Schizoporella tetragona: Ryland, 1968: 543

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 : 72-86

Schizoporella tetragona: Ryland, 1968: 543 .

DESCRIPTION (AMNH 966; CMRR 2247): Colony a broad, spreading, multiserial, unilaminar sheet in present material. Autozooids hexagonal to irregularly polygonal, large, convex, separated by deep grooves. Primary orifice as wide as long; poster equivalent to almost half total length, forming a broadly U­shaped sinus, occupying the whole proximal width of the anter below indistinct condyles. Two delicate, widely spaced, distal oral spines present in earliest ontogeny, lost as a low peristomial rim develops around distal and lateral borders of orifice. Frontal shield thick, vitreous, densely perforated by small round pores; thickening through ontogeny and becoming quite rugose as the rims of the pores are more pronounced; a prominent, stout, conical umbo developing immediately proximal to sinus, forming a large spike in some autozooids. Avicularia single or paired, the rounded, prominent cystid situated lateral or just distal to sinus; rostrum narrowly triangular, acute to frontal plane and directed distolaterally. Ovicell hyperstomial, prominent, about as wide as long, with highly arched aperture; imperforate centrally but with numerous pores laterally and around periphery; developing a low frontal umbo.

REMARKS: This appears to be the species described by Ryland (1968) from the Bosporus and attributed to. Schizoporella tetragon a ( Reuss, 1848). The shape of the primary orifice, the disposition of the avicularia and the thickly calcified umbonate frontal shield, noted by Ryland, are evident in this material. However, the equivalence with Reuss’ species needs reexamination, following the advent of SEM in bryozoan morphological research.

MEASUREMENTS (SKELETAL): AL 133 ± 10 µm, 112–145 (2, 12), AW 90 ± 8, 75–106 (2, 12), DO 428 ± 56, 310–507 (2, 20), OL 108 ± 8, 86–120 (2, 20), OW 114 ± 8, 95– 132 (2, 20), OvL 285 ± 24, 245–314 (2, 16), OvW 294 ± 20, 258–325 (2, 16), SL 18 ± 3, 12–22 (2, 20), SW 53 ± 4, 46–61 (2, 20), ZL 558 ± 49, 473–671 (2, 20), ZW 387 ± 40, 315–451 (2, 20).

GENUS ESCHARINA MILNE EDWARDS, 1836

Escharina vulgaris ( Moll, 1803)

Figure 32E–I

Eschara vulgaris var. a Moll, 1803: 55.

Lepralia botterii Heller, 1867: 106 .

Escharina vulgaris: Hayward and Ryland, 1999: 236 .

DESCRIPTION (AMNH 947, 967; CMRR 2248): Colonies irregular, multiserial, unilaminar patches. Autozooids oval to hexagonal, or irregularly polygonal, gently convex in early ontogeny, becoming rather flat as calcification thickens in later ontogeny, separated by shallow grooves. Primary orifice longer than wide, broadest at midlength; distal rim raised, thin; proximal border straight, with a short, rounded sinus medially, equivalent to one­quarter proximal width; condyles short and broad, tapering towards sinus, distinctly ridged. Five or six delicate oral spines present in periancestrular zooids, typically absent in later autozooids; peristome erect and thin, developed independently of the orifice rim, which remains distinct, extending distal to the spine bases, and proximally enclosing a smooth, cupped area below the sinus. Frontal shield finely granular, imperforate except for a few large, irregular marginal pores. Avicularia adventitious, single or paired, developed along lateral margins of autozooids; rostrum slender, acuminate, with a slight lateral curve, directed distally or distolaterally. Ovicell (not present in material described) spherical, finely granular, imperforate. Ancestrula about half the length of normal autozooids, oval, tatiform, with a narrow, smooth gymnocyst and indistinct cryptocystal rim bounded by about nine spines.

REMARKS: We can find no differences between Escharina vulgaris and the type specimen (UIIZ 217) of Lepralia botterii Heller, 1867 , from the northern Adriatic, illustrated here as fig. 32G.

DISTRIBUTION: On stones and shells, in coastal and shelf environments. Widespread and common throughout the Mediterranean, ranging south to Madeira and north along the western coasts of Britain and Ireland. Unknown in the North Sea.

MEASUREMENTS (SKELETAL): AL 106 ± 15 µm, 84–140 (2, 20), AW 67 ± 8, 54–83 (2, 20), DO 431 ± 49, 349–513 (2, 20), OL 103 ± 14, 83–126 (2, 20), OW 88 ± 6, 75–104 (2, 20), SL 18 ± 6, 10–18 (2, 20), SW 17 ± 5, 12–30 (2, 20), ZL 544 ± 37, 471–616 (2, 20), ZW416 ± 40, 353–517 (2, 20).

Escharina dutertrei protecta Zabala, Maluquer and Harmelin, 1993 Figure 33A–C

Escharina dutertrei protecta Zabala, Maluquer and Harmelin, 1993: 73 .

DESCRIPTION (AMNH 968): Colonies encrusting, multiserial, unilaminar sheets. Autozooids broadly hexagonal, convex, separat­ ed by distinct grooves. Primary orifice wider than long, poster a short, rounded sinus occupying just one­eighth of the proximal width of the anter; condyles short and broad, with denticulate edges. Six delicate spines around distal border of anter in early ontogeny, some or all of which become incorporated in development of conspicuous peristome, extending proximally from proximalmost spine pair, flaring outwards and frontally, enclosing and shielding the sinus. Close to proximal spine pair the peristome rim produced as stout, projecting processes, in many autozooids extending distally as a lobed rim, obscuring spine bases. Frontal shield nodular, vitreous, with indistinct marginal pores. Avicularia adventitious, paired, one each side of orifice, level with its distal half; cystid <0.05 mm long, with trough­like rostrum supporting base of a setiform mandible, 0.375 mm long, directed distomedially over frontal shield of distal autozooid. Ovicell wider than long, recumbent, with medially peaked aperture; imperforate, granular.

REMARKS: This taxon was only recently defined by Zabala et al. (1993), who distinguished it from the northeast Atlantic E. dutertrei haywardi . All previous Mediterranean records of E. dutertrei probably refer to this subspecies, which also occurs around the Azores, and it is probable that if Escharina dutertrei (Audouin) could be recognized, and a neotype accepted, the two subspecies introduced by Zabala et al. (1993), which appear to have distinct geographical Distribution patterns, could be accepted as full species.

DISTRIBUTION: See Remarks.

MEASUREMENTS (SKELETAL): DO 493 ± 40 µm, 422–611 (2, 20), OL 86 ± 13, 64–106 (2, 20), OW 86 ± 10, 69–106 (2, 20), OvL 170 ± 20, 150–190 (1, 3), OvW 276 ± 17, 263–301 (1, 4), SL 20 ± 3, 15–24 (2, 20), SW 13 ± 4, 8–23 (2, 20), ZL 565 ± 46, 503– 659 (2, 20), ZW 408 ± 34, 351–500 (2, 20).

FAMILY MARGARETTIDAE HARMER, 1957

GENUS MARGARETTA GRAY, 1843

Margaretta cereoides

( Ellis and Solander, 1786)

Figure 33D–G

Cellaria cereoides Ellis and Solander, 1786: 26 .

Margaretta cereoides: Gautier, 1962: 216 . Zabala and Maluquer, 1988: 155.

DESCRIPTION (AMNH 969; CMRR 2249): Colonies pink, erect, branching, jointed, to 8 cm high in present material; consisting of rigid, curving, round­sectioned internodes, each rising from a flexible, tubular, chitinous node (basis rami) with pronounced annulations. Each basis rami supporting single autozooid, which produces two zooid buds; thereafter, autozooids disposed in alternating whorls of four. Autozooids elongate, strongly convex, separated by distinct grooves. Primary orifice wider than long, D­shaped, visible at growing edge but in early ontogeny obscured by development of tall, cylindrical, longitudinally fluted peristome. Frontal shield thick, vitreous, densely perforated by large pores; distinctly larger ascopore just proximal to peristome base. In dried material, outer cuticle conspicuous. Reproduction and brooding not documented, except that reproductively active autozooids recognized by dimorphic peristomes, consisting of broadbased cone with tapered, distally curved apical aperture; chamber thus formed perhaps an external brood chamber. Branching adventitious, not dichotomous. Each basis rami arising from frontal surface of modified autozooid in which peristome sealed before complete development; basis rami budding at site of ascopore.

Tentacles light orange­pink, 25–28; lophophores bell­shaped, radially symmetrical, supported on long introverts.

DISTRIBUTION: Develops stiff clumps attached to a range of hard substrata, in shallow coastal waters, probably endemic to the Mediterranean.

MEASUREMENTS (SKELETAL): DO 903 ± 98 µm, 669–1097 (2, 23), OW 170 ± 23, 163– 231 (2, 22), OOW 219 ± 13, 199–245 (2, 20), OvL 608 ± 42, 533–719 (2, 20), ZL 1456 ± 170, 1177–1759 (2, 20), ZW 581 ± 54, 458–660 (2, 20). (POLYPIDE): IH 362 ± 104 µm, 260–580 (1, 10), LD 886 ± 60, 740–960 (1, 11), MD 37.3 ± 2.5, 35–40 (1, 3), TL 793 ± 63, 680–872 (1, 10).

FAMILY MYRIAPORIDAE GRAY, 1841

GENUS MYRIAPORA DE BLAINVILLE, 1830

Myriapora truncata ( Pallas, 1766)

Figure 34A–D

Millepora truncata Pallas, 1766: 249 .

Myriapora truncata: Gautier, 1962: 268 . Zabala

and Maluquer, 1988: 163.

DESCRIPTION (AMNH 970; CMRR 2250): Colonies red, erect, rigid, attached by encrusting base, branching irregularly; branches thick, cylindrical, of constant width, to 4 mm diameter, except thicker at base of colonies. Living specimens deep red. Autozooids in alternating whorls of seven to eight; individual boundaries indistinct, even at growing tips. Colony surface appearing uniformly finely granular, regularly perforat­ ed by small, round pores, with primary orifices of autozooids regularly spaced in strict quincuncial order. Primary orifice slightly longer than wide; anter more or less semicircular, comprising two­thirds orifice length; poster a short semiellipse below prominent, bluntly triangular condyles. No spines or peristome. Ovicell semiimmersed, its frontal surface scarcely projecting from colony frontal plane; more finely punctate than frontal shields of adjoining autozooids; orifice of brooding zooids dimorphic, nearly twice length and width of autozooid, with a proportionately broader poster.

Tentacles red, 27–29; lophophores paraboloid, supported on long introverts.

DISTRIBUTION: Widely distributed throughout the Mediterranean, on shell and hard substrata, in caves, on coralligenous deposits, from 30 m to at least 130 m depth, probably endemic to Mediterranean.

MEASUREMENTS (SKELETAL): DO 627 ± 59 µm, 519–723 (3, 20), OL 265 ± 11, 250– 283 (3, 25), OW 228 ± 11, 203–259 (3, 25), OOL 370 ± 25, 306–388 (2, 11), OOW 337 ± 15, 315–367 (2, 11), OvL 769 ± 55, 688– 891 (2, 10), OvW 820 ± 79, 701–915 (2, 11). (POLYPIDE): IH 460 ± 24 µm, 440–500 (1, 5), LD 1107 ± 83, 1040–1200 (1, 3), TL 1030 ± 37, 960–1060 (1, 6).

FAMILY LANCEOPORIDAE HARMER, 1957 GENUS CALYPTOTHECA HARMER, 1957

Calyptotheca rugosa Hayward, 1974

Figure 34E–G

Calyptotheca rugosa Hayward, 1974: 379 .

DESCRIPTION (AMNH 971): A single, small, multiserial, unilaminar colony of just 28 autozooids. Autozooids polygonal, convex, separated by distinct grooves. Primary orifice as wide as long; poster accounting for about one­quarter of total length and broadly triangular, occupying entire proximal width of anter. In absence of the operculum, very large, quadrangular condyles, orientated oblique to the disto­proximal axis, and a conspicuous inner shelf around the distal part of the anter, imparting distinctive outline to the orifice. Spines absent except in earliestformed autozooids, diminishing in number away from ancestrula; low, thick peristomial ridge encircling orifice. Frontal shield thick, nodular and vitreous, densely perforated by small round pores. Avicularium adventitious, suboral, rare; ovicell large, sutured, perforate (neither found in present material). Ancestrula identical to later autozooids; producing paired distolateral buds.

DISTRIBUTION: This species has not been reported since its original description, based on specimens from Chios, Aegean Sea.

MEASUREMENTS (SKELETAL): DO 502 ± 75 µm, 408–650 (1, 10), OL 140 ± 10, 127– 159 (1, 10), OW 143 ± 7, 135–154 (1, 10), ZL 654 ± 61, 550–742 (1, 10), ZW 442 ± 82, 362–635 (1, 10).

FAMILY CHEILOPORINIDAE BASSLER, 1936

GENUS HAGIOSYNODO S BISHOP AND HAYWARD, 1989

Hagiosynodos kirchenpaueri ( Heller, 1867) Figure 35E–H

Lepralia kirchenpaueri Heller, 1867: 105 .

Hippopodinella kirchenpaueri: Gautier, 1962:

179. Zabala and Maluquer, 1988: 136.

Hippopodinella lata (Busk) : Schmid, 1989: 47.

NEOTYPE (chosen here): UIIZ 244, bleached specimen on small gastropod.

DESCRIPTION (AMNH 966): Colonies encrusting, multiserial, unilaminar with local areas of self­overgrowth associated with rejuvenation. Autozooids oval to irregularly polygonal, separated by indistinct grooves. Primary orifice longer than wide, distinctive: anter widest at midlength, its rim constituting three­quarters of a complete ellipse; condyles short and bluntly pointed, directed medioproximally; poster with almost straight proximal border, and width three times its length; usually conspicuously wider than anter; rugose umbones locally present at proximolateral margins of orifice. No spines. Frontal shield gently convex, with a rugose, nodular surface, evenly punctured by pores. Ovicell recumbent, only slightly convex, calcification and perforation identical to frontal shield; closed by zooidal operculum. Small dietellae clearly visible at growing edges.

Tentacles light carmine, 12; lophophores campylonemidan.

REMARKS: We designate as lectotype Heller’s specimen illustrated here as fig. 35A, from syntype suite UIIZ 244. Through a comparison of morphology and biometrics, Schmid (1989) synonymised the type of Lepralia kirchenpaueri Heller , plus Miocene material from Austria, with Recent specimens of Hippopodinella lata from England. Although Schmid found similar ranges of measurements for autozooidal length and width, orifice length and width, and ovicell length and width, we prefer to retain kirchenpaueri and lata as separate species. Measurements for the two are generally similar, but we distinguish Hagiosynodos kirchenpaueri from H. latus ( Busk, 1856) by kirchenpaueri having on average larger dimensions (see fig. 35A–D and measurements below for H. kirchenpaueri and for H. latus from the British Isles, BMNH 2001.1.26.18– 2001.1.26.20), a proportionately broader poster, and the presence in H. latu s of a prominent distal lip in the primary orifice on which the operculum rests when closed and its absence in H. kirchenpaueri .

Seven of the eight specimens in the type suite of H. kirchenpaueri (UIIZ 244) occur on gastropod shells; the eighth is on an echinoid spine. The specimens from near Rovinj encrust rock. Hagiosynodos latus is a facultative symbiont of hermit crabs inhabiting gastropod shells (review in Taylor, 1994); the prevalence of H. kirchenpaueri on gastropod shells suggests that it, too, may be a facultative symbiont of hermit crabs. However, we do not know if the gastropods were alive or if the shells were inhabited by hermit crabs when encrusted by H. kirchenpaueri .

DISTRIBUTION: The species is known only from the Mediterranean.

MEASUREMENTS (SKELETAL):

Hagiosynodos hadros , new species Figure 36A–D

DIAGNOSIS: Encrusting colonies with relatively large autozooids, orifices, and spacing of pores in frontal shield; poster of orifice noticeably wider than anter.

HOLOTYPE: AMNH 972.

PARATYPES: AMNH 973; CMRR 2251– 2254.

ETYMOLOGY: The species is named from the Latin hadros (well developed, bulky) because of the large dimensions or its zooids relative to those of Hagiosynodos kirchenpaueri and H. latus .

DESCRIPTION: Colonies encrusting, multiserial, unilaminar with local areas of selfovergrowth associated with rejuvenation. Autozooids oval to irregularly polygonal, separated by indistinct grooves. Primary orifice longer than wide, distinctive: anter widest at midlength, its rim constituting threequarters of a complete ellipse; condyles short and bluntly pointed, directed medioproximally; poster with almost straight proximal border, and width three times its length. No spines. Frontal shield gently convex, with a rugose, nodular surface, evenly punctured by relatively large pores. Ovicell recumbent, only slightly convex, calcification and perforation identical to frontal shield; closed by zooidal operculum. Small dietellae clearly visible at growing edges.

Tentacles light carmine, 12; lophophores campylonemidan.

REMARKS: This species has significantly larger autozooids, orifices, and spacing of pores in the frontal shield than do H. kirchenpaueri and H. latus . Also, rugose umbones at the proximolateral margins of the orifice, common in H. kirchenpaueri and H. latus , are absent in the available material of H. hadros . It was found on gastropod and dead bivalve skeletons. There is no indication that the gastropods were alive or the shells inhabited by hermit crabs when encrusted by H. hadros .

MEASUREMENTS (SKELETAL): DO 425 ± 76 µm, 250–563 (5, 50), OL 135 ± 14, 106– 170 (5, 50), OW (anter) 88 ± 10, 66–114 (5, 50), OW (poster) 102 ± 11, 81–129 (5, 50), OvL 246 ± 26, 197–307 (5, 34), OvW 288 ± 22, 246–338 (5, 34), PS 48 ± 7, 32–73 (5, 50), ZL 559 ± 56, 471–745 (5, 50), ZW 351 ± 48, 240–472 (5, 50). (POLYPIDE): IH 114 ± 71 µm, 40–240 (1, 12), LD 510 ± 58, 440–600 (1, 10), MD 28.8 ± 2.5, 25–30 (1, 4), TL 520 ± 58, 440–640 (1, 10).

FAMILY CRYPTOSULIDAE VIGNEAUX, 1949

GENUS CRYPTOSULA CANU AND BASSLER, 1925

Cryptosula pallasiana ( Moll, 1803) Figure 36E, F

Eschara pallasiana Moll, 1803: 64 .

Cryptosula pallasiana Canu and Bassler, 1925: 33 . Hayward and Ryland, 1999: 194.

DESCRIPTION (AMNH 974; CMRR 2255): Colonies translucent white, encrusting, multiserial, unilaminar, developing thick, spreading, orange sheets. Autozooids rectangular to hexagonal, convex, separated by distinct sutures. Primary orifice longer than wide, post­ er shallowly concave below prominent, downcurved condyles, broader than anter and imparting a bell­shaped outline to the orifice. Operculum deep brown. Peristome developed as a low thickened rim, most prominent and slightly flared distally. Frontal shield thickly calcified, the epitheca glistening conspicuously in dried material; densely perforated by large round pores, with prominent nodular ridges developing between in later ontogeny. A stout median suboral umbo present in most autozooids; in some populations this position is occupied by an avicularium with broadly oval rostrum, acute to frontal plane and proximally directed, but it was not present in any of the Rovinj specimens. No ovicells, pale yellow embryos brooded internally.

Tentacles clear, 17; lophophores bellshaped, radially symmetrical, overlapped with adjacent lophophores up to 80 µm.

OCCURRENCE: Found only in the harbor and adjacent shore of Rt Kriz, 0–5 m depth.

DISTRIBUTION: Cryptosula pallasiana is widely distributed in the North Atlantic, from northern Norway throughout the Mediterranean and into the Black Sea, and from Nova Scotia to North Carolina. Around the British Isles it is a common littoral and shallow sublittoral species, as a characteristic component of the sessile fauna of kelp holdfasts and undersides of boulders. It is also a successful fouling species; it has been part of the fouling community of New Zealand since at least the1890s ( Gordon and Mawatari, 1992), and its presence in docks and harbors elsewhere around the world is probably attributable to ship borne dispersal. Schopf (1973) consid­

ered that populations lacking avicularia were characteristic of environmentally unstable habitats.

MEASUREMENTS (SKELETAL): AL 211 ± 14 µm, 191–240 (2, 20), AW 172 ± 10, 153– 193 (2, 20), DO 501 ± 105, 338–684 (2, 20), ZL 670 ± 62, 591–838 (2, 20), ZW 449 ± 66, 372–581 (2, 20). (POLYPIDE): IH 147 ± 35 µm, 100–200 (1, 6), LD 673 ± 50, 600– 740 (1, 6), MD 40 (1, 1), TL 650 ± 58, 560– 700 (1, 6).

FAMILY MICROPORELLIDAE HINCKS, 1879

GENUS FENESTRULINA JULLIEN, 1888

Fenestrulina malusii ( Audouin, 1826)

Figure 37A–D

Cellepora malusii Audouin, 1826: 239 . Savigny,

1809: pl. 8, fig. 8.

Fenestrulina malusii: Nielsen, 1981: 109 . Hay­

ward and Ryland, 1999: 299.

DESCRIPTION (AMNH 975–977; CMRR 2256): Colonies silvery white, encrusting, multiserial, unilaminar patches. Autozooids oval to hexagonal, convex, separated by well­marked ridges. Primary orifice wider than long, proximal edge straight, four short distal oral spines present. Frontal shield smooth, with a prominent median ascopore, distance between it and proximal orifice rim equivalent to orifice length. Ascopore rim transversely oval to reniform, thickened and raised, lumen crescentic, partly occluded by delicate denticles. Large round pores with thickened rims, also partly occluded and appearing cribriform, in single or double series around autozooid margins, and extending medially between ascopore and orifice. Ancestrula tatiform, about two­thirds length of normal autozooids, with oval opesia occupying over half total length, bordered by very narrow cryptocyst and about 10 spines. Ovicell recumbent on distal autozooid, wider than long, prominent; ectooecium membranous except for a thickened peripheral rim; endooecium smoothly calcified, imperforate, with distinct basal fluting.

Tentacles colorless, 13–15; lophophores bell­shaped, radially symmetrical.

Embryo color lemon yellow to medium orange.

DISTRIBUTION: Shallow sublittoral, and offshore to about 100 m, on a wide range of organic and inorganic substrata. Distributed throughout the Mediterranean, and northwards to the British Isles and western Norway.

MEASUREMENTS (SKELETAL): DO 484 ± 54 µm, 383–613 (2, 20), OL 104 ± 12, 79–134 (2, 20), OW 142 ± 15, 120–181 (2, 20), ZL 625 ± 89, 490–851 (2, 20), ZW 401 ± 45, 325–511 (2, 20). (POLYPIDE): IH 167 ± 33 µm, 140–240 (1, 9), LD 765 ± 106, 640– 900 (1, 4), MD 35–40 (1, 2), TL 660 ± 49, 600–700 (1, 4).

GENUS MICROPORELLA HINCKS, 1877

Microporella ciliata ( Pallas, 1766) Figure 37E–J

Eschara ciliata Pallas, 1766: 38 .

Microporella ciliata: Gautier, 1962: 172 . Hayward and Ryland, 1999: 296.

DESCRIPTION (AMNH 897, 913, 978–980; CMRR 2257): Colonies white though commonly colored by fouling algae, encrusting, multiserial, unilaminar sheets, typically silvery when dried. Autozooids oval to hexagonal, convex, separated by distinct grooves; 0.375–0.5 X 0.3 mm in present material. Primary orifice slightly wider than long, proximal border straight, with a minute condyle in each proximal corner; six slender spines equally spaced around distal and lateral rim. Frontal shield nodular, with few small, round pores, and fewer, more conspicuous marginal pores. Ascopore spaced from orifice by a distance equivalent to half orifice length; rim thickened, especially prominent in later ontogeny, lumen crescentic, with finely denticulate border. A single adventitious avicularium situated proximolateral to ascopore, on right or left; rostrum acute to frontal plane, directed distolaterally, broadly triangular with a distal groove to accommodate a setiform mandible, extending beyond tip of rostrum by twice its length; crossbar slender, complete, no columella or palate. Ovicell domed, hemispherical, aperture overarching primary orifice; surface uniformly nodular, imperforate. Vertical walls of autozooids with distinct basal pore chambers. Ancestrula tatiform, with oval opesia occupying about half total length, bordered by narrow cryptocyst and about nine spines.

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

Embryo color light to bright orange.

DISTRIBUTION: Microporella ciliata is common in shallow coastal waters, and is distributed in the northeast Atlantic from northern Norway to the Mediterranean, where it is widespread. Records from outside this geographical range require reexamination.

MEASUREMENTS (SKELETAL): AL 126 ± 15 µm, 101–148 (2, 15), AW 88 ± 13, 72–117 (2, 15), DO 382 ± 44, 311–454 (2, 20), OL 57 ± 6, 42–67 (2, 20), OW 85 ± 12, 73– 110 (2, 20), OvL 239 ± 8, 221–250 (1, 10), OvW 228 ± 9, 218–245 (1, 10), ZL 486 ± 35, 434–546 (2, 20), ZW 318 ± 38, 273– 404 (2, 20). (POLYPIDE): IH 130 ± 12 µm, 120–140 (1, 4), LD 402 ± 42. 360–500 (1, 9), TL 394 ± 38, 340–440 (1, 7).

SUPERFAMILY CELLEPOROIDEA JOHNSTON, 1838

FAMILY CELLEPORIDAE JOHNSTON, 1838

GENUS CELLEPORA LINNAEUS, 1767

Cellepora adriatica , new species Figure 38A–D

DIAGNOSIS: Primary orifice nearly circular, lacking condyles; peristome asymmetrical, suboral avicularium single, vertical; 1 to 3 pores on ovicell, close to apertural rim.

HOLOTYPE: AMNH 982.

PARATYPE: AMNH 983.

ETYMOLOGY: The species is named from its discovery in the Adriatic Sea.

DESCRIPTION: Colonies small, encrusting, unilaminar to multilaminar. Autozooids recumbent at unilaminar growing edge, erect elsewhere; broadly oval, convex, separated by distinct ridges. Frontal shield smoothly calcified, imperforate except for a few widely spaced marginal areolar pores. Primary orifice as wide as long, orbicular but not perfectly circular: the rim of the anter describing a flatter arc than the rim of the poster; no condyles visible. A single latero­proximal suboral avicularium present, with an inflated cystid tapered and umbonate distally; rostrum elliptical, perpendicular to plane of orifice, facing laterally; crossbar slender, complete, no palate. Peristome well developed, entirely enclosing the orifice and incorporat­ ing the avicularium, deepening in multilaminar parts of colony, where frontally budded autozooids are raised above peristomes of preceding autozooids; peristome rim characteristically asymmetrical, extending in a lobe around one side of the avicularium cystid. Ovicell prominent, raised above distal orifice rim; hemispherical, with a wide, highly arched aperture, close to the distal edge of which is a single large, round pore. The rim of the peristome extends across the frontal surface of the ovicell, encircling the pore. No vicarious avicularia. Vertical walls with small basal pore chambers.

REMARKS: Heller (1867) recorded Cellepora pumicosa from the Adriatic, together with two new species, C. hincksii and C. corticalis . It is possible that one of these may be the taxon described here as new, but Hell­ er did not figure any of these three species. Hayward (1974) described a new genus and species, Rhamphostomellina posidoniae , encrusting Posidonia leaves, from Chios, Aegean Sea. It is very similar to C. adriatica in its asymmetrical peristome, vertically orientated avicularium, orbicular orifice, and in its ovicell, which has 1 to 3 pores close to its aperture rim. It is clear that R. posidoniae should be assigned to Cellepora ; it differs from C. adriatica in its broader ovicell and smaller autozooids, and in the possession of a proximal process on the avicularium cystid, which defines a pseudospiramen within the peristome.

OCCURRENCE: Encrusting shells of dead gastropods.

MEASUREMENTS (SKELETAL): AL 128 ± 8 µm, 116–137 (1, 5), AW 87 ± 4, 83–94 (1, 5), DO 468 ± 87, 337–630 (1, 10), OL 148 ± 8, 134–158 (1, 10), OW 140 ± 11, 123– 156 (1, 10), OvL 147 ± 21, 124–192 (1, 10), OvW 222 ± 13, 198–238 (1, 10).

GENUS BUSKEA HELLER, 1867

Buskea nitida Heller, 1867

Figure 38E–G

Buskea nitida Heller, 1867: 89 .

Eschara quincuncialis Norman, 1867: 204 ; 1868: 222.

Buskea quincuncialis: Hayward and Ryland, 1979: 296 ; 1999: 350, figs. 162C, D, 164.

Buskea nitida: Zabala, 1986: 553 . Zabala and Maluquer, 1988: 156.

DESCRIPTION (AMNH 984, 985; CMRR 2258): Colonies white, erect, branching, slender, to 4 mm in present material. Branches cylindrical, consisting of alternating whorls of three or four autozooids, with almost constant diameter. Autozooids at growing tips elongate oval, convex, small: less than 0.4 mm long, including peristome. Primary orifice terminal, orbicular, with minute, medioproximal sinus, obscured by development of peristome in early ontogeny. No spines. Peristome with smooth, thickened rim, slightly flared, with a symmetrical, rounded, midproximal notch adjacent to which, on right or left, is a small adventitious avicularium, with distally hooked, triangular rostrum, acute to frontal plane and directed distolaterally. Frontal shield smooth, porcellanous, with very few, small, irregular marginal pores. Ovicell wider than long, with straight edged aperture, perforated by <10, large, round pores; becoming immersed in smooth calcification, only perforate frontal area remaining distinct. Frontal calcification thickens through ontogeny and the colony develops a uniformly smooth surface; autozooid boundaries are quite obscured and only rims of immersed peristomes, and circular perforate area marking ovicell, remain distinct.

REMARKS: Zabala (1986) argued that Buskea nitida is clearly identical to the northeast Atlantic B. quincuncialis (Norman, 1867) but both taxa were introduced in the same year and taxonomic priority has not been decided.

DISTRIBUTION: Buskea nitida was described from the Adriatic and has been recorded subsequently from numerous northern and western Mediterranean localities, and from the Atlantic coasts of southern Europe; as B. quincuncialis , it is also known from the Minch, West Scotland.

MEASUREMENTS (SKELETAL): DO 400 ± 80 µm, 286–530 (2, 13), OW 113 ± 9, 00–126 (2, 11), OvL 146 ± 27, 119–211 (2, 12), OvW 186 ± 18, 161–220 (2, 20), ZL 529 ± 40, 481–640 (3, 21), ZW 212 ± 19, 169– 238 (3, 15).

GENUS CELLEPORINA GRAY, 1848 Celleporina caminata ( Waters, 1879)

Figure 39A–D Cellepora retusa var. caminata Waters, 1879: 194 . Celleporina caminata: Gautier, 1962: 244 . Har­

melin, 1969b: 303. Zabala,1986: 563. Zabala and Maluquer, 1988: 158. Ristedt, 1996: 239.

DESCRIPTION (AMNH 986, 987; CMRR 2259): Colonies white to cream­colored, developing stout, frontally budded nodular or spherical growths. Autozooids erect with terminal orifice and tubular peristome, projecting visibly from colony surface. Frontal shield thickly calcified, vitreous and nodular; marginal pores developing tubular extensions to open around base of peristome. Primary orifice longer than wide, the deep, V­shaped sinus comprising about one­third its total length; peristome deep, tubular, rim lowest and broadly concave proximally, elsewhere often produced into short processes; three columnar avicularia traverse the length of peristome and project above rim: typically one middistally, two laterally; rostra terminal, acute to orifice plane, distal one directed distally, lateral pair distolaterally. Vicarious avicularia frequent, monomorphic, characteristic: cystid broadly conical, proximal portion of rostrum apical, spatulate distal portion directed basally, down side of the cystid; crossbar slender, without columella, palate with extensive foramen. Ovicell slightly wider than long, its frontal surface level with peristome rim, with a narrow, sickle­shaped area of membranous ectooecium, exposed entooecium vitreous, nodular, with a single series of large and irregular marginal pores.

Tentacles light orange, 14 –16; lophophores bell­shaped, radially symmetrical with short introverts to obliquely truncate with long introverts adjacent to excurrent chimneys.

Embryo color deep, bright red. Ancestrula described and figured by Ristedt (1996).

REMARKS: Celleporina caminata is similar to C. canariensis Aristegui, 1989 , but is distinguished by its larger, more robust colony form and larger autozooidal dimensions. The primary orifice, in particular, is larger than that of C. canariensis , with a proportionately deeper sinus, and the peristome rim of nonovicellate autozooids bears three, distinctly tubular avicularia.

DISTRIBUTION: This familiar species is common throughout the Mediterranean in shallow, coarse detrital habitats, encrusting stone, shell, hydroids, and other bryozoans.

It is probably endemic to the Mediterranean region.

MEASUREMENTS (SKELETAL): AAL 94 ± 10 µm, 77–123 (2, 20), AAW 65 ± 6, 52–77 (2, 20), DO 497 ± 87, 303–674 (2, 20), OL 195 ± 18, 169–229 (3, 30), OW 136 ± 14, 104–169 (3, 30), OvL 297 ± 21, 272–312 (1, 7), OvW 332 ± 27, 291–364 (1, 7), SL 23 ± 4, 17–33 (3, 30), SW 28 ± 4, 22–37 (3, 30), VAL 420 ± 42, 355–473 (2, 9), VAW 239 ± 29, 193–271. (POLYPIDE): IH 24 ± 37 µm, 0–120 (2, 19), LD 745 ± 74, 640– 840 (2, 20), MD 30 (1, 1), TL 615 ± 80, 500–800 (2, 20).

Celleporina canariensis Aristegui, 1989 Figure 39E–G

Figures

Fig. 32. A–D. Schizoporella cf. S. tetragona (AMNH 966). A. General aspect (0.5 mm). B. Autozooid (0.1 mm). C. Maternal zooid with ovicell (0.1 mm). D. Autozooidal orifice (0.05 mm). E–I. Escharina vulgaris. E. General aspect (AMNH 967; 0.5 mm). F. Autozooid (AMNH 967; 0.1 mm). G. Untreated autozooids with setae­like mandibles of avicularia (type specimen of Lepralia botterii, UIIZ 217; 0.1 mm). H. Oblique view of autozooidal orifice (AMNH 967; 0.05 mm). I. Ancestrula (lower right) and early autozooids (AMNH 947; 0.1 mm).

Fig. 33. A–C. Escharina dutertrei protecta (AMNH 968). A. General aspect of untreated colony (0.05 mm). B. Autozooid, with mandibular setae of vicarious avicularia crossed distal to orifice (0.1 mm). C. Maternal zooid with ovicell and setae as in Fig. 33B (0.1 mm). D–G. Margaretta cereoides (AMNH 969). D. Branch segment (1 mm). E. General aspect (0.5 mm). F. Autozooidal orifice (0.1 mm). G. Peristome­bearing zooid (0.2 mm).

Fig. 34. A–D. Myriapora truncata (AMNH 970). A. Truncated branch tip (1 mm). B. General aspect with autozooids and maternal zooids with embedded ovicells (0.5 mm). C. Autozooidal orifice (0.1 mm). D. Maternal zooid orifice (0.1 mm). E–G. Calyptotheca rugosa (AMNH 971). E. General aspect (0.5 mm). F. Autozooid (0.1 mm). G. Ancestrula (0.1 mm).

Fig. 35. A–D. Hagiosynodos latus (BMNH 2001.1.26.20; Western Channel, U.K.). A. General aspect (0.5 mm). B. Autozooids (0.1 mm). C. Maternal zooid with ovicell (0.1 mm). D. Autozooidal orifice (0.05 mm). E–H. Hagiosynodos kirchenpaueri.E. General aspect (lectotype, UIIZ 244; 0.5 mm). F. Autozooids (AMNH 966; 0.1 mm). G. Maternal zooid with ovicell (AMNH 966; 0.1 mm). H. Autozooidal orifice (AMNH 966; 0.05 mm).

Fig. 36. A–D. Hagiosynodos hadros. A. general aspect (holotype, AMNH 972; 0.5 mm). B. Autozooid (holotype, AMNH 972; 0.1 mm). C. Naternal zooid with ovicell (holotype, AMNH 972; 0.1 mm). D. Autozooidal orifice (paratype, AMNH 973; 0.05 mm). E, F. Cryptosula pallasiana (AMNH 974). E. General aspect (0.5 mm). F. Autozooid (0.1 mm).

Fig. 37. A–D. Fenestrulina malusii. A. General aspect (AMNH 975; 0.5 mm). B. Autozooid (AMNH 975; 0.1 mm). C. Maternal zooids with ovicells (AMNH 976; 0.1 mm). D. Ancestrula and early autozooids (AMNH 977; 0.1 mm). E–J. Microporella ciliata. E. Young colony encrusting Cellaria (AMNH 913; 0.5 mm). F. General aspect (AMNH 978; 0.5 mm). G. Ontogenetically young autozooid (AMNH 979; 0.1 mm). H. Maternal zooid with ovicell (AMNH 978; 0.1 mm). I. Autozooidal orifice (AMNH 980; 0.05 mm). J. Ancestrula (center) and early autozooids (AMNH 897; 0.1 mm)

Fig. 38. A–D. Cellepora adriatica. A. General aspect (holotype, AMNH 982; 0.5 mm). B. Oblique view of colony surface (holotype, AMNH 982; 0.1 mm). C. Maternal zooids, upper one with complete ovicell, lower with partially formed ovicell (holotype, AMNH 982; 0.1 mm). D. Autozooidal orifice (paratype, AMNH 983; 0.05 mm). E–G. Buskea nitida.E. General aspect (AMNH 984; 0.5 mm). F. Maternal zooid with partially submerged ovicell (AMNH 985; 0.1 mm). G. Two autozooidal orifices (AMNH 984; 0.05 mm).

Fig. 39. A–D. Celleporina caminata. A. general aspect (AMNH 986; 0.5 mm). B. oblique view of colony surface (AMNH 987; 0.1 mm). C. ontogenetically young autozooid (top right), maternal zooid with ovicell (left) and (bottom right) vicarious avicularium (AMNH 986; 0.1 mm). D. autozooidal orifice (AMNH 986; 0.05 mm). E–G. Celleporina canariensis (AMNH 988). E. general aspect (0.5 mm). F. maternal zooids with ovicells and (top left) vicarious avicularium (0.1 mm). G. autozooidal orifice (0.05 mm).