Smittia rouvillei Calvet, 1902: 57

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 : 48-53

Smittia rouvillei Calvet, 1902: 57 . Parasmittina rouvillei: Gautier, 1962: 199 . Parasmittina tropica: Hayward, 1974: 384 .

DESCRIPTION (AMNH 938, 939; CMRR 2230): Colony light yellow to orange, encrusting, multilaminar. Autozooids hexagonal to polygonal, in orderly arrangement at unilaminar growing edges, without common orientation in successive laminae; convex, separated by distinct ridges. Primary orifice about as wide as long; proximal border with short, anvil­shaped lyrula occupying about half its width; condyles conspicuous, round­ ed, downcurved, with denticulate, laminate structure, revealed by SEM, visible as a faint burr with light microscopy. Two or three distal oral spines present. Peristome developed proximal to spine bases, later extending over them, forming low, flared rim, produced midproximally resulting in pear­shaped secondary aperture. Frontal shield nodular, with single or double series of large marginal pores; in later ontogeny ridges of calcification extending between pores onto center of shield, the periphery then appearing reticulate. Avicularia adventitious, polymorphic, often abundant: small, triangular or parallel­sided avicularia proximal or proximolateral to orifice, directed proximally, triangular type potentially present elsewhere around peristome, often directed towards aperture; large avicularia, with distally rounded, parallel­sided rostrum half as long as autozooid frequent, budded proximolateral to orifice and directed proximally. Ovicell slightly longer than broad, globular, with a dense group of variably sized pores distally, the proximal frontal surface imperforate.

Tentacles light yellow, 12–15; lophophores bell­shaped, radially symmetrical between chimneys to obliquely truncate bordering chimneys.

REMARKS: The Mediterranean species of Parasmittina are in need of thorough taxonomic revision, including a review of type and other published materials, and this present attribution may have to be reconsidered. However, the material described here is the same species as that described from Chios ( Hayward, 1974) as P. tropica ( Waters, 1909) , a taxon then unrecorded for the Mediterranean.

It is almost certainly the species described by Gautier (1962) as P. rouvillei and is probably the same as that described by Canu and Bassler (1930) as Smittina rouvillei . Zabala and Maluquer (1988) considered that these and other citations referred to a single, variable species which they denoted P. tropica , referring to Harmer’s (1957) redefinition. However, it must be pointed out that Smittia tropica Waters, 1909 was described from the Sudanese Red Sea coast. Harmer’s (1957) concept of the species was based on specimens from Ghardaqa, India, Ceylon, the Torres Straits, Victoria, Adelaide, and the Siboga collections from the Indo­Malayan region, and it is clear that he did not reexamine Waters’ original material. Harmer’s (1957) illustrations of ’’ Smittina tropica ’’ depict four specimens from the Siboga collections, one from India and one from Victoria. It is doubtful that any of these corresponds to Waters’ species, or to any species present in the Mediterranean.

MEASUREMENTS (SKELETAL): AL (large) 290 ± 60, 175–398 (2, 12), AW (large) 104 ± 27, 65–154 (2, 12), AL (small) 109 ± 7, 101–122 (1, 8), AW (small) 73 ± 15, 49–93 (1, 8), DO 523 ± 74, 386–676 (2, 20), LW 52 ± 5, 47–62 (1, 9), OL 120 ± 14, 92–143 (2, 21), OW 119 ± 16, 81–143 (2, 21), ZL 687 ± 69, 566–816 (2, 20), ZW 411 ± 58, 323–569 (2, 20). (POLYPIDE): IH 120 ± 51 µm, 40–180 (1, 10), LD 640 ± 78, 560–800 (1, 10), MD 30–40 (1, 2), TL 646 ± 123, 440–820 (1, 10).

GENUS SMITTINA NORMAN, 1903

Smittina cervicornis ( Pallas, 1766) Figure 22A–C

Millepora cervicornis Pallas, 1766: 252 .

Eschara cervicornis: Milne Edwards, 1836: 19 .

Smittia cervicornis: Calvet in Jullien and Calvet, 1903: 151.

Porella cervicornis: Canu and Bassler, 1930: 54 . Lagaaij, 1952: 97. Gautier, 1962: 204.

Smittina cervicornis: Cook, 1968: 210 . Zabala and Maluquer, 1988: 121.

DESCRIPTION (AMNH 940; CMRR 2231): Colonies orange, erect, branching, to 30 mm in present material, narrow: maximum branch width 3 mm. Branches bilaminar, flatsectioned, twisting around long axis; each face of five or six alternating autozooid rows. Autozooids at growing tips of colony elongate, rectangular, convex, with distinct boundaries; commonly 0.625–0.75 x 0.375 mm. Primary orifice about as wide as long; proximal border straight, a deep lyrula, with straight edge and rounded corners, occupying nine­tenths of proximal width; condyles rounded quadrangular, conspicuous in early ontogeny. No spines. Frontal shield thickening steadily from earliest ontogeny, evenly pierced by large round pores. Peristome developing within one ontogenetic generation from growing edge, thickly calcified, cylindrical and projecting from colony surface, quite hiding primary orifice. Proximally it in­ corporates a prominent avicularium, with rounded rostrum, complete crossbar bearing a ligula, and supporting a semicircular mandible, proximally directed and normal to frontal plane. Ovicell not seen in present material; hyperstomial, hemispherical, perforate; obscured by peristome and completely immersed. In late ontogeny, frontal shield thickened to extent that frontal surface of colony becomes quite flat, peristomes and avicularia are immersed, and peristomial rims form circular secondary orifices; autozooids then have a broadly lingulate outline, each bordered by a distinct suture.

Tentacles light orange or yellowish orange, 14 –16; lophophores bell­shaped, radially symmetrical across most of branch surface, obliquely truncate adjacent to elongate chimney along branch margin.

DISTRIBUTION: Common throughout the Mediterranean in shallow coastal waters, ranging south to the Cape Verde Islands and Ghana.

MEASUREMENTS (SKELETAL): AL 111 ± 12 µm, 95–130 (2, 17), AW 101 ± 23, 72–159 (2, 17), DO 543 ± 67, 423–669 (2, 20), OL 164 ± 10, 149–181 (2, 17), OW 134 ± 9, 121–150 (2, 17), ZL 872 ± 54, 811–977 (2, 20), ZW452 ± 51, 386–543 (2, 20). (POL­ YPIDE): IH 158 ± 70 µm, 40–280 (1, 23), LDMn 556 ± 83, 460–620 (1, 3), LDMx 610 ± 79, 400–780 (1, 28), TLMn 462 ± 76, 340–560 (1, 16), TLMx 588 ± 114, 340– 820 (2, 23).

GENUS PRENANTIA GAUTIER, 1962

Prenantia cheilostoma ( Manzoni, 1869b) Figure 22D–H

Lepralia cheilostomata Manzoni, 1869b: 942 .

Smittia cheilostoma: Hincks, 1880: 349 .

Smittina cheilostoma: Gautier, 1962: 192 . Hayward and Ryland, 1999: 258.

Prenantia cheilostoma: Gautier, 1962: 193 .

DESCRIPTION (AMNH 932, 941, 942; CMRR 2232): Colonies orange, encrusting, multiserial, unilaminar sheets, light reddish purple when living. Autozooids hexagonal, gently convex in early ontogeny, later rather flat following ontogenetic thickening of the frontal shield; separated by shallow grooves. Primary orifice wider than long, proximal border with an anvil­shaped lyrula occupying about one­third total width; condyles small, rounded, indistinct. No oral spines. Peristome developing from earliest ontogeny, forming a low thickened rim completely encircling orifice, mid­proximal portion deeply U­shaped and channeled on its inner face. Frontal shield thick, vitreous, densely perforated with numerous small pores. Ancestrula with acute proximal end, spiramen, and circular orifice slightly smaller than orifice of autozooids and encircled by approximately 12 oral spines; all but apertural peristome covered when ancestrula is completely encircled by ring of autozooids. Ovicell wider than long, smooth surfaced, with many small frontal pores; developing a granular, peripheral ooecial cover continuous with the peristome.

REMARKS: Hincks (1880: 349) used the trivial name cheilostoma , which Manzoni, (1869b, p. 942) had originally spelled cheilostomata in the text when describing the new species, although the name as given by Manzoni on the plate (1869b: pl. 4, fig. 22) was cheilostoma . Hincks (1880) did not mention both spellings, and therefore his use of cheilostoma does not satisfy Article 24.2.3 of the ICZN Code for selection of one from among two or more spellings in an original work, but cheilostoma has become established in the literature to the virtual exclusion of cheilostomata for the species, which allows its preservation as ‘‘established usage’’ under Article 33.3.1 ( Ride et al., 1999).

DISTRIBUTION: Shallow sublittoral and offshore, often encrusting shell. Common throughout the Mediterranean, ranging northwards to the western English Channel and the Isle of Man.

MEASUREMENTS (skeletal): DO 474 ± 66 µm, 370–627 (2, 20), LW 62 ± 7, 54–81 (2, 20), OL 132 ± 19, 101–169 (2, 20), OW 142 ± 14, 120–171 (2, 20). OvL 214 ± 26, 178– 268 (1, 10), OvW 295 ± 20, 260–333 (1, 10), ZL 596 ± 49, 519–709 (2, 20), ZW 375 ± 30, 314–414 (2, 20).

GENUS PHOCEANA JULLIEN AND CALVET, 1903

Phoceana tubulifera ( Heller, 1867) Figure 23A–D

Eschara tubulifera Heller, 1867: 116 .

Smittina tubulifera: Harmelin, 1969b: 299 . Hay­

ward, 1974: 382.

DESCRIPTION (AMNH 943; CMRR 2233): Colonies light cream­tinted orange, erect, slender, branching dichotomously, the branches oval­sectioned, to 2 mm wide in present material. Each branch face with five alternating, longitudinal series of autozooids, broadening to seven prior to a dichotomy. Autozooids elongate, rectangular, separated by distinct grooves; frontal shield gently convex, finely granular, with few evenly scattered pores, each in a shallow pit. Primary orifice slightly longer than wide; a short, straight­edged lyrula, with cusped corners, occupying most of proximal edge. No spines. A deep tubular peristome developing in early ontogeny, curving frontally and projecting from frontal plane, completely hiding primary orifice. Secondary orifice more or less circular, with a thickened, slightly crenulate rim; a short, pointed tooth apparent midproximally within rim marks edge of a ridge descending inner, proximal face of peristome. No ovicells in present material.

DISTRIBUTION: This delicate species was described from the Adriatic ( Heller, 1867) and has been recorded on very few occasions since. It was reported from the coast of Tunisia by Canu and Bassler (1930), and has been described and illustrated from the Aegean on two occasions. It is perhaps restrict­ ed to the eastern Mediterranean; neither Gautier (1962) nor Zabala and Maluquer (1988) recorded it from the western Mediterranean.

MEASUREMENTS (SKELETAL): DO 513 ± 64 µm, 430–689 (2, 20), OL 148 ± 12, 118– 166 (2, 20), OW 149 ± 16, 128–190 (2, 20), ZL 784 ± 47, 725–892 (2. 20), ZW 357 ± 31, 314–438 (2, 20).

GENUS SMITTOIDEA OSBURN, 1952

Smittoidea reticulata

(J. Macgillivray, 1842)

Figure 23E–H

Lepralia reticulata J. Macgillivray, 1842: 467 .

Smittoidea reticulata: Hayward and Ryland, 1999: 264 .

DESCRIPTION (AMNH 913, 944): Colonies encrusting, multiserial, unilaminar sheets. Autozooids oval to hexagonal, convex, separated by distinct grooves; 0.35–0.5 X 0.25– 0.3 mm in present material. Primary orifice wider than long, distal rim with fine dentic­ ulation revealed by SEM, proximal edge with anvil­shaped lyrula occupying about half total width, condyles conspicuous, angular, downcurved. Two to four, most frequently three, distal oral spines. A single median suboral avicularium, with acute triangular rostrum, normal to frontal plane, proximally directed; crossbar slender, complete, palate with extensive foramen. Frontal shield nodular, bordered by a single series of large, distinct pores. Ovicell slightly longer than broad, aperture arched, with straight distal edge; about 20 small pores on distal and lateral surfaces. Ancestrula elongate oval, smoothly convex, tapered proximally; gymnocyst smooth; membranous frontal wall occupying half total length, surrounded by oval, crenulate rim bordered by four closely spaced distal spines and five spines uniformly spaced proximally; cryptocyst broadest proximally, opesia almost circular, two­thirds length of membrane.

REMARKS: The material described and illustrated here has a proportionately broader lyrula and longer avicularium than north European material figured by Hayward and Ryland (1999), but consists of small colonies only, in which ontogenetic variation may not be fully expressed.

DISTRIBUTION: This species is distributed in the eastern Atlantic from Arctic Norway to Morocco, and throughout the Mediterranean.

MEASUREMENTS (SKELETAL): AL 102 ± 14 µm, 70–128 (2, 20), AW 55 ± 10, 41–74 (2, 20), DO 349 ± 46, 276–449 (2, 20), OL 93 ± 11, 73–111 (3, 26), OW 106 ± 15, 79– 141 (3, 26), OvL 224 ± 26, 192–273 (1, 10), OvW 209 ± 17, 183–230 (1, 10), ZL 506 ± 40, 443–618 (2, 20), ZW 288 ± 34, 223–348 (2, 20).

FAMILY BITECTIPORIDAE MACGILLIVRAY, 1895

GENUS HIPPOPORINA NEVIANI, 1895

Hippoporina lineolifera ( Hincks, 1886) Figure 24

Schizoporella lineolifera Hincks, 1886: 267, 1887: 315 .

? Schizoporella marsupifera: Gautier, 1962: 144 .

LECTOTYPE (chosen here): BMNH reg. no. 1899.5.1.1072, Hincks collection, Adriatic; The Natural History Museum, London.

DESCRIPTION: Lectotype consisting of two fragments from single colony; the larger, mounted in wooden cavity slide, totaling 24 autozooids and an ancestrula; the smaller, Canada balsam mount of seven autozooids, prepared in 1929; all zooids damaged to some degree. Autozooids oval, convex, with distinct boundaries; frontal shield densely perforated by large round pores with thickened rims. Primary orifice slightly wider than long, proximal border shallowly concave between small condyles. No oral spines; proximal rim of orifice thickened, with median suboral umbo bearing small avicularium with semielliptical, proximally directed mandible. Ovicell, damaged in all examples, globose, recumbent upon distally succeeding zooid, with frontal pores evident in undamaged areas. Oval ancestrula with raised mural rim bordering membrane and indeterminate number of spines; broad, flat cryptocyst defining semielliptical opesia. Substratum apparently an alga frond.

REMARKS: We did not encounter this species; the lectotype is the only extant specimen. It is labeled in Hincks’ hand with its original taxonomic designation, and also ‘‘ = S. marsupifera Busk’’. The curious synonymy with the latter, described from a Challenger station off Marion Island, South Indian Ocean, may thus have originated with Hincks. Gautier (1962) had a single specimen, which he lost; he then repeated Hincks’ description of the species, attributing it to Busk’s taxon.

MEASUREMENTS (SKELETAL): OL 110 ± 10 µm (1, 10), OW 130 ± 8 (1, 10), ZL 470 ± 50 (1, 10), ZW 310 ± 40 (1, 10).

GENUS PENTAPORA FISCHER, 1807

Pentapora fascialis ( Pallas, 1766) Figure 25A–E

Figures

Fig. 22. A–C. Smittina cervicornis (AMNH 940). A. General aspect (0.5 mm). B. Autozooid (0.1 mm). C. Autozooidal orifice (0.05 mm). D–H. Prenantia cheilostoma.D. General aspect (AMNH 941; 0.5 mm). E. Maternal zooid with ovicell (AMNH 941; 0.1 mm). F. Autozooidal orifice (AMNH 942; 0.05 mm). G. Ancestrula (AMNH 932; 0.1 mm). H. Early autozooids surrounding orifice (center) of ancestrula covered by autozooids (AMNH 941; 0.1 mm).

Fig. 23. A–D. Phoceana tubulifera (AMNH 943). A. General aspect (0.5 mm). B. Autozooid (0.1 mm). C. Autozooidal primary orifice (0.05 mm). D. Oblique view of autozooidal secondary orifice with internal proximal ridge (0.05 mm). E–H. Smittoidea reticulata. E. General aspect (AMNH 944; 0.5 mm). F. Maternal zooid with ovicell (left) and (right) autozooid with orificial spines in place (AMNH 944). G. Autozooid with suboral avicularium (AMNH 944; 0.1 mm). H. Ancestrula (AMNH 913; 0.1 mm).

Fig. 24. Hippoporina lineolifera. Autozooid and maternal zooids with ovicells (left) and (right) ancestrula with early autozooids. (lectotype, BMNH reg. no. 1899.5.1.1072; 0.2 mm).

Fig. 25. A–E. Pentapora fascialis. A. General aspect (AMNH 945; 0.5 mm). B. Autozooid (AMNH 946; 0.1 mm). C. Maternal zooid with partially calcified ovicell and small suboral avicularium (AMNH 945; 0.1 mm). D. Autozooid with large suboral avicularium (AMNH 947; 0.1 mm). E. straight colony margin generated by kenozooids (AMNH 945; 0.5 mm). F–H. Schizomavella asymetrica (AMNH 948). F. General aspect (0.5 mm). G. Autozooid (right) and (center) maternal zooid with ovicell (0.1 mm). H. Autozooidal orifice (0.05 mm).