Ontogenetically young (top left) and (center) old autozooids (AMNH 1003; 0.5 mm). D. Ovicell (AMNH 1003; 0.05 mm). E. Autozooidal orifice (AMNH 1003; 0.05 mm). F–H. Rhynchozoon pseudodigitatum (AMNH 1004). F. General aspect (0.5 mm). G. Autozooids along colony margin (0.1 mm). H. Autozooidal orifice with orificial avicularium (0.05 mm).