Cellaria salicornioides Lamouroux, 1816: 127

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 : 36-44

Cellaria salicornioides Lamouroux, 1816: 127 . Cellaria salicornioides: Hayward and Ryland,

1998: 308.

DESCRIPTION (AMNH 923, 924; CMRR 2221): Colonies erect and branching, with slender, cylindrical internodes consisting of alternating whorls of two to five autozooids. Colonies, branches, and internodes may be as long as those of C. fistulosa , but tend to be far more diffuse and delicate, so that the two species are readily separated by the unaided eye. Autozooids oval to hexagonal or almost rectangular, with regular quincuncial arrangement, so that commonly none is in contact with those autozooids immediately distal and proximal to it. Opesia in distal third of zooid, semicircular, with crenulate distal rim and convex, slightly projecting proximal rim; short, bluntly tapered denticles just visible in proximal corners. Cryptocyst deeply concave, with well­marked longitudinal ridges especially prominent lateral to the opesia, grading into the autozooid boundary distally, but converging proximally to define a complete rim distinct from the zooid margin. Avicularia sporadic, distinctive: as large as an autozooid, with semicircular rostrum projecting at acute angle from frontal plane, supporting a conspicuous, brown mandible. Ovicell aperture a simple round opening distal to the opesia.

DISTRIBUTION: In shallow coastal waters; widespread throughout the Mediterranean, ranging northwards along the western coasts of Britain and Ireland, to Shetland.

MEASUREMENTS (SKELETAL): DO 306 ± 47 µm, 224–399 (2, 20), OpL 59 ± 4, 53–68 (2, 20), OpW 81 ± 7, 71–99 (2, 20), ZL 439 ± 33, 383–532 (2, 20), ZW 213 ± 17, 175– 240 (2, 20).

SUBORDER ASCOPHORA LEVINSEN, 1909 INFRAORDER ACANTHOSTEGA LEVINSEN, 1902 SUPERFAMILY CRIBRILINOIDEA HINCKS, 1879 FAMILY CRIBRILINIDAE HINCKS, 1879 GENUS COLLARINA JULLIEN, 1886

Collarina balzaci ( Audouin, 1826) Figure 16A–C

Flustra balzaci Audouin, 1826: 239 .

Cribrilina punctata: Hayward and Ryland, 1979: 56 .

Collarina balzaci: Zabala, 1986: 359 . Zabala and Maluquer, 1988: 105. Bishop, 1988: 749. Hayward and Ryland, 1998: 318.

DESCRIPTION (AMNH 912, 925, 926; CMRR 2222): Colonies encrusting, multiserial, unilaminar, small, irregular patches. Autozooids small, oval, with indistinct boundaries; gymnocyst rarely visible frontally. Frontal shield of four to seven pairs of costae; each costa with a prominent tubae­ form pelmatidium proximally, together forming a raised peripheral ring; two or three smaller pelmatidia along median fusion of costae. Intercostal pores large, typically one to three between each adjacent pair of costae. Apertural bar thickened, raised and mucronate. Orifice semicircular, wider than long; four oral spines, proximal pair elongate and curving over ovicell in fertile autozooids. Avicularia adventitious, single or paired, on each side of apertural bar, with short, triangular rostrum laterally directed. Ancestrula small, lacking adventitious avicularia, with fewer pairs of costae than subsequently budded autozooids. Ovicell recumbent on substratum, variably shorter or longer than wide, domed, with up to 20 tubaeform pores regularly distributed over its surface; a small avicularium at its distal edge, identical to the others, distally directed. Both autozooids and ovicell with large and conspicuous basal pore chambers.

REMARKS: The taxonomic status of this species was finally stabilized by Bishop (1988) through the selection of a neotype specimen.

DISTRIBUTION: The species is widely distributed throughout the Mediterranean, on stone, shell, red seaweeds and Posidonia , and ranges northwards to western Britain and the Faroe Isles.

MEASUREMENTS (SKELETAL): Al 79 ± 6 µm, 70–91 (1, 10), AW 53 ± 5, 47–61 (1, 10), DO 401 ± 61, 312–548 (4, 40), OL 59 ± 11, 38–80 (4, 32), OW 110 ± 14, 77–134 (4, 33), OvL 244 ± 27, 189–290 (3, 15), OvW 230 ± 12, 210–245 (3, 15), ZL 516 ± 51, 484–668 (4, 40), ZW 318 ± 32, 333– 436 (4, 40).

GENUS FIGULARIA JULLIEN, 1886

Lepralia figularis Johnston, 1847: 314 .

Figularia figularis: Hayward and Ryland, 1998:

338.

DESCRIPTION (AMNH 925): Colonies encrusting, multiserial, unilaminar. Autozooids rectangular to irregularly polygonal, convex, separated by deep grooves. Primary orifice about as wide as long, proximal border shallowly concave and frontally arched, between small, rounded condyles. No oral spines. Frontal surface a costate shield, comprising two­thirds total autozooid length, with a broad area of proximal gymnocyst extending distally as a narrower band on each side. Shield formed from 10–12 short costae, fused medially and at intervals laterally, leaving small, slitlike intercostal pores; each costa, additionally, with a single thick rimmed pelmatidium at its broad proximal (peripheral) end. Large vicarious avicularia, with spoonshaped mandible, and globular ovicells with a pair of large lateral fenestrae characteristic of this species, but neither was present in the small colony collected.

DISTRIBUTION: Figularia figularis is common in shallow coastal habitats throughout the Mediterranean, encrusting shell and other hard substrata. It ranges north to the southwest British Isles.

MEASUREMENTS (SKELETAL): DO 624 ± 61 µm, 530–730 (1, 10), OL 185 ± 14, 166– 210 (1, 10), OW 217 ± 19, 180–240 (1, 10), ZL 822 ± 64, 710–910 (1, 10), ZW 477 ± 51, 410–546 (1, 10).

GENUS PUELLINA JULLIEN, 1886

Puellina hincksi ( Friedl, 1917)

Figure 17A–C Cribrilina radiata var. hincksi Friedl, 1917: 236 . Cribrilaria hincksii (sic): Gautier, 1962: 111. Puellina (Cribrilaria) hincksi: Zabala and Malu­

quer, 1988: 107. Alvarez, 1994b: 134.

DESCRIPTION (AMNH 901, 927; CMRR 2223): Colonies encrusting, multiserial, unilaminar, small rounded patches. Autozooids oval to irregularly hexagonal, separated by deep grooves; gymnocyst clearly visible, smooth, imperforate; frontal shield of usually six or seven pairs of costae, each broadest and slightly umbonate proximally, the umbo bearing a minute pelmatidium; intercostal pores large and distinct, peripheral papilla pores the largest. Suboral lacuna transversely oval, large and conspicuous; apertural bar slightly thickened, first pair of costae often with a prominent umbo proximal to suboral lacuna. Primary orifice wider than long, bordered by five evenly spaced spines. Avicularium with small cystid wedged between autozooids; rostrum elongate, narrowly triangular, up to three­quarters length of auto­ zooids. Ovicell smooth, domed, with indistinct longitudinal ridge. Kenozooids sporadic.

REMARKS: First described from the Adriatic, this species was for long confounded with P. radiata ( Moll, 1803) and P. innominata ( Couch, 1844) , but its specific identity was finally confirmed by Gautier (1962), who reported it from the Marseilles region.

DISTRIBUTION: Puellina hincksi was described from Tunisia by Canu and Bassler (1930) and has recently been described and figured from west of Gibraltar ( Alvarez, 1994b). It is probably widely distributed within the Mediterranean.

MEASUREMENTS (SKELETAL): AL 323 ± 20 µm, 311–346 (1, 3), AW 153 ± 20, 130–168 (1, 3), DO 317 ± 39, 253–398 (2, 20), OL 47 ± 2, 41–50 (2, 17), OW 74 ± 3, 67–79 (2, 17), OvL 193 ± 14, 170–215 (1, 10), OvW 175 ± 14, 158–201 (1, 10), ZL 431 ± 49, 350–523 (2, 20), ZW 278 ± 35, 218– 337 (2, 20). (POLYPIDE): IH 0 µm (1, 1), LD 560–700 (1, 2), MD 20 (1, 1), TL 480–500 (1, 2).

SUPERFAMILY PATENICELLOIDEA BUSK, 1852 FAMILY SAVIGNYELLIDAE LEVINSEN, 1909 GENUS SAVIGNYELLA LEVINSEN, 1909

Savignyella lafontii ( Audouin, 1826) Figure 17D–F

Eucratea lafontii Audouin, 1826: 242 . Savigny, 1809: pl.13, figs. 21–25.

Savignyella lafontii: Gautier, 1962: 102 . Zabala and Maluquer, 1988: 145.

DESCRIPTION (AMNH 928; CMRR 2224): Colonies brown, erect, bushy tufts to 10 mm high in present material, of branching, uniserial chains of clavate autozooids. Autozooids 1–2 mm long, the proximal portion cylindrical, 0.08 mm wide, broadening to 0.12 mm at one­half to two­thirds total length; remaining distal portion 0.25 mm wide. Primary orifice terminal, D­shaped, with straight proximal edge, and seven short spines equally spaced around lateral and distal margins. A prominent median suboral avicularium partly obscuring orifice, its cylindrical cystid perpendicular to frontal plane; rostrum triangular, sharply hooked, orientat­ ed proximodistally and frontally directed. Frontal shield of broad distal portion of autozooid closely perforated by numerous round pores; a few smaller pores in single linear series on tubular proximal portion. Ovicell borne on distal rim of orifice, globular; ectooecium membranous except for a smooth, calcified disto­basal rim, entooecium calcified, with a coarsely reticulate surface. Each autozooid develops paired disto­basal buds from conspicuous tubular projections; new autozooids develop distal to a distinct chitinous joint. An additional septulum situated medially between the two daughter autozooids giving rise to delicate, cylindrical rhizoid.

Tentacles light brown, 18–22; lophophores bell­shaped, radially symmetrical, introverts short.

DISTRIBUTION: Savignyella lafontii has been reported on numerous occasions from the Adriatic, and its wider Distribution in the western Mediterranean was detailed by Gautier (1962). It has a practically circumglobal Distribution in warm temperate to tropical waters.

MEASUREMENTS (POLYPIDE): IH 96 ± 22 µm, 60–120 (1, 9), LD 772 ± 96, 650–960 (1, 11), MD 30–40 (1, 2), TL 741 ± 135, 440–920 (1, 9).

INFRAORDER HIPPOTHOOMORPHA GORDON, 1989

SUPERFAMILY HIPPOTHOOIDEA BUSK, 1859

FAMILY HIPPOTHOIDAE BUSK, 1859

GENUS HIPPOTHOA LAMOUROUX, 1821

Hippothoa divaricata Lamouroux, 1821 Figure 18A–E

Hippothoa divaricata Lamouroux, 1821: 82 . Ryland and Gordon, 1977: 19. Hayward and Ryland, 1999: 86.

DESCRIPTION (AMNH 929): Colony encrusting. Autozooids in branching, uniserial chains, diffuse or closely intergrown; clavate, with tubular proximal cauda and oval distal portion. Primary orifice longer than wide, poster comprising one­fifth total length, forming a short triangular sinus between distinct angular condyles. No spines. Frontal shield smoothly calcified, imperforate, with pronounced longitudinal median ridge, most prominent just proximal to the sinus. Dimor­ phic female zooids with slightly shorter cauda than autozooid, and broader orifice; ovicell hemispherical, imperforate, often with an umbo. Zooeciules of unknown function sporadic, with narrow, tubular cystid and minute elliptical orifice. Each autozooid producing a single distal bud; branching achieved by lateral budding, a maximum of two on each side, giving an X pattern when all four develop. Female zooids and zooeciules may arise at any of the five budding positions. Ancestrula oval, domed, with a complete gymnocystal frontal shield, and a small Dshaped orifice.

DISTRIBUTION: Hippothoa divaricata is widely distributed throughout the northeast Atlantic and the Mediterranean, on stones and shells, in shallow coastal habitats.

MEASUREMENTS (SKELETAL): CL 208 ± 33 µm, 163–256 (1, 7), OL 63 ± 4, 60–67 (1, 3), OW 52 ± 4, 49–56 (1, 3), ZL 503 ± 48, 421–577 (1, 7), ZW 169 ± 10, 151–183 (1, 7).

Hippothoa flagellum Manzoni, 1870 Figure 18F–I

Hippothoa flagellum Manzoni, 1870: 328 . Hayward and Ryland, 1999: 88.

DESCRIPTION (AMNH 929–931; CMRR 2225): Colonies uniserial traceries encrusting shell and other organic carbonates, form and extent dictated by space available and competition from other sessile organisms. Autozooids slender, clavate, with filiform proximal portion, or cauda, constituting up to three­quarters total length, and distal oval portion the rest. Primary orifice pear­shaped, broadest in its distal third, narrowest at proximal rim, which forms a deep, U­shaped sinus between short, pointed condyles. Branching a right­angled cross, each autozooid producing a single distal bud and paired lateral buds; female brooding zooids dimorphic, developed at lateral budding sites and rarely giving rise to further buds; broadly oval, abruptly tapered proximally, without a cauda, entire length slightly less than that of distal clavate portion of autozooid; orifice almost twice as wide as long, with broad, shallow sinus; ovicell imperforate. Ancestrula a circular, domed, kenozooidal structure, 0.3 mm diameter, budding a single oval autozooid, lacking the distinctive cauda.

DISTRIBUTION: This inconspicuous species has been reported from temperate to tropical seas throughout the world, in shallow coastal waters, almost invariably on shell and other biogenic carbonates.

MEASUREMENTS (SKELETAL): CL 244 ± 65 µm, 128–385 (2, 15), GL 255 ± 29, 227– 292 (1, 4), GW 157 ± 21, 125–169 (1, 4), OL 43 ± 7, 29–52 (2, 10), OW 41 ± 3, 37– 46 (2, 10), OvL 142 ± 8, 136–154 (1, 4), OvW 157 ± 21, 125–169 (1, 4), ZL 559 ± 67, 437–705 (2, 15), ZW 192 ± 17, 150– 222 (2, 15).

FAMILY CHORIZOPORIDAE VIGNEAUX, 1949 GENUS CHORIZOPORA HINCKS, 1879

Chorizopora brongniartii ( Audouin, 1826) Figure 19A–C

Flustra brongniartii Audouin, 1826: 240 . Savigny, 1809: pl. 10, fig. 6.

Chorizopora brongniartii: Hincks, 1880: 224 . Marcus, 1940: 214. Gautier, 1962: 126. Hayward and Ryland, 1979: 240. Zabala, 1986: 516. Zabala and Maluquer, 1988: 141. Hayward and Ryland, 1999: 100.

DESCRIPTION (AMNH 930, 932; CMRR 2226): Colonies clear, encrusting, multiserial, unilaminar, glassy, and often only apparent once dried. Autozooids oval, fusiform or irregular, only slightly convex; disjunct, adjacent autozooids linked by tubular extensions of basal pore chambers. Primary orifice Dshaped, with straight proximal border and indistinct condyles. No oral spines or peristome. Frontal shield finely granular, imperforate, transversely ridged or striated, and sometimes with a small median suboral umbo. Ancestrula with peristome and two spines on distal margin of orifice. Avicularia vicarious, monomorphic, typically one at distal end of each autozooid; cystid small and rounded, mandible triangular, acute to frontal plane, directed distally. Ovicell elongate, semielliptical, recumbent on substratum between distally succeeding autozooids; finely granular, imperforate, often developing a longitudinal frontal keel; partly incorporated within cystid of avicularium.

Autozooids may be closely packed, with linking tubules only just visible, or widely disjunct with intervening spaces infilled by a mosaic of irregular kenozooidal units, each linked to neighboring kenozooids or autozooids by tubular connections, and each with a small round frontal opesia.

Tentacles clear, 12–13; lophophores bellshaped, radially symmetrical; introverts long.

Embryo color bright orange.

DISTRIBUTION: Chorizopora brongniartii is a very common species in the Mediterranean, and on north European coasts, and is often abundant on every kind of organic substratum. It is possibly one of the few almost cosmopolitan species of cheilostomate bryozoans, having been reported, described, and illustrated from shelf environments throughout the world, with the exception of polar regions.

MEASUREMENTS (SKELETAL): AL 105 ± 14 µm, 80–130 (4, 20), AW 102 ± 14, 80–130 (4, 20), DO 367 ± 49, 300–380 (4, 20), KL 125 ± 25, 80–170 (4, 20), KW 90 ± 26, 50– 120 (4, 20), OL 57 ± 7, 40–70 (4, 20), OW 99 ± 10, 80–120 (4, 20), OvL 268 ± 27, 220–320 (4, 16), OvW 217 ± 17, 200–260 (4, 16), OOW 130 ± 20, 200–260 (4, 16), ZL 429 ± 23, 390–460 (4, 20), ZW 331 ± 33, 280–380 (4, 20). (POLYPIDE): IH 261 ± 41 µm, 200–320 (1, 11), LD 527 ± 51, 430– 580 (1, 11), MD 26 ± 3, 20–30 (1, 11), TL 457 ± 23, 420–500 (1, 11).

INFRAORDER UMBONULOMORPHA GORDON, 1989

SUPERFAMILY ADEONOIDEA BUSK, 1884

FAMILY ADEONIDAE BUSK, 1884

GENUS REPTADEONELLA BUSK, 1884

Reptadeonella violacea ( Johnston, 1847) Figure 19D–G

Figures

Fig. 16. A–C. Collarina balzaci. A. General aspect (AMNH 912; 0.5 mm). B. Autozooid (right) and (left) maternal zooid with ovicell (AMNH 925; 0.1 mm). C. Ancestrula and early autozooids (AMNH 926; 0.1 mm). D, E. Figularia figularis. (AMNH 925). D. General aspect (0.5 mm). E. Autozooid (0.1 mm).

Fig. 17. A–C. Puellina hincksi. A. General aspect (AMNH 901; 0.5 mm). B. Autozooid flanked by avicularia (AMNH 901; 0.1 mm). C. Two maternal zooids with ovicells (AMNH 927; 0.1 mm). D–F. Savignyella lafontii (AMNH 928). D. General aspect (0.5 mm). E. Distal portion of autozooid with aberrant, elongate, twisted suboral avicularium (0.1 mm). F. lateral view of distal end of zooid with intact oral spines and normal suboral avicularium (0.1 mm).

Fig. 18. A–E. Hippothoa divaricata (AMNH 929). A. General aspect (0.5 mm). B. Maternal zooid with ovicell (top) and (bottom) zooeciule budded from central autozooid (0.1 mm). C. Autozooidal orifice (0.05 mm). D. Ancestrula and early autozooids (0.5 mm). E. Ancestrula with D­shaped orifice (0.1 mm). F–I. Hippothoa flagellum. F. General aspect (AMNH 930; 0.5 mm). G. Maternal zooid with ovicell (AMNH 930; 0.1 mm). H. Autozooidal orifice (AMNH 931; 0.05 mm). I. Ancestrula and early autozooids (AMNH 929; 0.1 mm).

Fig. 19. A–C. Chorizopora brongniartii. A. General aspect (AMNH 930; 0.5 mm). B. Autozooid (right) and (center) maternal zooid with ovicell (AMNH 930; 0.1 mm). C. Ancestrula (AMNH 932; 0.1 mm). D–G. Reptadeonella violacea (AMNH 933). D. General aspect (0.5 mm). E. Autozooid with suboral avicularium (0.1 mm). F. Maternal zooid (0.1 mm). G. Ancestrular complex of six zooids diverging from center of figure (0.2 mm).