NORTHERN ADRIATIC BRYOZOA FROM THE VICINITY OF ROVINJ, CROATIA

PETER J. HAYWARD School of Biological Sciences, University of Wales Singleton Park, Swansea SA2 8PP, United Kingdom Honorary Research Fellow, Department of Zoology The Natural History Museum, London SW7 5BD, UK

FRANK K. MCKINNEY Research Associate, Division of Paleontology American Museum of Natural History Professor Emeritus, Department of Geology Appalachian State University, Boone, NC 28608

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 270, 139 pp., 63 ®gures, 1 table Issued June 24, 2002

Copyright ᭧ American Museum of Natural History 2002 ISSN 0003-0090 2 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

CONTENTS Abstract ...... 5 Introduction ...... 5 Materials and Methods ...... 7 Systematic Accounts ...... 10 Order Ctenostomata ...... 10 Nolella dilatata (Hincks, 1860) ...... 10 Walkeria tuberosa (Heller, 1867) ...... 10 Bowerbankia spp...... 11 Amathia pruvoti Calvet, 1911 ...... 12 Amathia vidovici (Heller, 1867) ...... 13 Order Cheilostomata ...... 14 Aetea sica (Couch, 1844) ...... 14 Aetea truncata (Landsborough, 1852) ...... 14 Scruparia ambigua (d'Orbigny, 1841) ...... 16 Pyripora sp. (Fleming, 1828) ...... 16 Callopora dumerilii (Audouin, 1826) ...... 16 Copidozoum tenuirostre (Hincks, 1880) ...... 18 Ellisina gautieri FernaÂndez Pulpeiro and Reverter Gil, 1993 ...... 18 Chartella tenella (Hincks, 1887) ...... 20 Bugula fulva Ryland, 1960 ...... 20 Beania cylindrica (Hincks, 1886) ...... 22 Beania hirtissima (Heller, 1867) ...... 24 Beania magellanica (Busk, 1852) ...... 24 Beania mirabilis Johnston, 1840 ...... 26 Synnotum aegyptiacum (Audouin, 1826) ...... 26 Scrupocellaria delilii (Audouin, 1826) ...... 27 Scrupocellaria scrupea Busk, 1851 ...... 27 Scrupocellaria scruposa (Linnaeus, 1758) ...... 27 Caberea boryi (Audouin, 1826) ...... 29 Calpensia nobilis (Esper, 1796) ...... 31 Rosseliana rosselii (Audouin, 1826) ...... 31 Mollia circumcincta (Heller, 1867) ...... 34 Cellaria ®stulosa (Linnaeus, 1758) ...... 34 Cellaria salicornioides Lamouroux, 1816 ...... 36 Collarina balzaci (Audouin, 1826) ...... 36 Figularia ®gularis (Johnston, 1847) ...... 38 Puellina hincksi (Friedl, 1917) ...... 38 Savignyella lafontii (Audouin, 1826) ...... 40 Hippothoa divaricata Lamouroux, 1821 ...... 40 Hippothoa ¯agellum Manzoni, 1870 ...... 42 Chorizopora brongniartii (Audouin, 1826) ...... 42 Reptadeonella violacea (Johnston, 1847) ...... 44 Adeonella pallasii (Heller, 1867) ...... 44 Escharoides mamillata (Wood, 1844) ...... 46 Umbonula ovicellata Hastings, 1944 ...... 46 Parasmittina rouvillei (Calvet, 1902) ...... 48 Smittina cervicornis (Pallas, 1766) ...... 49 Prenantia cheilostoma (Manzoni, 1869) ...... 49 Phoceana tubulifera (Heller, 1867) ...... 51 Smittoidea reticulata (J. MacGillivray, 1842) ...... 51 Hippoporina lineolifera (Hincks, 1886) ...... 53 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 3

Pentapora fascialis (Pallas, 1766) ...... 53 Schizomavella asymetrica (Calvet, 1927) ...... 56 Schizomavella cornuta (Heller, 1867) ...... 57 Schizomavella linearis (Hassall, 1841) ...... 59 Schizomavella mamillata (Hincks, 1880) ...... 59 Schizomavella rudis (Manzoni, 1869) ...... 61 Schizomavella subsolana, new species ...... 61 Watersipora complanata (Norman, 1864) ...... 63 Watersipora subovoidea (d'Orbigny, 1852) ...... 63 Schizobrachiella sanguinea (Norman, 1868) ...... 65 Schizoporella unicornis (Johnston in Wood, 1844) ...... 67 Schizoporella errata (Waters, 1878) ...... 67 Schizoporella dunkeri (Reuss, 1848) ...... 69 Schizoporella magni®ca Hincks, 1886 ...... 71 Schizoporella cf. S. tetragona (Reuss, 1848) ...... 72 Escharina vulgaris (Moll, 1803) ...... 72 Escharina dutertrei protecta Zabala, Maluquer and Harmelin, 1993 ...... 74 Margaretta cereoides (Ellis and Solander, 1786) ...... 74 Myriapora truncata (Pallas, 1766) ...... 76 Calyptotheca rugosa Hayward, 1974 ...... 76 Hagiosynodos kirchenpaueri (Heller, 1867) ...... 76 Hagiosynodos hadros, new species ...... 79 Cryptosula pallasiana (Moll, 1803) ...... 81 Fenestrulina malusii (Audouin, 1826) ...... 81 Microporella ciliata (Pallas, 1766) ...... 83 Cellepora adriatica, new species ...... 83 Buskea nitida Heller, 1867 ...... 85 Celleporina caminata (Waters, 1879) ...... 85 Celleporina canariensis Aristegui, 1989 ...... 86 Celleporina siphuncula, new species ...... 88 Celleporina tubulosa (Hincks, 1880) ...... 90 Turbicellepora avicularis (Hincks, 1860) ...... 90 Turbicellepora camera Hayward, 1978 ...... 92 Reteporella grimaldii (Jullien, 1903) ...... 92 Rhynchozoon neapolitanum Gautier, 1962 ...... 94 Rhynchozoon pseudodigitatum Zabala and Maluquer, 1988 ...... 96 Rhynchozoon revelatus, new species ...... 96 Schizotheca ®ssa (Hincks, 1856) ...... 98 Schizotheca serratimargo (Hincks, 1886) ...... 98 Order Cyclostomata ...... 101 Tubulipora liliacea (Pallas, 1766) ...... 101 Tubulipora plumosa Harmer, 1898 ...... 103 Exidmonea triforis (Heller, 1867) ...... 104 Exidmonea coerula (Harmelin, 1976) ...... 107 Annectocyma arcuata (Harmelin, 1976) ...... 107 Annectocyma cf. A. major (Johnston, 1847) ...... 109 Annectocyma sp...... 109 Entalophoroecia de¯exa (Couch, 1844) ...... 110 Entalophoroecia robusta Harmelin, 1976 ...... 110 Plagioecia patina (Lamarck, 1816) ...... 112 Plagioecia sarniensis (Norman, 1864) ...... 114 Eurystrotos compacta (Norman, 1867) ...... 116 Diplosolen obelium (Johnston, 1838) ...... 117 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Frondipora verrucosa (Lamouroux, 1821) ...... 119 Crisia ®stulosa Heller, 1867 ...... 120 Crisia ramosa Harmer, 1891 ...... 122 Crisia recurva Heller, 1867 ...... 122 Disporella hispida (Fleming, 1828) ...... 123 Patinella radiata (Audouin, 1826) ...... 127 Acknowledgments ...... 128 References ...... 129 Appendix 1 ...... 138 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 5

ABSTRACT One hundred six species of Bryozoa collected from the northern Adriatic in the vicinity of Rovinj, Croatia, are distributed among the orders Ctenostomata (8 species), Cheilostomata (79 species), and Cyclostomata (19 species). Ctenostomes are underrepresented in the collections relative to the two orders with calci®ed colonies. Five of the cheilostome species are new: Hagiosynodos hadros n. sp., Schizomavella subsolana n. sp., Cellepora adriatica n. sp., Cel- leporina siphuncula n. sp., and Rhynchozoon revelatus n. sp. (previously referred to as Rhyn- chozoon sp. II Hayward). Seven species named by Heller (1867) are stabilized by selection of lectotypes (Beania hirtissima, Adeonella pallasii, Hagiosynodos kirchenpaueri, Exidmonea triforis, Crisia recurva) and neotypes (Mollia circumcincta, Schizomavella cornuta) from Hell- er's collection in the University of Innsbruck Institute of Zoology. Lectotypes are designated for the Adriatic species Hippoporina lineolifera (Hincks, 1886) and for Schizomavella mam- illata (Hincks, 1880). Beania cylindrica (Hincks, 1886) and Schizoporella asymetrica (Calvet, 1927) are recognized as species rather than as subspeci®c units. The species-rich cheilostome genus Schizoporella Hincks, 1877, which contains some of the most widely known fouling bryozoans, is designated a nomen protectum. The species name Smittina cheilostoma (Man- zoni, 1869) is preserved as established usage.

INTRODUCTION lin, 1993, 2000), and functional morphology (McKinney, 1988, 1989, 1991a, 1991b; M. J. The diverse and abundant Bryozoa of the McKinney, 1997). The abundance and diver- Adriatic Sea have attracted attention since sity of bryozoans in the Adriatic Sea along the mid-19th century, when Heller (1867) the Istrian coast north and south of Rovinj, produced the ®rst taxonomic account of the as well as their central role in some of the fauna, soon supplemented by Hincks (1886, benthic biocoenoses, has long warranted a 1887). Since then numerous publications monograph of the bryozoans of the region. have extended or modi®ed the list of species The fauna described and illustrated here is present, reported new occurrences of species located within the northern Adriatic, which previously reported from the Adriatic, and occupies a shallow platform north of two increasingly during the latter part of the 20th successively deeper basins in the middle and century, used the bryozoans as subjects for southern Adriatic (Buljan and Zore-Arman- ecological and geological studies. Extended systematic description and illus- da, 1976). The northern Adriatic has a slight- tration of the species described from the ly less diverse fauna than may be found far- Adriatic has been limited. Very few species ther south (Gamulin-Brida, 1979). However, either in taxonomic works or in other studies it is of great interest because of the increas- of Adriatic bryozoans have been illustrated ing ecological stresses due to the interaction in suf®cient detail that they can be recog- of variations in circulation and growing hu- nized with con®dence, other than the limited man impact on the small volume of water fauna inhabiting Italian lagoons (Occhipinti spread across the area (e.g. Degobbis et al., Ambrogi, 1981). 1979; Degobbis, 1989; Marchetti et al., The present study is a monograph of the 1989; Stachowitsch, 1991). (The northern species known to occur in the vicinity of Adriatic as a whole averages only 35 m Rovinj, Croatia, along and just offshore of depth; Buljan and Zore-Armanda, 1976.) The the Istrian Peninsula (®g. 1). Rovinj histori- shallow, usually oligotrophic environment of cally has been a center where bryozoans the northeastern Adriatic is an important have been noted, including production of model for understanding Cenozoic deposits faunal lists (Friedl, 1925) and studies of re- to the north and west of the present Adriatic cruitment (McKinney and McKinney, 1993, (SenesÏ, 1988), and its benthic communities, 2002), growth and reproduction (Friedl, some of which are dominated by calci®ed 1925), ecology (NikolicÂ, 1954, 1959a, bryozoans, are important in understanding 1959b, 1960; Strenger and Splechtna, 1978; what may be persistent differences between McKinney, 1992, 2000; McKinney and Jak- past and present benthic faunas in oligotro- 6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 1. Map of the northeastern Adriatic Sea in the vicinity of Rovinj, Croatia, encompassing the area from which specimens in this study were collected. 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 7 phic and in mesotrophic to eutrophic marine ders and cobbles derived from them, and, lo- waters. cally, caves extending into the rock. The oth- er general natural environment consists of a MATERIALS AND METHODS gently sloping plain of sediment, largely sand but locally dominated by mud, that is at a COLLECTION AND REPOSITORY OF MATERIAL water depth of only a few meters adjacent to Bryozoan samples were collected by some parts of the mainland but that declines dredging and by hand collecting. Dredging westward to an extensive area of 30±40 m was accomplished by towing a ``Muslar'' depth, with a few deeper depressions. In ad- dredge behind the Burin, a boat operated by dition, surfaces of a few arti®cial substrata in the Center for Marine Research±Rovinj the Rovinj harbor and in Lim Channel were (CMRR). The dredge was towed for approx- sampled. imately 20 minutes each time, sampling a Locality 1. Shore near west end of north side of path approximately 1 m wide by 500 m long. Punta Corrente, cobbles (hand collected), 0±1 Most hand collections were made by SCU- m depth 45Њ03.8ЈN, 13Њ37.5ЈE). 23 September BA divers, led by Andrej Jaklin of the 1990, 12 June 1997 CMRR. Shore collections were made by F.K. Locality 2. South side, mouth of Lim Channel, McKinney and M.J. McKinney. rock (SCUBA), 2±3 m depth (45Њ06.9ЈN, Collections are not equally representative 13Њ37.0ЈE). 12 October 1990 of each site. Some sites were visited and col- Locality 3. Cave in south side of Sv. Ivan Island, lected intensely as many as eight times, oth- near landing site on the island (SCUBA), 3±5 ers were visited only once. Some of the m depth (45Њ02.4ЈN, 13Њ37.1ЈE). 9 September 1987 SCUBA collections were focused speci®cal- Locality 4. West side of Banjole Island (top of ly on large, calci®ed colonies of certain spe- boulder at entrance to cave (SCUBA), 3.5 m cies. Small and inconspicuous species that depth 45Њ 03.9ЈN, 13Њ36.5ЈE). 3 July 1998 can be seen well only under a microscope Locality 5. West and south sides of Banjole Is- would have been missed in such situations, land, rock wall (SCUBA), 5±20 m depth whereas they were much more likely to be (45Њ03.9ЈN, 13Њ36.5ЈE). 14 September 1987, 7 encountered during the close scrutiny of sur- June 1988, 20 September 1990, 9 October faces in the dredged material. 1990, 8 June 1995 All material collected was examined live Locality 6. Lim Channel, approximately 1 km in the CMRR within a week of collection, from mouth, rock ledge (SCUBA), 5±10 m depth (45Њ07.5ЈN, 13Њ37.6ЈE). 24 September then preserved in formalin, alcohol, or by 1987 drying. Other than the Heller types, which Locality 7. Northwest tip of Sv. Ivan Island, are in the University of Innsbruck Institute ¯anks and undersides of boulders (SCUBA), 8± of Zoology (UIIZ), and specimens of Hagio- 10 m depth (45Њ02.4ЈN, 13Њ37.1ЈE). 2 October synodos latus and Hippoporina lineolifera, 1990 which are in the collections of the Natural Locality 8. Northwest tip of Sv. Ivan Island, rock History Museum, London (BMNH), each wall (SCUBA), 10±30 m depth (45Њ02.4ЈN, specimen illustrated here by scanning elec- 13Њ37.1ЈE). 20 June 1988 tron microscopy is in the Bryozoan Collec- Locality 9. Northwest corner of Pelago Island, tions of the American Museum of Natural rock (SCUBA), 10±20 m depth (45Њ 03.2ЈN, History (AMNH). Indicated paratypes of 13Њ36.6ЈE). 20 June 1988 Locality 10. Cave in west side of Banjole Island new species and other reference materials are (SCUBA), 10±11 m depth (45Њ 03.9ЈN, deposited in the RudÅer BosÏkovic Institute 13Њ36.5ЈE). 7 June 1988 Center for Marine Research±Rovinj. Locality 11. Lim Channel, approximately 5 km from mouth, rock (SCUBA), 12±20 m depth COLLECTING LOCALITIES (45Њ07.4ЈN, 13Њ39.7ЈE). 10 June 1988 Locality 12. Approximately 100 m west of Stur- Two general types of natural environments ago Island, ¯anks and undersides of boulders were sampled. The underwater continuations (SCUBA), 20 m depth (45Њ03.2ЈN, 13Њ36.8ЈE). of the shore and islands in the vicinity of 23 June 1988 Rovinj consist of limestone rock walls, boul- Locality 13. West and south sides of Banjole Is- 8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

land, rock wall (SCUBA), 25±30 m depth Ivan Island, sand bottom (dredge), 35 m depth (45Њ03.9ЈN, 13Њ36.5ЈE). 20 September 1990, 8 (45Њ00.8ЈN, 13Њ39.2ЈE). 13 November 1997 June 1995 Locality 29. Approximately 400 m north-north- Locality 14. Northwest corner of Pelago Island, west of Pelago Island, 44±61 m depth isolated boulders on sand bottom (SCUBA), (45Њ02.4ЈN, 13Њ36.5ЈE). 13 November 1997, 9 30±50 m depth (45Њ03.2ЈN, 13Њ36.6ЈE). 20 June July 1998 1990 Locality 30. CMRR oceanographic station SJ-311, Locality 15. Leso Bay, Rovinj, midway between offshore from Pula, sand bottom (SCUBA), 45 Figarola Island and mainland, sand bottom m depth (44Њ37.2ЈN, 13Њ 40.0ЈE). 4±6 June (dredge), 14±17 m depth (45Њ05.4ЈN, 1988, 25 September 1990 13Њ37.4ЈE). 9 July 1998 Locality 31. Channel between Sv. Ivan Island and Locality 16. West side of Banjole Island, sand Pelago Island, sand bottom (dredge), 50±60 m platform (SCUBA), 17.5 m depth (45Њ03.9ЈN, depth (45Њ02.3N, 13Њ36.7ЈE). 3 June 1988, 23 13Њ36.5ЈE). 3 July 1998 June 1988, 3 June 1997 Locality 17. Approximately 1 km north of Figar- Locality 32. Harbor by CMRR, concrete wall ola Island, 500 m off mainland, sand bottom (SCUBA), 0±5 m depth (45Њ04.9ЈN, (dredge), 20±22 m depth (45Њ05.6ЈN, 13Њ38.15ЈE). 3 July 1998 13Њ36.7ЈE). 3 June 1997, 13 November 1997 Locality 33. Mussel-culture station, Lim Channel, Locality 18. Approximately 200 m offshore of Rt ropes, tiles, and wooden frame of platform Kriz, sand bottom (dredge), 20 m depth (SCUBA), 0±2 m depth; pulsating salinity (45Њ03.8ЈN, 13Њ37.2ЈE). 26 May 1988 changes due to actively mixing marine and Locality 19. Channel between Banjole Island and fresh spring waters (45Њ07.4ЈN, 13Њ 40.9ЈE). 24 Sv. Katarina Island, 150 m from S. Catarina, September 1987, 10 June 1988 Locality 33a. Shell debris from mussel-culture sand bottom (dredge), 22 m depth (45Њ04.1ЈN, station, collected from channel ¯oor immedi- 13Њ37.2ЈE). 20 June 1988 ately below. 24 September 1987 Locality 20. Approximately 3 km north of Rovinj, Locality 34. Outer dock, Rovinj Harbor, concrete 300 m offshore, sand bottom (dredge), 25 m block (grapple hook), 4 m depth (45Њ04.4ЈN, depth (45Њ05.7ЈN, 13Њ36.5ЈE). 29 September 13Њ37.6ЈE). 12 November 1997 1987, 21 September 1990, 6 June 1995 Locality 35. Approximately 9 km west of Istrian Locality 21. Approximately 1 km offshore of Rt Coast, ®ne sand bottom (Van Veen grab), 24.5 Kriz, sand bottom (dredge), 29 m depth m depth. Specimens provided by A. Jaklin. (45Њ03.8ЈN, 13Њ36.7ЈE). 15 June 1988, 4 Octo- (45Њ26.1ЈN, 13Њ3.8ЈE). 25 September 1991 ber 1990, 12 October 1990 Locality 36. Mud with low proportion of skeletal Locality 22. Channel between Sv. Ivan Island and detritus (probably SCUBA), 35 m depth (45Њ Pelago Island sand bottom (dredge), 30±42 m 01.1ЈN, 12Њ59.7ЈE). 30 October 1990 depth (45Њ02.3N, 13Њ36.7ЈE). 15 June 1988, 2 Locality 37. Approximately 560 m west of Figar- October 1990, 8 June 1995, 9 July 1998 ola, ®ne sand bottom (dredge), 20±30 m depth Locality 23. Sediment adjacent to west side of (45Њ5.3ЈN, 36Њ36.8ЈE). 27 June 1984 base of Banjole Island, sand bottom (SCUBA), 31 m depth (45Њ03.9ЈN, 13Њ36.5ЈE). 3 July 1998 SPECIES DISTRIBUTIONS Locality 24. Approximately 200 m southwest of Banjole, sand bottom (dredge), 32 m depth Any taxonomic survey risks incomplete- (45Њ03.8ЈN, 13Њ34.2ЈE). 23 June 1988 ness, and incompleteness in benthic marine Locality 25. Approximately 3 km south of Sv. surveys of topographically complex areas is Ivan Island, sand bottom (dredge), 33±34 m virtually assured because sampling is limited depth (450Њ0.7ЈN, 13Њ37.1ЈE). 9 July 1998 and because locally unique environments Locality 26. Channel between Banjole Island and may be missed completely. In addition, some Sv. Katarina Island, midway; sand bottom organisms are simply more dif®cult to see (dredge), 33±35 m depth (45 04.1 N, Њ Ј than are others. It is likely that many cteno- 13Њ36.9ЈE). 2 October 1990, 4 October 1990 Locality 27. ''Cellaria meadow,'' west of Banjole stome bryozoans are present in the region in- Island, mud bottom (SCUBA), 35 m depth cluded in this paper but were not found by (45Њ04.0ЈN, 13Њ36.3ЈE). 29 September 1987, 7 us because of their small size, inconspicuous June 1988, 9 October 1990, 3 June 1997, 18 growth habits, neutral color, and tendency to November 1997, 3 July 1998, 7 July 1998, 15 grow within tangled mats of algae and hy- July 1998 droids. Some were spotted in the laboratory Locality 28. Approximately 4 km southeast of Sv. when their tentacles were protruded to feed- 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 9 ing positions above the mats, but any others AL: Avicularium length (type de®ned in within the mats that did not feed while in the text of species description) ®eld of view of the microscope would have AW: Avicularium width (type de®ned in text been missed completely. of species description) Meadows of the marine grass Posidonia CL: Cauda length were formerly present in the region of Rov- DO: Distance between midpoints of adja- cent ori®ces inj but died off in the early 1970s (D. Za- KL: Kenozooid length vodnik, 1977; N. Zavodnik, 1983). Such KW: Kenozooid width marine grasses carry a diverse bryozoan LW: Lyrula width fauna further south in the Adriatic and else- ML: Mandible length (avicularia) where in the Mediterranean (Harmelin, MW: Mandible width (avicularia) 1973a; Nicoletti et al., 1995) and part of that OL: Ori®ce length fauna is restricted to growth on the grasses OOL: Ovicellate ori®ce length (where differ- (e.g., Electra posidoniae Gautier, 1954) or ent from normal ori®ce width) occurs preferentially on them (e.g., Platonea OOW: Ovicellate ori®ce width (where differ- stoechas Harmelin, 1976). Posidonia cur- ent from normal ori®ce width) rently is extending its range back up the OpL: Opesia length eastern Adriatic coast toward Rovinj (N. Za- OpW: Opesia width vodnik, personal commun.). Should fully OvL: Ovicell length OvW: Ovicell width developed meadows of Posidonia and other OW: Ori®ce width grasses again occur there, it is likely that PS: Distance between pores in frontal locally absent epiphytic bryozoans will also shield reappear. SOD: Secondary ori®ce diameter Our collections inevitably underrepresent VAL: Vicarious avicularium length environments and depth distributions of VAW: Vicarious avicularium width many of the bryozoan species that we found. ZL: Autozooid length as seen on colony Where species apparently have notable re- surface strictions in depth or environment, we men- ZW: Autozooid width as seen on colony sur- tion this in a note on Occurrence in the spe- face. cies descriptions. For all other species, oc- Measurements on cyclostome skeletons currence data are summarized in appendix 1. are identi®ed by these acronyms and abbre- viations: MEASUREMENTS AND COUNTS AD: Aperture diameter Some zooidal measurements were made ADMN: Minimum aperture diameter optically at ϫ50 using a Wild M8 stereomi- ADMX: Maximum aperture diameter AlvD: Alveoli diameter croscope with an ocular micrometer. Optical AS: Distance between midpoints of adja- measurements were made in units of 10 ␮m. cent apertures Most measurements were made through ste- ASW: Distance between midpoints of adja- reomicroscopes ®tted with a digital camera cent apertures within a row or fascicle and recorded electronically with National In- BrD: Branch diameter or width stitute of Health image analysis software. Gap: Distance between edges of adjacent Each measurement or count is given in the fascicles text as mean plus or minus standard devia- GL: Length of in¯ated portion of gonozooid tion, observed range, and (enclosed in paren- GW: Width of in¯ated portion of gonozooid theses) number of specimens used and total OsD: Ooeciopore diameter number of measurements or counts made. OsDMN: Ooeciopore minimum diameter Except where noted, measurements are given OsDMX: Ooeciopore maximum diameter in microns. Measurements and counts for PD: Peristome diameter (exterior) at outer end cheilostome skeletons are identi®ed by the RS: Distance between centers of adjacent following acronyms and abbreviations: zooecial rows or fascicles. AAL: Adventitious avicularium length Measurements and counts of polypide AAW: Adventitious avicularium width structures are identi®ed by these acronyms: 10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

IH: Introvert height of autozooid slender, ®liform, ca. 0.06 mm LD: Lophophore diameter wide, of variable length, linked with preced- LDMn: Minimum lophophore diameter ing autozooid. Buds developed around edges LDMx: Maximum lophophore diameter of expanded base of tube, developing as ®- MD: Mouth diameter liform proximal portions of new autozooids. MDMn: Minimum mouth diameter MDMx: Maximum mouth diameter Tentacles clear; number reported to vary TL: Tentacle length between 16 and 20, but 18±22 in present ma- TLMn: Minimum tentacle length terial. Lophophores bell-shaped, radially TLMx: Maximum tentacle length. symmetrical. DISTRIBUTION: Nolella dilatata is common We have not provided complete synony- in shallow coastal waters throughout the mies for each species; reference to the orig- northeast Atlantic and the Mediterranean, but inal description is given, together with ref- it is inconspicuous and easily overlooked. It erence to signi®cant sources relating either to encrusts a variety of substrata but is most the Adriatic or to the Mediterranean in gen- frequent on other sessile invertebrates, in- eral. More extended synonymies are provid- cluding bryozoans. ed for formerly taxonomically confused spe- cies, and for all species synonymic histories FAMILY WALKERIIDAE HINCKS, 1880 may be found in the major sources cited (emend. Bassler, 1953) (e.g., Prenant and Bobin, 1956; Hayward, 1985; Hayward and Ryland, 1985a, 1998, GENUS WALKERIA FLEMING, 1823 1999). Walkeria tuberosa (Heller, 1867) For the Cheilostomata, we follow the fam- Figure 2B, C ily to suborder working classi®cation of Den- nis P. Gordon (1989, personal commun.). Valkeria tuberosa Heller 1867: 128. Prenant and This is the classi®cation that is being devel- Bobin, 1956: 255. oped for eventual use in the revised Treatise Walkeria tuberosa: Zabala and Maluquer, 1988: on invertebrate paleontology. Classi®cation 70. of the Ctenostomata and Cyclostomata is lit- DESCRIPTION (AMNH 888; CMRR 2201): tle modi®ed from that of Hayward (1985) Colonies thin, adherent kenozooidal stolons, and Hayward and Ryland (1985a), respec- 35±50 ␮m wide, with clumps of erect, cylin- tively. drical autozooids at intervals of 2±3 mm. At each such point the stolon buds on each side SYSTEMATIC ACCOUNTS a fan of ¯at, quadrangular kenozooids, 50± CLASS GYMNOLAEMATA ALLMAN, 1856 100 ␮m diameter, each giving rise to a single ORDER CTENOSTOMATA BUSK, 1852 autozooid; groups consist of up to 30 auto- zooids on each side of the stolon. Autozooids FAMILY NOLELLIDAE HARMER, 1915 cylindrical, but with a square-sectioned distal GENUS NOLELLA GOSSE, 1855 portion; abruptly constricted proximally and Nolella dilatata (Hincks, 1860) thus appearing shortly stalked; up to 0.6 mm Figure 2A long in present material, with maximum width 0.2 mm. The small kenozooids also Farrella dilatata Hincks, 1860: 279. bud further stolonal kenozooids and the col- Nolella dilatata Nordgaard, 1918: 14. Hayward, ony develops a ramifying form, creeping 1985: 90. over the substratum. DESCRIPTION (AMNH 887; CMRR 2200): Tentacles 8, clear; lophophores radially Colony encrusting. Autozooids clumped or symmetrical, slightly concave-sided conical. diffuse, each consisting of an erect, cylindri- OCCURRENCE: Epizoic on other bryozoans, cal portion, 1.0±2.5 mm long in present ma- especially Cellaria and Amathia. terial, of more or less constant width, ca. DISTRIBUTION: Originally described from 0.25 mm, not narrowed at base, but instead the Adriatic, W. tuberosa has been recorded broadened to an expanded, ¯at base adherent sporadically from elsewhere in the Mediter- to substratum. Proximal, encrusting portion ranean, from the Red Sea (Waters, 1910) and 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 11

Fig. 2. A. Nolella dilatata (AMNH 887; 0.4 mm). B, C. Walkeria tuberosa. B. Single clump of zooids (AMNH 888; 0.1 mm). C. Three stolon-connected clumps of zooids on branch of Cellaria (AMNH 888; 0.3 mm). (In this and subsequent ®gures, length of the scale bar will be indicated in parentheses.) from the Indo-Mayalan region (Harmer, lantic region, and it is not certain even 1915; SileÂn, 1942). whether any of the ®ve species known from MEASUREMENTS (CYSTID): ZL 527 Ϯ 42 north European coasts are the same as those ␮m, 480±560 (1, 7), LD 342 Ϯ 20, 320±360 recorded from northeast American coasts. (1, 10), MD 20 (1, 1), TL 344 Ϯ 18, 320± Three species have been reported from Med- 380 (1, 10). iterranean localities, namely Bowerbankia pustulosa (Ellis and Solander, 1786) and FAMILY VESICULARIIDAE HINCKS, 1880 Bowerbankia imbricata (Adams, 1798), both GENUS BOWERBANKIA FARRE, 1837 of which may develop dense shrubby colo- Figure 3 nies, and B. gracilis Leidy, 1855, which typ- The taxonomy of Bowerbankia poses ically grows as diffuse, creeping colonies of problems outside of the temperate north At- small autozooids, individually spaced along 12 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 3. Nonpreserved ctenostomes. A. Small Bowerbankia inhabiting Tubulipora skeleton. B. Bow- erbankia-like form with large lophophore. C. Stolonate ctenostomate with campylonemidan lophophore of 10 tentacles. D. Robust stolonate ctenostomate with 16±17 tentacles. (0.5 mm) a very slender, inconspicuous stolon. Bow- Tubulipora (®g. 3A). The lophophore erbankia caudata Hincks, 1880, character- was a radially symmetrical bell of eight ized by proximally tapered, or ``caudate'' au- tentacles; this is consistent with B. tozooids, has for long been synonymized gracilis, but there are no other grounds with B. gracilis, yet two genetic species of for identifying it with this taxon. ``B. gracilis'' are present in the Swansea re- 2. A large species (®g. 3B) with a short gion of South Wales (Hayward, 1985), and cylindrical collar and an elongate radi- it is probable that several others exist within ally symmetrical bell of eight tentacles. the wide geographic range accorded to this 3. A slightly larger epizoic species (®g. taxon. Pending the results of current molec- 3C) with a diffuse colony form of dis- ular genetic investigations into the taxonomy persed zooids on an unseen stolon. The of Bowerbankia, species can only be distin- lophophore was a bilaterally symmet- guished by such characters as the shape of rical, campylonemidan bell with 10 the lophophore and the number of tentacles, tentacles. embryo color, breeding period, and colony Finally, an unidenti®ed stolonate cteno- organization. Unfortunately, species with ``B. stomate (®g. 3D) with a large lophophore of gracilis'' morphology are cryptic in habit at least 16 tentacles and a ¯ared collar to and usually only become apparent during the which much debris adheres must remain later stages of sample sorting. Three small completely unattributed: no species of Bow- species of stoloniferan ctenostomates were erbankia has such a large lophophore and it present in the Rovinj collections; each was is unclear to which genus this species be- photographed, but for none is there suf®cient longs. morphological or other data to be certain of its taxonomic identity, and they are here at- GENUS AMATHIA LAMOUROUX, 1812 tributed, tentatively, only to the genus Bow- Amathia pruvoti Calvet, 1911 erbankia, as follows: Figure 4A 1. A small species with a short cylindrical Amathia pruvoti Calvet, 1911: 59. Hayward, collar, inhabiting empty zooid tubes of 1985: 136. Chimonides, 1987: 336. 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 13

Fig. 4. A. Amathia pruvoti (AMNH 889; 0.5 mm). B. Amathia vidovici (UIIZ 366; 0.5 mm).

DESCRIPTION (AMNH 889; CMRR 2202): stressing the signi®cance of growth pattern Colonies sparse, erect tufts, to 2 cm in pre- for discriminating between them. sent material, of stiff, kenozooidal stolons, DISTRIBUTION: Amathia pruvoti is wide- colorless at extremities but light brown ba- spread and common in shallow coastal wa- sally; attached by an encrusting portion of ters throughout the Mediterranean, and on stolon, without additional rhizoids. Stolon of the southwest coasts of the British Isles. constant width, ca. 0.175 mm, not tapering distally, bifurcating at intervals of 3±6 mm; Amathia vidovici (Heller, 1867) slightly curved prior to the dichotomy, one Figure 4B ramus de¯ected at approximately 40Њ to main Valkeria vidovici Heller, 1867: 128. axis, subsidiary ramus at ca. 60Њ and alter- Amathia vidovici: Prenant and Bobin, 1956: 283. nating left and right along the major axis. DESCRIPTION: A single small tuft of this Autozooids cylindrical, closely adherent to species was collected, 7.5 mm high, com- stolon, arranged in tight biserial groups, each prising two stolon bifurcations and six au- originating proximal to the dichotomy at tozooid groups. Stolon 0.2 mm wide; auto- one-third to one-half distance between it and zooid groups each consisting of a single preceding dichotomy. Six to 10 pairs of au- proximal autozooid and about eight pairs, all tozooids in each group, with a single, un- about the same height. Each group forming paired proximal autozooid; forming a half a complete spiral of 360Њ around the stolon, spiral of about 90Њ around stem, with auto- usually commencing just proximal to a bi- zooid length decreasing successively distally. furcation and enclosing bases of the two REMARKS: Chimonides (1987) provided a daughter kenozooids; one group of auto- detailed analysis of colony morphology in zooids was situated midway between two this and several other species of Amathia, successive bifurcations, a characteristic of 14 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 the species noted by Prenant and Bobin Tentacles clear, 10; lophophores broadly (1956). Basal attachment of A. vidovici re- conical, radially symmetrical. mains quite unknown (Chimonides, 1987). DISTRIBUTION: Occurs on a wide variety of REMARKS: We could not locate any type organic and inorganic substrata, in shallow material of Valkeria vidovici in the UIIZ col- coastal waters throughout the Mediterranean. lections; instead, we illustrate part of a spec- Aetea sica has been recorded from most of imen of A. vidovici from an unspeci®ed lo- the world's seas, but its actual Distribution is cality in the Adriatic, in the collections of the probably more limited. University of Innsbruck Institute of Zoology MEASUREMENTS (POLYPIDE): IH 60 ␮m (1, (specimen 366). Although little material of 1), LD 200±280 (2, 2), TL 250±420 (2, 2). this species was collected, its morphology is recognizably that of A. vidovici, which was Aetea truncata (Landsborough, 1852) named for material from the Adriatic (Heller, Figure 5D±F 1867). Anguinaria truncata Landsborough, 1852: 288. DISTRIBUTION: Amathia vidovici is widely Aetea truncata: Gautier, 1962: 29. Hayward and distributed in shallow coastal waters of the Ryland 1998: 104. Mediterranean, and ranges north to Roscoff, Brittany. It has been recorded in the western DESCRIPTION (AMNH 892, 893; CMRR Atlantic form Colombia to Massachusetts, 2204): Colonies white, uniserial, continuous- from the Cape Verde Isles and from east Af- ly or intermittently encrusting. Adnate sto- rica, a broad geographical range which per- loniform portion of autozooid particularly haps requires reinvestigation. slender, as is basal dilatation of erect distal portion. Relatively short distal portion of au- ORDER CHEILOSTOMATA BUSK, 1852 tozooid ®nely and densely punctate, lacking SUBORDER INOVICELLATA JULLIEN, 1888 any annulations. Colonies collected 26 May 1988 offshore from Rt Kriz had irregular un- SUPERFAMILY AETEOIDEA SMITT, 1867a attached growth form reported in other spe- FAMILY AETEOIDAE SMITT, 1867a cies of Aetea (Hayward and Ryland, 1998) GENUS AETEA LAMOUROUX, 1812 when associated with hydroid and bryozoan turfs. They were loosely intergrown with a Aetea sica (Couch, 1844) tuft of Savignyella lafontii. Stoloniform por- Figure 5A±C tions were up to 0.125 mm long but just 0.025 mm wide, broadening abruptly to 0.07 Hippothoa sica Couch, 1844: 102. mm at base of distal portion, which was re- Aetea sica: Alvarez, 1990: 24. Hayward and Ry- land, 1998: 102. ¯ected at an angle of about 120Њ to long axis of stoloniform portion. Distal portion, ac- DESCRIPTION (AMNH 890, 891; CMRR commodating the polypide, was 0.45±0.5 2203): Colonies forming delicate, branching mm long, cylindrical proximally and slightly traceries over biogenic substrata, especially broadened to a maximum width of 0.12 mm shell, with autozooids in branching linear at midlength. Opesia occupying about half of chains. Encrusting proximal portion of au- total length of this part of autozooid. Distal- tozooid consisting of a ®liform part of vari- ly, autozooid abruptly truncate, bearing a ter- able length, but just 0.05 mm wide, broad- minal operculum. Branching varied: new sto- ening abruptly to a substantial fusiform part loniform buds arising most frequently from 0.5 mm long and 0.2 mm wide, from which basal dilatation of autozooid, either basally the erect distal portion arises. This also varies or laterally, or from basal (i.e., abfrontal) sur- in length, to 1.5 mm in present material, with face of distal portion of autozooid, or even a basal width of 0.1 mm, gently broadening from various points on stolonifom parts of to about 0.15 mm at the distal end. Opesia autozooids. occupying distal third of erect portion. Cal- Tentacles clear, 11±12, usually 12; lopho- ci®cation thin and delicate, that of encrusting phore broadly conical, radially symmetrical. portion smooth and imperforate, erect por- REMARKS: Geometry of adnate and erect tion ®nely but distinctly annulate. portions of autozooids, together with punc- 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 15

Fig. 5. A±C. Aetea sica. A. Group of zooids with adnate portion covered by thin sponge (AMNH 890; 0.5 mm). B. Erect portion of zooid, including opesia with terminal ori®ce (AMNH 891; 0.1 mm). C. Annulae around base of erect portion of zooid (AMNH 890; 0.1 mm) D±F. Aetea truncata. D. Group of zooids encrusting radula-scored shell surface (AMNH 892; 0.5 mm). E. Erect portion of zooid, including opesia and inclined terminal ori®ce (AMNH 893; 0.1 mm). F. Smooth base of erect portion of zooid (AMNH 893; 0.1 mm). 16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 tuation and lack of annulations suf®ce to dis- often epifaunal in habit, but is rather incon- tinguish this species from all other Mediter- spicuous and frequently overlooked. ranean species of Aetea. Balduzzi et al. (1991) noted that the super®cially similar A. SUBORDER MALACOSTEGA LEVINSEN, 1902 lepadiformis Waters has a very few, but dis- SUPERFAMILY MEMBRANIPOIDEA BUSK, 1852 tinct annulations immediately proximal to the FAMILY ELECTRIDAE D'ORBIGNY, 1851 opesia GENUS PYRIPORA D'ORBIGNY, 1852 OCCURRENCE: This species was common to abundant at most stations where it occurred, Pyripora sp. (Fleming, 1828) encrusting dead shell and as an epizooite, es- Figure 6B, C pecially of other bryozoans. DESCRIPTION (AMNH 895): Colony en- DISTRIBUTION: Aetea truncata has been re- crusting, runner-like, inconspicuous. Auto- ported from temperate and tropical seas zooids in uniserial chains, branching irregu- throughout the world but, especially follow- larly; clavate, with a slender, ®liform proxi- ing the work of Balduzzi et al. (1991), it mal portion broadening at about half total must be considered probable that such a wide length to an oval distal portion. Calci®cation Distribution encompasses more than one spe- gymnocystal, smooth; opesia oval, occupy- cies. ing three-quarters length of distal portion, MEASUREMENTS (POLYPIDE): IH 46 36 Ϯ bordered by a very narrow, indistinct cryp- m, 20±120 (4, 17), LD 421 57, 280±530 ␮ Ϯ tocystal rim. No spines or polymorphs. Each (4, 29), MD 21.7 2.5, 20±25 (3, 9), TL Ϯ autozooid giving rise to a single distal bud, 299 55, 160±440 (4, 27). Ϯ and may also produce paired lateral buds, al- lowing branching of the chains. SUBORDER SCRUPARIINA SILEÂ N, 1941 REMARKS: The single colony of Pyripora SUPERFAMILY SCRUPARIOIDEA BUSK, 1852 in our material differs from P. catenularia FAMILY SCRUPARIIDAE GRAY, 1848 (Fleming, 1828), the species previously re- GENUS SCRUPARIA OKEN, 1815 ported from the northeastern Atlantic and Mediterranean, in lacking a well-developed Scruparia ambigua (d'Orbigny, 1841) cryptocystal rim around the opesia, having a Figure 6A proportionally longer cauda, and having Eucratea ambigua d'Orbigny, 1841: pl. 3, ®gs proximolateral rather than more distolateral 13±17. budding. Scruparia ambigua: Hayward and Ryland, 1998: MEASUREMENTS (SKELETAL): OpL 155 Ϯ 12 108. ␮m, 29±168 (1, 10), OpW 81 Ϯ 8, 70±93 (1, 10), ZL 548 Ϯ 50, 480±621 (1, 10), ZW DESCRIPTION (AMNH 894): Colony erect, 155 Ϯ 9, 141±170 (1, 10). branching, uniserial chains of horn-shaped autozooids, budded from encrusting chains SUBORDER FLUSTRINA SMITT, 1867b of identical autozooids. Frontal surface of au- SUPERFAMILY CALLOPOROIDEA NORMAN, tozooid with an oval opesia occupying half 1903 its length, bounded by a thin raised rim. Em- bryos brooded in dimorphic zooids with a FAMILY CALLOPORIDAE NORMAN, 1903 distinctive bivalved brood chamber (absent GENUS CALLOPORA GRAY, 1848 from present material). Callopora dumerilii (Audouin, 1826) OCCURRENCE: A portion of one colony, Figure 6D±G comprising two encrusting autozooids and two erect shoots, of three and one complete Flustra dumerilii Audouin, 1826: 240. autozooids, encrusting Cellaria ®stulosa Callopora dumerilii; Hayward and Ryland, 1998: (Linnaeus, 1758). 160. DISTRIBUTION: This rather inconspicuous DESCRIPTION (AMNH 896±898; CMRR species is widely distributed throughout the 2205): Colonies encrusting, multiserial, uni- Mediterranean, and has been reported from laminar patches. Ancestrula approximately temperate and tropical seas worldwide. It is 225±250 ␮m long, with 10 peripheral spines; 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 17

Fig. 6. A. Scruparia ambigua (AMNH 894; 0.5 mm). B, C. Pyripora sp. B. Group of zooids; note proximo-medial budding (AMNH 895; 0.5 mm). C. Single zooid (AMNH 895; 0.1 mm). D±G. Callo- pora dumerilii. D. General aspect (AMNH 896; 0.5 mm). E. Maternal zooid with ovicell (AMNH 897; 0.1 mm). F. Autozooid (AMNH 896; 0.1 mm). G. Ancestrula (AMNH 898; 0.1 mm). 18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 number of spines reduced and autozooids en- as a coarsely granular rim, narrowed slightly larged over about 5±8 generations. Auto- at distal end of autozooid, opesia constituting zooids elongate oval, broadest proximally, almost three-quarters total autozooid length. separated by distinct grooves. Opesia oval, Spines absent, except in ®rst three or four bordered by a thin crenulate mural rim, en- astogenetic generations. Avicularia interzooi- closing a narrow shelf of ®nely granular dal, about half length of autozooid; mandible cryptocyst, widest at the proximal end; gym- slender, distally directed, lodged within a nocyst smooth, imperforate. Four distal oral narrow, parallel-sided rostrum formed from spines present, the proximalmost pair robust incurved gymnocystal calci®cation, pivoting and conspicuous. A single adventitious avi- on two pronounced condyles, proximal to cularium (rarely two) present on the gym- which there is a broad shelf of cryptocyst. nocyst of most autozooids, with short, co- Ovicell longer than wide, domed, with a dis- lumnar cystid; rostrum acute to frontal plane, tinctive coarsely nodular surface. Ancestrula triangular, distally hooked, usually directed oval, with minimal development of crypto- proximally; with a narrow cryptocyst bor- cyst, bordered by (?) eight slender spines; au- dering the proximal opesia, but no crossbar. tozooids of the ®rst three or four astogenetic Ovicell hyperstomial, prominent, more or generations occasionally with single spines less spherical, but narrowed towards arched proximally, and two or more distally, all de- aperture; ectooecium membranous except for veloped on the gymnocyst. a basal peripheral rim, entooecium coarsely DISTRIBUTION: On stones and shells, in nodular. Distal to an ovicell, avicularium dis- shallow coastal habitats; C. tenuirostre is placed laterally, with a distolateral orienta- widely distributed in the Mediterranean and tion. north to the western English Channel. Tentacles clear, 11±12; lophophores bell- MEASUREMENTS (SKELETAL): AL 229 Ϯ 31 shaped, radially symmetrical. ␮m, 167±298 (2, 15), AW 123 Ϯ 14, 94±155 DISTRIBUTION: Common throughout the (2. 15), DO 376 Ϯ 63, 256±490 (3, 30), OpL temperate northeast Atlantic and Mediterra- 315 Ϯ 29, 268±381 (3, 30), OpW 205 Ϯ 24, nean, on stone, shell and other organic sub- 162±256 (3, 30), OvL 177 Ϯ 9, 162±190 (2, strata, in shallow coastal habitats. 7), OvW 190 Ϯ 14, 176±217 (2, 7), ZL 484 MEASUREMENTS (SKELETAL): DO 368 Ϯ 52 Ϯ 56, 391±560 (3, 30), ZW: 322 Ϯ 26, 279± ␮m, 285±497 (3, 30), OpL 294 Ϯ 30, 222± 379 (3, 30). 350 (3, 25), OpW 208 Ϯ 16, 180±240 (3, 25), OvL 177 Ϯ 17, 156±226 (3, 20), OvW GENUS ELLISINA NORMAN, 1903 172 Ϯ 21, 129±216 (3, 20), ZL 507 Ϯ 37, Ellisina gautieri FernaÂndez Pulpeiro and 424±590 (3, 25), ZW 307 Ϯ 25, 260±357 (3, Reverter Gil, 1993 25). (POLYPIDE): IH 30±80 ␮m (1, 2), LD 373 Figure 7E±G Ϯ 31, 340±400 (1, 3), MD 20 (1, 1), TL 287 Ϯ 23, 260±300 (1, 3). Ellisina gautieri FernaÂndez Pulpeiro and Reverter Gil, 1993: 98. Hayward and Ryland, 1998: 192. GENUS COPIDOZOUM HARMER, 1926 DESCRIPTION (AMNH 901): Colonies en- Copidozoum tenuirostre (Hincks, 1880) crusting, multiserial, unilaminar patch. Au- Figure 7A±D tozooids oval, separated by distinct grooves. Frontal surface almost entirely membranous, Membranipora tenuirostre Hincks, 1880: 70. the extensive opesia bordered by a very nar- Copidozoum tenuirostre: Prenant and Bobin, row rim of granular cryptocyst, and with a 1966: 257. Hayward and Ryland, 1998: 180. minimal area of smooth gymnocyst just vis- DESCRIPTION (AMNH 899, 900): Colonies ible at proximal end of autozooid. No spines. encrusting, multiserial, unilaminar. Auto- Avicularia interzooidal, budded from distal zooids elongate oval, distinctly separated by pore chamber of most autozooids; cystid well-marked grooves. Frontal surface of au- rounded quadrangular, ca. 0.125 mm long, tozooid almost entirely membranous; gym- rostrum triangular, acute to frontal plane, di- nocyst reduced to a smooth, proximal and rected distolaterally. Two or more avicularia proximolateral border; cryptocyst developed may occur at the division of autozooid rows, 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 19

Fig. 7. A±D. Copidozoum tenuirostre. A. General aspect (AMNH 899; 0.5 mm). B. Autozooid (AMNH 899; 0.1 mm). C. Maternal zooid with ovicell (AMNH 899; 0.1 mm). D. Ancestrula and early autozooids (AMNH 900; 0.1 mm). E±G. Ellisina gautieri (AMNH 901). E. General aspect (0.5 mm). F. Autozooids (left) and maternal zooids with ovicells bearing distal avicularia (0.1 mm). G. Ancestrula and early autozooids (0.1 mm). 20 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 or where regular spacing of autozooids is dis- distal tips of some branches. Ovicell hemi- turbed. Ovicell slightly longer than wide, spherical, completely immersed, conspicuous domed, recumbent on substratum between through its white calci®cation. distally succeeding autozooids; smooth and Tentacles colorless, 13±15; lophophores imperforate, with a distinct, longitudinal bell-shaped, radially symmetrical. frontal ridge and intimately associated with Light orange embryos present in material distal avicularium; closed by zooidal oper- from locality 21, 15 June 1988. culum. DISTRIBUTION: Chartella tenella is an en- REMARKS: This species was ®rst noted demic Mediterranean species, common in from Marcella by Gautier (1962), as E. cf. shallow coastal habitats, and offshore to 200 levata, and its identity was only recently es- m (Prenant and Bobin, 1966). It has been tablished by FernaÂndez Pulpeiro and Revert- widely reported throughout the western Med- er Gil (1993). iterranean and the Adriatic. DISTRIBUTION: E. gautieri is known from MEASUREMENTS (SKELETAL): AL 314 ␮m the Atlantic coast of northwest Spain and (1, 1), AW 221 (1, 1), DO 444 Ϯ 84, 301± from the western English Channel; the single 632 (2, 20), OL 76 Ϯ 8, 62±92 (1, 10), OW small colony described here represents its 134 Ϯ 8, 121±148 (1, 10), ZL 685 Ϯ 81, ®rst record from the eastern Mediterranean. 585±841 (2, 20), ZW 250 Ϯ 36, 184±321 (2, MEASUREMENTS (SKELETAL): AL 158 Ϯ 22 20). (POLYPIDE): IH 201 Ϯ 74 ␮m, 60±340 ␮m, 131±198 (2, 20), AW 140 Ϯ 20, 113± (2, 24), LD 595 Ϯ 94, 440±700 (2, 27), MD 184 (2, 20), DO 359 Ϯ 36, 296±424 (2, 20), 20 (1, 1), TL 534 Ϯ 62, 400±650 (2, 24). OpL 346 Ϯ 27, 304±387 (2, 20), OpW 205 Ϯ 15, 182±246 (2, 20), ZL 521 Ϯ 42, 465± SUPERFAMILY BUGULOIDEA GRAY, 1848 596 (2, 20), ZW 296 Ϯ 14, 274±325 (2, 20). FAMILY BUGULIDAE GRAY, 1848 GENUS BUGULA OKEN, 1815 SUPERFAMILY FLUSTROIDEA FLEMING, 1828 FAMILY FLUSTRIDAE FLEMING, 1828 Bugula fulva Ryland, 1960 Figure 8C±E GENUS CHARTELLA GRAY, 1848 Bugula fulva Ryland, 1960: 86. Hayward and Ry- Chartella tenella (Hincks, 1887) land, 1998: 218. Figure 8A, B DESCRIPTION (AMNH 903, 904; CMRR Flustra tenella Hincks, 1887: 313. 2207): Colonies erect, branching, developing Chartella tenella: Gautier, 1962: 48. as stiff, open fans, to 10 mm in present ma- Termino¯ustra tenella: Prenant and Bobin, 1966: 191. terial but recorded up to 30 mm. Branches unilaminar, dividing by regular dichotomy, DESCRIPTION (AMNH 902; CMRR 2206): consisting of two longitudinal autozooid se- Colonies erect, ¯abellate, bilaminar, lightly ries, increasing to four just prior to each di- calci®ed, to 20 mm in present material; di- chotomy. Autozooids elongate, narrow, fron- viding irregularly at varying intervals, de- tal dimensions ca. 0.6 ϫ 0.2 mm, slightly veloping a narrowly lobed, frondose form. tapered proximally; frontal surface entirely Autozooids rectangular, ¯at, 0.65 ϫ 0.25 membranous. Usually three large spines at mm. Frontal surface entirely membranous, each distal corner, sometimes ®ve on en- vertical walls lightly calci®ed; a short, thin closed autozooids proximal to a dichotomy, and very inconspicuous spine present at each the distalmost spine on the outer corner es- distal corner. Avicularia vicarious, at bifur- pecially robust. Most autozooids with a lat- cation of autozooid rows; cystid rectangular, eral avicularium, attached on outer edge of 0.25 ϫ 0.2 mm, thickly calci®ed, with sem- frontal membrane just below proximal-most ielliptical rostrum slightly acute to frontal spine; length slightly greater than the auto- plane, directed distally or just oblique to dis- zooid width, but variable and locally less tal axis. Kenozooids ®lling spaces between than width on some autozooids; beak gently autozooids where bifurcating branches di- downcurved. Ovicell large, globular, with a verge and developed as tubular extensions at wide, arched aperture; entooecium with ®ne 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 21

Fig. 8. A, B. Chartella tenella. A. General aspect (AMNH 902; 0.5 mm). B. Autozooids and vi- carious avicularium (right center; AMNH 902; 0.5 mm). C±E. Bugula fulva. C. Entire colony (AMNH 903; 1 mm). D. Biserial branch (AMNH 904; 0.5 mm). E. Detailed view of avicularia, oral spines, and ovicells (AMNH 904; 0.1 mm). 22 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 radiating striations. Ancestrula distinctive: 405 (2, 20), OvL 212 Ϯ 16. 185±242 (2, 20), very elongate, slender, trumpet-shaped; tu- OvW 194 Ϯ 9, 180±209 (2, 20), ZL 577 Ϯ bular proximal portion to 2.5 mm in present 40, 500±660 (2, 20), ZW 178 Ϯ 20, 130± material, with oval distal opesia bordered by 220 (4, 40). (POLYPIDE): IH 40 ␮m (1.10), LD six distal and three proximal spines, some- 450 Ϯ 54, 360±540 (1, 10), MD 20 (1, 1), times with an additional lateral pair between. TL 460 Ϯ 51, 400±560 (1, 10). Tubular rhizoids budded from proximal re- gion of ancestrula anchor the colony. FAMILY BEANIIDAE CANU AND BASSLER, 1927 REMARKS: Ancestrulae were present in ev- GENUS BEANIA JOHNSTON, 1840 ery colony in 1997 and 1998 samples from the 35 m deep Cellaria meadow west of Ban- Beania cylindrica (Hincks, 1886) jole Island. They were remarkable for their Figure 9A, B length, resembling the basal kenozooids of Diachoris hirtissima forma cylindrica Hincks, colonies of Caulibugula, which caused this 1886: 263. local population to be incorrectly identi®ed Beania hirtissima: Gautier, 1962: 95. Prenant and in an ecological study (McKinney and Jaklin, Bobin, 1966: 557. 2000). Both Prenant and Bobin (1966) and Beania hirtissima forma cylindrica: Gautier, 1962: 96. Hayward and Ryland (1998) ®gured the an- cestrula of B. fulva, and the former noted that DESCRIPTION (AMNH 905; CMRR 2208): it might be very much longer than an auto- Autozooids elongate, narrow, commonly 1.0 zooid, although none was reported to be as ϫ 0.25 mm; tapered proximally, truncate dis- long as those described here. In other re- tally, with the terminal operculum orientated spects, however, the morphology of the an- perpendicular to frontal plane. Frontal sur- cestrulae from Rovinj is identical to that de- face entirely membranous, bordered by long, scribed previously and the exceptionally thin spines: six erect spines evenly spaced elongate, tubular proximal portion apparently around distolateral rim of operculum, eight re¯ects a phenotypic response to microhabi- to ten pairs of spines along margins of frontal tat characteristics. The pattern of autozooids membrane, slightly incurved; additional, at a bifurcation mostly conforms to the Type equally long or longer, spines developed on 5 of Harmer (1923), with zooid F enclosed lateral and basal surfaces of autozooids, proximal to the bifurcation, and zooid G ad- aligned with those on frontal margins. Each jacent to the axil. However, Hayward and autozooid linked to six neighbors by short, Ryland (1998) note that this is inconstant and tubular connections: one proximal and two that some dichotomies within a colony may proximolateral, all closely spaced, and three be achieved by simple splitting of the four on basal wall midway along the length of the autozooid series. In one of the Rovinj spec- autozooid. imens this had occurred immediately distal Colonies are erect tufts, to 12 mm, loosely to the ancestrula, and the ®rst bud developed and irregularly branched. Primary branches as another very elongate autozooid, reinforc- of ®ve to eight alternating autozooid series ing the specimen's super®cial resemblance to de®ning a convex frontal surface encom- Caulibugula. passing two-thirds circumference of branch. Tentacles clear, 14±15; lophophores bell- Primary branches divide dichotomously at ir- shaped, all obliquely truncate. regular intervals; minor branches of two to DISTRIBUTION: Bugula fulva has been three autozooid series originating at axil of patchily recorded from the southwest British dichotomies or along branch margins. Isles and parts of the Mediterranean, low in Abfrontal surface concave, with the space it the littoral zone and in shallow coastal wa- encloses crossed by basal spines of auto- ters. It is also associated with ports and har- zooids. Septula on basal autozooid walls give bors, and its wide occurrence in the western rise to tubular kenozooidal rhizoids which Atlantic, from Brazil to Maine, perhaps sug- pass proximally to attach colony to substra- gests an exotic origin as a fouling species. tum. MEASUREMENTS (CYSTID): AL 196 Ϯ 19 Tentacles clear, 26; lophophores bell- ␮m, 170±235 (2, 19), DO 228 Ϯ 34, 276± shaped, radially symmetrical. 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 23

Fig. 9. A, B. Beania cylindrica. A. General aspect (AMNH 905; 0.5 mm). B. Zooid, with frontal- margin and basal spine rows (AMNH 905; 0.1 mm). C±E. Beania hirtissima. C. General aspect, auto- zooids and (lower left) avicularium (lectotype, UIIZ 34; 0.5 mm). D. Distal end of zooid, with horizontal operculum (lectotype, UIIZ 34; 0.1 mm). E. Autozooid with lateral stalked avicularium (AMNH 906; 0.1 mm). 24 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

REMARKS: Material described here con- er's specimen illustrated here as ®g. 9C and forms to the colony morphotype often de- 9D (UIIZ specimen 34, one of two speci- noted Beania hirtissima form or var. cylin- mens in the collection indicated as Heller's drica. However, as discussed below, B. hir- types of the species). The lectotype has the tissima has very different autozooid shape features attributed to B. robusta, described and orientation plus scattered vicarious avi- originally from Algiers (Hincks, 1881) as cularia, which are absent in B. cylindrica. Diachoris hirtissima forma robusta. DISTRIBUTION: Beania cylindrica has been DISTRIBUTION: Beania hirtissima has been accorded a wide geographical Distribution, accorded a wide geographical Distribution, in shallow, warm temperate to tropical seas in shallow, warm temperate to tropical seas throughout the world, which possibly needs throughout the world, which needs to be re- to be re-examined. examined in light of the characteristics of the MEASUREMENTS (SKELETAL): DO 549 Ϯ 92 lectotype. As B. robusta, this rather substan- ␮m,, 377±792 (2, 10), OL 139 Ϯ 19, 117± tial Beania has been reported from a number 176 (2, 7), OW 138 Ϯ 8, 120±149 (2, 6), ZL of localities in the northwestern Mediterra- 993 Ϯ 97, 900±1204 (2, 10), ZW 381 Ϯ 34, nean, and also from Tunisia (Gautier, 1962). 327±443 (2, 10). (POLYPIDE): IH 602 Ϯ 80 It does not seem to have been recorded from ␮m, 500±700 (1, 10), LD 995 Ϯ 72, 880± east of the Adriatic. 1100 (1, 8), MD 40 (1, 1), TL962 Ϯ 29, MEASUREMENTS (SKELETAL): AL 216 Ϯ 14 920±1020 (1, 10). ␮m, 199±230 (1, 4), AW 170 Ϯ 4, 166±176 (1, 4), DO 499 Ϯ 65, 394±600 (1, 11), OL Beania hirtissima (Heller, 1867) 161 Ϯ 17, 141±194 (1, 10), OW 200 Ϯ 13, Figure 9C±E 183±222(1, 10).

Diachoris hirtissima Heller, 1867: 94. Beania magellanica (Busk, 1852) Diachoris hirtissima forma robusta Hincks, 1881: 133. Figure 10A, B Beania robusta: Gautier, 1962: 99. Zabala and Diachoris magellanica Busk, 1852: 54. Maluquer, 1988: 101. Beania magellanica: Gautier, 1962: 97. Prenant and Bobin, 1966: 555. LECTOTYPE (chosen here): UIIZ 34. DESCRIPTION (AMNH 906; CMRR 2209): DESCRIPTION (AMNH 907; CMRR 2210): Colonies multiserial, unilaminar sheets of Colonies forming repent sheets, supported closely spaced, suberect autozooids, each above and anchored to substratum by short, linked to six neighbors by a ring of six short tubular, kenozooidal rhizoids. Autozooids connecting tubes around proximal end. Au- large, elongate oval, ca. 0.8 mm long in pre- tozooids broadest proximally, narrowest at sent material, membranous frontal surface the distal end; 0.8±1.0 mm long. Membra- with a smooth rim, lacking spines. Distal end nous frontal surface bordered by stout spines: of autozooid rounded, edge of the operculum three in a line along distal edge of the ter- ®tting along distal rim; proximal end gently minal operculum and two on each side, all tapered. Autozooids disjunct, linked by tu- especially long and straight, to 0.5 mm; ®ve bular connecting processes, 0.2 mm long; erect spines on each lateral margin, straight each autozooid with six tubes: one midprox- and successively shorter proximally, each imal, two proximolateral, and three distobas- with a shorter, thinner spine at its base, curv- al, the latter equally spaced in an arc in distal ing medially over frontal margin. Basal sur- third of the autozooid. Conspicuous, paired face of colony with three to eight irregularly avicularia present, attached to lateral walls tapered and curved, thin processes on each close to distal series of connecting tubes; pe- autozooid. dunculate, with bird's head form, rostrum ta- Avicularia infrequent; pedunculate, at- pered, curved, and distally hooked. At rest tached to lateral wall of autozooid level with mandible lying in frontal plane of autozooid, proximal corner of operculum; globose, with with rostrum directed distally. Ovicell an in- a broadly triangular, distally hooked rostrum. conspicuous, partly immersed, distal cap, not REMARKS: We designate as lectotype Hell- seen in present material. 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 25

Fig. 10. A, B. Beania magellanica (AMNH 907). A. General aspect (0.5 mm). B. Autozooid with paired lateral stalked avicularia (0.1 mm). C, D. Beania mirabilis. (AMNH 908) C. Frontal view of single zooid (0.1). D. Lateral view of single zooid (0.1 mm). E, F. Synnotum aegyptiacum (AMNH 909). E. Uniserial erect branch (0.5 mm). F. Single zooid (0.1 mm). G±I. Scrupocellaria delilii (AMNH 910). G. Frontal surface of two biserial branches (0.5 mm). H. Reverse surface of untreated branch; vibracular setae visible (0.5). I. Autozooid with scutum, single outer and paired inner distal spines, and lateral avicularium (0.1 mm). 26 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Twenty-seven tentacles (based on single Beania mirabilis has been described also count). from Indo-Paci®c localities, but these records REMARKS: This is the most distinctive require reexamination. Beania species occurring in the Mediterra- MEASUREMENTS (CYSTID): ZL (distal) 677 nean region. Ϯ 45 ␮m, 620±780 (2, 13), ZW (distal) 280 DISTRIBUTION: Widely distributed through- (1, 1). (POLYPIDE): IH 295 Ϯ 59 ␮m, 229± out the Mediterranean, in shallow coastal wa- 343 (2, 3), LD 642 Ϯ 130, 491±806 (2, 4), ters, most frequently on biogenic substrata. TL 532 Ϯ 128, 395±689 (2, 9). Beania magellanica has been reported from throughout the warm temperate to tropical FAMILY EPISTOMIIDAE GREGORY, 1893 Atlantic, as well as from the western Paci®c. MEASUREMENTS (SKELETAL): AL 439 Ϯ 53 GENUS SYNNOTUM PIEPER, 1881 ␮m, 342±528 (3, 16), AW 151 Ϯ 14, 118± 173 (2, 15), DO 586 Ϯ 88, 438±689 (2, 15), Synnotum aegyptiacum (Audouin, 1826) OL 110 Ϯ 12, 95±126 (1, 5), OW 158 Ϯ 16, Figure 10E, F 143±184 (1, 5), ZL 713 Ϯ 55, 634±799 (3, Loricaria aegyptiaca Audouin, 1826: 243; Savig- 13), ZW 440 Ϯ 33, 373±478 (3, 13). (POLYP- ny, 1809: pl.13, ®gs 4(1)±4(5). IDE): LD 1140 ␮m (1, 1), TL 980 (1, 1). Synnotum aegyptiacum: Prenant and Bobin, 1966: 461. Zabala and Maluquer, 1988: 100. Beania mirabilis Johnston, 1840 Figure 10C, D DESCRIPTION (AMNH 909; CMRR 2212): Beania mirabilis Johnston, 1840: 272. Alvarez, Colony erect, branching, jointed, to 7.5 mm 1990: 26. Hayward and Ryland, 1998: 244. in present material. Autozooids slender, fu- siform, narrowest proximally, broadest dis- DESCRIPTION (AMNH 908; CMRR 2211): tally; arranged in back-to-back pairs, each Colonies formed from uniserial chains of au- separated from preceding pair by a slender, tozooids only lightly attached to substratum, tubular chitinous node. Frontal surface en- often largely free. Each autozooid consisting tirely membranous, with a terminal opercu- of a ®liform proximal portion comprising lum slightly acute to frontal plane. Squat, more than half its length, and a boat-shaped sessile avicularia present on disto-basal sur- distal portion, broadest proximally and ta- faces of some autozooids, others bearing pered towards the terminal operculum. Cal- larger, pedunculate, bird's head type with ci®cation thin and translucent, the frontal short, hooked mandible. Embryos brooded in surface entirely membranous. Four short distinctive globular, dimorphic zooids (not spines are spaced around the operculum and found in present material). Each autozooid of another 10 pairs of equally short, thin, and straight or only slightly curved spines dis- a pair giving rise to a single bud disto-ba- tributed along the margins of the frontal sally, which traverses the basal wall junction membrane. Each autozooid budding its suc- of the succeeding autozooid pair as a slender cessor from the basal wall, in the proximal tube, then expands distal to them to form third of the broad portion. Between this bud- broad distal portion of new autozooid. Au- ding point and the ®liform portion is a pair tozooid thus consisting of a proximal tube of short, tubular processes; on rare occasions comprising two-thirds its total length and a one of these may develop as a zooid bud, fusiform distal portion bearing the frontal allowing colony to branch, but most auto- membrane. The node between each auto- zooids uniserially arranged. zooid pair, consequently, consisting of four Tentacles clear, 19±21; lophophores bell- closely juxtaposed tubes. shaped, radially symmetrical. DISTRIBUTION: Widespread and common DISTRIBUTION: Probably common in coast- throughout the Mediterranean, in shallow al habitats throughout the northeast Atlantic coastal waters, and recorded from warm tem- and the Mediterranean, on a wide variety of perate and tropical seas worldwide. organic and inorganic substrata, but its in- MEASUREMENTS (SKELETAL): ZL 500 Ϯ 41 conspicuous colonies are easily overlooked. ␮m, 430±577 (2, 21). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 27

FAMILY CANDIDAE D'ORBIGNY, 1851 the geographical ranges of more than one GENUS SCRUPOCELLARIA LEVINSEN, 1909 species. MEASUREMENTS (SKELETAL): DO 323 Ϯ 47 Scrupocellaria delilii (Audouin, 1826) ␮m, 245±413 (2. 20), OpL 217 Ϯ 21, 178± Figure 10G±I 243 (2, 20), OpW 137 Ϯ 15, 107±161 (2, 20), ZL 425 Ϯ 31, 383±503 (2, 20), ZW 219 Crisia delilii Audouin, 1826: 242. Savigny, 1809: Ϯ 16, 188±251 (2, 20). pl. 12, ®g. 3. Scrupocellaria macandrei: Heller, 1867: 87. Scrupocellaria delilii: Gautier, 1962: 86. Prenant Scrupocellaria scrupea Busk, 1851 and Bobin, 1966: 435. Figure 11A±E

DESCRIPTION (AMNH 910; CMRR 2213): Scrupocellaria scrupea Busk, 1851: 83. Hayward Colonies erect, branching dichotomously, to and Ryland, 1998: 274. 20 mm high in present material. Branching DESCRIPTION (AMNH 911±913; CMRR frequency variable, at 2-mm intervals in 2214): Colonies to 18 mm in present mate- proximal regions of colony, to 4 mm or more rial, dichotomously branching at frequent in- towards distal portion; each ramus originat- tervals, giving dense, bushy form. Opesia oc- ing as a bipartite, tubular chitinous joint. cupying half or less of total autozooid length; Branches ¯at, consisting of two alternating each autozooid with three spines on outer longitudinal autozooid series; opesiae of ad- distal angle, two on the inner. Scutum arises jacent autozooids overlapping for about one- close to base of proximal-most inner spine; ®fth of total length. Autozooids ca 0.5 ϫ lobe reniform, not pointed distally, covering 0.25 mm; opesia oval, comprising more than almost half the area of the opesia. Prominent half total frontal length, bordered by ®nely lateral avicularium present on each auto- granular cryptocystal rim. Two spines pre- zooid, with hooked rostrum, but no frontal sent, one on each distal corner of opesia, avicularia. Basal vibraculum present at prox- erect, slender and curved, converging medi- imal end of each autozooid, aligned disto- ally above operculum. A small scutum pre- proximally, and two at axil of each dichoto- sent proximal to inner distal spine, with a my. Ovicell broader than long, inclined to- slender stalk curving over frontal membrane; wards median longitudinal axis of the oval, or slightly crescentic, consisting largely branch; lightly calci®ed, with a single, irreg- of a proximal lobe, with only a minimal lobe ular foramen close to aperture. distal to stalk. Each autozooid with a prom- Tentacles clear, 11; lophophores bell- inent lateral avicularium, its cystid half as shaped, radially symmetrical. long as opesia, rostrum 0.15 mm long, sharp- DISTRIBUTION: In shallow coastal habitats ly hooked distally and projecting distinctly throughout the Mediterranean, and north to from branch margin. Small frontal avicularia the southwest British Isles. present on some autozooids, proximal to MEASUREMENTS (SKELETAL): DO 319 Ϯ 66 opesia, with short, triangular rostrum perpen- ␮m, 246±459 (2, 20), OpL 166 Ϯ 16, 137± dicular to branch axis. Ovicell short, broad, 195 (2, 20), OpW 109 Ϯ 9, 95±129 (2, 20), tilted towards branch axis, imperforate. A ZL 378 Ϯ 31, 326±435 (2, 20), ZW 201 Ϯ small vibraculum on abfrontal surface of 17, 177±233 (2, 20). (POLYPIDE): IH 13 Ϯ 23 each autozooid, with setal groove straight ␮m, 0±40 (1, 4), LD 273 Ϯ 31, 240±300 (1, and extending halfway to middle of the au- 4), TL 327 Ϯ 23, 300±340 (1, 4). tozooid; two such vibracula in axil of each dichotomy. Scrupocellaria scruposa (Linnaeus, 1758) DISTRIBUTION: Scrupocellaria delilii has Figure 11F±J been reported from the Atlantic coasts of Spain, south to Madeira, and from numerous Sertularia scruposa Linnaeus, 1758: 815. localities in the Mediterranean. It has also Scrupocellaria scruposa: Gautier, 1962: 92. Hay- ward and Ryland, 1998: 276. been recorded from the Red Sea, the Indian Ocean, Australia and the West Paci®c, but DESCRIPTION (AMNH 909, 914±916; this wider Distribution possibly encompasses CMRR 2215): Colonies erect, branching 28 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 11. A±E. Scrupocellaria scrupea. A. Frontal surface of untreated branches (AMNH 911; 0.5 mm). B. Frontal surface of bleached branch (AMNH 912; 0.5 mm). C. Reverse surface of treated branch (AMNH 912; 0.5 mm). D. Autozooids with scutum, distal spines, and lateral avicularia (AMNH 911; 0.1 mm). E. Maternal zooids with ovicells (AMNH 913; 0.1 mm). F±J. Scrupocellaria scruposa. F. Frontal surface of untreated branches (AMNH 914; 0.5 mm). G. Frontal surface of bleached branch (AMNH 909; 0.5 mm). H. Reverse surface of treated branch (AMNH 915; 0.5 mm). I. Autozooid with distal spines, lateral avicularium, and vibraculum with seta (AMNH 916; 0.1 mm). J. Maternal zooids with ovicells (AMNH 915; 0.1 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 29 tufts, to 20 mm in present material; branches longitudinal series, or three just prior to di- dividing dichotomously, each ramus sup- chotomies, de®ning the frontal surface of the ported by a bipartite, chitinous joint. Branch- branch, commonly 0.35 ϫ 0.2 mm. Frontal es ¯at, consisting of two alternating auto- planes of the two autozooid series angled at zooid series, opesiae of adjacent autozooids about 90Њ to each other; longitudinal axis of not or scarcely overlapping. Autozooids each autozooid slightly oblique to branch elongate, slender, commonly 0.5 ϫ 0.2 mm, axis, inclining medially. Frontal surfaces of with narrowly oval opesia occupying half or alternate autozooids slightly overlapped; less of total frontal length; opesia with a nar- opesia and operculum together constituting row border of cryptocystal calci®cation. four-®fths of total frontal length, bordered Three spines present at outer distal corner of proximally and laterally by broad, ®nely autozooid, two at inner corner, all erect, granular cryptocyst, no frontal gymnocystal straight and slender. No scutum. Lateral av- calci®cation present. Three distal oral spines icularia variably sized, from one-third to one- present: two on inner distal angle of auto- half length of opesia; rostrum perpendicular zooid and one, very much longer, on the out- to longitudinal branch axis, projecting con- er distal angle. Much of opesia obscured by spicuously, with pronounced distal hook. A thick, oval scutum, attached by a thick stalk small, distally directed frontal avicularium just proximal to inner distal spines; its distal present distal to ovicell, which is globular, edge straight, abutting with a blunt process longer than wide, with a highly arched ap- projecting from opposite, outer edge of the erture and drop-shaped frontal fenestra. Bas- opesia, and de®ning the proximal border of al vibraculum small, at outer proximal corner the operculum, which is terminal in position of autozooid; cystid parallel to longitudinal and tilted basally at a marked angle to frontal branch axis, with short setal groove only plane. A small adventitious avicularium on slightly oblique to it. Two vibracula present the inner distal angle of many autozooids, at axil between each dichotomy. with short triangular mandible, distally di- DISTRIBUTION: Scrupocellaria scruposa is rected; frequently, a much larger avicularium common in shallow coastal waters in the en- proximal to dichotomy, with broadly trian- tire northeast Atlantic region, from Iceland gular rostrum frontally directed and almost to the Cape Verde Islands, and throughout perpendicular to frontal plane. Ovicell re- the Mediterranean. cumbent on distally succeeding autozooid, MEASUREMENTS (SKELETAL): DO 364 Ϯ 89 inclined medially; broader than long, with ␮m, 273±501 (2, 19), OpL 209 Ϯ 29, 170± highly arched aperture bordered by broad 274 (2, 16), OpW 127 Ϯ 20, 106±191 (2, band of uncalci®ed ectooecium. Abfrontal 16), OvL 168 Ϯ 17, 151±184 (2, 4), OvW surface of colony distinctive: basal wall of 183 Ϯ 9, 171±193 (2, 4), ZL 491 Ϯ 37, 428± each autozooid obscured by a large, proxi- 543 (2, 15), ZW 183 Ϯ 20, 137±212 (2, 16). mally tapered vibraculum; seta 1.5 mm long, ®nely toothed along one edge, ®tting into a GENUS CABEREA LAMOUROUX, 1816 narrow groove extending from outer distal Caberea boryi (Audouin, 1826) edge of vibraculum, proximally and medial- ly, contributing to a pronounced ridge ex- Figure 12A±E tending length of branch. Thin kenozooidal Crisia boryi Audouin, 1826: 242. Savigny, 1809: rhizoids arising from vibracular chambers, pl. 12 ®gs 4.1±4.6. passing proximally along median abfrontal Caberea boryi: Gautier, 1962: 93. Prenant and surface of each branch and coalescing as Bobin, 1966: 449. Hayward and Ryland, 1998: thick bundles at base of colony. 252. Tentacles clear, 13; lophophores campy- DESCRIPTION (AMNH 917±919; CMRR lonemidan. 2216): Colonies stiff, erect fans, to 8 mm DISTRIBUTION: Caberea boryi is wide- high in present material, with cylindrical spread and abundant in shallow inshore wa- branches dividing dichotomously at regular ters throughout the Mediterranean, and rang- intervals; without visible joints at the di- es northwards to the southwest British Isles, chotomies. Autozooids in two alternating and southwards to South Africa. It has been 30 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 12. A±E. Caberea boryi.A.Frontal surface of untreated branches (AMNH 917; 0.5 mm). B. Serrated vibracular setae (AMNH 917; 0.1 mm). C. Ancestrula (AMNH 918; 0.1 mm). D. Reverse surface of treated branch with conspicuous proximally tapered vibraculae (AMNH 919; 0.1 mm). E. Frontal surface of branch, with autozooid (right) and (left) maternal zooid with ovicell and distal avi- cularium (AMNH 917; 0.1 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 31 recorded from the Red Sea and the Indian shaped, radially symmetrical, supported on Ocean, and from the West Paci®c. very long introverts, commonly overlapped MEASUREMENTS (SKELETAL): DO 223 Ϯ 34 with adjacent lophophores up to 300 ␮m. ␮m, 172±310 (3, 28), OvL 122 Ϯ 9, 110± REMARKS: This common and conspicuous 137 (1, 6), OvW 156 Ϯ 14, 142±181 (1, 6), Mediterranean species is suf®ciently distinc- ZL 357 Ϯ 40, 265±435 (3, 29), ZW 147 Ϯ tive to be recognized with the unaided eye. 11, 132±169 (1, 9). (POLYPIDE): IH 32 Ϯ 7 Poluzzi and Coppa (1991) described growth ␮m, 20±40 (1, 9), LDMn 438 Ϯ 27, 400± and regeneration in C. nobilis, in relation to 460 (1, 6), LDMx 497 Ϯ 37, 440±540 (1, 9), spatial competition. McKinney and Jaklin MD 20 Ϯ 0 (1, 5), TL 443 Ϯ 55, 360±540 (1993) reported ephemeral populations of (1, 10). free-lying colonies composed of single zo- oid-thick sheets, up to several centimeters in SUPERFAMILY MICROPOROIDEA GRAY, 1846 diameter, off the Istrian Peninsula 30 to 50 FAMILY MICROPORIDAE GRAY, 1848 km south-southwest of Pula. Ristedt (1991) GENUS CALPENSIA JULLIEN, 1888 described the ancestrula and early astogeny, but sexual reproduction has not been de- Calpensia nobilis (Esper, 1796) scribed. Figure 13A±C OCCURRENCE: This species encrusts virtu- Cellepora nobilis Esper, 1796: 145. ally every type of nontoxic ®rm to hard sub- Membranipora bifoveolata Heller, 1867: 95. stratum, including rock, shell debris, pottery, Calpensia nobilis: Gautier, 1962: 59. Zabala, glass, other bryozoans, thecate ascidians, 1986: 289. Zabala and Maluquer, 1988: 90. andÐelsewhere in the Adriatic and Mediter- Hayward and Ryland, 1998: 292. ranean (Poluzzi and Coppa, 1991)Ðmarine DESCRIPTION (AMNH 981, 1008; CMRR grasses. 2217): Colonies widely spreading, multiser- DISTRIBUTION: The species is widespread ial, ¯at, light-colored grayish-brown sheets, throughout the Mediterranean and occurs often exceeding 10 ϫ 10 cm; through suc- sparsely northwards to the Gulf of St. Malo, cessive phases of regeneration and over- and on the northwest African coast. growth may form thick cylinders and nodules MEASUREMENTS (SKELETAL): DO 530 Ϯ 93 on every kind of hard substratum. Auto- ␮m, 366±684 (2, 20), OpL 96 Ϯ 8, 80±107 zooids simple rectangles or hexagons with a (2, 20), OpW 150 Ϯ 17, 119±174 (2, 20), thick, densely perforated cryptocyst, a small ZL 817 Ϯ 70, 719±935 (2, 20), ZW 383 Ϯ terminal opesia, equivalent to about 12% to- 36, 322±439 (2, 20). (POLYPIDE): IH 495 Ϯ tal autozooid length and coincident with the 283 ␮m, 220±1200 (4, 21), LD 955 Ϯ 128, operculum, and paired opesiules almost as 480±1140 (4, 40), MD 34 Ϯ 8, 20±50 (3, 9), large as opesia. Outer epitheca glistens in TL 654 Ϯ 63, 560±820 (4, 18). dried material. First asexual zooid budded from distal end of ancestrula but with growth GENUS ROSSELIANA JULLIEN, 1888 direction oriented 120Њ with respect to an- Rosseliana rosselii (Audouin, 1826) cestrula and lying beside ancestrula, estab- Figure 14A±E lishing early colony growth in direction op- posite the polarity of the ancestrula (®g. Flustra rosselii Audouin, 1826: 240. 13C). Occasional reparative autozooidal bud- Membranipora rosselii: Hincks, 1880: 166. ding. No polymorphs. Rosseliana rosselii: Canu and Bassler, 1925: 17. Cystids are occupied by a succession of up Hayward and Ryland, 1998: 294. Rosseliana ``sp. nov.'' Gautier, 1962: 63. to four generations of polypides, and the brown body remains of previous polypides DESCRIPTION (AMNH 1053±1055; CMRR are stored in the proximal portions of zooids. 2218): Colonies multiserial encrusting, ¯at, Proximal portions of large colonies are se- beige colored. Autozooids arched oval dis- nescent and commonly are bright green in tally and tapered proximally (proximal shape color due to a thin growth of chlorophyte determined by shape of immediately preced- algae. ing zooids), with boundaries marked by Tentacles clear, 19±22; lophophores bell- prominently raised lateral walls around 32 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 13. A±C. Calpensia nobilis. A. General aspect (AMNH 981; 0.5 mm). B. Autozooid (AMNH 981; 0.1 mm). C. Ancestrula and early autozooids (AMNH 1008; 0.2 mm). D±H. Mollia circumcincta. D. General aspect (AMNH 920; 0.5 mm). E. Lectotype, untreated, with several kenozooids at colony margin (upper right; UIIZ 116; 0.5 mm). F. Lectotype, autozooid (UIIZ 116; 0.1 mm). G. Maternal zooid with ovicell, ¯anked by autozooids (AMNH 920; 0.1 mm). H. Ancestrula and early autozooids (AMNH 921; 0.1 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 33

Fig. 14. A±E. Rosseliana rosselii. A. General aspect (AMNH 1053; 0.5 mm). B. Autozooid (AMNH 1053; 0.1 mm). C. Maternal zooid with largely immersed ovicell (AMNH 1054; 0.1 mm). D. Growing edge of colony (AMNH 1055; 0.10 mm). E. Ancestrula and early autozooids (AMNH 1055; 0.2 mm). arched distal end. Gymnocyst absent; cryp- (length 0.70±0.82 mm) and distinguished an tocyst ¯at to gently arched transversely, with unnamed new species for colonies with coarse-grained or ®nely tuberculate surface. smaller zooids (length 0.40±0.68 mm). We Opesia terminal, semielliptical with very encountered only colonies with the smaller slightly concave proximal edge, equivalent to zooids, essentially equivalent in size to the about 40% total autozooid length. Ancestrula zooids of colonies attributed to R. rosselii similar to autozooids; ®rst three asexually around Britain (Hayward and Ryland, 1998: budded zooids distal and distolateral; pri- 294). Prenant and Bobin (1966: 346±348) mary astogenetic zone of change only three noted the two forms recognized by Gautier or four generations Ovicell small, largely im- but indicated that they seem to co-occur eco- mersed, apparent as shallow crescentic cap at logically and geographically and could not at distal end of zooid. the time be separated into two distinct spe- Tentacles clear, 15; lophophores bell- cies. Although only one of the forms has shaped, radially symmetrical away from col- been found by us in the northern Adriatic, ony margin between chimneys, obliquely suggesting that the geographic ranges of the truncate adjacent to chimneys and colony two ``forms'' or perhaps species are not fully margin; introverts long. coincident, we have preferred to retain the REMARKS: Gautier (1962: 63±64) used R. name R. rosselii for the northern Adriatic rosselii for Rosseliana with large zooids specimens. 34 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

DISTRIBUTION: This species is widespread ology, labeled as ``Caleschara patellaria var. in the eastern temperate Atlantic, from the circumcincta Heller. Adria (S. Heller) 9837.'' Cape Verde Islands to Shetland, and through- There is no annotation that this specimen is out the Mediterranean. one of Heller's original types, but we illus- MEASUREMENTS (SKELETAL): DO 452 Ϯ 46 trate it here as at least an indication of long ␮m, 363±520 (2, 20), OpL 246 Ϯ 24, 218± consistency in concept of the species and 308 (2, 20), OpW 227 Ϯ 27, 185±277 (2, designate it as neotype for the species. If 20), ZL 591 Ϯ 30, 533±668 (2, 20), ZW 375 eventually discovered to be a specimen on Ϯ 45, 307±457 (2, 20). (POLYPIDE): IH 366 which Heller originally described Membran- Ϯ 127 ␮m, 200±720 (2, 22), LDMn 560 Ϯ ipora circumcincta, it should be redesignated 62, 400±630 (2, 13), LDMx 622 Ϯ 72, 420± as lectotype. 840 (2, 24), MD 24 Ϯ 3, 20±30 (2, 10), DISTRIBUTION: This pretty species was ®rst TLMn 476 Ϯ 62, 340±590 (2, 23), TLMx described from the Adriatic (Heller, 1867) 588 Ϯ 113, 400±790 (2, 25). and has been reported from very few addi- tional localities. It is known from the Siculo- GENUS MOLLIA LAMOUROUX, 1816 Tunisian shelf and the northwest Mediterra- Mollia circumcincta (Heller, 1867) nean (Gautier, 1962), but does not seem to Figure 13D±H range outside of the Mediterranean. MEASUREMENTS (SKELETAL): DO 336 Ϯ 36 Membranipora circumcincta Heller, 1867: 96. ␮m, 264±412 (2, 20), OpL 86 Ϯ 14, 62±107 Mollia circumcincta: Gautier, 1962: 61. Zabala (2, 20), OpW 120 Ϯ 19, 84±159 (2, 20), OvL and Maluquer, 1988: 92. 102 Ϯ 10, 82±115 (2, 17), OvW 174 Ϯ 12, NEOTYPE (chosen here): UIIZ 116. 156±206 (2, 17), ZL 410 Ϯ 35, 371±479 (2, DESCRIPTION (AMNH 920, 921; CMRR 20), ZW 267 Ϯ 24, 234±338 (2, 20). 2219): Colonies light orange, encrusting, multiserial, unilaminar sheets. Autozooids SUPERFAMILY CELLARIOIDEA FLEMING, 1828 oval, 0.375±0. 50 mm; disjunct, each linked FAMILY CELLARIIDAE FLEMING, 1828 to its neighbors by about 12 short, tubular communication organs. Frontal body wall GENUS CELLARIA ELLIS AND SOLANDER, 1786 membranous, underlain for about two-thirds Cellaria ®stulosa (Linnaeus, 1758) of its extent by a vitreous, granular crypto- Figure 15A±E cyst; opesia semielliptical, the proximal edge straight or slightly convex, and gently arched Eschara ®stulosa Linnaeus, 1758: 804. frontally, with indistinct opesiular indenta- Cellaria ®stulosa: Hayward and Ryland, 1998: 306. tions at each corner. Vertical walls of the au- tozooid develop a crenulate mural rim, which DESCRIPTION (AMNH 922; CMRR 2220): is especially raised distally, forming a cowl Colonies erect, branching, consisting of rig- above the distal edge of the opesia. Spines id, cylindrical internodes formed from alter- absent, except in ®rst two or three astoge- nating whorls of three to six autozooids, netic generations. Ovicell hemispherical, par- linked by ¯exible, chitinous nodes at which tially immersed, with a crenulate vertical dichotomous branching occurs. Internodes wall bordering a vitreous, nodular cryptocys- 2±9 mm long, typically about 5 mm, in pre- tal frontal surface. No polymorphs other than sent material, with diameter 0.45±0.7 mm, kenozooids, which may occur in clusters. typically about 0.55 mm. Colonies attach by Tentacles clear, 15; lophophores bell- bundles of tubular rhizoids developed from shaped, radially symmetrical away from col- the frontal cuticles of the proximal auto- ony margin between chimneys, obliquely zooids of the colony. Autozooids diamond- truncate adjacent to chimneys and colony shaped or hexagonal in outline; successive margin; introverts long. autozooids of longitudinal rows commonly REMARKS: Specimens of Mollia circum- but not always in contact along common cincta collected from the Rovinj area are transversely linear border and completely conspeci®c with bryozoan specimen 116 in bounded laterally by the two adjacent offset the University of Innsbruck Institute of Zo- rows of autozooids; less commonly succes- 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 35

Fig. 15. A±E. Cellaria ®stulosa (AMNH 922). A. Typical surface of young branch, distal joints not yet decalci®ed (0.5 mm). B. Branch with swollen area of maternal zooids (bottom of ®gure) and (top left) only one distal joint decalci®ed (0.5 mm). C. Branch segment with several distal autozooids with proximal perforations from which non-calci®ed rhizooidal kenozooids emerged (0.5 mm). D. Autozooids (bottom of ®gure) with proximal perforations for kenozooids, small avicularium (center), and maternal zooids with brood chambers (top of ®gure; 0.1 mm). E. Avicularium separating two maternal zooids with calci®ed plates restricting brood chamber apertures (0.1 mm). F±K. Cellaria salicornioides. F. Typical surface of young branch, distal joints not yet decalci®ed (AMNH 923; 0.5 mm). G. Autozooids, one with proximal perforation for kenozooid (AMNH 923; 0.1 mm). H. Base of laterally budded branch (AMNH 923; 0.2 mm). I. Avicularium (AMNH 924; 0.1 mm). J. Polymorphic zooid from which lateral branch arose (AMNH 923; 0.2 mm). K. Lateral pro®le of avicularium (left) (AMNH 924; 0.1 mm). 36 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 sive zooids in longitudinal rows separated by two species are readily separated by the un- line marking contact between the two adja- aided eye. Autozooids oval to hexagonal or cent offset rows of autozooids. Frontal mem- almost rectangular, with regular quincuncial brane distinct and glistening in living mate- arrangement, so that commonly none is in rial; in cleaned specimens, cryptocystal fron- contact with those autozooids immediately tal shield concave between prominent ridges distal and proximal to it. Opesia in distal marking boundaries of autozooids. Calci®- third of zooid, semicircular, with crenulate cation ®nely nodular; opesia wider than long, distal rim and convex, slightly projecting equivalent to one-sixth zooid length, distal proximal rim; short, bluntly tapered denticles rim arched and ®nely beaded, proximal rim just visible in proximal corners. Cryptocyst convex, with a blunt, frontally curved den- deeply concave, with well-marked longitu- ticle in each proximal corner. A pair of well- dinal ridges especially prominent lateral to marked longitudinal cryptocystal ridges ap- the opesia, grading into the autozooid bound- parent on each side of the opesia, extending ary distally, but converging proximally to de- proximal to it, converging but not meeting. ®ne a complete rim distinct from the zooid Avicularia about one-third length of auto- margin. Avicularia sporadic, distinctive: as zooids, rounded-quadrangular, with a narrow, large as an autozooid, with semicircular ros- semielliptical opesia; mandible narrowly trum projecting at acute angle from frontal crescentic, supported on an arched rostrum, plane, supporting a conspicuous, brown man- directed distally or distolaterally. Ovicell ap- dible. Ovicell aperture a simple round open- erture small and rounded, situated immedi- ing distal to the opesia. ately distal to opesia, partially occluded in DISTRIBUTION: In shallow coastal waters; older parts of colonies by calci®ed plate ex- widespread throughout the Mediterranean, tending from proximal edge, as has been not- ranging northwards along the western coasts ed elsewhere for C. salicornioides (Hayward of Britain and Ireland, to Shetland. and Ryland, 1998). MEASUREMENTS (SKELETAL): DO 306 Ϯ 47 Tentacles clear, 13; lophophores bell- ␮m, 224±399 (2, 20), OpL 59 Ϯ 4, 53±68 shaped, radially symmetrical. (2, 20), OpW 81 Ϯ 7, 71±99 (2, 20), ZL 439 DISTRIBUTION: Common in shallow, coastal Ϯ 33, 383±532 (2, 20), ZW 213 Ϯ 17, 175± waters throughout the Mediterranean, and 240 (2, 20). northwards to the British Isles, western Nor- way and Iceland. SUBORDER ASCOPHORA LEVINSEN, 1909 MEASUREMENTS (SKELETAL): DO 314 Ϯ 23 INFRAORDER ACANTHOSTEGA LEVINSEN, 1902 m, 252±345 (2, 16), OpL 71 8, 60±87 ␮ Ϯ SUPERFAMILY CRIBRILINOIDEA HINCKS, 1879 (2, 20), OpW 107 Ϯ 8, 97±126 (2, 17), ZL 564 Ϯ 21, 510±601 (2, 20), ZW 262 Ϯ 14, FAMILY CRIBRILINIDAE HINCKS, 1879 241±283 (2, 15). (POLYPIDE): IH 67 Ϯ 27 ␮m, GENUS COLLARINA JULLIEN, 1886 20±100 (1, 6), LD 534 Ϯ 38, 490±580 (1, 7) Collarina balzaci (Audouin, 1826) MD 30 (1, 1), TL 437 Ϯ 29, 400±480 (1, 6). Figure 16A±C Cellaria salicornioides Lamouroux, 1816 Flustra balzaci Audouin, 1826: 239. Figure 15F±K Cribrilina punctata: Hayward and Ryland, 1979: 56. Cellaria salicornioides Lamouroux, 1816: 127. Collarina balzaci: Zabala, 1986: 359. Zabala and Cellaria salicornioides: Hayward and Ryland, Maluquer, 1988: 105. Bishop, 1988: 749. Hay- 1998: 308. ward and Ryland, 1998: 318.

DESCRIPTION (AMNH 923, 924; CMRR DESCRIPTION (AMNH 912, 925, 926; 2221): Colonies erect and branching, with CMRR 2222): Colonies encrusting, multi- slender, cylindrical internodes consisting of serial, unilaminar, small, irregular patches. alternating whorls of two to ®ve autozooids. Autozooids small, oval, with indistinct Colonies, branches, and internodes may be boundaries; gymnocyst rarely visible frontal- as long as those of C. ®stulosa, but tend to ly. Frontal shield of four to seven pairs of be far more diffuse and delicate, so that the costae; each costa with a prominent tubae- 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 37

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 ®gularis. (AMNH 925). D. General aspect (0.5 mm). E. Au- tozooid (0.1 mm). 38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 form pelmatidium proximally, together form- small, rounded condyles. No oral spines. ing a raised peripheral ring; two or three Frontal surface a costate shield, comprising smaller pelmatidia along median fusion of two-thirds total autozooid length, with a costae. Intercostal pores large, typically one broad area of proximal gymnocyst extending to three between each adjacent pair of costae. distally as a narrower band on each side. Apertural bar thickened, raised and mucro- Shield formed from 10±12 short costae, nate. Ori®ce semicircular, wider than long; fused medially and at intervals laterally, four oral spines, proximal pair elongate and leaving small, slitlike intercostal pores; each curving over ovicell in fertile autozooids. costa, additionally, with a single thick Avicularia adventitious, single or paired, on rimmed pelmatidium at its broad proximal each side of apertural bar, with short, trian- (peripheral) end. Large vicarious avicularia, gular rostrum laterally directed. Ancestrula with spoonshaped mandible, and globular small, lacking adventitious avicularia, with ovicells with a pair of large lateral fenestrae fewer pairs of costae than subsequently bud- characteristic of this species, but neither was ded autozooids. Ovicell recumbent on sub- present in the small colony collected. stratum, variably shorter or longer than wide, DISTRIBUTION: Figularia ®gularis is com- domed, with up to 20 tubaeform pores reg- mon in shallow coastal habitats throughout ularly distributed over its surface; a small av- the Mediterranean, encrusting shell and other icularium at its distal edge, identical to the hard substrata. It ranges north to the south- others, distally directed. Both autozooids and west British Isles. ovicell with large and conspicuous basal pore MEASUREMENTS (SKELETAL): DO 624 Ϯ 61 chambers. ␮m, 530±730 (1, 10), OL 185 Ϯ 14, 166± REMARKS: The taxonomic status of this 210 (1, 10), OW 217 Ϯ 19, 180±240 (1, 10), species was ®nally stabilized by Bishop ZL 822 Ϯ 64, 710±910 (1, 10), ZW 477 Ϯ (1988) through the selection of a neotype 51, 410±546 (1, 10). specimen. DISTRIBUTION: The species is widely dis- GENUS PUELLINA JULLIEN, 1886 tributed throughout the Mediterranean, on Puellina hincksi (Friedl, 1917) stone, shell, red seaweeds and Posidonia, and Figure 17A±C ranges northwards to western Britain and the Faroe Isles. Cribrilina radiata var. hincksi Friedl, 1917: 236. MEASUREMENTS (SKELETAL): Al 79 Ϯ 6 Cribrilaria hincksii (sic): Gautier, 1962: 111. ␮m, 70±91 (1, 10), AW 53 Ϯ 5, 47±61 (1, Puellina (Cribrilaria) hincksi: Zabala and Malu- quer, 1988: 107. Alvarez, 1994b: 134. 10), DO 401 Ϯ 61, 312±548 (4, 40), OL 59 Ϯ 11, 38±80 (4, 32), OW 110 Ϯ 14, 77±134 DESCRIPTION (AMNH 901, 927; CMRR (4, 33), OvL 244 Ϯ 27, 189±290 (3, 15), 2223): Colonies encrusting, multiserial, uni- OvW 230 Ϯ 12, 210±245 (3, 15), ZL 516 Ϯ laminar, small rounded patches. Autozooids 51, 484±668 (4, 40), ZW 318 Ϯ 32, 333± oval to irregularly hexagonal, separated by 436 (4, 40). deep grooves; gymnocyst clearly visible, smooth, imperforate; frontal shield of usually GENUS FIGULARIA JULLIEN, 1886 six or seven pairs of costae, each broadest Figularia ®gularis (Johnston, 1847) and slightly umbonate proximally, the umbo Figure 16D, E bearing a minute pelmatidium; intercostal pores large and distinct, peripheral papilla Lepralia ®gularis Johnston, 1847: 314. pores the largest. Suboral lacuna transversely Figularia ®gularis: Hayward and Ryland, 1998: oval, large and conspicuous; apertural bar 338. slightly thickened, ®rst pair of costae often DESCRIPTION (AMNH 925): Colonies en- with a prominent umbo proximal to suboral crusting, multiserial, unilaminar. Autozooids lacuna. Primary ori®ce wider than long, bor- rectangular to irregularly polygonal, convex, dered by ®ve evenly spaced spines. Avicu- separated by deep grooves. Primary ori®ce larium with small cystid wedged between au- about as wide as long, proximal border shal- tozooids; rostrum elongate, narrowly trian- lowly concave and frontally arched, between gular, up to three-quarters length of auto- 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 39

Fig. 17. A±C. Puellina hincksi. A. General aspect (AMNH 901; 0.5 mm). B. Autozooid ¯anked 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). 40 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 zooids. Ovicell smooth, domed, with Frontal shield of broad distal portion of au- indistinct longitudinal ridge. Kenozooids tozooid closely perforated by numerous sporadic. round pores; a few smaller pores in single REMARKS: First described from the Adri- linear series on tubular proximal portion. atic, this species was for long confounded Ovicell borne on distal rim of ori®ce, glob- with P. radiata (Moll, 1803) and P. innom- ular; ectooecium membranous except for a inata (Couch, 1844), but its speci®c identity smooth, calci®ed disto-basal rim, entooecium was ®nally con®rmed by Gautier (1962), calci®ed, with a coarsely reticulate surface. who reported it from the Marseilles region. Each autozooid develops paired disto-basal DISTRIBUTION: Puellina hincksi was de- buds from conspicuous tubular projections; scribed from Tunisia by Canu and Bassler new autozooids develop distal to a distinct (1930) and has recently been described and chitinous joint. An additional septulum situ- ®gured from west of Gibraltar (Alvarez, ated medially between the two daughter au- 1994b). It is probably widely distributed tozooids giving rise to delicate, cylindrical within the Mediterranean. rhizoid. MEASUREMENTS (SKELETAL): AL 323 Ϯ 20 Tentacles light brown, 18±22; lophophores ␮m, 311±346 (1, 3), AW 153 Ϯ 20, 130±168 bell-shaped, radially symmetrical, introverts (1, 3), DO 317 Ϯ 39, 253±398 (2, 20), OL short. 47 Ϯ 2, 41±50 (2, 17), OW 74 Ϯ 3, 67±79 DISTRIBUTION: Savignyella lafontii has (2, 17), OvL 193 Ϯ 14, 170±215 (1, 10), been reported on numerous occasions from OvW 175 Ϯ 14, 158±201 (1, 10), ZL 431 Ϯ the Adriatic, and its wider Distribution in the 49, 350±523 (2, 20), ZW 278 Ϯ 35, 218± western Mediterranean was detailed by Gau- 337 (2, 20). (POLYPIDE): IH 0 ␮m (1, 1), LD tier (1962). It has a practically circumglobal 560±700 (1, 2), MD 20 (1, 1), TL 480±500 Distribution in warm temperate to tropical (1, 2). waters. MEASUREMENTS (POLYPIDE): IH 96 Ϯ 22 SUPERFAMILY PATENICELLOIDEA BUSK, 1852 ␮m, 60±120 (1, 9), LD 772 Ϯ 96, 650±960 FAMILY SAVIGNYELLIDAE LEVINSEN, 1909 (1, 11), MD 30±40 (1, 2), TL 741 Ϯ 135, 440±920 (1, 9). GENUS SAVIGNYELLA LEVINSEN, 1909 Savignyella lafontii (Audouin, 1826) INFRAORDER HIPPOTHOOMORPHA GORDON, 1989 Figure 17D±F SUPERFAMILY HIPPOTHOOIDEA BUSK, 1859 Eucratea lafontii Audouin, 1826: 242. Savigny, 1809: pl.13, ®gs. 21±25. FAMILY HIPPOTHOIDAE BUSK, 1859 Savignyella lafontii: Gautier, 1962: 102. Zabala GENUS HIPPOTHOA LAMOUROUX, 1821 and Maluquer, 1988: 145. Hippothoa divaricata Lamouroux, 1821 DESCRIPTION (AMNH 928; CMRR 2224): Figure 18A±E Colonies brown, erect, bushy tufts to 10 mm high in present material, of branching, uni- Hippothoa divaricata Lamouroux, 1821: 82. Ry- serial chains of clavate autozooids. Auto- land and Gordon, 1977: 19. Hayward and Ry- land, 1999: 86. zooids 1±2 mm long, the proximal portion cylindrical, 0.08 mm wide, broadening to DESCRIPTION (AMNH 929): Colony en- 0.12 mm at one-half to two-thirds total crusting. Autozooids in branching, uniserial length; remaining distal portion 0.25 mm chains, diffuse or closely intergrown; clavate, wide. Primary ori®ce terminal, D-shaped, with tubular proximal cauda and oval distal with straight proximal edge, and seven short portion. Primary ori®ce longer than wide, spines equally spaced around lateral and dis- poster comprising one-®fth total length, tal margins. A prominent median suboral av- forming a short triangular sinus between dis- icularium partly obscuring ori®ce, its cylin- tinct angular condyles. No spines. Frontal drical cystid perpendicular to frontal plane; shield smoothly calci®ed, imperforate, with rostrum triangular, sharply hooked, orientat- pronounced longitudinal median ridge, most ed proximodistally and frontally directed. prominent just proximal to the sinus. Dimor- 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 41

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 ori®ce (0.05 mm). D. Ancestrula and early autozooids (0.5 mm). E. Ancestrula with D-shaped ori®ce (0.1 mm). F±I. Hippothoa ¯agellum. F. General aspect (AMNH 930; 0.5 mm). G. Maternal zooid with ovicell (AMNH 930; 0.1 mm). H. Autozooidal ori®ce (AMNH 931; 0.05 mm). I. Ancestrula and early autozooids (AMNH 929; 0.1 mm). 42 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 phic female zooids with slightly shorter cau- mm diameter, budding a single oval auto- da than autozooid, and broader ori®ce; ovi- zooid, lacking the distinctive cauda. cell hemispherical, imperforate, often with an DISTRIBUTION: This inconspicuous species umbo. Zooeciules of unknown function spo- has been reported from temperate to tropical radic, with narrow, tubular cystid and minute seas throughout the world, in shallow coastal elliptical ori®ce. Each autozooid producing a waters, almost invariably on shell and other single distal bud; branching achieved by lat- biogenic carbonates. eral budding, a maximum of two on each MEASUREMENTS (SKELETAL): CL 244 Ϯ 65 side, giving an X pattern when all four de- ␮m, 128±385 (2, 15), GL 255 Ϯ 29, 227± velop. Female zooids and zooeciules may 292 (1, 4), GW 157 Ϯ 21, 125±169 (1, 4), arise at any of the ®ve budding positions. OL 43 Ϯ 7, 29±52 (2, 10), OW 41 Ϯ 3, 37± Ancestrula oval, domed, with a complete 46 (2, 10), OvL 142 Ϯ 8, 136±154 (1, 4), gymnocystal frontal shield, and a small D- OvW 157 Ϯ 21, 125±169 (1, 4), ZL 559 Ϯ shaped ori®ce. 67, 437±705 (2, 15), ZW 192 Ϯ 17, 150± DISTRIBUTION: Hippothoa divaricata is 222 (2, 15). widely distributed throughout the northeast Atlantic and the Mediterranean, on stones FAMILY CHORIZOPORIDAE VIGNEAUX, 1949 and shells, in shallow coastal habitats. GENUS CHORIZOPORA HINCKS, 1879 MEASUREMENTS (SKELETAL): CL 208 Ϯ 33 Chorizopora brongniartii (Audouin, 1826) ␮m, 163±256 (1, 7), OL 63 Ϯ 4, 60±67 (1, Figure 19A±C 3), OW 52 Ϯ 4, 49±56 (1, 3), ZL 503 Ϯ 48, 421±577 (1, 7), ZW 169 Ϯ 10, 151±183 (1, Flustra brongniartii Audouin, 1826: 240. Savig- 7). ny, 1809: pl. 10, ®g. 6. Chorizopora brongniartii: Hincks, 1880: 224. Marcus, 1940: 214. Gautier, 1962: 126. Hay- Hippothoa ¯agellum Manzoni, 1870 ward and Ryland, 1979: 240. Zabala, 1986: Figure 18F±I 516. Zabala and Maluquer, 1988: 141. Hayward and Ryland, 1999: 100. Hippothoa ¯agellum Manzoni, 1870: 328. Hay- ward and Ryland, 1999: 88. DESCRIPTION (AMNH 930, 932; CMRR 2226): Colonies clear, encrusting, multiserial, DESCRIPTION (AMNH 929±931; CMRR unilaminar, glassy, and often only apparent 2225): Colonies uniserial traceries encrusting once dried. Autozooids oval, fusiform or ir- shell and other organic carbonates, form and regular, only slightly convex; disjunct, adja- extent dictated by space available and com- cent autozooids linked by tubular extensions petition from other sessile organisms. Auto- of basal pore chambers. Primary ori®ce D- zooids slender, clavate, with ®liform proxi- shaped, with straight proximal border and in- mal portion, or cauda, constituting up to distinct condyles. No oral spines or peri- three-quarters total length, and distal oval stome. Frontal shield ®nely granular, imper- portion the rest. Primary ori®ce pear-shaped, forate, transversely ridged or striated, and broadest in its distal third, narrowest at prox- sometimes with a small median suboral imal rim, which forms a deep, U-shaped si- umbo. Ancestrula with peristome and two nus between short, pointed condyles. spines on distal margin of ori®ce. Avicularia Branching a right-angled cross, each auto- vicarious, monomorphic, typically one at dis- zooid producing a single distal bud and tal end of each autozooid; cystid small and paired lateral buds; female brooding zooids rounded, mandible triangular, acute to frontal dimorphic, developed at lateral budding sites plane, directed distally. Ovicell elongate, and rarely giving rise to further buds; broadly semielliptical, recumbent on substratum be- oval, abruptly tapered proximally, without a tween distally succeeding autozooids; ®nely cauda, entire length slightly less than that of granular, imperforate, often developing a lon- distal clavate portion of autozooid; ori®ce al- gitudinal frontal keel; partly incorporated most twice as wide as long, with broad, shal- within cystid of avicularium. low sinus; ovicell imperforate. Ancestrula a Autozooids may be closely packed, with circular, domed, kenozooidal structure, 0.3 linking tubules only just visible, or widely 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 43

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 ®gure (0.2 mm). 44 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 disjunct with intervening spaces in®lled by a proximal edge; surrounded by a short, thick mosaic of irregular kenozooidal units, each peristome surmounted by a transversely oval linked to neighboring kenozooids or auto- secondary ori®ce. No oral spines. Frontal zooids by tubular connections, and each with shield ®nely granular, thickening through on- a small round frontal opesia. togeny, with a single series of slit-like mar- Tentacles clear, 12±13; lophophores bell- ginal pores; a small round spiramen medial- shaped, radially symmetrical; introverts long. ly, distal to it a single adventitious avicular- Embryo color bright orange. ium, with short, triangular rostrum acute to DISTRIBUTION: Chorizopora brongniartii is frontal plane, distally directed and extending a very common species in the Mediterranean, onto proximal part of peristome. No ovicells; and on north European coasts, and is often embryos brooded in modi®ed gonozooids, abundant on every kind of organic substra- broader than autozooids, with proportionate- tum. It is possibly one of the few almost cos- ly larger, broader ori®ce. On settlement larva mopolitan species of cheilostomate bryozo- metamorphoses into a radially symmetrical ans, having been reported, described, and il- complex of six ancestrular zooids, readily lustrated from shelf environments throughout seen in young colonies. the world, with the exception of polar re- Tentacles colorless, 17±18; lophophores gions. slightly obliquely truncate. MEASUREMENTS (SKELETAL): AL 105 Ϯ 14 DISTRIBUTION: This is a warm temperate ␮m, 80±130 (4, 20), AW 102 Ϯ 14, 80±130 species, distributed in the east Atlantic from (4, 20), DO 367 Ϯ 49, 300±380 (4, 20), KL the southwest British Isles to West Africa, 125 Ϯ 25, 80±170 (4, 20), KW 90 Ϯ 26, 50± and abundantly throughout the Mediterra- 120 (4, 20), OL 57 Ϯ 7, 40±70 (4, 20), OW nean. Also present in the warm temperate to 99 Ϯ 10, 80±120 (4, 20), OvL 268 Ϯ 27, tropical west Atlantic, and on the Paci®c 220±320 (4, 16), OvW 217 Ϯ 17, 200±260 coasts of North America. (4, 16), OOW 130 Ϯ 20, 200±260 (4, 16), MEASUREMENTS (SKELETAL): AL 112 Ϯ 21 ZL 429 Ϯ 23, 390±460 (4, 20), ZW 331 Ϯ ␮m, 75±146 (2, 15), AW 64 Ϯ 11, 48±84 (2, 33, 280±380 (4, 20). (POLYPIDE): IH 261 Ϯ 15), DO 439 Ϯ 53, 283±490 (2, 20), OL 70 41 ␮m, 200±320 (1, 11), LD 527 Ϯ 51, 430± Ϯ 9, 58±93 (2, 15), OW 102 Ϯ 8, 91±115 580 (1, 11), MD 26 Ϯ 3, 20±30 (1, 11), TL (2, 15), OOL 86 Ϯ 20, 63±99 (1, 3), OOW 457 Ϯ 23, 420±500 (1, 11). 151 Ϯ 3, 148±154 (1, 3), ZL 633 Ϯ 64, 516± 737 (2, 15), ZW 367 Ϯ 41, 312±449 (2, 15). INFRAORDER UMBONULOMORPHA (POLYPIDE): IH 210 Ϯ 78 ␮m, 60±280 (1, 6), GORDON, 1989 LD 606 Ϯ 132, 400±740 (1, 10), MD 28 Ϯ SUPERFAMILY ADEONOIDEA BUSK, 1884 4, 20±30 (1, 6), TL 520 Ϯ 51, 440±560 (1, FAMILY ADEONIDAE BUSK, 1884 5).

GENUS REPTADEONELLA BUSK, 1884 FAMILY ADEONELLIDAE GREGORY, 1893 Reptadeonella violacea (Johnston, 1847) GENUS ADEONELLA BUSK, 1884 Figure 19D±G Adeonella pallasii (Heller, 1867) Lepralia violacea Johnston, 1847: 325. Figure 20A±E Reptadeonella violacea: Hayward and Ryland, Eschara pallasii Heller, 1867: 115. 1999: 186. Schizoporella pallasii: Hincks, 1886: 268. DESCRIPTION (AMNH 933; CMRR 2227): Adeonella polystomella: Zabala and Maluquer, 1988: 144. Colonies encrusting, multiserial, extensive È unilaminar sheets, deep purple when living, Adeonella pallasii:Unsal and d'Hondt. 1979: 622. Hayward, 1988: 171. with a pale border of actively growing au- tozooids. Autozooids pyriform to regularly LECTOTYPE (chosen here): UIIZ 169. hexagonal, with distinct boundaries, ¯at or DESCRIPTION (AMNH 934±936; CMRR with a slight median concavity; 0.5±0.62 ϫ 2228): Colonies light orange, erect, bilami- 0.3±0.5 in present material. Primary ori®ce nar, branching dichotomously, developing a semicircular, with straight or slightly convex ¯at, two-dimensional structure; to 20 mm 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 45

Fig. 20. A±E. Adeonella pallasii. A. General aspect (lectotype, UIIZ 169; 1 mm). B. Untreated autozooids (lectotype, UIIZ 169; 0.2 mm). C. Ontogenetically young autozooids at branch tip (AMNH 934; 0.1 mm). D. Autozooidal ori®ce (AMNH 935; 0.05 mm). E. Autozooid (AMNH 936; 0.1 mm). F±H. Escharoides mamillata (AMNH 895). F. General aspect (0.2 mm). G. Autozooid (0.1 mm). H. Maternal zooid with ovicell, ¯anked by two avicularia (0.1 mm). 46 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 high in present material; maximum branch Aperture slightly broader than long; distally width 2.5 mm. Autozooids oval to hexago- with a broad, straight-edged suboral shelf, nal, broadest distally, tapered proximally; bordered by four to six short thick spines; bordered by distinct raised sutures. Frontal proximally with a tall, medially pointed mu- shield convex, ®nely granular; pores large cro, with a short, square denticle on its inner and distinct in early ontogeny, later con®ned face, ¯anked on each side by a short pointed to margins of shield. Primary ori®ce as wide denticle. Frontal shield thick, with distinctive as long, proximal edge with two pronounced nodular surface, bordered by densely spaced, cusps de®ning a narrow, U-shaped sinus. round marginal pores. Avicularia adventi- Secondary ori®ce transversely oval, becom- tious, lateral oral, paired, with acute trian- ing more nearly circular in later ontogeny; gular rostrum, slightly acute to frontal plane, peristomial bridge slender, de®ning proxi- distally directed and with a gentle lateral mally a broad oval spiramen through which curve towards the autozooid midline. Ovicell the primary ori®ce is clearly visible. Avicu- hemispherical, with arched, straight-edged laria small, single or paired, proximolateral aperture; surface ®nely nodular, identical to to secondary ori®ce, with bluntly triangular that of frontal shield, with small, indistinct rostrum directed medially onto peristomial pores scattered around its periphery. bridge. Ontogenetic thickening of frontal DISTRIBUTION: Escharoides mamillata is shield producing a prominent umbo proxi- common in Pliocene deposits of southeast molateral to secondary ori®ce on each side, England and the Low Countries but has been and another proximal to spiramen. reported on only few occasions from Recent REMARKS: Heller's specimen chosen here habitats. It is known from two localities off as lectotype and illustrated as Figure 20A northwest Britain, and from the Gulf of Mar- and 20B is one of two specimens in the col- seille, but has not been reported from else- lections of the UIIZ indicated as Heller's where in the northeast Atlantic or the Med- types of the species. iterranean. DISTRIBUTION: This distinctive species is MEASUREMENTS (SKELETAL): AL 188 Ϯ 19 perhaps endemic to the eastern Mediterra- ␮m, 151±228 (2, 20), AW 111 Ϯ 7, 97±121 nean. It is common in the eastern Adriatic (2, 20), DO 607 Ϯ 55, 510±704 (2, 20), OL and the Aegean. Its most westerly record 125 Ϯ 14, 92±149 (2, 19), OW 150 Ϯ 9, seems to be that of Hincks (1886), from Na- 135±65 (2, 19), OvL 286 Ϯ 34, 224±335 (2, ples, which has still to be substantiated. 20), OvW 410 Ϯ 38, 354±478 (2, 20), ZL MEASUREMENTS (SKELETAL): AL 71 Ϯ 9 664 Ϯ 60, 560±795 (2, 20), ZW 514 Ϯ 47, ␮m, 54±82 (2, 16), AW 45 Ϯ 8, 32±58 (2, 438±625 (2, 20). 16), DO 379 Ϯ 27, 331±419 (2, 20), OL 96 Ϯ 7, 84±108 (1, 10), OW 82 Ϯ 6, 70±89 (2, FAMILY UMBONULIDAE CANU, 1904 13), ZL 537 Ϯ 30, 471±583 (2, 20), ZW 399 GENUS UMBONULA HINCKS, 1880 Ϯ 36, 326±462 (2, 20). Umbonula ovicellata Hastings, 1944 Figure 21A±C SUPERFAMILY LEPRALIELLOIDEA VIGNEAUX, 1949 Umbonula ovicellata Hastings, 1944: 280. Hay- FAMILY ROMANCHEINIDAE JULLIEN, 1888 ward and Ryland, 1999: 106. GENUS ESCHAROIDES MILNE EDWARDS, 1836 DESCRIPTION (AMNH 937): Colonies or- Escharoides mamillata (Wood, 1844) ange encrusting, multiserial, unilaminar sheets. Autozooids hexagonal to irregularly Figure 20F±H polygonal, strongly convex, separated by Lepralia mamillata Wood, 1844: 19. thin raised sutures. Aperture slightly longer Escharoides mamillata: Hayward and Ryland, than wide, large, operculum clearly visible 1999: 118. within, its rim developed as a thin, wavy DESCRIPTION (AMNH 895; CMRR 2229): peristome, especially prominent laterally. Colonies spreading, multiserial, unilaminar Midproximally each autozooid with a small sheets. Autozooids oval to hexagonal, steeply suboral avicularium, the oval rostrum orien- convex, separated by well-marked grooves. tated perpendicularly to plane of aperture and 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 47

Fig. 21. A±C. Umbonula ovicellata (AMNH 937). A. General aspect (0.5 mm). B. Ori®ces of ontogenetically young autozooids with suboral avicularia (0.2 mm). C. Ontogenetically mature maternal zooids with ovicells (0.1 mm). D±G. Parasmittina rouvillei. D. General aspect (AMNH 938; 0.5 mm). E. Autozooids (upper right), maternal zooids with ovicells (lower left), and variable-sized vicarious avicularia (AMNH 938; 0.2 mm). F. Autozooidal ori®ce (AMNH 939; 0.05 mm). G. Ori®cial condyle (AMNH 939; 0.01 mm). 48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 facing distally. Frontal shield imperforate burr with light microscopy. Two or three dis- centrally, but with about a dozen deep, mar- tal oral spines present. Peristome developed ginal areolae, from between which prominent proximal to spine bases, later extending over ribs converge on the suboral avicularium, the them, forming low, ¯ared rim, produced mid- cystid of which becomes distinctly umbonate proximally resulting in pear-shaped second- as calci®cation thickens. Ovicell (absent in ary aperture. Frontal shield nodular, with sin- present material) broader than long, ¯attened gle or double series of large marginal pores; frontally, with numerous pores. in later ontogeny ridges of calci®cation ex- Tentacles light orange, 18; lophophores tending between pores onto center of shield, bell-shaped, radially symmetrical to slightly the periphery then appearing reticulate. Av- obliquely truncate. icularia adventitious, polymorphic, often Embryos bright yellow, observed 7 June abundant: small, triangular or parallel-sided 1988. avicularia proximal or proximolateral to ori- DISTRIBUTION: Lower shore and shallow ®ce, directed proximally, triangular type po- sublittoral, on algae, sea grasses and shells. tentially present elsewhere around peristome, Widely distributed in the Mediterranean, often directed towards aperture; large avicu- ranging north to southwest England. laria, with distally rounded, parallel-sided MEASUREMENTS (SKELETAL): AL 109 Ϯ 14 rostrum half as long as autozooid frequent, ␮m, 84±131 (2, 20), AW 111 Ϯ 15, 92±140 budded proximolateral to ori®ce and directed (2, 20), OL 181 Ϯ 17, 156±215 (2, 20), OW proximally. Ovicell slightly longer than 194 Ϯ 14, 162±219 (2, 20), OvL 179 Ϯ 24, broad, globular, with a dense group of vari- 127±203 (1, 10), OvW 210 Ϯ 16, 187±242 ably sized pores distally, the proximal frontal (1, 10), ZL 617 Ϯ 52, 536±701 (2, 20), ZW surface imperforate. 407 Ϯ 41, 328±489 (2, 20). (POLYPIDE): IH Tentacles light yellow, 12±15; lopho- 118 Ϯ 35 ␮m, 60±180 (1, 9), LD 924 Ϯ 134, phores bell-shaped, radially symmetrical be- 640±1080 (1, 9), TLMn 782 Ϯ 200, 400± tween chimneys to obliquely truncate bor- 960 (1, 6), TLMx: 886 Ϯ 190, 400±1050 (1, dering chimneys. 9). REMARKS: The Mediterranean species of Parasmittina are in need of thorough taxo- INFRAORDER LEPRALIOMORPHA nomic revision, including a review of type GORDON, 1989 and other published materials, and this pre- SUPERFAMILY SMITTINOIDEA LEVINSEN, 1909 sent attribution may have to be reconsidered. FAMILY SMITTINIDAE LEVINSEN, 1909 However, the material described here is the same species as that described from Chios GENUS PARASMITTINA OSBURN, 1952 (Hayward, 1974) as P. tropica (Waters, Parasmittina rouvillei (Calvet, 1902) 1909), a taxon then unrecorded for the Med- Figure 21D±G iterranean. It is almost certainly the species described Smittia rouvillei Calvet, 1902: 57. by Gautier (1962) as P. rouvillei and is prob- Parasmittina rouvillei: Gautier, 1962: 199. ably the same as that described by Canu and Parasmittina tropica: Hayward, 1974: 384. Bassler (1930) as Smittina rouvillei. Zabala DESCRIPTION (AMNH 938, 939; CMRR and Maluquer (1988) considered that these 2230): Colony light yellow to orange, en- and other citations referred to a single, vari- crusting, multilaminar. Autozooids hexago- able species which they denoted P. tropica, nal to polygonal, in orderly arrangement at referring to Harmer's (1957) rede®nition. unilaminar growing edges, without common However, it must be pointed out that Smittia orientation in successive laminae; convex, tropica Waters, 1909 was described from the separated by distinct ridges. Primary ori®ce Sudanese Red Sea coast. Harmer's (1957) about as wide as long; proximal border with concept of the species was based on speci- short, anvil-shaped lyrula occupying about mens from Ghardaqa, India, Ceylon, the Tor- half its width; condyles conspicuous, round- res Straits, Victoria, Adelaide, and the Si- ed, downcurved, with denticulate, laminate boga collections from the Indo-Malayan re- structure, revealed by SEM, visible as a faint gion, and it is clear that he did not reexamine 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 49

Waters' original material. Harmer's (1957) il- corporates a prominent avicularium, with lustrations of ''Smittina tropica'' depict four rounded rostrum, complete crossbar bearing specimens from the Siboga collections, one a ligula, and supporting a semicircular man- from India and one from Victoria. It is doubt- dible, proximally directed and normal to ful that any of these corresponds to Waters' frontal plane. Ovicell not seen in present ma- species, or to any species present in the Med- terial; hyperstomial, hemispherical, perfo- iterranean. rate; obscured by peristome and completely MEASUREMENTS (SKELETAL): AL (large) immersed. In late ontogeny, frontal shield 290 Ϯ 60, 175±398 (2, 12), AW (large) 104 thickened to extent that frontal surface of Ϯ 27, 65±154 (2, 12), AL (small) 109 Ϯ 7, colony becomes quite ¯at, peristomes and 101±122 (1, 8), AW (small) 73 Ϯ 15, 49±93 avicularia are immersed, and peristomial (1, 8), DO 523 Ϯ 74, 386±676 (2, 20), LW rims form circular secondary ori®ces; auto- 52 Ϯ 5, 47±62 (1, 9), OL 120 Ϯ 14, 92±143 zooids then have a broadly lingulate outline, (2, 21), OW 119 Ϯ 16, 81±143 (2, 21), ZL each bordered by a distinct suture. 687 Ϯ 69, 566±816 (2, 20), ZW 411 Ϯ 58, Tentacles light orange or yellowish orange, 323±569 (2, 20). (POLYPIDE): IH 120 Ϯ 51 14±16; lophophores bell-shaped, radially ␮m, 40±180 (1, 10), LD 640 Ϯ 78, 560±800 symmetrical across most of branch surface, (1, 10), MD 30±40 (1, 2), TL 646 Ϯ 123, obliquely truncate adjacent to elongate chim- 440±820 (1, 10). ney along branch margin. DISTRIBUTION: Common throughout the GENUS SMITTINA NORMAN, 1903 Mediterranean in shallow coastal waters, ranging south to the Cape Verde Islands and Smittina cervicornis (Pallas, 1766) Ghana. Figure 22A±C MEASUREMENTS (SKELETAL): AL 111 Ϯ 12 Millepora cervicornis Pallas, 1766: 252. ␮m, 95±130 (2, 17), AW 101 Ϯ 23, 72±159 Eschara cervicornis: Milne Edwards, 1836: 19. (2, 17), DO 543 Ϯ 67, 423±669 (2, 20), OL Smittia cervicornis: Calvet in Jullien and Calvet, 164 Ϯ 10, 149±181 (2, 17), OW 134 Ϯ 9, 1903: 151. 121±150 (2, 17), ZL 872 Ϯ 54, 811±977 (2, Porella cervicornis: Canu and Bassler, 1930: 54. 20), ZW452 Ϯ 51, 386±543 (2, 20). (POL- Lagaaij, 1952: 97. Gautier, 1962: 204. YPIDE): IH 158 Ϯ 70 ␮m, 40±280 (1, 23), Smittina cervicornis: Cook, 1968: 210. Zabala and Maluquer, 1988: 121. LDMn 556 Ϯ 83, 460±620 (1, 3), LDMx 610 Ϯ 79, 400±780 (1, 28), TLMn 462 Ϯ DESCRIPTION (AMNH 940; CMRR 2231): 76, 340±560 (1, 16), TLMx 588 Ϯ 114, 340± Colonies orange, erect, branching, to 30 mm 820 (2, 23). in present material, narrow: maximum branch width 3 mm. Branches bilaminar, ¯at- GENUS PRENANTIA GAUTIER, 1962 sectioned, twisting around long axis; each Prenantia cheilostoma (Manzoni, 1869b) face of ®ve or six alternating autozooid rows. Figure 22D±H Autozooids at growing tips of colony elon- gate, rectangular, convex, with distinct Lepralia cheilostomata Manzoni, 1869b: 942. boundaries; commonly 0.625±0.75 x 0.375 Smittia cheilostoma: Hincks, 1880: 349. mm. Primary ori®ce about as wide as long; Smittina cheilostoma: Gautier, 1962: 192. Hay- proximal border straight, a deep lyrula, with ward and Ryland, 1999: 258. Prenantia cheilostoma: Gautier, 1962: 193. straight edge and rounded corners, occupying nine-tenths of proximal width; condyles DESCRIPTION (AMNH 932, 941, 942; rounded quadrangular, conspicuous in early CMRR 2232): Colonies orange, encrusting, ontogeny. No spines. Frontal shield thicken- multiserial, unilaminar sheets, light reddish ing steadily from earliest ontogeny, evenly purple when living. Autozooids hexagonal, pierced by large round pores. Peristome de- gently convex in early ontogeny, later rather veloping within one ontogenetic generation ¯at following ontogenetic thickening of the from growing edge, thickly calci®ed, cylin- frontal shield; separated by shallow grooves. drical and projecting from colony surface, Primary ori®ce wider than long, proximal quite hiding primary ori®ce. Proximally it in- border with an anvil-shaped lyrula occupying 50 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 22. A±C. Smittina cervicornis (AMNH 940). A. General aspect (0.5 mm). B. Autozooid (0.1 mm). C. Autozooidal ori®ce (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 ori®ce (AMNH 942; 0.05 mm). G. Ancestrula (AMNH 932; 0.1 mm). H. Early autozooids surrounding ori®ce (center) of ancestrula covered by autozooids (AMNH 941; 0.1 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 51

about one-third total width; condyles small, DESCRIPTION (AMNH 943; CMRR 2233): rounded, indistinct. No oral spines. Peri- Colonies light cream-tinted orange, erect, stome developing from earliest ontogeny, slender, branching dichotomously, the forming a low thickened rim completely en- branches oval-sectioned, to 2 mm wide in circling ori®ce, mid-proximal portion deeply present material. Each branch face with ®ve U-shaped and channeled on its inner face. alternating, longitudinal series of autozooids, Frontal shield thick, vitreous, densely per- broadening to seven prior to a dichotomy. forated with numerous small pores. Ances- Autozooids elongate, rectangular, separated trula with acute proximal end, spiramen, and by distinct grooves; frontal shield gently con- circular ori®ce slightly smaller than ori®ce of vex, ®nely granular, with few evenly scat- autozooids and encircled by approximately tered pores, each in a shallow pit. Primary 12 oral spines; all but apertural peristome ori®ce slightly longer than wide; a short, covered when ancestrula is completely encir- straight-edged lyrula, with cusped corners, cled by ring of autozooids. Ovicell wider occupying most of proximal edge. No spines. than long, smooth surfaced, with many small A deep tubular peristome developing in early frontal pores; developing a granular, periph- ontogeny, curving frontally and projecting eral ooecial cover continuous with the peri- from frontal plane, completely hiding pri- stome. mary ori®ce. Secondary ori®ce more or less REMARKS: Hincks (1880: 349) used the circular, with a thickened, slightly crenulate trivial name cheilostoma, which Manzoni, rim; a short, pointed tooth apparent mid- (1869b, p. 942) had originally spelled chei- proximally within rim marks edge of a ridge lostomata in the text when describing the descending inner, proximal face of peri- new species, although the name as given by stome. No ovicells in present material. Manzoni on the plate (1869b: pl. 4, ®g. 22) DISTRIBUTION: This delicate species was was cheilostoma. Hincks (1880) did not described from the Adriatic (Heller, 1867) mention both spellings, and therefore his use and has been recorded on very few occasions of cheilostoma does not satisfy Article 24.2.3 since. It was reported from the coast of Tu- of the ICZN Code for selection of one from nisia by Canu and Bassler (1930), and has among two or more spellings in an original been described and illustrated from the Ae- work, but cheilostoma has become estab- gean on two occasions. It is perhaps restrict- lished in the literature to the virtual exclusion ed to the eastern Mediterranean; neither Gau- of cheilostomata for the species, which al- tier (1962) nor Zabala and Maluquer (1988) lows its preservation as ``established usage'' recorded it from the western Mediterranean. under Article 33.3.1 (Ride et al., 1999). MEASUREMENTS (SKELETAL): DO 513 Ϯ 64 DISTRIBUTION: Shallow sublittoral and off- ␮m, 430±689 (2, 20), OL 148 Ϯ 12, 118± shore, often encrusting shell. Common 166 (2, 20), OW 149 Ϯ 16, 128±190 (2, 20), throughout the Mediterranean, ranging north- ZL 784 Ϯ 47, 725±892 (2. 20), ZW 357 Ϯ wards to the western English Channel and 31, 314±438 (2, 20). the Isle of Man. MEASUREMENTS (skeletal): DO 474 Ϯ 66 GENUS SMITTOIDEA OSBURN, 1952 ␮m, 370±627 (2, 20), LW 62 Ϯ 7, 54±81 (2, Smittoidea reticulata 20), OL 132 Ϯ 19, 101±169 (2, 20), OW 142 (J. Macgillivray, 1842) Ϯ 14, 120±171 (2, 20). OvL 214 Ϯ 26, 178± Figure 23E±H 268 (1, 10), OvW 295 Ϯ 20, 260±333 (1, 10), ZL 596 Ϯ 49, 519±709 (2, 20), ZW 375 Lepralia reticulata J. Macgillivray, 1842: 467. Ϯ 30, 314±414 (2, 20). Smittoidea reticulata: Hayward and Ryland, 1999: 264. GENUS PHOCEANA JULLIEN AND CALVET, 1903 DESCRIPTION (AMNH 913, 944): Colonies Phoceana tubulifera (Heller, 1867) encrusting, multiserial, unilaminar sheets. Figure 23A±D Autozooids oval to hexagonal, convex, sep- Eschara tubulifera Heller, 1867: 116. arated by distinct grooves; 0.35±0.5 ϫ 0.25± Smittina tubulifera: Harmelin, 1969b: 299. Hay- 0.3 mm in present material. Primary ori®ce ward, 1974: 382. wider than long, distal rim with ®ne dentic- 52 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 23. A±D. Phoceana tubulifera (AMNH 943). A. General aspect (0.5 mm). B. Autozooid (0.1 mm). C. Autozooidal primary ori®ce (0.05 mm). D. Oblique view of autozooidal secondary ori®ce 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 ori®cial spines in place (AMNH 944). G. Autozooid with suboral avicularium (AMNH 944; 0.1 mm). H. Ancestrula (AMNH 913; 0.1 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 53

ulation revealed by SEM, proximal edge with DESCRIPTION: Lectotype consisting of two anvil-shaped lyrula occupying about half to- fragments from single colony; the larger, tal width, condyles conspicuous, angular, mounted in wooden cavity slide, totaling 24 downcurved. Two to four, most frequently autozooids and an ancestrula; the smaller, three, distal oral spines. A single median su- Canada balsam mount of seven autozooids, boral avicularium, with acute triangular ros- prepared in 1929; all zooids damaged to trum, normal to frontal plane, proximally di- some degree. Autozooids oval, convex, with rected; crossbar slender, complete, palate distinct boundaries; frontal shield densely with extensive foramen. Frontal shield nod- perforated by large round pores with thick- ular, bordered by a single series of large, dis- ened rims. Primary ori®ce slightly wider than tinct pores. Ovicell slightly longer than long, proximal border shallowly concave be- broad, aperture arched, with straight distal tween small condyles. No oral spines; prox- edge; about 20 small pores on distal and lat- imal rim of ori®ce thickened, with median eral surfaces. Ancestrula elongate oval, suboral umbo bearing small avicularium with smoothly convex, tapered proximally; gym- semielliptical, proximally directed mandible. nocyst smooth; membranous frontal wall oc- Ovicell, damaged in all examples, globose, cupying half total length, surrounded by recumbent upon distally succeeding zooid, oval, crenulate rim bordered by four closely with frontal pores evident in undamaged ar- spaced distal spines and ®ve spines uniform- eas. Oval ancestrula with raised mural rim ly spaced proximally; cryptocyst broadest bordering membrane and indeterminate num- proximally, opesia almost circular, two-thirds ber of spines; broad, ¯at cryptocyst de®ning length of membrane. semielliptical opesia. Substratum apparently REMARKS: The material described and il- an alga frond. lustrated here has a proportionately broader REMARKS: We did not encounter this spe- lyrula and longer avicularium than north Eu- cies; the lectotype is the only extant speci- ropean material ®gured by Hayward and Ry- men. It is labeled in Hincks' hand with its land (1999), but consists of small colonies original taxonomic designation, and also `` ϭ only, in which ontogenetic variation may not S. marsupifera Busk''. The curious synony- be fully expressed. my with the latter, described from a Chal- DISTRIBUTION: This species is distributed in lenger station off Marion Island, South In- the eastern Atlantic from Arctic Norway to dian Ocean, may thus have originated with Morocco, and throughout the Mediterranean. Hincks. Gautier (1962) had a single speci- MEASUREMENTS (SKELETAL): AL 102 Ϯ 14 men, which he lost; he then repeated Hincks' ␮m, 70±128 (2, 20), AW 55 Ϯ 10, 41±74 (2, description of the species, attributing it to 20), DO 349 Ϯ 46, 276±449 (2, 20), OL 93 Busk's taxon. Ϯ 11, 73±111 (3, 26), OW 106 Ϯ 15, 79± MEASUREMENTS (SKELETAL): OL 110 Ϯ 10 141 (3, 26), OvL 224 Ϯ 26, 192±273 (1, 10), ␮m (1, 10), OW 130 Ϯ 8 (1, 10), ZL 470 Ϯ OvW 209 Ϯ 17, 183±230 (1, 10), ZL 506 Ϯ 50 (1, 10), ZW 310 Ϯ 40 (1, 10). 40, 443±618 (2, 20), ZW 288 Ϯ 34, 223±348 (2, 20). GENUS PENTAPORA FISCHER, 1807 Pentapora fascialis (Pallas, 1766) FAMILY BITECTIPORIDAE MACGILLIVRAY, 1895 Figure 25A±E GENUS HIPPOPORINA NEVIANI, 1895 Eschara fascialis Pallas, 1766: 42. Hippoporina lineolifera (Hincks, 1886) Millepora foliacea Ellis and Solander, 1786: 133. Figure 24 Eschara foliacea: Busk, 1854: 89. Lepralia foliacea: Hincks, 1880: 300. Schizoporella lineolifera Hincks, 1886: 267, Hippodiplosia foliacea: Canu and Bassler, 1930: 1887: 315. 50. ? Schizoporella marsupifera: Gautier, 1962: 144. Pentapora fascialis: Zabala, 1986: 401. Hayward and Ryland, 1999: 200. LECTOTYPE (chosen here): BMNH reg. no. 1899.5.1.1072, Hincks collection, Adriatic; DESCRIPTION (AMNH 945±947; CMRR The Natural History Museum, London. 2234): Colonies red, erect, rigid, heavily cal- 54 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

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).

ci®ed; consisting of bilaminar plates, divid- togeny, when individual autozooid boundar- ing and anastomosing at irregular intervals, ies become obscured. Ovicell elongate oval, forming a three-dimensional coralliform with scattered frontal pores, visible only in structure. Deep orange to red-orange when early ontogeny, soon obscured by thickening living. Autozooids elongate, oval to hexag- calci®cation. onal; distinct when newly budded, each sep- Tentacles light orange, 17±19; lopho- arated by thick, raised ridges. Primary ori®ce phores bell-shaped, radially symmetrical narrowly bell-shaped, longer than wide, with away from colony edge, obliquely truncate straight or slightly convex proximal margin along colony margins and locally (at chim- between small, triangular, downcurved con- neys?) on colony surface. dyles. No oral spines. A small suboral avi- REMARKS: The colony habit is the narrow cularium sporadically present, missing from fronded, open form characteristic of Medi- many autozooids; rostrum short, oval, slight- terranean populations. This is the largest ly acute to frontal plane, proximally directed; and most conspicuous calci®ed bryozoan in occasionally replaced by a dimorphic avicu- the Adriatic, with colonies up to 0.5 m high larium with slightly spatulate rostrum, 0.2 and 0.3 m in diameter in the vicinity of Sv. mm long, normal to frontal plane, proximally Ivan Island before the anoxic events of the directed. Frontal shield convex, evenly per- late 1980s. Colonies established after the forated; thickening rapidly from earliest on- anoxic event had grown to exceed 15 cm in togeny, with ridges developing between the diameter by summer 1998, although reestab- pores, fusing to form a reticulate shield lishment of large colonies may require many which continues to thicken through later on- more years due to exponential decrease in 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 55

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 calci®ed 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 ori®ce (0.05 mm). 56 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 growth rate with increased size of colonies GENUS SCHIZOMAVELLA CANU AND (Cocito et al., 1998a). Recovery of P. fas- BASSLER, 1917 cialis from the northern Adriatic anoxic Schizomavella asymetrica (Calvet, 1927) events of the late 1980s has been much Figure 25F±H slower than 3.5-year recovery of the species at a shallower site in the Ligurian Sea fol- Schizoporella auriculata var. asymetrica Calvet, lowing a severe storm in the early 1990s 1927: 20. Gautier, 1962: 134. (Cocito et al., 1998b). Where little disturbed Calyptotheca triarmata Hayward, 1974: 381, ®g. by humans, P. fascialis normally grows in 5b. exposed positions; large colonies offshore Schizomavella cuspidata (in part): Reverter-Gil and FernaÂndez-Pulpeiro, 1995: 265 (®g. 4G of Rovinj commonly are found attached to only). Hayward and Ryland, 1999: ®g. 131D gorgonians or ¯exible bryozoans, which re- only. duces wave and current fragmentation. In other areas of the Mediterranean, P. fasci- DESCRIPTION (AMNH 948): Colonies en- alis grows in exposed positions on rock in crusting sheets, multiserial, unilaminar or depths as shallow as 11 m, although single with frontally budded patches. Autozooids severe storms can eliminate such shallow rectangular to irregularly polygonal, rather populations (Cocito et al., 1998b). In areas ¯at, separated by distinct, raised sutures. Pri- frequented by divers, colonies are less fre- mary ori®ce wider than long, its greatest quent, smaller, and more cryptic (Sala et al., width at midlength; proximal border with a 1996). shallow, U-shaped median sinus, occupying OCCURRENCE: With the exception of two one-®fth of its width, accentuated by raised localities (18, 19) approximately 20 m deep shoulders and broad, deep condyles which where colony fragments were dredged, Pen- extend from the edges of the sinus, almost to tapora fascialis was found at 29 m and great- lateral rim of ori®ce. Two or three indistinct er depths (summer thermocline is at about 29 distal oral spine bases visible in some auto- m depth; MarinkovicÂ, 1959), growing on zooids, but most apparently lack spines en- rock walls or more commonly on cobbles or tirely. Frontal shield thick, nodular, perforat- on gorgonian stalks. Large colonies, over ed by large, well-spaced pores, the marginal 30ϫ20ϫ20 cm, were especially common on pores especially conspicuous. Avicularium gorgonian stalks attached to cobbles and median suboral, on a small prominence, boulders on the sand ¯oor northeast of Pe- close to proximal ori®ce rim but distinct lago Island at 30 m and greater depth. from it; monomorphic, the rostrum parallel DISTRIBUTION: Found from the Hebrides to frontal plane, proximally directed, small, and the western coasts of Britain and Ireland broadly oval, with a complete crossbar but southwards, and throughout the Mediterra- no palate. Ovicell distinctive: hyperstomial, nean. Characteristic of current-swept coarse recumbent on distally succeeding autozooid grounds. but projecting well above frontal plane; MEASUREMENTS (SKELETAL): AL(normal) broader than long, the frontal surface pierced 93 Ϯ 19 ␮m, 63±132 (2, 20), AW(normal) by as many as 20 small pores, developing a 71 Ϯ 15, 43±101 (2, 20), AL(spatulate) 335 spiky, distal ooecial cover. Ϯ 29, 287±372 (1, 6), AW(spatulate) 188 Ϯ REMARKS: Elevation of Calvet's (1927) va- 15, 163±206 (1, 6), DO 532 Ϯ 102, 383±757 riety asymetrica to full species rank is (2, 20), OL 197 Ϯ 26, 164±247 (2, 20), OW prompted by Reverter-Gil and FernaÂndez- 165 Ϯ 20, 132±199 (2, 20), OvL 363 Ϯ 20, Pulpeiro (1995), who published a micrograph 320±383 (1, 10), OvW 315 Ϯ 17, 294±345 showing the primary ori®ce in a specimen (1, 10), ZL 815 Ϯ 70, 658±966 (2, 20), ZW which they stated to be identical to material 451 Ϯ 40, 383±539 (2, 20). (POLYPIDE): IH labeled ``var. asymetrica'' in Calvet's hand. 152 Ϯ 95 ␮m, 60±440 (3, 19), LD 735 Ϯ It is unquestionably the same species as that 121, 540±920 (3, 25), MDMn 28 Ϯ 5, 20± described and ®gured here. Reverter Gil and 35 (2, 11), MDMx 34 Ϯ 7, 20±40 (2, 11), FernaÂndez Pulpeiro (1995) considered var. TLMn 639 Ϯ 135, 380±920 (3, 23), TLMx asymetrica to fall within the range of varia- 694 Ϯ 116, 400±920 (3, 24). tion displayed by S. cuspidata (Hincks, 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 57

1880) as recently rede®ned (Hayward and 10), OW 98 Ϯ 5, 86±104 (1, 10), OvL 278 Thorpe, 1995). However, the primary ori®ce Ϯ 17, 250±306 (1, 10), OvW 280 Ϯ 28, of that species differs subtly in shape from 240±320 (1, 10), SL 11 Ϯ 2, 9±15 (1, 10), that of var. asymetrica, being more rounded SW 20 Ϯ 2, 18±26 (1, 10), ZL 511 Ϯ 27, laterally, and the avicularia are dissimilar in 472±548 (1, 10), ZW 359 Ϯ 20, 324±382 (1, the two species. In S. cuspidata the avicular- 10). ium is always elongate oval and is polymor- phic; the proximal rim of the rostrum is con- Schizomavella cornuta (Heller, 1867) tinuous with the proximal rim of the primary Figure 26A±D ori®ce; ®nally, in all avicularia there is a sub- stantial palate with a characteristic trifoliate Lepralia cornuta Heller, 1867: 110. foramen. In S. asymetrica the avicularium is Schizoporella auriculata var. cuspidata Hincks, monomorphic, shortly oval and lacks a pal- 1880: 261. ate; further, it is distant from, and quite un- Schizomavella cuspidata: Hayward and Ryland, attached to, the rim of the primary ori®ce. 1999: 286, ®g. 131A±C (not 131D). The frontal shield of S. cuspidata has fewer, NEOTYPE (chosen here): UIIZ 270. smaller pores than that of S. asymetrica, and DESCRIPTION (AMNH 895, 949; CMRR the ovicell is far less prominent, with fewer, 2235): Colonies cream-colored, encrusting, larger pores, while the projecting ovicell of multiserial, unilaminar to multilaminar, often S. asymetrica bears up to 20 small pores developing thick nodular formations. Auto- frontally. zooids rectangular in newly grown, unilam- Calvet (1927) characterized var. asymetri- inar colonies, usually irregularly polygonal ca especially by the suboral avicularium, in multilaminar colonies developed through which he noted to be umbonate and laterally frontal budding; individual boundaries gen- displaced with reference to the sinus. Gautier erally distinct, as narrow, raised ridges. Pri- (1962) noted that in specimens from the mary ori®ce characteristic: wider than long, western Mediterranean, the avicularium was either symmetrically or asymmetrically de- with greatest width at midlength; proximal veloped. In the present material the avicular- border with a short, symmetrical, U-shaped ium is consistently median suboral in posi- sinus de®ned by short lobes of the proximal tion, but in specimens of what is certainly ori®ce rim, proximal corners of ori®ce deep- the same species from Chios, Aegean Sea, ly notched; condyles short, broad and thick, some autozooids display asymmetrical de- extending from, and emphasizing, the sinus velopment of the avicularium, while in others into the proximal corners of the ori®ce. Two a median avicularium may be accompanied to six distal oral spines present in early on- by one or two lateral suboral avicularia. togeny, commonly obscured during later on- These specimens were described as Calyp- togeny. Ancestrula small, tatiform, opesia totheca triarmata by Hayward (1974), and surrounded by nine marginal spines. Avicu- reexamination of the holotype specimen larium suboral, proximally directed, with ros- (NHM 1973.4.4.4) shows C. triarmata to be trum almost parallel to frontal plane; dimor- a junior subjective synonym of Schizomav- phic: short, oval, ca. 0.06 mm long, or ca. ella asymetrica (Calvet, 1927). 0.2 mm long, narrowly spoon-shaped, with DISTRIBUTION: The species was ®rst de- slightly concave sides and gently expanded scribed from the vicinity of Monaco (Calvet, distal end. In both types of avicularia, prox- 1927); Gautier (1962) recorded specimens imal rim of rostrum continuous with outer from the same region and also from the Gulf rim of sinus, and a thick cross bar with a of Marseille and Tunisia. The specimen il- stout columella divides proximal portion of lustrated by Hayward and Ryland (1999, ®g. narrow palatal foramen to form a distinctive 131D) is a Norman collection specimen Y shape. Ovicell broader than long, ¯attened (NHM 1911.10.1.1538) from an unknown frontally, with few (ca. 5) large, irregular British locality. pores immediately distal to the arched aper- MEASUREMENTS (SKELETAL): DO 378 Ϯ 46 ture; developing a nodular ooecial cover in ␮m, 330±445 (1, 10), OL 88 Ϯ 5, 82±97 (1, later ontogeny. 58 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 26. A±D. Schizomavella cornuta. A. General aspect (neotype, UIIZ 270; 0.5 mm). B. Ori®ce of autozooid (neotype, UIIZ 270; 0.05). C. autozooids and kenozooid (bottom right), with variable-sized suboral avicularia (AMNH 949; 0.1 mm). D. Maternal zooids with ovicells (AMNH 895; 0.1 mm). E± G. Schizomavella linearis (AMNH 950). E. General aspect (0.5 mm). F. Autozooids and (bottom center) maternal zooid with ovicell (0.1 mm). G. autozooidal ori®ce (0.05 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 59

Embryo color medium orange to dark, gular to polygonal, rather ¯at, vertical walls brick-red. distinct; frontal shield nodular, with conspic- REMARKS:AsSchizomavella cuspidata, uous marginal perforations, and few, large this species has only recently been rede- pores medially. Primary ori®ce as wide as scribed and distinguished from S. auriculata long, broadest at midlength; sinus short, U- (Hassall), with which it was for long con- shaped, occupying about half width of prox- fused (Hayward and Thorpe, 1995). How- imal border; condyles short, broad and ever, as recognized by Friedl (1917: 272), rounded. Two or three slender distal oral part of Heller's material of Lepralia cornuta spines present. Avicularia single or paired, in the University of Innsbruck collections has lateroproximal to ori®ce; rostrum narrowly af®nities with Schizomavella auriculata (re- acuminate, typically directed medially or ferred to as Smittina auriculata by Heller). proximo-medially, occasionally distally. We designate UIIZ specimen 270, illustrated Large adventitious avicularia sporadic, char- here as ®g. 26A and 26B, as neotype of Le- acteristic: cystid domed, occupying entire pralia cornuta because it may not have been frontal shield of bearing autozooid, mandible in Heller's possession when he named the narrowly crescentic, 0.25 mm wide. Ovicells species. If eventually discovered to be a broader than long, ¯attened frontally, with specimen on which Heller originally de- numerous irregular pores and an umbonate scribed Lepralia cornuta, it should be rede- ooecial cover. signated as lectotype. UIIZ 270 is conspeci®c DISTRIBUTION: This species is common on with S. cuspidata, and Lepralia cornuta is north European coasts. It has been recorded senior objective synonym of the two. previously in the Mediterranean but is per- DISTRIBUTION: It is common on northwest haps far less frequent on southern European European coasts, although its geographical coasts. It was the least common of the de- and ecological Distribution is still incom- scribed species of Schizomavella found at pletely known. In the Mediterranean its oc- Rovinj. currence is even more imprecisely described MEASUREMENTS (SKELETAL): AL(normal) through confusion with numerous other spe- 107 Ϯ 9 ␮m, 88±121 (2, 21), AW(normal) cies of Schizomavella. The ovicell ®gured by 88 Ϯ 10, 70±105 (2, 21), DO 412 Ϯ 78, 315± Hayward and Ryland (1999, ®g. 131D) is not 576 (2, 20), OL 108 Ϯ 10, 90±122 (2, 20), that of S. cuspidata, but rather S. asymetrica, OW 91 Ϯ 8, 81±106 (2, 20), OvL 256 Ϯ 22, here redescribed for the ®rst time. 236±301 (1, 10), OvW 297 Ϯ 30, 282±329 MEASUREMENTS (SKELETAL): AL(normal) (1, 10), SL 24 Ϯ 4, 19±29 (2, 20), SW 26 Ϯ 63 Ϯ 9 ␮m, 48±81 (2, 20), AW(normal) 68 7, 16±38 (2, 20), ZL 491 Ϯ 47, 426±586 (2, Ϯ 8, 54±84 (2, 20), AL(spatulate) 149 Ϯ 16, 20), ZW 333 Ϯ 50, 265±461 (2, 20). 131±175 (1, 6), AW(spatulate) 65 Ϯ 9, 55± 76 (1, 6), DO 409 Ϯ 66, 317±526 (2, 20), Schizomavella mamillata (Hincks, 1880) OL 86 Ϯ 6, 75±98 (2, 20), OW 102 Ϯ 4, Figure 27A±C 94±104 (2, 20), SL 20 Ϯ 2, 16±24 (2, 20), Schizoporella linearis var. mamillata Hincks, SW 27 Ϯ 4, 21±34 (2, 20), ZL 458 Ϯ 39, 1880: 248. 383±510 (2, 20), ZW 347 Ϯ 52, 244±460 (2, Schizomavella mamillata: Gautier, 1958: 199. Za- 20). bala, 1986: 475. Zabala and Maluquer, 1988: 132. Schizomavella linearis (Hassall, 1841) Schizomavella linearis var. mamillata: Gautier, 1962: 143. Harmelin, 1969a: 1201. Figure 26E±G DESCRIPTION (AMNH 951, 952; CMRR Lepralia linearis Hassall, 1841: 368. 2237): Colonies orange, encrusting, multi- Schizomavella linearis: Gautier, 1962: 140. Za- serial, multilaminar, developing a massive bala, 1986: 473. Zabala and Maluquer, 1988: 132. Hayward and Thorpe, 1995: 671. mamillate form. Autozooids rectangular to irregularly polygonal, in regular series at un- DESCRIPTION (AMNH 950; CMRR 2236): ilaminar growing edges, randomly orientated Colonies encrusting, multiserial, unilaminar in areas of vigorous frontal budding; bound- to multilaminar sheets. Autozooids quadran- aries always distinct, marked by thin raised 60 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 27. A±C. Schizomavella mamillata. A. General aspect (AMNH 951; 0.5 mm). B. Autozooids (AMNH 952; 0.1 mm). C. Autozooidal ori®ce (AMNH 952; 0.05 mm). D±H. Schizomavella rudis. D. General aspect (AMNH 953; 0.5 mm). E. Maternal zooid with ovicell (AMNH 953; 0.1 mm). F. Autozooid (AMNH 953; 0.1 mm). G. Autozooidal ori®ce (AMNH 954; 0.05 mm). H. Ancestrula and early autozooids (AMNH 932; 0.1 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 61 sulci. Primary ori®ce longer than wide, ta- rous and with small areolae, lacking suboral pered proximally, midproximal border with a umbo; autozooidal ori®ce roughly equidi- short, symmetrical rounded notch; thick, fac- mensional, lacking distal spines and ori®cial eted condyles on each side of notch de®ne a peristome, proximal sinus relatively wide deep, V-shaped sinus. Oral spines absent to and de®ned by essentially smooth condyles. four, most commonly two. Peristome devel- Suboral, small, oval, laterally or proximally oped as a thin, inconspicuous rim, most pro- orientated adventitious avicularium present nounced, and slightly ¯ared, on each side of on some autozooids. Ovicells prominent, sinus. Frontal shield convex, nodular and vit- emergent, perforated by pores distributed in reous, evenly perforated. Avicularium diag- a semi-lunar region partially encircling a nostic: median suboral, monomorphic, di- proximal, pore-free region adjacent to the or- rected proximally, slightly acute to frontal i®ce. plane; rostrum elongate triangular, its tip Tentacles 13±15; lophophores bell-shaped, downcurved and ®nely denticulate; crossbar radially symmetrical midway between chim- complete, with median columella, palate with neys to strongly obliquely truncate (some trifoliate foramen. Ovicell hyperstomial, campylonemidan) at chimneys and colony prominent, about as wide as long, with 10± margins; adjacent lophophores overlapped up 15 large pores frontally. to at least 50 ␮m. Tentacles light orange, 14±15; lopho- REMARKS: Reverter-Gil and FernaÂndez- phores bell-shaped, radially symmetrical be- Pulpeiro (1995) characterized S. rudis as usu- tween chimneys and obliquely truncate ad- ally having pores on the ovicell con®ned to jacent to chimneys. a central, essentially circular area, but occa- REMARKS: We here designate as lectotype sionally having them distributed in a semi- BMNH specimen 1899.5.1.1059, Hincks lunar region partially encircling a proximal, Collection, from Algiers. pore-free region adjacent to the ori®ce. All DISTRIBUTION: Apparently widespread and specimens of S. rudis examined from the vi- common throughout the Mediterranean, on cinity of Rovinj have the semilunar distri- shallow, coarse detrital grounds. bution of pores. MEASUREMENTS (SKELETAL): AL 191 Ϯ 26 OCCURRENCE: Encrusts diverse bivalve ␮m, 153±247 (2, 210), AW 127 Ϯ 17, 104± substrata. 151 (2, 21), DO 520 Ϯ 90, 351±664 (2, 20), DISTRIBUTION: Gautier (1962) recorded this OL 106 Ϯ 11, 94±141 (2, 20), OW 100 Ϯ species from numerous localities in the west- 8, 88±116 (2, 20), SL 28 Ϯ 6, 19±36 (2, 20), ern Mediterranean, from 20±80 m depth, SW 32 Ϯ 4, 25±42 (2, 20), ZL 690 Ϯ 70, most often encrusting biogenic carbonates. 563±822 (2, 20), ZW 398 Ϯ 45, 351±502 (2, MEASUREMENTS (SKELETAL): AL 57 Ϯ 11 20). (POLYPIDE): IH 110 Ϯ 37 ␮m, 60±180 ␮m, 50±70 (3, 15), AW 52 Ϯ 13, 40±90 (3, (1, 7), LDMn 478 Ϯ 65, 420±660 (1, 12), 15), DO 377 Ϯ 64, 270±480 (3, 30), OL 101 LDMx 500 Ϯ 68, 420±660 (1, 12), MD 26 Ϯ 9, 80±110 (3, 15), OW 100 Ϯ 10, 80±120 Ϯ 2, 25±30 (1, 4), TLMn 514 Ϯ 38, 470± (3, 15), ZL 530 Ϯ 62, 440±620 (3, 15), ZW 580 (1, 8), TLMx 540 Ϯ 66, 480±680 (1, 8). 360 Ϯ 55, 300±480 (3, 15). (POLYPIDE): IH 165 Ϯ 74 ␮m, 100±310 (2, 15), LDMn 556 Schizomavella rudis (Manzoni, 1869) Ϯ 82, 440±720 (2, 12), LDMx 608 Ϯ 113, Figure 27D±H 460±780 (2, 18), MD 25.7 Ϯ 2.6, 25±30 (2, 16), TLMn 462 Ϯ 98, 300±600 (2, 13), Lepralia rudis Manzoni, 1869a: 18. TLMx 649 Ϯ 134, 400±840 (2, 16). Schizomavella rudis: Gautier, 1962: 146. Zabala, 1986: 478. Zabala and Maluquer, 1988: 132. Reverter-Gil and FernaÂndez-Pulpeiro, 1995: Schizomavella subsolana, new species 261. Figure 28A±D

DESCRIPTION (AMNH 932, 953, 954 HOLOTYPE: AMNH 955. CMRR 2238): Colonies cream-colored, or- PARATYPES: AMNH 956, 901; CMRR ange, or red; multiserial, encrusting, unilam- 2239±2240. inar. Autozooidal frontal wall uniformly po- DIAGNOSIS: Primary ori®ce nearly equidi- 62 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 28. A±D. Schizomavella subsolana. (holotype, AMNH 955) A. General aspect (0.5 mm). B. Maternal zooids with ovicells and (top) ontogenetically young zooids (0.1 mm). C. Autozooidal ori®ce of young zooid with incompletely formed avicularium (0.05 mm). D. Ovicell (0.05 mm). E, F. Waters- ipora complanata (AMNH 957). E. General aspect (0.5 mm). F. Autozooid (0.1 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 63 mensional, with shallow sinus; three to ®ve ␮m, 74±115 (2, 20), AW 95 Ϯ 12, 78±130 distal oral spines; single small suboral avi- (2, 20), DO 366 Ϯ 42, 274±433 (3, 30), OL cularium with elongate triangular rostrum, 85 Ϯ 6, 78±100 (2, 12), OW 88 Ϯ 8, 78± steeply inclined to plane of ori®ce. 104 (2, 20), OvL 201 Ϯ 27, 155±248 (3, 30), ETYMOLOGY: The species name is from the OvW 205 Ϯ 15, 178±239 (3, 30), SL 18 Ϯ Latin sub (under) and sol (sun) in recognition 7, 9±34 (2, 12), SW 50 Ϯ 6, 35±56 (2, 12), of its discovery along a coast that is a pop- ZL 431 Ϯ 30, 386±490 (3, 30), ZW 309 Ϯ ular summer holiday destination. 36, 275±339 (3, 30). DESCRIPTION: Colonies cream-colored, small, rounded, multiserial, unilaminar FAMILY WATERSIPORIDAE VIGNEAUX, 1949 patches. Autozooids in regular, radiating GENUS WATERSIPORA NEVIANI, 1895 lines, oval to hexagonal, separated by deep Watersipora complanata (Norman, 1864) grooves. Frontal shield steeply convex, dis- Figure 28E, F tinctly umbonate distally, umbo incorporat- ing median, suboral avicularium; rugose, Lepralia complanata Norman, 1864: 85. evenly perforated by small, round pores. Pri- Watersipora complanata: Hayward and Ryland, mary ori®ce slightly wider than long, a shal- 1999: 192. low sinus occupying more than half its prox- DESCRIPTION (AMNH 957; CMRR 2241): imal width, between short, inconspicuous Colonies brown, spreading multiserial, uni- condyles. Three to ®ve (commonly four) dis- laminar sheets. Autozooids oval to hexago- tal oral spines present; peristome developed nal, ¯at or slightly convex, separated by dis- as a low rim, extending, on each side, from tinct raised ridges. Primary ori®ce as wide as proximal-most spine to avicularium, deepest long, more or less bell-shaped, with lateral proximally. Avicularium almost perpendicu- rims curving inwards towards small, down- lar to plane of ori®ce, rostrum elongate tri- curved condyles just above the proximal bor- angular. Ovicell slightly broader than long, der, which is straight-edged but arched fron- ¯attened frontally, with 20±30 irregular tally. No spines. Ori®ce rim thickens in later pores; initially smooth, but developing a ontogeny, forming pronounced distal and coarse ooecial cover, co-extensive with the proximal lips. Frontal shield smooth, vitre- frontal shield of distally succeeding auto- ous, glistening, densely perforated by small zooid. round pores, except for a small area imme- REMARKS: The taxonomy of north Euro- diately proximal to ori®ce. No avicularia. No pean species of Schizomavella was reviewed ovicells. and revised by Hayward and Thorpe (1995), DISTRIBUTION: Watersipora complanata is who redescribed and illustrated ®ve species. probably widely distributed in the Mediter- Subsequently, new species have been de- ranean. although it has been reported on rel- scribed from both north European and Med- atively few occasions. Outside of the Medi- iterranean localities (Reverter-Gil and Fer- terranean, it is known only from the Scilly naÂndez-Pulpeiro, 1995, 1997), and the tax- Isles, southwest England, where it is com- onomy of some west Mediterranean species mon on the lower shore, encrusting a wide has also been clari®ed. However, the system- range of substrata. atics of the genus in the Mediterranean as a MEASUREMENTS (SKELETAL): DO 471 Ϯ 75 whole remains in need of revision. This new ␮m, 344±709 (3, 30), OL 121 Ϯ 13, 100± species differs from all of those recently de- 166 (3, 30), OW 171 Ϯ 22, 135±211 (3, 30), scribed or redescribed; its distinctive suite of ZL 692 Ϯ 66, 588±813 (3, 30), ZW 390 Ϯ characters includes the primary ori®ce, with 61, 277±503 (3, 30). an especially broad sinus, three to ®ve oral spines, and the comparatively large, frontally Watersipora subovoidea (d'Orbigny, 1852) directed avicularium. It does not seem to Figure 29A, B have been noted in any of the literature treat- Cellepora ovoidea Audouin, 1826: 238. Savigny, ing ``varieties'' of Mediterranean Schizomav- 1809: pl. 8, ®g. 1. ella species (e.g., Gautier, 1962). Cellepora subovoidea d'Orbigny, 1852: 402. MEASUREMENTS (SKELETAL): AL 88 Ϯ 11 Lepralia cucullata Busk, 1854: 81. 64 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 29. A, B. Watersipora subovoidea (AMNH 958). A. General aspect (0.5 mm). B. Autozooid (0.1 mm). C±F. Schizobrachiella sanguinea. C. General aspect of surface with bryozoan-calci®ed tubes that enclosed commensal hydroid stolons along some colony boundaries (AMNH 959; 0.5 mm). D. Autozooid (AMNH 960; 0.1 mm). E. Maternal zooid with ovicell (AMNH 950; 0.1 mm). F. Calci®ed butes that enclosed stolons of commensal hydroid (AMNH 959; 0.1 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 65

Watersipora subovoidea: Gautier, 1962: 183. Za- they refer to an inadequately discriminated bala, 1986: 396. species complex. DISTRIBUTION: Throughout the Mediterra- DESCRIPTION (AMNH 958; CMRR 2242): nean, and for some unknown distance be- Colonies spreading, multiserial, unilaminar, yond, depending upon resolution of the tax- deep purple to black sheets to erect bilaminar onomy of the species or species complex. plates (not seen in present material), often MEASUREMENTS (SKELETAL): DO 682 Ϯ 91 with zooidal boundaries and colonial orga- ␮m, 500±820 (2, 20), OL 239 Ϯ 1 2, 219± nization dif®cult to recognize in living spec- 265 (2, 20), OW 249 Ϯ 14, 216±266 (2, 20), imens. Autozooids with thick, deep purple to SL 56 Ϯ 11, 39±78 (2, 20), SW 127 Ϯ 14, black epitheca, and deep black operculum. 96±148 (2, 20), ZL 998 Ϯ 114, 772±1236 (2, Autozooidal skeletons with densely and uni- 20), ZW 514 Ϯ 65, 378±673 (2, 20). (POL- formly perforated cryptocystidean frontal YPIDE): IH 492 Ϯ 81 ␮m, 340±700 (1, 17), shield. Primary ori®ce slightly wider than LD 1261 Ϯ 178, 1060±1620 (1, 12), MDMn long, with robustly thickened rim and broad, 35 Ϯ 6, 30±40 (1, 4), MDMx 42.5 Ϯ 5, 40± U-shaped sinus occupying half of width of 50 (1, 4), TL 1154 Ϯ 129, 900±1400 (1, 16), proximal edge between pointed, upturned TN 25 Ϯ 1.3, 23±28 (1, 16). condyles. Tentacles bright red (carmine), 23±26; SUPERFAMILY SCHIZOPORELLOIDEA lophophores bell-shaped, radially symmetri- JULLIEN, 1882 cal, supported on long introverts. FAMILY SCHIZOPORELLIDAE JULLIEN, 1882 REMARKS: All of our material formed jet GENUS SCHIZOBRACHIELLA CANU AND black, unilaminar sheets encrusting rock, BASSLER, 1920 with the black color extending all the way to Schizobrachiella sanguinea (Norman, 1868) the growing edge, and with zooidal bound- Figure 29C±F aries and opercula indistinguishable under the microscope when alive. Soule and Soule Hemeschara sanguinea Norman, 1868: 222. (1975) and Gordon (1989) discussed some of Schizoporella sanguinea: Hincks, 1880: 252. Harmer, 1902: 303. the tortuous synonymy of a group of con- Schizobrachiella sanguinea: Canu and Bassler, fused watersiporids that includes ``W. subo- 1930: 32. Gautier, 1962: 128. Zabala, 1986: voidea.'' Gordon (1989) justi®ed use of the 460. Zabala and Maluquer, 1988: 130. Hayward epithet subtorquata in preference to subo- and Ryland, 1999: 222. voidea for the widespread fouling species DESCRIPTION (AMNH 959, 960; CMRR that occurs in the Paci®c ocean. Our speci- 2243): Colonies with bright red cuticle, en- mens exhibit several differences from the crusting, multiserial, unilaminar to multilam- characteristics of W. subtorquata as de- inar, occasionally as erect, folded plates only scribed and illustrated in Gordon (1989) and partly attached to the substratum; deep red Gordon and Mawatari (1992). The New Zea- when living, dull brown when dead and land W. subtorquata has smaller (0.74±0.85 dried. Autozooids rectangular to irregularly mm length), orange lophophores (very light- polygonal, ¯at or slightly convex, separated colored in Gordon and Mawatari, 1992, Plate by thin, raised sutures. Primary ori®ce wider 1D) comprising 24 tentacles; colonies grade than long; proximal border with a short, U- from black or dark gray in colony centers to shaped sinus occupying about one-third its dull orange peripherally, with broad orange width, de®ned by prominent shoulders which margins; and the species is a common fouler intersect on each side with broad, angular in harbors and marinas, whereas specimens condyles, such that ori®ce appears proximal- that we assign to W. subovoidea were found ly tridentate. No oral spines or peristome. only on rock in clean water well away from Frontal shield densely perforated by large harbors. W. subtorquata and W. subovoidea round pores, each bordered by a thickened, may be two names applied to a single, widely raised rim. Avicularia adventitious, mono- distributed and variable species, or more morphic, sporadic: typically developed on likely, as implied by Soule and Soule (1975) autozooid margins, adjacent or distal to ori- 66 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

®ce; small, Ͻ 0.1 mm long, inconspicuous, GENUS SCHIZOPORELLA HINCKS, 1877 rostrum acute or rounded, orientation vari- (nomen protectum) able. Ovicell recumbent on succeeding au- Schizoporella Hincks, 1877, is apparently tozooid, domed and prominent, slightly wid- a junior subjective synonym of Multiporina er than long; densely perforated by small d'Orbigny, 1852, but the younger name is round pores and appearing coarsely tuber- valid according to section 23.9.1 of the In- culate, closed by autozooid operculum. ternational Code of Zoological Nomenclature Tentacles bright red to purple, 14±20 but (Ride et al., 1999: 28). with smaller range within individual colo- The genus Multiporina was established by nies; lophophores bell-shaped, radially sym- d'Orbigny (1852: 445) for his new species metrical between chimneys, obliquely trun- Multiporina ostracites (d'Orbigny, 1852: cate adjacent to chimneys; adjacent lopho- 445) from Oligocene deposits of Montmar- phores overlapped up to 100 ␮m. tre, Paris, France as type species. d'Orbigny Embryos dark purple to black. Ancestrula gave only a diagnosis with no illustration, and astogeny were described by Friedl and the diagnosis was considered subse- (1925). quently to be insuf®cient for the genus to be REMARKS: Many colonies are infested with recognizable, for example, Bassler (1935: a commensal stoloniferous hydrozoan (Hal- 152, 1953: G235). The name Multiporina has ocoryne epizoica Hadzi, 1917, according to not been used to our knowledge since 1899 Gautier, 1962: 394). The relationship is ap- (except to note that it should be suppressed: parently obligate for the hydrozoan (Gautier, Canu, 1908; Buge, 1975; Gordon, 1989, or 1962) but not for the host species. is unrecognizable: Bassler, 1935, 1953). DISTRIBUTION: This distinctive species is Schizoporella was established by Hincks widespread throughout the Mediterranean, (1877: 527), with Lepralia unicornis John- encrusting organic carbonates in particular, ston in Wood (1844: 19) from the Pliocene and ranges north to the western approaches Crag deposits of East Anglia as type species. to the English Channel, and to southwest Johnston (1847: 320, pl. 57, ®g. 1), often is Cornwall, where it is rare. Harmelin et al. cited as the publication in which Lepralia un- (1989) note the similarity between S. san- icornis was named. Recognition of the 1844 guinea, type species of Schizobrachiella publication is important not only for the date Canu and Bassler, 1920, and species of the and any implications of priority with respect warm temperate to tropical genus Calypto- to other names, but also because the 1847 theca Harmer, 1957. Three species of Calyp- publication describes living, not fossil ma- totheca have been described from Mediter- terial, which potentially is of importance in ranean localities in recent years (Hayward, characterizing the species. 1974; Harmelin et al., 1989), although one, The genus name Schizoporella has been C. triarmata Hayward, is here shown to be used in many more than 25 works (probably a synonym of Schizomavella asymetrica thousands of works) published by at least 10 (Calvet). authors during the past 50 years. Species of MEASUREMENTS (SKELETAL): DO 514 Ϯ 78 Schizoporella are prominent among marine ␮m, 410±715 (2, 20), OL 162 Ϯ 15, 135± fouling faunas (e.g., Geraci and Relini, 1970; 188 (2, 20), OW 182 Ϯ 19, 153±209 (2, 20), Occhipinti Ambrogi, 1981; Brock, 1985), OvL 318 Ϯ 60, 236±392 (2, 6), OvW 364 Ϯ and in many instances they cover a high pro- 30, 327±400 (2, 6), SL 16 Ϯ 3, 10±20 (2, portion of the substratum (Sutherland, 1977, 20), SW 31 Ϯ 4, 25±37 (2, 20), ZL 474 Ϯ 1978). The genus is species rich, found wide- 43, 386±548 (2, 20), ZW 670 Ϯ 46, 598±759 ly in the Atlantic and Paci®c Oceans, ranges (2, 20). (POLYPIDE): IH 108 Ϯ 58 ␮m, 10± from shallow equatorial waters (e.g., Cook, 260 (4, 38), LDMn 674 Ϯ 113, 500±900 (4, 1985) to the highest latitudes (e.g., Kluge, 43), LDMx 705 Ϯ 131, 500±1080 (4, 51), 1975), is diverse and abundant in temperate MD 38 Ϯ 2, 30±40 (4, 20), TLMn499 Ϯ 95, waters (e.g., Maturo, 1957; Gautier, 1962; 380±780 (4, 30), TLMx 596 Ϯ 139, 380± Winston, 1982; Zabala, 1986; Zabala and 1000 (4, 38). Maluquer, 1988; Gordon, 1989; Hayward 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 67 and Ryland, 1999) and has an abundant fossil convex, separated by deep grooves. Primary record (e.g., Ziko, 1985; Spencer and Camp- ori®ce slightly wider than long, with a deep bell, 1987; Poluzzi et al., 1988; Moissette V-shaped sinus occupying about half width and Saint Martin, 1995). Consequently it is of proximal border; condyles small but dis- among the most frequently identi®ed and re- tinctive, with acute upturned corners. No corded bryozoan genera by marine biologists spines. Frontal shield with conspicuous epi- in general, not just by bryozoan specialists. theca, glistening in dried material; densely Schizoporella is noted as an exemplar in perforated by numerous small pores. Adven- studies of benthic ecology (e.g., Buss, 1981; titious avicularia lateral to ori®ce, single or Lindberg and Stanton, 1989; Hurlbut, 1991; paired, situated adjacent to sinus; cystid McKinney, 1992; Herrera and Jackson, 1996; swollen, rostrum acute to frontal plane, dis- Cocito et al., 2000), in comparative studies tolaterally directed, supporting a triangular of feeding and related functions (Winston, mandible. Ovicell hyperstomial, globose; im- 1977, 1978; M. J. McKinney, 1997), in evo- perforate centrally, but developing a reticu- lutionary research (Schopf and Dutton, late secondary calci®cation which appears 1976), and among diverse other topics. porous; small pores present marginally, as- Canu (1908: 86) examined d'Orbigny's sociated with conspicuous basal ¯uting. An- holotype of Multiporina ostracites and cestrula oval, tatiform, about half size of nor- placed it in synonymy with Schizoporella un- mal autozooids, at least ®ve spines around icornis, giving preference to both the trivial distal end of opesia, producing single distal name unicornis and the generic name Schi- bud. zoporella. Buge (1975: 50, pl. 4, ®g. 3) also DISTRIBUTION: Schizoporella unicornis is examined, and illustrated, d'Orbigny's holo- distributed in the northeast Atlantic region type of Multiporina ostracites, agreeing with from western Norway to the western Medi- Canu in considering it to be synonymous terranean, and does not appear to have been with Schizoporella unicornis. Probably os- reported previously from the Adriatic. tracites is not the same species as unicornis, Around the British Isles it is common in lit- inasmuch as cheilostome species seldom toral and shallow sublittoral habitats, espe- have a geological range suf®ciently long to cially among kelp holdfasts and on the un- encompass both the Pliocene and Oligocene. dersides of boulders. Even if Multiporina ostracites and Schizo- MEASUREMENTS (SKELETAL): AL 119 Ϯ 14 porella unicornis are not synonymous, they ␮m, 94±134 (2, 8), AW 90 Ϯ 14, 73±111 (2, are closely related within the same genus. 8), DO, 370 Ϯ 70, 250±445 (2, 20), OL 102 This interpretation is based on similarity of Ϯ 9, 85±119, 2, 21), OW 104 Ϯ 6, 98±111 zooidal characters such as texture of perfo- (2, 20), OvL 259 Ϯ 25, 218±3±19 (2, 14), rations of the frontal wall; placement and OvW 282 Ϯ 16, 264±321 (2, 14), SL 27 Ϯ shape of the ori®ce; and placement, shape, 6, 17±38 (2, 21), SW 49 Ϯ 7, 33±62 (2, 21), size, and orientation of the single adventi- ZL 488 Ϯ 36, 421±550 (2, 20), ZW 327 Ϯ tious avicularium per zooid. We here accept 59, 244±428 (2, 20). that Schizoporella is the valid name, and Multiporina should be suppressed. Schizoporella errata (Waters, 1878) Figure 30F±I Schizoporella unicornis Lepralia errata Waters, 1878: 11. (Johnston in Wood, 1844) Schizoporella errata: Hayward and Ryland, 1999: Figure 30A±E 212. Lepralia unicornis Johnston, in Wood, 1844: 19. DESCRIPTION (AMNH 962; CMRR 2244): Schizoporella unicornis Hincks, 1877: 320. Hay- ward and Ryland, 1999: 220. Colonies purple-brown with orange growing margins, multilaminar predominantly DESCRIPTION (AMNH 961): Colony en- through frontal budding, often substantial. crusting, multiserial, unilaminar to multilam- Autozooids irregularly polygonal and ex- inar, developing extensive sheets. Autozooids tremely variable in both size and shape; ori- rectangular to irregularly polygonal, strongly entation inconstant in multilaminar colonies. 68 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 30. A±E. Schizoporella unicornis (AMNH 961). A. general aspect (0.5 mm). B. Autozooid (0.1 mm). C. Maternal zooid with ovicell (0.1 mm). D. Autozooidal ori®ce (0.05 mm). E. Broken ancestrula (bottom) and ®rst-budded autozooid (0.1 mm). F±I. Schizomavella errata (AMNH 962). F. General aspect of region with frontally budded autozooids (0.5 mm). G. Autozooid (left) and (right) kenozooid (0.1 mm). H. Maternal zooid with ovicell (0.1 mm). I. Autozooidal ori®ce (0.05 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 69

Ori®ce slightly longer than wide; the anter inhabitants is the apparent cause of mainte- semielliptical, widest proximally; condyles nance of the large hollow centers as branches short and rounded, lodged in proximolateral extend and divide distally. corners and rather inconspicuous; sinus a DISTRIBUTION: This is a readily recognized, deep, symmetrical U-shape, occupying about widely distributed, and common fouling spe- half proximal width of ori®ce. No spines. cies. It is common throughout the Mediter- Frontal shield convex, regularly and closely ranean and ¯ourishes on man-made substrata perforated by large, round pores, the inter- as a persistent and sometimes troublesome vening calci®cation thickening as sharp ridg- fouling organism. It is present in similar sit- es. Avicularia sporadic, developed proximo- uations in warm temperate to subtropical wa- laterally to the ori®ce, the rostrum sharply ters worldwide, and is a vigorously invasive tapered, directed distolaterally. Ovicell glob- species. Gordon and Mawatari (1992) record ular, with scattered pores, calci®cation thick- its recent spread in New Zealand ports. ening and pores deepening in later ontogeny. MEASUREMENTS (SKELETAL): AL 181 Ϯ 8 Tentacles light orange, 16±19; lopho- ␮m, 173±195 (1, 10), AW 137 Ϯ 15, 117± phores bell-shaped, radially symmetrical, 164 (1, 10), DO 456 Ϯ 85, 311±627 (2, 20), slightly obliquely truncate bordering chim- OL 124 Ϯ 18, 103±151 (2, 20), OW 128 Ϯ neys. 10, 112±146 (2, 20), SL 29 Ϯ 4, 22±36 (2, Embryos bright orange. 20), SW 64 Ϯ 5, 53±72 (2, 20), ZL 570 Ϯ REMARKS: S. errata is a proli®c fouling or- 48, 495±676 (2, 20), ZW 384 Ϯ 78, 288±518 ganism; it covers almost all the available (2, 20). (POLYPIDE): IH 149 Ϯ 82 ␮m, 40± hard human-constructed substrata in parts of 320 (4, 35), LDMn 578 Ϯ 128, 360±780 (1, the Rovinj harbor (locality 34) and the mus- 33), LDMx 648 Ϯ 150, 360±900 (4, 49), sel-culture station in Lim Channel (locality TLMn 527 Ϯ 81, 400±720 (4, 35), TLMx 33). The structure in Rovinj harbor is ex- 576 Ϯ 131, 400±900 (4, 40). posed to vigorous hydrodynamic conditions, whereas the Lim Channel locality is protect- Schizoporella dunkeri (Reuss, 1848) ed, with very low kinetic energy conditions. Figure 31A±E Colony morphology at the harbor locality is massive, with irregular knobby surfaces and Cellepora dunkeri Reuss, 1848: 90. centimeter-wide, millimeter-high cavities in Schizoporella unicornis form longirostris Hincks, which sediment or other extraneous objects 1886: 266. may be immured; colonies are more densely Schizoporella longirostris: Gautier, 1962: 151. Hayward and Ryland, 1979: 173 (part). Zabala, grown and lack such cavities at the mussel- 1986: 489. Zabala and Maluquer, 1988: 133. culture station. At the mussel-culture station, Schizoporella dunkeri: Hayward and Ryland, two morphologies predominate: relatively 1995: 39, 1999: 210. smooth-surfaced multilaminar encrustations, and proli®c, commonly hollow, erect branch- DESCRIPTION (AMNH 891, 963; CMRR es. These differences in growth habits be- 2245): Colonies orange, multiserial, unilam- tween the two localities parallel the mor- inar to multilaminar, developing extensive, phologies that the species exhibits between spreading sheets. Autozooids large, rectan- high-energy and low-energy localities in the gular to irregularly polygonal; convex, sep- Ligurian Sea along the Italian coast (Cocito arated by distinct, narrow sutures. Primary et al., 2000). However, the hollow centers of ori®ce about as wide as long, proximal bor- colonies found by Cocito et al. were gener- der with a narrow sinus medially, its sides ally very narrow and re¯ect growth around often straight and in parallel, proximal ex- soft-bodied erect organisms such as hy- tremity more or less rounded; condyles mas- droids. This may also be a cause for the ini- sive, faceted, prominent and emphasizing si- tiation of erect growth at the Lim Channel nus. No oral spines; peristome developed as mussel-culture station, but extensive portions a thickened ridge distally and laterally in lat- of the erect branches there have broad, open er ontogeny. Frontal shield ®nely granular, centers that house vagrant organisms such as densely perforated by small round pores, polychaetes and decapods. Movement of the bounded by thickened rims; a low median 70 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 31. A±E. Schizoporella dunkeri. A. General aspect (AMNH 891; 0.5 mm). B. Autozooid with large adventitious avicularium adjacent to ori®ce (AMNH 891; 0.1 mm). C. Untreated specimen with mandibles and polypide rudiment preserved in avicularia (AMNH 963; 0.1 mm). D. Maternal zooid with ovicell (AMNH 891; 0.1 mm). E. Autozooidal ori®ce (AMNH 963; 0.05 mm). F±J. Schizoporella magni®ca. F. General aspect (AMNH 964; 0.5 mm). G. Autozooid (AMNH 964; 0.1 mm). H. Maternal zooid with ovicell (AMNH 964; 0.1 mm). I. Autozooidal ori®ce (AMNH 964; 0.05 mm). J. Ancestrula (lower left) and early autozooids (AMNH 965; 0.1 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 71 suboral umbo developing in later ontogeny. Schizoporella magni®ca Hincks, 1886 Avicularia characteristic: monomorphic, Figure 31F±J proximolateral to ori®ce, single or paired, Schizoporella magni®ca Hincks, 1886: 268. Za- generally distolaterally directed; broadest at bala, 1986: 491. Zabala and Maluquer, 1988: the thickened crossbar, the rostrum abruptly 133. Hayward and Ryland, 1999: 216. narrowed distal to it and tapered to a thin, blunt tip; frequently Ͼ0.2 mm long; mandi- DESCRIPTION (AMNH 964, 965; CMRR ble slender, acuminate. Additional, larger av- 2246): Colonies cream-colored, encrusting, icularia may be budded elsewhere on auto- multiserial, unilaminar to multilaminar. Au- zooid frontal surface when frontal expansion tozooids hexagonal to irregularly polygonal, of colony occurs. Ovicell recumbent on dis- convex, separated by distinct grooves. Pri- tally succeeding autozooid, globular and mary ori®ce longer than wide, sinus a nar- prominent, calci®cation and perforation uni- row, U-shaped slit equivalent to one-third form with that of the frontal shield, often de- length of anter, occupying about half proxi- veloping a frontal umbo. mal width; condyles rounded, faceted and Tentacles light orange, 16±20; lopho- conspicuous. Oral spines generally absent phores bell-shaped, radially symmetrical be- except in earliest colony astogeny, decreas- tween chimneys, obliquely truncate and lo- ing in number distally from three or four in cally scalloped adjacent to chimneys; overlap zooids budded from ancestrula; no peri- between adjacent lophophores up to 200 ␮m. stome. Frontal shield thick, tuberculate, Embryo color dark orange. Ancestrula and densely perforated by small round pores. An- early astogeny ®rst described by Friedl cestrula ®nely perforate, with eight oral (1925). spines around large D-shaped ori®ce, with REMARKS: The type specimen is a Miocene straight proximal margin; ori®ce wider than fossil from the Vienna basin. Hincks (1886) long, maximum width at about midlength. introduced the taxon longirostris as a variety Avicularia paired (rarely, single), situated lat- of the north European species S. unicornis eral to ori®ce, usually distal to the sinus, (Johnston in Wood) and as S. longirostris it some autozooids with an additional distola- was subsequently reported from all parts of terally directed adventitious avicularium the Mediterranean, and northwards to the proximal to ori®ce; cystid rounded, promi- nent, rostrum narrowly triangular, acute to Isles of Scilly. Hayward and Ryland (1995) frontal plane, directed distally or slightly dis- ®nally established the synonymy of S. dunk- tolaterally; crossbar slender, complete. Ovi- eri (Reuss) and S. longirostris (Hincks). cell recumbent on frontal shield of distally DISTRIBUTION: In addition to the Distribu- succeeding autozooid, elongate oval, domed tion cited above (Remarks), Schizoporella and prominent; texture and perforation sim- dunkeri is abundant throughout the Adriatic, ilar to that of frontal shield, but a median in shallow detritic environments. apical area imperforate, and often developed MEASUREMENTS (SKELETAL): AL 301 Ϯ 45 as a longitudinal ridge or umbo. ␮m, 204±395 (3, 30), AW 138 Ϯ 26, 97±182 DISTRIBUTION: This species is widespread (3, 30), DO 609 Ϯ 97, 390±790 (3, 30), OL and common throughout the Mediterranean 160 Ϯ 11, 133±176 (3, 29), OW 136 Ϯ 13, and reaches its northern limit off the south- 110±162 (3, 29), OvL 301 Ϯ 19, 282±319 west British Isles. (1, 3), OvW 358 Ϯ 23, 332±372 (1, 3), SL MEASUREMENTS (SKELETAL): AL 161 Ϯ 19 42 Ϯ 8, 31±53 (2, 9), SW 26 Ϯ 8, 19±39 (2, ␮m, 131±194 (2, 20), AW 109 Ϯ 15, 90±144 9), ZL 739 Ϯ 69, 599±827 (3, 28), ZW 644 (2, 20), DO 436 Ϯ 52, 325±575 (2, 20), OL Ϯ 94, 489±813 (3, 28). (POLYPIDE): IH 234 126 Ϯ 9, 108±145 (2, 20), OW 97 Ϯ 9, 74± Ϯ 59 ␮m, 140±350 (3, 20), LDMn 808 Ϯ 111 (2, 20), OvL 380 Ϯ 28, 334±446 (2, 20), 100, 620±1000 (4, 39), LDMx 876 Ϯ 116, OvW 304 Ϯ 27, 243±334 (2, 20), SL 35 Ϯ 620±1200 (4, 43), MD 37 Ϯ 4, 30±45 (3, 6, 25±45 (2, 20), SW 24 Ϯ 4, 15±32 (2, 20), 32), TLMn 718 Ϯ 63, 600±850 (3, 12), ZL 540 Ϯ 36, 494±641 (2, 20), ZW 389 Ϯ TLMx884 Ϯ 187, 680±1280 (4, 19). 31, 330±442 (2, 20). 72 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Schizoporella cf. S. tetragona (Reuss, 1848) GENUS ESCHARINA MILNE EDWARDS, 1836 Figure 32A±D Escharina vulgaris (Moll, 1803) Schizoporella tetragona: Ryland, 1968: 543. Figure 32E±I Eschara vulgaris var. a Moll, 1803: 55. DESCRIPTION (AMNH 966; CMRR 2247): Lepralia botterii Heller, 1867: 106. Colony a broad, spreading, multiserial, uni- Escharina vulgaris: Hayward and Ryland, 1999: laminar sheet in present material. Autozooids 236. hexagonal to irregularly polygonal, large, convex, separated by deep grooves. Primary DESCRIPTION (AMNH 947, 967; CMRR ori®ce as wide as long; poster equivalent to 2248): Colonies irregular, multiserial, uni- almost half total length, forming a broadly laminar patches. Autozooids oval to hexag- U-shaped sinus, occupying the whole proxi- onal, or irregularly polygonal, gently convex mal width of the anter below indistinct con- in early ontogeny, becoming rather ¯at as dyles. Two delicate, widely spaced, distal calci®cation thickens in later ontogeny, sep- oral spines present in earliest ontogeny, lost arated by shallow grooves. Primary ori®ce as a low peristomial rim develops around dis- longer than wide, broadest at midlength; dis- tal and lateral borders of ori®ce. Frontal tal rim raised, thin; proximal border straight, shield thick, vitreous, densely perforated by with a short, rounded sinus medially, equiv- small round pores; thickening through onto- alent to one-quarter proximal width; con- geny and becoming quite rugose as the rims dyles short and broad, tapering towards si- of the pores are more pronounced; a promi- nus, distinctly ridged. Five or six delicate oral spines present in periancestrular zooids, nent, stout, conical umbo developing imme- typically absent in later autozooids; peri- diately proximal to sinus, forming a large stome erect and thin, developed indepen- spike in some autozooids. Avicularia single dently of the ori®ce rim, which remains dis- or paired, the rounded, prominent cystid sit- tinct, extending distal to the spine bases, and uated lateral or just distal to sinus; rostrum proximally enclosing a smooth, cupped area narrowly triangular, acute to frontal plane below the sinus. Frontal shield ®nely granu- and directed distolaterally. Ovicell hypersto- lar, imperforate except for a few large, irreg- mial, prominent, about as wide as long, with ular marginal pores. Avicularia adventitious, highly arched aperture; imperforate centrally single or paired, developed along lateral mar- but with numerous pores laterally and around gins of autozooids; rostrum slender, acumi- periphery; developing a low frontal umbo. nate, with a slight lateral curve, directed dis- REMARKS: This appears to be the species tally or distolaterally. Ovicell (not present in described by Ryland (1968) from the Bos- material described) spherical, ®nely granular, porus and attributed to. Schizoporella tetra- imperforate. Ancestrula about half the length gona (Reuss, 1848). The shape of the pri- of normal autozooids, oval, tatiform, with a mary ori®ce, the disposition of the avicularia narrow, smooth gymnocyst and indistinct and the thickly calci®ed umbonate frontal cryptocystal rim bounded by about nine shield, noted by Ryland, are evident in this spines. material. However, the equivalence with REMARKS: We can ®nd no differences be- Reuss' species needs reexamination, follow- tween Escharina vulgaris and the type speci- ing the advent of SEM in bryozoan morpho- men (UIIZ 217) of Lepralia botterii Heller, logical research. 1867, from the northern Adriatic, illustrated MEASUREMENTS (SKELETAL): AL 133 Ϯ 10 here as ®g. 32G. ␮m, 112±145 (2, 12), AW 90 Ϯ 8, 75±106 DISTRIBUTION: On stones and shells, in (2, 12), DO 428 Ϯ 56, 310±507 (2, 20), OL coastal and shelf environments. Widespread 108 Ϯ 8, 86±120 (2, 20), OW 114 Ϯ 8, 95± and common throughout the Mediterranean, 132 (2, 20), OvL 285 Ϯ 24, 245±314 (2, 16), ranging south to Madeira and north along the OvW 294 Ϯ 20, 258±325 (2, 16), SL 18 Ϯ western coasts of Britain and Ireland. Un- 3, 12±22 (2, 20), SW 53 Ϯ 4, 46±61 (2, 20), known in the North Sea. ZL 558 Ϯ 49, 473±671 (2, 20), ZW 387 Ϯ MEASUREMENTS (SKELETAL): AL 106 Ϯ 15 40, 315±451 (2, 20). ␮m, 84±140 (2, 20), AW 67 Ϯ 8, 54±83 (2, 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 73

Fig. 32. A±D. Schizoporella cf. S. tetragona (AMNH 966). A. General aspect (0.5 mm). B. Auto- zooid (0.1 mm). C. Maternal zooid with ovicell (0.1 mm). D. Autozooidal ori®ce (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 setae-like mandibles of avicularia (type specimen of Lepralia botterii, UIIZ 217; 0.1 mm). H. Oblique view of autozooidal ori®ce (AMNH 967; 0.05 mm). I. Ancestrula (lower right) and early autozooids (AMNH 947; 0.1 mm). 74 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

20), DO 431 Ϯ 49, 349±513 (2, 20), OL 103 ␮m, 422±611 (2, 20), OL 86 Ϯ 13, 64±106 Ϯ 14, 83±126 (2, 20), OW 88 Ϯ 6, 75±104 (2, 20), OW 86 Ϯ 10, 69±106 (2, 20), OvL (2, 20), SL 18 Ϯ 6, 10±18 (2, 20), SW 17 Ϯ 170 Ϯ 20, 150±190 (1, 3), OvW 276 Ϯ 17, 5, 12±30 (2, 20), ZL 544 Ϯ 37, 471±616 (2, 263±301 (1, 4), SL 20 Ϯ 3, 15±24 (2, 20), 20), ZW416 Ϯ 40, 353±517 (2, 20). SW 13 Ϯ 4, 8±23 (2, 20), ZL 565 Ϯ 46, 503± 659 (2, 20), ZW 408 Ϯ 34, 351±500 (2, 20). Escharina dutertrei protecta Zabala, Maluquer and Harmelin, 1993 FAMILY MARGARETTIDAE HARMER, 1957 Figure 33A±C GENUS MARGARETTA GRAY, 1843 Escharina dutertrei protecta Zabala, Maluquer Margaretta cereoides and Harmelin, 1993: 73. (Ellis and Solander, 1786) Figure 33D±G DESCRIPTION (AMNH 968): Colonies en- crusting, multiserial, unilaminar sheets. Au- Cellaria cereoides Ellis and Solander, 1786: 26. tozooids broadly hexagonal, convex, separat- Margaretta cereoides: Gautier, 1962: 216. Zabala ed by distinct grooves. Primary ori®ce wider and Maluquer, 1988: 155. than long, poster a short, rounded sinus oc- DESCRIPTION (AMNH 969; CMRR 2249): cupying just one-eighth of the proximal Colonies pink, erect, branching, jointed, to 8 width of the anter; condyles short and broad, cm high in present material; consisting of with denticulate edges. Six delicate spines rigid, curving, round-sectioned internodes, around distal border of anter in early onto- each rising from a ¯exible, tubular, chitinous geny, some or all of which become incor- node (basis rami) with pronounced annula- porated in development of conspicuous peri- tions. Each basis rami supporting single au- stome, extending proximally from proximal- tozooid, which produces two zooid buds; most spine pair, ¯aring outwards and fron- thereafter, autozooids disposed in alternating tally, enclosing and shielding the sinus. Close whorls of four. Autozooids elongate, strongly to proximal spine pair the peristome rim pro- convex, separated by distinct grooves. Pri- duced as stout, projecting processes, in many mary ori®ce wider than long, D-shaped, vis- autozooids extending distally as a lobed rim, ible at growing edge but in early ontogeny obscuring spine bases. Frontal shield nodular, obscured by development of tall, cylindrical, vitreous, with indistinct marginal pores. Av- longitudinally ¯uted peristome. Frontal icularia adventitious, paired, one each side of shield thick, vitreous, densely perforated by ori®ce, level with its distal half; cystid Ͻ0.05 large pores; distinctly larger ascopore just mm long, with trough-like rostrum support- proximal to peristome base. In dried materi- ing base of a setiform mandible, 0.375 mm al, outer cuticle conspicuous. Reproduction long, directed distomedially over frontal and brooding not documented, except that re- shield of distal autozooid. Ovicell wider than productively active autozooids recognized by long, recumbent, with medially peaked ap- dimorphic peristomes, consisting of broad- erture; imperforate, granular. based cone with tapered, distally curved api- REMARKS: This taxon was only recently cal aperture; chamber thus formed perhaps de®ned by Zabala et al. (1993), who distin- an external brood chamber. Branching ad- guished it from the northeast Atlantic E. du- ventitious, not dichotomous. Each basis rami tertrei haywardi. All previous Mediterranean arising from frontal surface of modi®ed au- records of E. dutertrei probably refer to this tozooid in which peristome sealed before subspecies, which also occurs around the complete development; basis rami budding at Azores, and it is probable that if Escharina site of ascopore. dutertrei (Audouin) could be recognized, and Tentacles light orange-pink, 25±28; loph- a neotype accepted, the two subspecies intro- ophores bell-shaped, radially symmetrical, duced by Zabala et al. (1993), which appear supported on long introverts. to have distinct geographical Distribution DISTRIBUTION: Develops stiff clumps at- patterns, could be accepted as full species. tached to a range of hard substrata, in shal- DISTRIBUTION: See Remarks. low coastal waters, probably endemic to the MEASUREMENTS (SKELETAL): DO 493 Ϯ 40 Mediterranean. 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 75

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 ori®ce (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 ori®ce (0.1 mm). G. Peristome-bearing zooid (0.2 mm). 76 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

MEASUREMENTS (SKELETAL): DO 903 Ϯ 98 Ϯ 15, 315±367 (2, 11), OvL 769 Ϯ 55, 688± ␮m, 669±1097 (2, 23), OW 170 Ϯ 23, 163± 891 (2, 10), OvW 820 Ϯ 79, 701±915 (2, 231 (2, 22), OOW 219 Ϯ 13, 199±245 (2, 11). (POLYPIDE): IH 460 Ϯ 24 ␮m, 440±500 20), OvL 608 Ϯ 42, 533±719 (2, 20), ZL (1, 5), LD 1107 Ϯ 83, 1040±1200 (1, 3), TL 1456 Ϯ 170, 1177±1759 (2, 20), ZW 581 Ϯ 1030 Ϯ 37, 960±1060 (1, 6). 54, 458±660 (2, 20). (POLYPIDE): IH 362 Ϯ 104 ␮m, 260±580 (1, 10), LD 886 Ϯ 60, FAMILY LANCEOPORIDAE HARMER, 1957 740±960 (1, 11), MD 37.3 Ϯ 2.5, 35±40 (1, GENUS CALYPTOTHECA HARMER, 1957 3), TL 793 Ϯ 63, 680±872 (1, 10). Calyptotheca rugosa Hayward, 1974 FAMILY MYRIAPORIDAE GRAY, 1841 Figure 34E±G GENUS MYRIAPORA DE BLAINVILLE, 1830 Calyptotheca rugosa Hayward, 1974: 379.

Myriapora truncata (Pallas, 1766) DESCRIPTION (AMNH 971): A single, Figure 34A±D small, multiserial, unilaminar colony of just Millepora truncata Pallas, 1766: 249. 28 autozooids. Autozooids polygonal, con- Myriapora truncata: Gautier, 1962: 268. Zabala vex, separated by distinct grooves. Primary and Maluquer, 1988: 163. ori®ce as wide as long; poster accounting for about one-quarter of total length and broadly DESCRIPTION (AMNH 970; CMRR 2250): triangular, occupying entire proximal width Colonies red, erect, rigid, attached by en- of anter. In absence of the operculum, very crusting base, branching irregularly; branch- large, quadrangular condyles, orientated es thick, cylindrical, of constant width, to 4 oblique to the disto-proximal axis, and a con- mm diameter, except thicker at base of col- spicuous inner shelf around the distal part of onies. Living specimens deep red. Auto- the anter, imparting distinctive outline to the zooids in alternating whorls of seven to ori®ce. Spines absent except in earliest- eight; individual boundaries indistinct, even formed autozooids, diminishing in number at growing tips. Colony surface appearing away from ancestrula; low, thick peristomial uniformly ®nely granular, regularly perforat- ridge encircling ori®ce. Frontal shield thick, ed by small, round pores, with primary ori- nodular and vitreous, densely perforated by ®ces of autozooids regularly spaced in strict small round pores. Avicularium adventitious, quincuncial order. Primary ori®ce slightly suboral, rare; ovicell large, sutured, perforate longer than wide; anter more or less semi- (neither found in present material). Ancestru- circular, comprising two-thirds ori®ce length; la identical to later autozooids; producing poster a short semiellipse below prominent, paired distolateral buds. bluntly triangular condyles. No spines or DISTRIBUTION: This species has not been peristome. Ovicell semiimmersed, its frontal reported since its original description, based surface scarcely projecting from colony fron- on specimens from Chios, Aegean Sea. tal plane; more ®nely punctate than frontal MEASUREMENTS (SKELETAL): DO 502 Ϯ 75 shields of adjoining autozooids; ori®ce of ␮m, 408±650 (1, 10), OL 140 Ϯ 10, 127± brooding zooids dimorphic, nearly twice 159 (1, 10), OW 143 Ϯ 7, 135±154 (1, 10), length and width of autozooid, with a pro- ZL 654 Ϯ 61, 550±742 (1, 10), ZW 442 Ϯ portionately broader poster. 82, 362±635 (1, 10). Tentacles red, 27±29; lophophores parab- oloid, supported on long introverts. FAMILY CHEILOPORINIDAE BASSLER, 1936 DISTRIBUTION: Widely distributed through- GENUS HAGIOSYNODOS BISHOP AND out the Mediterranean, on shell and hard sub- HAYWARD, 1989 strata, in caves, on coralligenous deposits, from 30 m to at least 130 m depth, probably Hagiosynodos kirchenpaueri (Heller, 1867) endemic to Mediterranean. Figure 35E±H MEASUREMENTS (SKELETAL): DO 627 Ϯ 59 Lepralia kirchenpaueri Heller, 1867: 105. ␮m, 519±723 (3, 20), OL 265 Ϯ 11, 250± Hippopodinella kirchenpaueri: Gautier, 1962: 283 (3, 25), OW 228 Ϯ 11, 203±259 (3, 25), 179. Zabala and Maluquer, 1988: 136. OOL 370 Ϯ 25, 306±388 (2, 11), OOW 337 Hippopodinella lata (Busk): Schmid, 1989: 47. 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 77

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 ori®ce (0.1 mm). D. Maternal zooid ori®ce (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). 78 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

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 ori®ce (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. Auto- zooidal ori®ce (AMNH 966; 0.05 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 79

NEOTYPE (chosen here): UIIZ 244, encrust rock. Hagiosynodos latus is a facul- bleached specimen on small gastropod. tative symbiont of hermit crabs inhabiting DESCRIPTION (AMNH 966): Colonies en- gastropod shells (review in Taylor, 1994); the crusting, multiserial, unilaminar with local prevalence of H. kirchenpaueri on gastropod areas of self-overgrowth associated with re- shells suggests that it, too, may be a facul- juvenation. Autozooids oval to irregularly tative symbiont of hermit crabs. However, we polygonal, separated by indistinct grooves. do not know if the gastropods were alive or Primary ori®ce longer than wide, distinctive: if the shells were inhabited by hermit crabs anter widest at midlength, its rim constituting when encrusted by H. kirchenpaueri. three-quarters of a complete ellipse; condyles DISTRIBUTION: The species is known only short and bluntly pointed, directed medi- from the Mediterranean. oproximally; poster with almost straight MEASUREMENTS (SKELETAL): proximal border, and width three times its length; usually conspicuously wider than an- ter; rugose umbones locally present at prox- imolateral margins of ori®ce. No spines. Frontal shield gently convex, with a rugose, nodular surface, evenly punctured by pores. Ovicell recumbent, only slightly convex, cal- ci®cation 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 Hell- er's specimen illustrated here as ®g. 35A, from syntype suite UIIZ 244. Through a comparison of morphology and biometrics, Hagiosynodos hadros, new species Schmid (1989) synonymised the type of Le- Figure 36A±D pralia kirchenpaueri Heller, plus Miocene material from Austria, with Recent speci- DIAGNOSIS: Encrusting colonies with rela- mens of Hippopodinella lata from England. tively large autozooids, ori®ces, and spacing Although Schmid found similar ranges of of pores in frontal shield; poster of ori®ce measurements for autozooidal length and noticeably wider than anter. width, ori®ce length and width, and ovicell HOLOTYPE: AMNH 972. length and width, we prefer to retain kir- PARATYPES: AMNH 973; CMRR 2251± chenpaueri and lata as separate species. 2254. Measurements for the two are generally sim- ETYMOLOGY: The species is named from ilar, but we distinguish Hagiosynodos kir- the Latin hadros (well developed, bulky) be- chenpaueri from H. latus (Busk, 1856) by cause of the large dimensions or its zooids kirchenpaueri having on average larger di- relative to those of Hagiosynodos kirchen- mensions (see ®g. 35A±D and measurements paueri and H. latus. below for H. kirchenpaueri and for H. latus DESCRIPTION: Colonies encrusting, multi- from the British Isles, BMNH 2001.1.26.18± serial, unilaminar with local areas of self- 2001.1.26.20), a proportionately broader overgrowth associated with rejuvenation. poster, and the presence in H. latusofa Autozooids oval to irregularly polygonal, prominent distal lip in the primary ori®ce on separated by indistinct grooves. Primary or- which the operculum rests when closed and i®ce longer than wide, distinctive: anter wid- its absence in H. kirchenpaueri. est at midlength, its rim constituting three- Seven of the eight specimens in the type quarters of a complete ellipse; condyles short suite of H. kirchenpaueri (UIIZ 244) occur and bluntly pointed, directed medioproxi- on gastropod shells; the eighth is on an echi- mally; poster with almost straight proximal noid spine. The specimens from near Rovinj border, and width three times its length. No 80 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 36. A±D. Hagiosynodos hadros. A. general aspect (holotype, AMNH 972; 0.5 mm). B. Auto- zooid (holotype, AMNH 972; 0.1 mm). C. Naternal zooid with ovicell (holotype, AMNH 972; 0.1 mm). D. Autozooidal ori®ce (paratype, AMNH 973; 0.05 mm). E, F. Cryptosula pallasiana (AMNH 974). E. General aspect (0.5 mm). F. Autozooid (0.1 mm). spines. Frontal shield gently convex, with a H. hadros. It was found on gastropod and rugose, nodular surface, evenly punctured by dead bivalve skeletons. There is no indica- relatively large pores. Ovicell recumbent, tion that the gastropods were alive or the only slightly convex, calci®cation and per- shells inhabited by hermit crabs when en- foration identical to frontal shield; closed by crusted by H. hadros. zooidal operculum. Small dietellae clearly MEASUREMENTS (SKELETAL): DO 425 Ϯ 76 visible at growing edges. ␮m, 250±563 (5, 50), OL 135 Ϯ 14, 106± Tentacles light carmine, 12; lophophores 170 (5, 50), OW (anter) 88 Ϯ 10, 66±114 (5, campylonemidan. 50), OW (poster) 102 Ϯ 11, 81±129 (5, 50), REMARKS: This species has signi®cantly OvL 246 Ϯ 26, 197±307 (5, 34), OvW 288 larger autozooids, ori®ces, and spacing of Ϯ 22, 246±338 (5, 34), PS 48 Ϯ 7, 32±73 pores in the frontal shield than do H. kir- (5, 50), ZL 559 Ϯ 56, 471±745 (5, 50), ZW chenpaueri and H. latus. Also, rugose um- 351 Ϯ 48, 240±472 (5, 50). (POLYPIDE): IH bones at the proximolateral margins of the 114 Ϯ 71 ␮m, 40±240 (1, 12), LD 510 Ϯ ori®ce, common in H. kirchenpaueri and H. 58, 440±600 (1, 10), MD 28.8 Ϯ 2.5, 25±30 latus, are absent in the available material of (1, 4), TL 520 Ϯ 58, 440±640 (1, 10). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 81

FAMILY CRYPTOSULIDAE VIGNEAUX, 1949 ered that populations lacking avicularia were GENUS CRYPTOSULA CANU AND BASSLER, 1925 characteristic of environmentally unstable habitats. Cryptosula pallasiana (Moll, 1803) MEASUREMENTS (SKELETAL): AL 211 Ϯ 14 Figure 36E, F ␮m, 191±240 (2, 20), AW 172 Ϯ 10, 153± 193 (2, 20), DO 501 Ϯ 105, 338±684 (2, 20), Eschara pallasiana Moll, 1803: 64. ZL 670 Ϯ 62, 591±838 (2, 20), ZW 449 Ϯ Cryptosula pallasiana Canu and Bassler, 1925: 33. Hayward and Ryland, 1999: 194. 66, 372±581 (2, 20). (POLYPIDE): IH 147 Ϯ 35 ␮m, 100±200 (1, 6), LD 673 Ϯ 50, 600± DESCRIPTION (AMNH 974; CMRR 2255): 740 (1, 6), MD 40 (1, 1), TL 650 Ϯ 58, 560± Colonies translucent white, encrusting, mul- 700 (1, 6). tiserial, unilaminar, developing thick, spread- ing, orange sheets. Autozooids rectangular to FAMILY MICROPORELLIDAE HINCKS, 1879 hexagonal, convex, separated by distinct su- GENUS FENESTRULINA JULLIEN, 1888 tures. Primary ori®ce longer than wide, post- Fenestrulina malusii (Audouin, 1826) er shallowly concave below prominent, downcurved condyles, broader than anter Figure 37A±D and imparting a bell-shaped outline to the or- Cellepora malusii Audouin, 1826: 239. Savigny, i®ce. Operculum deep brown. Peristome de- 1809: pl. 8, ®g. 8. veloped as a low thickened rim, most prom- Fenestrulina malusii: Nielsen, 1981: 109. Hay- inent and slightly ¯ared distally. Frontal ward and Ryland, 1999: 299. shield thickly calci®ed, the epitheca glisten- DESCRIPTION (AMNH 975±977; CMRR ing conspicuously in dried material; densely 2256): Colonies silvery white, encrusting, perforated by large round pores, with prom- multiserial, unilaminar patches. Autozooids inent nodular ridges developing between in oval to hexagonal, convex, separated by later ontogeny. A stout median suboral umbo well-marked ridges. Primary ori®ce wider present in most autozooids; in some popu- than long, proximal edge straight, four short lations this position is occupied by an avi- distal oral spines present. Frontal shield cularium with broadly oval rostrum, acute to smooth, with a prominent median ascopore, frontal plane and proximally directed, but it distance between it and proximal ori®ce rim was not present in any of the Rovinj speci- equivalent to ori®ce length. Ascopore rim mens. No ovicells, pale yellow embryos transversely oval to reniform, thickened and brooded internally. raised, lumen crescentic, partly occluded by Tentacles clear, 17; lophophores bell- delicate denticles. Large round pores with shaped, radially symmetrical, overlapped thickened rims, also partly occluded and ap- with adjacent lophophores up to 80 ␮m. pearing cribriform, in single or double series OCCURRENCE: Found only in the harbor around autozooid margins, and extending and adjacent shore of Rt Kriz, 0±5 m depth. medially between ascopore and ori®ce. An- DISTRIBUTION: Cryptosula pallasiana is cestrula tatiform, about two-thirds length of widely distributed in the North Atlantic, from normal autozooids, with oval opesia occu- northern Norway throughout the Mediterra- pying over half total length, bordered by nean and into the Black Sea, and from Nova very narrow cryptocyst and about 10 spines. Scotia to North Carolina. Around the British Ovicell recumbent on distal autozooid, wider Isles it is a common littoral and shallow sub- than long, prominent; ectooecium membra- littoral species, as a characteristic component nous except for a thickened peripheral rim; of the sessile fauna of kelp holdfasts and un- endooecium smoothly calci®ed, imperforate, dersides of boulders. It is also a successful with distinct basal ¯uting. fouling species; it has been part of the foul- Tentacles colorless, 13±15; lophophores ing community of New Zealand since at least bell-shaped, radially symmetrical. the1890s (Gordon and Mawatari, 1992), and Embryo color lemon yellow to medium its presence in docks and harbors elsewhere orange. around the world is probably attributable to DISTRIBUTION: Shallow sublittoral, and off- ship borne dispersal. Schopf (1973) consid- shore to about 100 m, on a wide range of 82 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

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 auto- zooids (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 ori®ce (AMNH 980; 0.05 mm). J. Ancestrula (center) and early autozooids (AMNH 897; 0.1 mm) 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 83 organic and inorganic substrata. Distributed Tentacles clear, 12±13; lophophores bell- throughout the Mediterranean, and north- shaped, radially symmetrical. wards to the British Isles and western Nor- Embryo color light to bright orange. way. DISTRIBUTION: Microporella ciliata is com- MEASUREMENTS (SKELETAL): DO 484 Ϯ 54 mon in shallow coastal waters, and is distrib- ␮m, 383±613 (2, 20), OL 104 Ϯ 12, 79±134 uted in the northeast Atlantic from northern (2, 20), OW 142 Ϯ 15, 120±181 (2, 20), ZL Norway to the Mediterranean, where it is 625 Ϯ 89, 490±851 (2, 20), ZW 401 Ϯ 45, widespread. Records from outside this geo- 325±511 (2, 20). (POLYPIDE): IH 167 Ϯ 33 graphical range require reexamination. ␮m, 140±240 (1, 9), LD 765 Ϯ 106, 640± MEASUREMENTS (SKELETAL): AL 126 Ϯ 15 900 (1, 4), MD 35±40 (1, 2), TL 660 Ϯ 49, ␮m, 101±148 (2, 15), AW 88 Ϯ 13, 72±117 600±700 (1, 4). (2, 15), DO 382 Ϯ 44, 311±454 (2, 20), OL 57 Ϯ 6, 42±67 (2, 20), OW 85 Ϯ 12, 73± GENUS MICROPORELLA HINCKS, 1877 110 (2, 20), OvL 239 Ϯ 8, 221±250 (1, 10), OvW 228 Ϯ 9, 218±245 (1, 10), ZL 486 Ϯ Microporella ciliata (Pallas, 1766) 35, 434±546 (2, 20), ZW 318 Ϯ 38, 273± Figure 37E±J 404 (2, 20). (POLYPIDE): IH 130 Ϯ 12 ␮m, Eschara ciliata Pallas, 1766: 38. 120±140 (1, 4), LD 402 Ϯ 42. 360±500 (1, Microporella ciliata: Gautier, 1962: 172. Hay- 9), TL 394 Ϯ 38, 340±440 (1, 7). ward and Ryland, 1999: 296. SUPERFAMILY CELLEPOROIDEA DESCRIPTION (AMNH 897, 913, 978±980; JOHNSTON, 1838 CMRR 2257): Colonies white though com- FAMILY CELLEPORIDAE monly colored by fouling algae, encrusting, JOHNSTON, 1838 multiserial, unilaminar sheets, typically sil- GENUS CELLEPORA LINNAEUS, 1767 very when dried. Autozooids oval to hexag- onal, convex, separated by distinct grooves; Cellepora adriatica, new species 0.375±0.5 ϫ 0.3 mm in present material. Pri- Figure 38A±D mary ori®ce slightly wider than long, proxi- DIAGNOSIS: Primary ori®ce nearly circular, mal border straight, with a minute condyle in lacking condyles; peristome asymmetrical, each proximal corner; six slender spines suboral avicularium single, vertical; 1 to 3 equally spaced around distal and lateral rim. pores on ovicell, close to apertural rim. Frontal shield nodular, with few small, round HOLOTYPE: AMNH 982. pores, and fewer, more conspicuous marginal PARATYPE: AMNH 983. pores. Ascopore spaced from ori®ce by a dis- ETYMOLOGY: The species is named from its tance equivalent to half ori®ce length; rim discovery in the Adriatic Sea. thickened, especially prominent in later on- DESCRIPTION: Colonies small, encrusting, togeny, lumen crescentic, with ®nely dentic- unilaminar to multilaminar. Autozooids re- ulate border. A single adventitious avicular- cumbent at unilaminar growing edge, erect ium situated proximolateral to ascopore, on elsewhere; broadly oval, convex, separated right or left; rostrum acute to frontal plane, by distinct ridges. Frontal shield smoothly directed distolaterally, broadly triangular calci®ed, imperforate except for a few widely with a distal groove to accommodate a seti- spaced marginal areolar pores. Primary ori- form mandible, extending beyond tip of ros- ®ce as wide as long, orbicular but not per- trum by twice its length; crossbar slender, fectly circular: the rim of the anter describing complete, no columella or palate. Ovicell a ¯atter arc than the rim of the poster; no domed, hemispherical, aperture overarching condyles visible. A single latero-proximal primary ori®ce; surface uniformly nodular, suboral avicularium present, with an in¯ated imperforate. Vertical walls of autozooids cystid tapered and umbonate distally; ros- with distinct basal pore chambers. Ancestrula trum elliptical, perpendicular to plane of or- tatiform, with oval opesia occupying about i®ce, facing laterally; crossbar slender, com- half total length, bordered by narrow cryp- plete, no palate. Peristome well developed, tocyst and about nine spines. entirely enclosing the ori®ce and incorporat- 84 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

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 ori®ce (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 ori®ces (AMNH 984; 0.05 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 85

ing the avicularium, deepening in multilam- DESCRIPTION (AMNH 984, 985; CMRR inar parts of colony, where frontally budded 2258): Colonies white, erect, branching, autozooids are raised above peristomes of slender, to 4 mm in present material. Branch- preceding autozooids; peristome rim charac- es cylindrical, consisting of alternating teristically asymmetrical, extending in a lobe whorls of three or four autozooids, with al- around one side of the avicularium cystid. most constant diameter. Autozooids at grow- Ovicell prominent, raised above distal ori®ce ing tips elongate oval, convex, small: less rim; hemispherical, with a wide, highly than 0.4 mm long, including peristome. Pri- arched aperture, close to the distal edge of mary ori®ce terminal, orbicular, with minute, which is a single large, round pore. The rim medioproximal sinus, obscured by develop- of the peristome extends across the frontal ment of peristome in early ontogeny. No surface of the ovicell, encircling the pore. No spines. Peristome with smooth, thickened vicarious avicularia. Vertical walls with rim, slightly ¯ared, with a symmetrical, small basal pore chambers. rounded, midproximal notch adjacent to REMARKS: Heller (1867) recorded Celle- which, on right or left, is a small adventitious pora pumicosa from the Adriatic, together avicularium, with distally hooked, triangular with two new species, C. hincksii and C. cor- rostrum, acute to frontal plane and directed ticalis. It is possible that one of these may distolaterally. Frontal shield smooth, porcel- be the taxon described here as new, but Hell- lanous, with very few, small, irregular mar- er did not ®gure any of these three species. ginal pores. Ovicell wider than long, with Hayward (1974) described a new genus and straight edged aperture, perforated by Ͻ10, species, Rhamphostomellina posidoniae, en- large, round pores; becoming immersed in crusting Posidonia leaves, from Chios, Ae- smooth calci®cation, only perforate frontal gean Sea. It is very similar to C. adriatica in area remaining distinct. Frontal calci®cation its asymmetrical peristome, vertically orien- thickens through ontogeny and the colony tated avicularium, orbicular ori®ce, and in its develops a uniformly smooth surface; auto- ovicell, which has 1 to 3 pores close to its zooid boundaries are quite obscured and only aperture rim. It is clear that R. posidoniae rims of immersed peristomes, and circular should be assigned to Cellepora; it differs perforate area marking ovicell, remain dis- from C. adriatica in its broader ovicell and tinct. REMARKS: Zabala (1986) argued that Bus- smaller autozooids, and in the possession of kea nitida is clearly identical to the northeast a proximal process on the avicularium cystid, Atlantic B. quincuncialis (Norman, 1867) but which de®nes a pseudospiramen within the both taxa were introduced in the same year peristome. and taxonomic priority has not been decided. OCCURRENCE: Encrusting shells of dead DISTRIBUTION: Buskea nitida was described gastropods. from the Adriatic and has been recorded sub- MEASUREMENTS (SKELETAL): AL 128 Ϯ 8 sequently from numerous northern and west- ␮m, 116±137 (1, 5), AW 87 Ϯ 4, 83±94 (1, ern Mediterranean localities, and from the 5), DO 468 Ϯ 87, 337±630 (1, 10), OL 148 Atlantic coasts of southern Europe; as B. Ϯ 8, 134±158 (1, 10), OW 140 Ϯ 11, 123± quincuncialis, it is also known from the 156 (1, 10), OvL 147 Ϯ 21, 124±192 (1, 10), Minch, West Scotland. OvW 222 Ϯ 13, 198±238 (1, 10). MEASUREMENTS (SKELETAL): DO 400 Ϯ 80 ␮m, 286±530 (2, 13), OW 113 Ϯ 9, 00±126 GENUS BUSKEA HELLER, 1867 (2, 11), OvL 146 Ϯ 27, 119±211 (2, 12), Buskea nitida Heller, 1867 OvW 186 Ϯ 18, 161±220 (2, 20), ZL 529 Ϯ Figure 38E±G 40, 481±640 (3, 21), ZW 212 Ϯ 19, 169± Buskea nitida Heller, 1867: 89. 238 (3, 15). Eschara quincuncialis Norman, 1867: 204; 1868: GENUS CELLEPORINA GRAY, 1848 222. Buskea quincuncialis: Hayward and Ryland, Celleporina caminata (Waters, 1879) 1979: 296; 1999: 350, ®gs. 162C, D, 164. Figure 39A±D Buskea nitida: Zabala, 1986: 553. Zabala and Ma- Cellepora retusa var. caminata Waters, 1879: 194. luquer, 1988: 156. Celleporina caminata: Gautier, 1962: 244. Har- 86 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

melin, 1969b: 303. Zabala,1986: 563. Zabala It is probably endemic to the Mediterranean and Maluquer, 1988: 158. Ristedt, 1996: 239. region. MEASUREMENTS (SKELETAL): AAL 94 Ϯ 10 DESCRIPTION (AMNH 986, 987; CMRR ␮m, 77±123 (2, 20), AAW 65 Ϯ 6, 52±77 2259): Colonies white to cream-colored, de- (2, 20), DO 497 Ϯ 87, 303±674 (2, 20), OL veloping stout, frontally budded nodular or 195 Ϯ 18, 169±229 (3, 30), OW 136 Ϯ 14, spherical growths. Autozooids erect with ter- 104±169 (3, 30), OvL 297 Ϯ 21, 272±312 minal ori®ce and tubular peristome, project- (1, 7), OvW 332 Ϯ 27, 291±364 (1, 7), SL ing visibly from colony surface. Frontal 23 Ϯ 4, 17±33 (3, 30), SW 28 Ϯ 4, 22±37 shield thickly calci®ed, vitreous and nodular; (3, 30), VAL 420 Ϯ 42, 355±473 (2, 9), marginal pores developing tubular extensions VAW 239 Ϯ 29, 193±271. (POLYPIDE): IH 24 to open around base of peristome. Primary Ϯ 37 ␮m, 0±120 (2, 19), LD 745 Ϯ 74, 640± ori®ce longer than wide, the deep, V-shaped 840 (2, 20), MD 30 (1, 1), TL 615 Ϯ 80, sinus comprising about one-third its total 500±800 (2, 20). length; peristome deep, tubular, rim lowest and broadly concave proximally, elsewhere Celleporina canariensis Aristegui, 1989 often produced into short processes; three col- Figure 39E±G umnar avicularia traverse the length of peri- stome and project above rim: typically one Celleporina canariensis Aristegui, 1989: 147. middistally, two laterally; rostra terminal, DESCRIPTION (AMNH 988; CMRR 2260): acute to ori®ce plane, distal one directed dis- Colonies cream-colored or light yellow, en- tally, lateral pair distolaterally. Vicarious av- crusting, multilaminar, forming small pisi- icularia frequent, monomorphic, characteris- form nodules, to 3 mm diameter in present tic: cystid broadly conical, proximal portion material. Autozooids small, with smoothly of rostrum apical, spatulate distal portion di- calci®ed frontal shield and large, conspicu- rected basally, down side of the cystid; cross- ous marginal pores. Primary ori®ce longer bar slender, without columella, palate with than wide, with a short, U-shaped sinus mid- extensive foramen. Ovicell slightly wider proximally and rounded, prominent con- than long, its frontal surface level with peri- dyles. No oral spines. Peristome deep, tubu- stome rim, with a narrow, sickle-shaped area lar, obscuring ori®ce, its rim shallowly of membranous ectooecium, exposed entooe- notched midproximally; peristomial avicular- cium vitreous, nodular, with a single series ia paired, lateral, rostrum terminal, elliptical, of large and irregular marginal pores. ca. 0.09 mm long, with complete crossbar, Tentacles light orange, 14±16; lopho- bluntly triangular, directed distolaterally. phores bell-shaped, radially symmetrical Distal to each avicularium, peristome rim is with short introverts to obliquely truncate produced as a short, blunt, medially directed with long introverts adjacent to excurrent process. Vicarious avicularia sparse, dimor- chimneys. phic: elliptical, 0.2 mm long, with complete Embryo color deep, bright red. Ancestrula crossbar, thickened columella, no palate; or described and ®gured by Ristedt (1996). larger, to ca. 0.4 mm, with broadly spatulate REMARKS: Celleporina caminata is similar rostrum, lacking palate, angled at 45Њ to mi- to C. canariensis Aristegui, 1989, but is dis- nute proximal opesia and directed basally. tinguished by its larger, more robust colony Ovicell about as wide as long, frontal surface form and larger autozooidal dimensions. The level with peristome rim; entooecial tabula primary ori®ce, in particular, is larger than occupying most of the frontal surface, bor- that of C. canariensis, with a proportionately dered by a single row of pores. Aperture deeper sinus, and the peristome rim of non- edged by a broad band of smooth ectooe- ovicellate autozooids bears three, distinctly cium, often produced medially into a short tubular avicularia. umbo. DISTRIBUTION: This familiar species is Tentacles light yellow, 12; lophophores common throughout the Mediterranean in ¯aring conical, radially symmetrical. shallow, coarse detrital habitats, encrusting REMARKS: This species is most similar to stone, shell, hydroids, and other bryozoans. C. caminata, a common Mediterranean spe- 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 87

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 ori®ce (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 ori®ce (0.05 mm). 88 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 cies which develops robust nodular colonies. proximal border, between short, broad con- It is distinguished from that species princi- dyles, its depth equivalent to one-®fth total pally by its smaller ori®ce and proportion- ori®ce length. No oral spines. Peristome en- ately shorter sinus. Further, the peristome rim circles whole of primary ori®ce, developing in C. caminata typically bears three colum- as a tall cylinder, its rim slightly ¯ared and nar avicularia with elliptical rostra. Aristegui produced into a variable number of short (1989) described only spatulate vicarious av- lobes. A small avicularium present midprox- icularia in his type suite but both these and imally on peristome rim, originating from a the smaller elliptical type occur in single col- septulum proximolateral to ori®ce, either onies of the material described here. right or left, traversing peristome as a slender DISTRIBUTION: Celleporina canariensis is tube and expanding into an ovate cystid on presently known from several localities its distal edge; rostrum acute to ori®ce plane, around the Canary Isles, and from caves proximally directed, oval, with a ®nely den- along the Catalan coast. ticulate distal rim, crossbar complete, slen- MEASUREMENTS (SKELETAL): AAL 98 Ϯ 10 der, mandible semielliptical. Vicarious avi- ␮m, 77±113 (3, 28), AAW 55 Ϯ 7, 47±72 cularia frequent, 0.25±0.3 mm long; rostrum (3, 8), DO 354 Ϯ 53, 259±452 (3, 30), OL broadened distally and deeply cupped, but 130 Ϯ 6, 119±139 (5, 22), OW 103 Ϯ 5, 89± not spatulate, crossbar complete, slender, 112 (5, 22), OvL 145 Ϯ 18, 116±174 (3, 12), without a columella; palate simply a narrow OvW 218 Ϯ 18, 189±224 (3, 12), SL 23 Ϯ shelf bordering a foramen occupying most of 5, 15±36 (5, 22), SW 23 Ϯ 4, 15±31 (5, 22), length and breadth of rostrum. Ovicell hemi- VAL 267 Ϯ 49, 213±308 (2, 3), VAW 166 spherical; a broad tabula occupying most of Ϯ 41, 142±213 (2, 3). (POLYPIDE): IH 0 ␮m visible frontal surface, with a thickened ec- (1, 2), LD 880 (1, 10), TL 560±680 (1, 2). tooecial rim and a single marginal series of large pores; aperture wide and arched, open- Celleporina siphuncula, new species ing into proximal portion of peristome. Figure 40A±D Above the tabula lateral lobes of peristome curve medially and abut, forming a suture as DIAGNOSIS: Colonies pisiform, small; ori- peristome lengthens into a tube. ®ce widely sinuate; peristome long, tubular; Tentacles cream-colored, 14±16; lopho- peristomial avicularium single, proximo-me- phores bell- to funnel-shaped, radially sym- dial. metrical between chimneys, obliquely trun- HOLOTYPE: AMNH 989. cate adjacent to chimneys. PARATYPES: AMNH 990±994; CMRR Embryo color pale orange. 2261. REMARKS: This tiny species is distin- ETYMOLOGY: The species name is from the guished from other Celleporina species in Latin sipho (bent tube) and ±culum (suf®x the northeast Atlantic-Mediterranean region denoting diminutive) because of the shape of by its widely sinuate ori®ce, long tubular the small peristomial avicularium. peristome and single, proximo-median peri- DESCRIPTION: Colonies cream colored, pi- stomial avicularium. siform, to 4.5 mm diameter in present ma- OCCURRENCE: It was abundant on Cellaria terial. Autozooids closely packed, oval in ®stulosa in the 35 m-deep meadow west of outline with a smooth, convex frontal shield, Banjole Island, together with Celleporina with a few indistinct marginal pores; com- caminata. monly 0.3 mm in length, with a cylindrical MEASUREMENTS (SKELETAL): AAL 63 Ϯ 9 peristome up to 0.375 mm long. Autozooid ␮m, 51±82 (3, 11), AAW 45 Ϯ 8, 34±65 (3, boundaries distinct only in the smallest col- 10), DO 295 Ϯ 47, 206±389 (3, 30), OL 89 onies, of fewer than 20 autozooids; subse- Ϯ 9, 79±101 (3, 5)m OW 93 Ϯ 5, 86±101 quent astogeny proceeds through fronto-lat- (3, 6), OvL 136 Ϯ 15, 116±167 (3, 18), OvW eral budding, with peristomes projecting per- 215 Ϯ 15, 194±242 (3, 18), SL 19 Ϯ 2, 16± pendicularly to colony surface. Primary ori- 22 (3, 7), SW 54 Ϯ 3, 50±59 (3, 7), VAL ®ce slightly wider than long; a shallow, 244 Ϯ 24, 216±272 (2, 4), VAW 159 Ϯ 14, u-shaped sinus occupies two-thirds of the 142±176 (2, 4). (POLYPIDE): IH 202 Ϯ 27 ␮m, 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 89

Fig. 40. A±D. Celleporina siphuncula. A. General aspect (paratype, AMNH 990; 0.5 mm). B. Au- tozooids and maternal zooids with ovicells (holotype, AMNH 989; 0.1 mm). C. Vicarious avicularia (holotype, AMNH 989; 0.1 mm). D. Autozooidal ori®ce (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 ori®ce (0.05 mm). 90 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

160±240 (1, 10), LD 634 Ϯ 74, 500±740 (1, 10), DO 354 Ϯ 37, 301±405 (1, 10), OL 130 10), MD 30 (1, 2), TL 610 Ϯ 50, 540±680 (1, 2), OW 103±109 (1, 2), AL 314±318 (1, (1, 10). 2), AW 238±258 (1, 2).

Celleporina tubulosa (Hincks, 1880) GENUS TURBICELLEPORA RYLAND, 1963 Figure 40E±G Turbicellepora avicularis (Hincks, 1860) Cellepora costazii var. tubulosa Hincks, 1880: Figure 41A±D 412. Cellepora avicularis Hincks, 1860: 278, 1880: Celleporina hassallii var. tubulosa: Zabala and 406. Maluquer, 1988: 159. Turbicellepora avicularis: Hayward and Ryland, Celleporina tubulosa: Hayward and Ryland, 1999: 336. 1999: 330. DESCRIPTION (AMNH 996±998; CMRR DESCRIPTION (AMNH 995): Colony pisi- 2262): Colonies orange, with variable form, form. Primary autozooidal ori®ce transverse- from low mounds and nodules to stout, ir- ly oval with broad, shallow sinus occupying regularly lobed or erect forms with tapered entire proximal width of anter. Tall, tubular branches; to 15 mm high in present material. peristome slightly ¯ared apically, narrowly Autozooids broad, convex, with boundaries transversely oval at aperture; avicularia visible only in few, frontally budded individ- paired, situated laterally on peristome rim, uals; frontal shield smooth, with a single se- with small oval rostra acute to ori®ce plane, ries of few, conspicuous marginal pores. Pri- directed laterally and with mandibles facing mary ori®ce slightly longer than wide, a each other. Ovicell with a peripheral row of deep, V-shaped proximal sinus comprising large pores, opening into peristome at about about one-quarter total length, ¯anked by halfway along its length. Vicarious avicularia thick, rounded condyles; no spines. Peri- broadly spatulate, with an extensive palatal stome low, absent midproximally, adjacent to foramen and a very slender crossbar lacking lateral suboral avicularium with laterally di- a columella. rected, elongate triangular rostrum, hooked REMARKS: A single specimen of this spe- distally, oblique to ori®ce plane; crossbar cies was found, but it is suf®ciently distinct stout, with distinct columella. Vicarious av- from other European species of Celleporina icularia characteristic, to 0.6 mm long, to be readily recognized. It is especially dis- broadly spatulate to more nearly parallel-sid- tinguished by its primary ori®ce. ed, rounded distally; always with an exten- DISTRIBUTION: This rare species has been sive palate, and a thick crossbar with a stout reported reliably on just three previous oc- columella often bi®d distally. A second, less casions. The type locality is unknown and frequently occurring vicarious avicularium the type specimen lost, but Hayward and Ry- has a broadly triangular, distally hooked ros- land (1999) described and ®gured a specimen trum and thick crossbar with equally thick from the Scilly Isles, southwest England, columella. Ovicell broader than long, smooth which closely matched Hincks' (1880) orig- surfaced, with 10±15 small round pores. inal description and ®gure. Zabala and Ma- Tentacles light orange, 16±18; lopho- luquer (1988) ®gured what is clearly the phores bell-shaped, radially symmetrical same species from the Catalan coast. The midway between chimneys to obliquely trun- specimen listed by Gautier (1962) from Si- cate, supported by long introverts. cily was said to be identical in morphology Embryo color bright, medium orange. and measurements to C. hassallii (Johnston) DISTRIBUTION: This is the most commonly except for an elongate peristome; no mention occurring species of Turbicellepora in the was made of the distinctive primary ori®ce northeast Atlantic region, distributed from and the positioning of the peristomial avi- northern Norway and southern Iceland cularia, and it is thus not certain that the southwards to the Mediterranean, where it is specimen belonged to C. tubulosa. widespread in shallow coastal waters. MEASUREMENTS (SKELETAL): AAL 78 Ϯ 9 MEASUREMENTS (SKELETAL): AAL 140 Ϯ ␮m, 69±93 (1, 10), AAW 37 Ϯ 3, 33±44 (1, 19 ␮m, 113±172 (3, 19), AAW 96 Ϯ 13, 62± 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 91

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 ori®ce (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). 92 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

120 (3, 19), DO 441 Ϯ 102, 262±624 (3, 30), cent Atlantic coast, described originally from OL 132 Ϯ 16, 110±167 (3, 26), OW 128 Ϯ the Aegean Sea (Hayward, 1978) and sub- 10, 108±145 (3, 26), SL 43 Ϯ 7, 26±54 (3, sequently reported from the Bay of Cadiz 26), SW 69 Ϯ 16, 47±103 (3, 26), VAL 448 and the Strait of Gibraltar (Harmelin and Ϯ 40, 412±537 (3, 10), VAW 276 Ϯ 26, d'Hondt, 1992). 248±338 (3, 10). (POLYPIDE): IH 383 Ϯ 51 MEASUREMENTS (SKELETAL): AAL 155 Ϯ ␮m, 320±460 (1, 10), LDMn 655 Ϯ 74, 24 ␮m, 124±186 (2, 9), AAW 133 Ϯ 22, 540±800 (1, 14), LDMx 695 Ϯ 59, 600±800 106±169 (2, 9), DO 389 Ϯ 70, 252±503 (3, (1, 17), MD 30 Ϯ 3, 25±35 (1, 8), TLMn 29), OL 112 Ϯ 8, 96±128 (3, 25), OW 119 554 Ϯ 38, 500±620 (1, 10), TLMx 707 Ϯ Ϯ 11, 101±136 (3, 25), SL 42 Ϯ 5, 35±53 140. 600±960 (1, 10). (2, 12), SW 88 Ϯ 9, 74±105 (2, 12), VAL 456 Ϯ 47, 394±518 (3, 7), VAW 320 Ϯ 28, Turbicellepora camera Hayward, 1978 267±394 (3, 7), ZL 483 Ϯ 28, 446±512 (1, Figure 41E±H 5), ZW333 Ϯ 28, 301±360 (1, 5).

Turbicellepora camera Hayward 1978: 570, ®gs. FAMILY PHIDOLOPORIDAE GABB AND 4N±P, 5T, 10. HORN, 1862 DESCRIPTION (AMNH 999; CMRR 2263): GENUS RETEPORELLA BUSK, 1884 Colonies orange, initially small, domed nod- ules, some of the present material developing Reteporella grimaldii (Jullien, 1903) the slender, cylindrical, branching growth Figure 42A±F characteristic of the species. Autozooids Retepora grimaldii Jullien, in Jullien and Calvet, oval, convex, with boundaries visible only in 1903: 62. newly budded individuals; smoothly calci®ed Sertella septentrionalis Harmer, 1933: 620. Gau- frontal shield bordered by few, small mar- tier, 1962: 233. Geraci, 1975: 244. Zabala, ginal pores. Primary ori®ce slightly longer 1986: 544. Zabala and Maluquer, 1988: 154. than broad, with a deep, V-shaped proximal Reteporella septentrionalis: Hayward and Ryland, sinus comprising almost half total length; 1996: 109, 1999: 374. Reteporella grimaldii: Reverter-Gil and FernaÂn- condyles broad and thick, no spines. A low, dez-Pulpeiro, 1999: 1416. thickened peristome developing in early on- togeny; lowest midproximally, adjacent to DESCRIPTION (AMNH 1000±1002; CMRR proximolateral suboral avicularium, with el- 2264): Colonies orange, erect, brittle, con- liptical, round-ended rostrum, oblique to or- sisting of bilaminar fenestrate plates, com- i®ce plane and laterally directed. Vicarious plexly infolded and fused into rigid three-di- avicularia characteristic: rostrum broadly mensional structure; inwards-facing surfaces spoon-shaped, 0.25±0.3 mm long, cupped bearing autozooids, outer faces consisting of distally; crossbar slender, with an indistinct sheets of kenozooids. Colony size and archi- median columella; completely lacking a pal- tecture variable, probably dependent upon ate and in cleaned material appearing as an microhabitat; a single colony from locality open chamber. Additional adventitious avi- 12 measuring 18 ϫ 14 ϫ 13 cm, but larger cularia sporadic, abundant in some areas of in situ specimens reported up to about 70 cm colony, identical to suboral type. Ovicell diameter and 35 cm thick perhaps consisted hemispherical, smooth, with few (Ͻ10) large, of several closely intergrown colonies. Fe- irregular pores. nestrulae elongate oval, commonly 1.5 ϫ 0.5 REMARKS: The present material does not mm; trabeculae of three to ®ve alternating differ from the Aegean type specimen (Hay- longitudinal autozooid series. Autozooids at ward, 1978); the shape of the ori®ce, the dis- growing edge elongate, rectangular, convex, tinctive vicarious avicularia and the prolif- with indistinct boundaries. Frontal shield erating adventitious avicularia together dis- smooth, with few, large marginal pores tinguish T. camera from all other Mediter- around proximal margins. Primary ori®ce ranean species of Turbicellepora. broader than long; distal border ®nely den- DISTRIBUTION: The species is apparently ticulate, proximal border shallowly concave endemic to the Mediterranean and the adja- between indistinct condyles. Single pair of 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 93

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). 94 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 oral spines present, one on each side of ori- 380±500 (1, 23), TLMn 324 Ϯ 22, 300±360 ®ce, persisting in late ontogeny. Peristome (1, 5), TLMx 447 Ϯ 25, 400±520 (1, 23). developing in early ontogeny, obscuring pri- mary ori®ce; deep, slightly ¯ared frontally, GENUS RHYNCHOZOON HINCKS, 1895 with conspicuous medio-proximal pore, Rhynchozoon neapolitanum Gautier, 1962 linked to peristome rim by clear groove. In- Figure 43A±E ner face of peristome coarsely denticulate, thickened spine bases prominent in each Rhynchozoon neapolitanum Gautier, 1962: 243. Zabala, 1986: 531. Zabala and Maluquer, 1988: proximal corner. Avicularia adventitious. 149. Frontal avicularia dimorphic: short, elliptical, 0.06 mm long, normal to frontal plane, with DESCRIPTION (AMNH 968, 1003; CMRR varying orientation; or larger, with narrowly 2265): Colonies cream-colored to tan, en- triangular, distally hooked rostrum, ca. 0.3 crusting, multilaminar; developing frontally mm long, orientated perpendicular to frontal budded, massive nodular forms, frequently plane and very conspicuous. Single large, exceeding 3 ϫ 3 ϫ 3 cm. Autozooids with transversely orientated avicularia in proxi- vitreous, nodular frontal shield bordered by mal, abfrontal axils of fenestrulae while short a single series of conspicuous marginal elliptical type are common across abfrontal pores. Primary ori®ce as wide as long; distal surfaces. Ovicell slightly longer than wide, and lateral rim with large, rounded, closely ¯attened frontally, with narrow ®ssure ex- spaced denticulations; proximal border with tending for about half its length; aperture short, narrow U-shaped sinus accentuated by overhung by short, straight-edged labellum. stout, faceted condyles. No oral spines. Peri- Tentacles light orange, 12; lophophores stome thick, deepening through ontogeny; distally ¯aring conical, radially symmetrical distal and lateral border developing short to slightly obliquely truncate, commonly knobs; proximal border characteristically with a pair of stout columnar processes ¯ank- overlapped with adjacent tentacles up to 60 ing a deep median embayment; at base of ␮m. inner surface of one process, a thick angular Embryo color deep, bright reddish-orange. uncinate lateral process projects into the lu- REMARKS: Colonies are small and delicate men of the peristome. Median suboral avi- in quiet environments such as within the Cel- cularium present in some autozooids with laria turf west of Banjole Island (®g. 42A), uncinate process at its proximal end; rostrum but overall colony size and branch width are broadly triangular, hooked distally, acute to much more robust in more exposed, less con- plane of ori®ce and facing laterally. Frontal ®ned environments (®g. 42B). avicularia often frequent in later ontogeny; DISTRIBUTION: Offshore, typically on hard characteristic: with voluminous cystid occu- substrata. This species is common in the pying more than half frontal surface of bear- northeast Atlantic region, from the White ing autozooid, rostrum elongate triangular, Sea, the Kara Sea, Iceland, and western acute to frontal plane, directed proximally or Greenland south to the Mediterranean, where proximolaterally. Ovicell short and broad; it seems to occur in shallower habitats than ¯at frontally, with a transversely oval area of to the north. Hayward and Ryland (1996) membranous ectooecium; aperture straight could not distinguish Mediterranean from edged, with a short notch in each corner. North Atlantic specimens. Tentacles cream-colored, 13±15; lopho- MEASUREMENTS (SKELETAL): AAL(large) phores distally ¯ared conical, radially sym- 324 Ϯ 77 ␮m, 192±494 (3, 30), AAW(large) metrical to obliquely truncate. 202 Ϯ 45, 133±297 (3, 26), AL(fenestrulae) Embryo color pale orange. 248 Ϯ 23, 331±386 (1, 10), DO 273 Ϯ 41, REMARKS: The largest colonies often in- 199±359 (4, 40), OW 97 Ϯ 11, 79±122 (3, corporate tubular channels opening over the 30), OvL 193 Ϯ 25, 149±248 (3, 24), OvW surface, suggesting they had grown in asso- 173 Ϯ 16, 143±208 (3, 24), ZL 353 Ϯ 17, ciation with hydroids (as in Rhynchozoon 331±386 (1, 10), ZW 187 Ϯ 13, 160±202 (1, larreyi Auct.; see Ristedt and Schuhmacher, 10). (POLYPIDE): IH ϳ0 ␮m, LD 450 Ϯ 35, 1985). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 95

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. Ontogenet- ically young (top left) and (center) old autozooids (AMNH 1003; 0.5 mm). D. Ovicell (AMNH 1003; 0.05 mm). E. Autozooidal ori®ce (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. Auto- zooidal ori®ce with ori®cial avicularium (0.05 mm). 96 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

DISTRIBUTION: This common and distinc- DISTRIBUTION: Mediterranean; this current tive species is widely distributed throughout record represents the ®rst occurrence of the the Mediterranean. species in the eastern Mediterranean. MEASUREMENTS (SKELETAL): DO 447 Ϯ 79 MEASUREMENTS (SKELETAL): AAL 85 Ϯ 11 ␮m, 301±635 (2, 20), OL 106 Ϯ 6, 93±117 ␮m, 71±96 (1, 6), AAW 56 Ϯ 11, 39±74 (1, (2, 20), OW 125 Ϯ 4, 117±131 (2, 20), OvL 6), DO 284 Ϯ 27, 229±317 (1, 11), SOD 134 196 Ϯ 16, 173±213 (2, 20), OvW 287 Ϯ 20, Ϯ 13, 112±163 (1, 22), ZL 401 Ϯ 44, 330± 258±311 (2, 20), SL 14 Ϯ 3, 11±20 (2, 20), 471 (1, 11), ZW 273 Ϯ 27, 229±317 (1, 11). SW 25 Ϯ 5, 19±36 (2, 20), ZL 614 Ϯ 54, 513±733 (2, 20), ZW 388 Ϯ 50, 292±462 (2, Rhynchozoon revelatus, new species 20). (POLYPIDE)IH69Ϯ 42 ␮m, 1±120 (1, Figure 44A±D 10), LD 709 Ϯ 61, 600±800 (1, 10), MD 30 Ϯ 4, 25±35 (1, 4), TL 663 Ϯ 61, 560±780 Rhynchozoon species 2: Hayward, 1974: 389. Za- bala, 1986: 528. Zabala and Maluquer, 1988: (1, 10). 149.

Rhynchozoon pseudodigitatum DIAGNOSIS: Primary ori®ce wide, D- Zabala and Maluquer, 1988 shaped, with widely spaced, ®ne denticles Figure 43F±H along distal rim; oral spines lacking; median suboral avicularium with distally-facing ros- Rhynchozoon lobulatum: Gautier, 1962: 242. trum acute to frontal plane, accompanied by Rhynchozoon pseudodigitatum Zabala and Malu- quer, 1988: 147. blunt uncinate process; dimorphic suboral avicularium. DESCRIPTION (AMNH 1004): A single, HOLOTYPE: AMNH 1005. small, encrusting, multiserial, unilaminar col- PARATYPES: AMNH 1006, 1007, 1056; ony. Primary ori®ce wider than long, the an- CMRR 2266±2269. ter denticulate and the poster shallowly con- ETYMOLOGY: The species name is from the cave between small, rounded condyles. With- Latin revelatus (unveil). in the peristome is a broad, straight-edged DESCRIPTION: Colonies cream-colored, en- uncinate process, the free corner of which is crusting, multilaminar, developing domed, produced and may abut against opposing side mamillate nodules, with mammillae forming of peristome wall, de®ning a pseudospira- incipient erect branches. Autozooids with men. Lateral suboral avicularium, with thick, vitreous frontal shield, bordered by a hooked rostrum acute to plane of ori®ce and single series of large marginal pores. Primary laterally directed; soon obscured by a thick- ori®ce wider than long, D-shaped; distal rim ened peristome, with a lobed rim developing with ®ne, widely spaced denticles, proximal into knobbed processes projecting over the rim straight, small rounded condyles just vis- lumen. Frontal shield nodular, vitreous, with ible in proximolateral corners. No oral some nodules developed as pronounced spines. Peristome a thick, low rim, not ob- knobs. Small adventitious avicularia with scuring ori®ce; median suboral avicularium slender, triangular rostrum, distributed spo- present on most autozooids, with distally radically on frontal shields of autozooids, hooked, triangular rostrum orientated acute with variable orientation. to frontal plane and facing distolaterally. Ad- REMARKS: The shape of the primary ori®ce jacent to proximal part of avicularium, peri- and the lobed peristome rim are typical of R. stome rim is shallowly notched, and a short, pseudodigitatum as described and ®gured by blunt uncinate process projects from base of Zabala and Maluquer (1988), who consid- avicularium cystid; apical part of cystid var- ered that the single specimen from the Gulf iably developed as a stout umbo. Suboral av- of Marseille attributed to the poorly de®ned icularium dimorphic, in most autozooids ros- R. lobulatum Waters by Gautier (1962) was trum length less than width of ori®ce; mas- the same species. sive type with broadly triangular, distally OCCURRENCE: The single, small colony hooked rostrum, Ͼ0.3 mm long. In smaller which appears to be attributable to this spe- avicularia, crossbar complete and thickened, cies was encrusting shell. but larger type lacks a columella; there is no 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 97

Fig. 44. A±D. Rhynchozoon revelatus. A. General aspect (holotype, AMNH 1005; 0.5 mm). B. Autozooids and (bottom right) maternal zooid with ovicell (holotype, AMNH 1005; 0.1 mm). C. Au- tozooidal ori®ce (holotype, AMNH 1005; 0.05 mm). D. Ancestrula (center) and early autozooids (par- atype, AMNH 1056). E±H. Schizotheca ®ssa. E. Entire, untreated colony (AMNH 942; 0.5 mm). F. Maternal zooid with normal ovicell (AMNH 1009; 0.1 mm). G. Oblique view from distal end of maternal zooid with ovicell bearing distal adventitious avicularium (AMNH 1010; 0.1 mm). H. Ontogenetically young, untreated autozooids (AMNH 942; 0.1 mm). 98 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 palate. No frontal adventitious avicularia. luquer, 1988: 150. Hayward and Ryland, 1999: Ovicell elongate oval, ¯at frontally, with a 382. broadly arched aperture; projecting vertically DESCRIPTION (AMNH 942, 1009, 1010; above peristome rim; smooth surfaced, fron- CMRR 2270): Colonies translucent, encrust- tal ectooecium almost entirely membranous. ing, multiserial, unilaminar, small, rounded In large colonies, greater than 2 cm high and patches. Autozooids oval to hexagonal, wide, basal surfaces largely unattached to the slightly convex, with indistinct boundaries. substratum. Frontal shield thickly calci®ed, ®nely gran- Ancestrula 0.25 mm long with transverse- ular, becoming coarser in later ontogeny; ly oval opesia occupying half frontal surface, with few (1±3) indistinct marginal pores. Pri- bordered by undetermined number of spines. mary ori®ce orbicular, with denticulate distal Two to four spines present in ®rst three to rim and small, U-shaped proximal sinus. four astogenetic generations only. Peristome well developed, cylindrical, pro- Tentacles clear, 12±13 in zooids closest to jecting well above frontal shield; six long, ancestrula, 14±15 in subsequent zooids; slender spines evenly spaced around distal lophophores bell-shaped, radially symmetri- and lateral borders (four in ovicellate zo- cal midway between chimneys, grading to oids); midproximally, rim bears short, U- strongly obliquely truncate, campylonemi- shaped notch, continued on inner distal face dan, in some instances scalloped adjacent to of peristome as deep groove. Avicularia vi- chimneys. carious, cystid about as large as an auto- OCCURRENCE: Diverse shell substrata; zooid; rostrum triangular, directed distally. common on the thecate ascidian Microcos- Ovicell elongate oval, recumbent on distally mus. succeeding autozooid, smooth surfaced, im- DISTRIBUTION: Originally recorded from perforate, with elongate frontal ®ssure; at Chios, Aegean Sea, this species has remained ®rst prominent, later immersed in thickened undescribed although it has been reported calci®cation; some with avicularium on distal from several other localities in the Mediter- surface. ranean. It was extremely common in the DISTRIBUTION: Often common on biogenic carbonates, S. ®ssa is widespread throughout Rovinj region. the Mediterranean, in shallow detritic envi- MEASUREMENTS (SKELETAL): AL(large) 384 ronments. It ranges north to the southwest 72 m, 222±474 (3, 12), AW(large) 257 Ϯ ␮ British Isles, and south to the Cape Verde Ϯ 74, 136±386 (3, 10), AL(small) 146 Ϯ 12, Islands. 132±167 (1, 10), AW(small) 102 Ϯ 9, 92± MEASUREMENTS (SKELETAL): AL 376 Ϯ 106 117 (1, 10), DO 472 Ϯ 97, 334±691 (3, 30), ␮m, 207±528 (3, 9), AW 225 Ϯ 52, 171±322 OL 118 Ϯ 7, 106±133 (3, 30), OW 162 Ϯ (3, 9), DO 333 Ϯ 30, 293±395 (3, 25), OL 12, 140±187 (3, 30), OvL 190 Ϯ 24, 148± 78 Ϯ 5, 72±86 (1, 10), OW 83 Ϯ 7, 74±92 220 (3, 6), OvW 259 Ϯ 34, 196±286 (3, 6), (1, 10), OvL 183 Ϯ 27, 149±242 (2, 16), ZL 658 Ϯ 83, 556±770 (1, 10), ZW 404 Ϯ OvW 211 Ϯ 18, 189±249 (2, 16), ZL 436 Ϯ 19, 370±431 (1, 10). (POLYPIDE): IH 116 Ϯ 44, 364±510 (3, 22), ZW 290 Ϯ 20, 251± 59 ␮m, 40±300 (2, 24), LDMn 608 Ϯ 70, 322 (3, 22). 500±740 (2, 23), LDMx 692 Ϯ 166, 500± 1140 (2, 30), MD 25 (1, 1), TLMn 592 Ϯ Schizotheca serratimargo (Hincks, 1886) 55, 420±680 (2, 25). Figures 45A±H, 46 Schizoporella serratimargo Hincks, 1886: 268. GENUS SCHIZOTHECA HINCKS, 1877 Schizotheca serratimargo: Zabala and Maluquer, 1988: 148. Schizotheca ®ssa (Hincks, 1856) Figure 44E±H DESCRIPTION (AMNH 1011±1013; CMRR 2271): Colonies orange, erect, bilaminate, Lepralia ®ssa Busk, 1856: 308. branching and anastomosing, developing rig- Schizotheca ®ssa Hincks, 1880: 284. Gautier, id, three-dimensional, reticulate structures; 1962: 223. Zabala, 1986: 543. Zabala and Ma- branches of variable width (3.6 Ϯ 1.0 mm, 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 99

Fig. 45. A±H. Schizotheca serratimargo. A. Ontogenetically young zooids at branch tip (AMNH 1011; 0.5 mm). B. Ontogenetically mature maternal zooids with ovicells (AMNH 1012; 0.5 mm). C. Ontogenetically young maternal zooid with incipient ovicell (AMNH 1011; 0.1 mm). D. Branch margin with vicarious avicularia along edge (AMNH 1013; 0.5 mm). E. Completely formed ovicells of maternal zooids (AMNH 1012; 0.1 mm). F. Ontogenetically old autozooids with adventitious avicularia (AMNH 1011; 0.1 mm). G, Vicarious avicularium (AMNH 1011; 0.1 mm). H. Autozooidal ori®ce (AMNH 1012; 0.05 mm). 100 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 46. Schizotheca serratimargo. A. Tentacle length and height of introvert with respect to distance from edge of branch. B. Lophophore asymmetry plotted as ratio of maximum tentacle length to minimum tentacle length with respect to distance from edge of branch.

N ϭ 221), dividing frequently and irregular- denticulate inner rim; poster shallowly con- ly. Autozooids hexagonal to irregularly po- cave with an indistinct median sinus, empha- lygonal, becoming more rounded in later on- sized and deepened by thick, prominent con- togeny but remaining distinct, each bounded dyles extending on each side from proxi- by thin, raised sutures. Primary ori®ce longer molateral corner to edge of sinus. Two wide- than wide; anter semielliptical, with ®nely ly spaced distal oral spines present, persisting 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 101 in late ontogeny. Frontal shield thickly cal- CLASS STENOLAEMATA BORG, 1926 ci®ed, convex, nodular with few (Ͻ6) irreg- ORDER CYCLOSTOMATA BUSK, 1852 ularly distributed marginal pores. Adventi- SUBORDER TUBULIPORINA tious avicularia sporadic, often absent over MILNE EDWARDS, 1838 young areas of the colony while frequent on FAMILY TUBULIPORIDAE JOHNSTON, 1838 older portions; rostrum elongate triangular, distally hooked, acute to frontal plane and GENUS TUBULIPORA LAMARCK, 1816 with variable orientation. Large vicarious av- Tubulipora liliacea (Pallas, 1766) icularia present along branch margins, cystid Figures 47A±D, 48 as large as an autozooid, rostrum almost as long, elongate triangular and distally hooked; Millepora liliacea Pallas, 1766: 248. Tubulipora liliacea: Harmer, 1898: 90. Harmelin, palate with a triangular foramen, crossbar 1976: 171. Hayward and Ryland,1985a: 74. Za- thick but without a columella. Ovicell hy- bala and Maluquer, 1988: 176. perstomial, longer than wide, rather small; aperture widely open, extending distally as a DESCRIPTION (AMNH 1014±1016; CMRR triangular ®ssure; ooecial cover obscures ovi- 2272): Colonies encrusting to semierect, cell in later ontogeny. commonly extending as two multiserial lobes Tentacles light orange, 12±15, usually 14; diverging laterally from axis of ancestrula, lophophores bell-shaped, radially symmetri- up to at least 8 mm in the vicinity of Rovinj; cal with uniform tentacle lengths away from usually violet-colored. Autozooids with long branch margins, grading to strongly oblique- peristomes, longest in the most cryptic mi- ly truncate with increasingly unequal tentacle croenvironments. Peristomes isolated in early lengths and longer introverts along branch astogenetic portion of colony, then organized margins (®g. 46), which serve as elongate into single-row fascicles radiating from cen- excurrent strips; lophophores also obliquely ters of lobes. Peristomes contiguous and ap- pressed within fascicles but in some, inde- truncate immediately adjacent to chimneys, pendent at distal ends. Individual fascicles which develop on broadest bifoliate surfaces constituted by 2 to 8 peristomes in present (McKinney, 1989). material and elsewhere in the Mediterranean DISTRIBUTION: Described from the Adriat- (Harmelin, 1976), but up to 12 in British ic, where it is common on hard substrata in specimens (Hayward and Ryland, 1985a). shallow inshore habitats, S. serratimargo has Spacing between fascicles variable. Auto- also been found to be abundant on the Cat- zooidal apertures oval where isolated and alan coast (Zabala, 1986) and is probably elongate polygonal where connate. widely distributed throughout the Mediter- Gonozooids distinctly in¯ated, centered in ranean. There is a single record of it from lobes, more densely perforated and with larg- the western English Channel (Hayward and er pseudopores than autozooids. More than Ryland, 1999). one ooeciostome present in some lobes, sug- MEASUREMENTS (SKELETAL): AAL 177 Ϯ gesting multiple con¯uent or contiguous 37 ␮m, 134±252 (2, 15), AAW 103 Ϯ 28, gonozooids. Ooeciostomes located distally, 73±177 (2, 15), DO 356 Ϯ 36, 295±439 (2, adjacent to base of second or third peristome, 20), OL 110 Ϯ 7, 99±125 (2, 20), OW 96 Ϯ usually located against side of fascicle facing 5, 88±105 (2, 20), OvL 171 Ϯ 13, 154±204 distal end of lobe, curved, opening laterally (2, 20), OvW 174 Ϯ 10, 160±193 (2, 20), SL or downward, toward surface of gonozooid; 11 Ϯ 2, 7±17 (2, 20), SW 21 Ϯ 4, 15±28 (2, ooeciostomes constricted at some point to 20), VAL 531 Ϯ 58, 437±656 (2, 19), VAW about the diameter of autozooids, then ter- 335 Ϯ 38, 267±402 (2, 19), ZL 502 Ϯ 55, minating in a broadly ¯aring oval hood. 421±641 (2, 20), ZW 345 Ϯ 19, 310±379 (2, Protoecium hemispherical, about 120 ␮m 20). (POLYPIDE): IH 125 Ϯ 63 ␮m, 30±240 diameter, giving rise to laterally oriented, (6, 52), LDMn 560 Ϯ 107, 420±800 (4, 37), elongate tubular portion of ancestrula. Pro- LDMx 638 Ϯ 143, 380±940 (6, 62), TLMn toecial cone gradually widening, generally 472 Ϯ 88, 360±640 (6, 42), TLMx 689 Ϯ curved right or left and consisting of about 186, 400±1060 (6, 53). four generations of autozooids with isolated 102 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 47. A±D. Tubulipora liliacea. A. General aspect (AMNH 1014; 0.5 mm). B. Position of ooe- ciostome (center) (AMNH 1015; 0.2 mm). C. Ooeciopore (AMNH 1014; 0.1 mm). D. Ancestrula and early autozooids (AMNH 1016; 0.2 mm). E, F. Tubulipora plumosa (AMNH 1017). E. General aspect (0.5 mm). F. Damaged ooeciostome (center) and (right) fascicle of autozooids (0.01 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 103

truncate due to increasingly unequal tentacle lengths toward chimneys in colony centers (®g. 48A), although tentacles are arrayed to maintain roughly circular shapes (®g. 48B). OCCURRENCE: The species is widely dis- tributed, occurring on bivalve shells, bryo- zoans, rocks, and Microcosmus. In the pre- sent material, colonies on bivalve shells, rocks, and Pentapora fascialis usually are adnate, while those on Cellaria and Micro- cosmus usually are semierect. Specimens that occur on bivalve shells are overwhelmingly located on the shell interiors, indicating a strong preference for cryptic microenviron- ments (Harmelin, 1976; McKinney, 2000). DISTRIBUTION: T. liliacea is widespread in the eastern North Atlantic and Mediterra- nean; it has also been reported from the Ba- rents Sea (Kluge, 1962, 1975) and from the northeastern coast of North America (Os- burn, 1912), though these more distant re- cords need to be con®rmed. Harmelin (1976) recorded the species at depths up to 60 m in the western Mediterranean, and found that variations in the robustness of colonies and orientation of the ooeciostome were related to the physical environment. MEASUREMENTS (SKELETAL): ADMN 142 Ϯ 17 ␮m, 120±180 (4, 40), ADMX 169 Ϯ 19, 120±200 (3, 30), Gap 327 Ϯ 35, 240±420 (4, 31), OsDMN 213 Ϯ 23, 200±240 (2, 3), Fig. 48. Tubulipora liliacea. A. Ontogenetic OsDMX 328 Ϯ 32, 270±360 (3, 6). (POLYP- increase in tentacle length: Filled circles plot max- IDE): IH 0 Ϯ 0 ␮m (6, 67), LMN 503 Ϯ 144, imum tentacle length within lophophores (Y ϭ 277.244 ϩ 0.445X) and open circles plot mini- 260±880 (6, 41), LMX 520 Ϯ 133, 320±920 mum tentacle lengths (Y ϭ 266.548 ϩ 0.235X). (6, 43), MD 25 Ϯ 0 (2, 2), TLMn 462 Ϯ B. Ontogenetic increase in lophophore diameter: 121, 160±720 (6, 54), TLMx 601 Ϯ 204, Filled circles plot maximum lophophore diameters 300±1080 (6, 59). (Y ϭ 306.178 ϩ 0.285X) and open circles plot minimum diameters (Y ϭ 269.182 ϩ 0.314X). Tubulipora plumosa Harmer, 1898 Figure 47E, F Tubulipora plumosa Harmer, 1898: 105. Harme- peristomes. At distal end of protoecial cone, lin, 1976: 177. Hayward and Ryland, 1985a: 80. budding rate increased rapidly, particularly Zabala, 1986: 672. Zabala and Maluquer, 1988: along lateral margins (establishing two lat- 176. erally diverging lobes), and transversely ori- Tubulipora ¯abellaris Johnston, 1847: 274. Busk, ented fascicles of peristomes appearing 1875: 25 (pars, ®de Harmer). abruptly, establishing astogenetic zone of Tubulipora ®mbria: Hincks, 1880: 448. repetition. DESCRIPTION (AMNH 1017): Colonies en- Tentacles clear but locally colored by pur- crusting, white, extending as one or more ple or brown granules, 11; lophophores short, multiserial lobes. Autozooids in one- small, conical, radially symmetrical along to two-row fascicles; up to at least 18 com- growing margin to campylonemidan, pro- pletely connate autozooids per fascicle. Au- gressively larger and more strongly obliquely tozooids relatively large, elongate polygonal 104 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 to semipolygonal in cross section. Autozooi- differ from the Middle Jurassic type species dal fascicles diverging to right and left from of Idmonea Lamouroux 1821, which is pre- axis of lobe. Fascicles highest at inner ends, dominantly encrusting and has elliptical where composed of oldest autozooids, grad- branch cross sections in the occasional erect ually diminishing to zero height at outer portions. Additionally, species of Exidmonea ends, in circum-colony budding zone. lack proximally directed kenozooids on the Gonozooids located distally within lobes, ¯at abfrontal surface of the branches and their brood chambers ramifying between sev- therefore differ from Idmidronea Canu and eral adjacent fascicles. Ooeciopore located at Bassler, 1920, which has them. As noted by distal end of a centrally located rami®cation. Taylor and Voigt (1992) Mongereau did not Ooeciopores adjacent to second or third peri- choose a type species from among the spe- stome within fascicle, with broadly ¯ared, cies that he listed for Exidmonea in 1969, so highly elongate oval hoods oriented away the ®rst valid use of Exidmonea was by Da- from surface of brood chamber. Colony sur- vid et al. (1972), when the type species Ex- faces, including fascicles, ®nely but clearly idmonea atlantica AUCT. was ®rst speci®ed. corrugated, with abundant circular to distally tapered triangular pseudopores. Surfaces of Exidmonea triforis (Heller, 1867) brood chambers poorly corrugated, with very Figures 49A±H, 50 abundant pseudopores. Distance between pseudopores on roofs of brood chambers ap- Idmonea triforis Heller, 1867: 120. Waters, 1879: 271. Waters, 1922: 12. Friedl, 1917: 276. proximately equaling pseudopore diameter. Idmonea meneghinii Heller, 1867: 120. Tentacles clear, 11; lophophores bell- Idmonea concava: Waters, 1879: 271. shaped, radially symmetrical to obliquely Idmonea marionensis: Waters, 1879: 270. truncate and campylonemidan, grading from Tubulipora atlantica: Harmer, 1915: 124. Cook, small along colony margin to largest and 1968: 230. most strongly obliquely truncate bordering Idmidronea atlantica: Osburn, 1947: 5. Harmelin, chimneys in colony centers. 1976: 182. Zabala, 1986: 658. REMARKS: Authorship of this species is Exidmonea atlantica: David et al., 1972: 84. Mc- commonly attributed to Thompson in Harm- Kinney, 1991a: 264. McKinney, 1991b: 437. er (1898), because Harmer adopted the trivial LECTOTYPE (chosen here): UIIZ 321. name plumosa that was used by Thompson DESCRIPTION (AMNH 1018, 1019; CMRR for specimens from Ireland sent to Johnston 2273): Delicate erect planar or nearly planar (cited in Johnston, 1847). However, Harmer fan-shaped, white colonies of dichotomous (1898) based his description on specimens branches. Branches subtriangular in cross from the coast of Devon, leading Ryland section, with ¯at abfrontal surface along (1963: 7) to designate one of Harmer's spec- which the zooids originated and keeled fron- imens from Devon, housed in the Museum tal surface. Autozooids divergent at low an- of Zoology, Cambridge, as lectotype. gle from abfrontal surface toward frontal sur- DISTRIBUTION: T. plumosa is known from face; those emergent near branch midline be- the eastern North Atlantic, from Norway to ing longest. Autozooids in single-row con- the Mediterranean. nate fascicles, each fascicle extending across MEASUREMENTS (SKELETAL): ADMN 181 Ϯ half of frontal surface, from central keel to 20 ␮m, 160±220 (1, 10), ADMX 194 Ϯ 21, 160±220 (1, 10), Gap 420 Ϯ 85, 230±530 lateral edge. Typically four, less commonly (1, 10), OsDMN 230 (1), OsDMX: 440 (1). three or ®ve, autozooids within a row. Fas- cicles typically curved proximally near lat- (POLYPIDE): IH 0 Ϯ 0 ␮m (1, 10), LD 424 Ϯ 82, 320±556 (1, 10), TLMn 358 Ϯ 112, 200± eral branch margin. Peristomes curved, bend- 540 (1, 10), TLMx 402 Ϯ 112, 200±600 (1, ing away from the branch midline. Peri- 11). stomes near branch midline longer, usually extending laterally beyond those closer to GENUS EXIDMONEA DAVID, MONGEREAU, AND lateral branch margin. POUYET, 1972 Gonozooids centered on frontal branch Mongereau (1969) introduced the name surfaces, most commonly situated immedi- Exidmonea for erect idmoneid species that ately proximal to branch bifurcations Brood 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 105

Fig. 49. A±H. Exidmonea triforis. A. General aspect (lectotype, UIIZ 321; 2 mm). B. Gonozooid (lectotype, UIIZ 321; 0.5 mm). C. Recurved ooeciostome and terminal ooeciopore (lectotype, UIIZ 321; 0.05 mm). D. Reverse surface of colony (AMNH 1018; 0.5 mm). E. Pseudopores and growth lines on reverse surface (AMNH 1018; 0.1 mm). F. Colony with two gonozooids at branch bifurcations (AMNH 1019; 0.5 mm). G. Reverse surface of broken colony base with kenozooids (AMNH 1018; 0.2 mm). H. Transverse break near colony base with kenozooids (above) and (below) autozooids (AMNH 1018; 0.02 mm). I, J. Exidmonea coerula (AMNH 1020). I. Frontal surface (0.5 mm). J. Reverse surface (0.5 mm). 106 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

lophophores, those from branch center ex- tending laterally farther than those at branch margins, with their highly obliquely truncate outer end oriented toward abfrontal side. They therefore generate a uni®ed colonial feeding current that draws nutrient-bearing water from the abfrontal side of branches and ejects ®ltered water along the frontal surface (McKinney, 1991a, 1991b). REMARKS: We designate as lectotype Hell- er's specimen illustrated here as ®g. 49A±C (UIIZ 321). This species has long been re- ferred to as Idmonea atlantica Forbes in Johnston, 1847, or as I. atlantica auctt. When David et al. (1972: 84) speci®ed Exidmonea atlantica AUCT. as type species of Exidmo- nea, they established a new species, separate from Idmonea atlantica Forbes in Johnston, Fig. 50. Exidmonea triforis. Relationship be- 1847. Although David et al. (1972) listed tween maximum tentacle length and maximum Miocene (Burdigalian) specimens as the ma- lophophore diameters within autozooids. Auto- terial in hand when they established the new zooids in row closest to branch axis indicated by species, they did not designate a holotype. crosses, and those progressively farther from axis However, type material exists for two of and closer to lateral margin of branch indicated Heller's (1867) species that have precedence successively by pluses, asterisks, triangles, and for at least the Recent Idmonea atlantica open circles. Relationship between tentacle length and lophophore diameter (marked by regression auctt. of the Adriatic and probably beyond line) is Y ϭ 0.008X1.810, and is generated by (Mediterranean, Atlantic). Specimens of Ex- change in lophophore shape from broadly ¯ared, idmonea from the vicinity of Rovinj encom- almost discoidal, at branch margin to more nar- pass the attributes present in Heller's speci- rowly conical nearer branch axis, where lopho- mens of Idmonea meneghinii (UIIZ 320) and phores are campylonemidian. Idmonea triforis. We prefer to use the name triforis for the species, since most of our specimens are gracile rather than robust. chambers engul®ng lower portions of peri- The lectotype of E. triforis has proximally stomes in two to four fascicles on each side directed kenozooids on the proximal end of of branch; ooeciopore situated at the end of its reverse surface. The proximal end of the short, approximately 180Њ-curved ooeciosto- specimen is slightly expanded and apparently me with smaller diameter than peristomes of is broken from just above its base of attach- typical autozooids. Ooeciopores typically ment, which is not preserved. In other spec- opening downward, toward branch surface. imens, the base of attachment is composed Ooeciostomes typically located within distal largely of kenozooids that lap a short dis- portion of brood chamber, against distal side tance up around the base of the erect portion. of a fascicle of autozooidal peristomes. Beyond the short distance of kenozooidal Tentacles clear, 10; lophophores of zooids thickening of the proximal end, the reverse located along branch keel highly asymmet- side has no kenozooids. rical and campylonemidian, with longest ten- Robustness of colonies is strongly in¯u- tacles on the side closest to the branch center. enced by environment and microenviron- Size of lophophores and constituent tentacles ment. In a study of growth habits of E. tri- decreased, degree of asymmetry progressive- foris across several environments, from ob- ly decreased, and ¯are of the tentacles pro- scure parts of caves to relatively exposed gressively increased, in autozooids closer to shallow terrigenous and rocky substrata, Har- lateral branch margin (®g. 50). Due to the melin (1973b) found length between bifur- shapes, sizes, placement, and orientation of cations to be highly variable, and bifurcation 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 107 angle, width of ooeciostome, width of typically curved proximally near the lateral branches, and number of zooecia per fascicle branch margin. Most peristomes curved, bent to be moderately variable. Several of these away from branch midline. Proximal por- variable branch characteristics interact to tions of autozooids and their peristomes lon- produce more irregular, delicate colonies in gest for zooids along the central keel and environments such as caves and more geo- shortest for those along lateral edge of metrically regular, though often smaller, col- branch. Individual peristomes may alternate- onies with more densely spaced, thicker ly expand or contract, and their diameters are branches in the more exposed environments. less constant than are those in E. atlantica. Size of gonozooids, and especially size of Peristome and aperture diameters decreased apertures and spacing between adjacent rows slightly toward the abfrontal surface. of apertures, were stable across environ- REMARKS: Harmelin (1976) reported typi- ments. cally ®ve to six peristomes per fascicle, with DISTRIBUTION: E. triforis is widespread on a range from three to eight in the original rock and skeletal substrata in the Mediterra- specimens that he collected from the Medi- nean Sea and cold temperate Atlantic Ocean. terranean near Marseilles. No gonozooids MEASUREMENTS (SKELETAL): AD 83 Ϯ 8 were found in the Adriatic specimens ex- ␮m, 70±100 (3, 30), PD 105 Ϯ 9, 90±120 amined, but Harmelin's (1976) specimens (3, 30), ASW 201 Ϯ 24, 160±240 (3, 30), had frontally centered, elongate gonozooids, RS 405 Ϯ 48, 300±500 (3, 30), GL 1408 Ϯ with in¯ated brood chambers that extended 284, 1020±2120 (3, 12), GW 793 Ϯ 253, across three to ®ve fascicles of zooids. Brood 480±1340 (3, 12), OsD 76 Ϯ 13, 60±100 (2, chambers most commonly are situated im- 7), BW 537 Ϯ 126, 360±800 (3, 20). (POL- mediately proximal to branch bifurcations, YPIDE): IH 0 Ϯ 0 ␮m (7, 70), MD 19 Ϯ 2.5, though some extend from the parent branch 15±20 (2, 4), LDMn 309 Ϯ 80, 200±420 (4, onto the proximal portions of both descen- 22), LDMx 339 Ϯ 63, 200±460 (7, 80), dant branches. Ooeciopores are located at TLMn 203 Ϯ 43, 95±320 (7. 64), TLMx 308 about midlength of brood chambers, on the Ϯ 96, 90±560 (7, 88). distal side of an autozooidal peristome, at the end of short, recurved, transversely com- Exidmonea coerula (Harmelin, 1976) pressed ooeciostome. The outer side of each Figure 49I, J recurved ooeciostome is prolonged as a short hood partially occluding the ooeciopore. Idmidronea coerula Harmelin, 1976: 185. DISTRIBUTION: The species is previously DESCRIPTION (AMNH 1020; CMRR 2274): known only from the Mediterranean. Erect, fan-shaped colonies of dichotomous MEASUREMENTS (SKELETAL): AD 126 Ϯ 9 branches that curve gently laterally or fron- ␮m, 110±140 (2, 20), PD 144 Ϯ 9, 130±160 tally. Colonies less compact and branches (2, 20), ASW 270 Ϯ 51, 180±400 (2, 20), more robust than for Exidmonea atlantica. RS 515 Ϯ 46, 440±600 (2, 20), BW 635 Ϯ Branch cross section subtriangular, with ¯at 85, 480±780 (2, 20). abfrontal surface along which zooids origi- nated and with keeled frontal surface. Dis- FAMILY ANNECTOCYMIDAE tally convex growth lines on abfrontal sur- HAYWARD AND RYLAND, 1985b face re¯ect former positions of branch tips. GENUS ANNECTOCYMA Peristomes of zooids arranged in well-devel- HAYWARD AND RYLAND, 1985b oped to ragged transverse fascicles, each lo- cated within one half of the frontal surface. Annectocyma arcuata (Harmelin, 1976) Complete rows consist of usually four, lo- Figure 51A±D cally ®ve, peristomes, with distal tips isolated Diaperoecia arcuata Harmelin, 1976: 89. Zabala, and bent in various directions. Locally, au- 1986: 620. tozooidal rows less well de®ned, with fasci- Annectocyma arcuata: Zabala and Maluquer, cles of three or two peristomes, and less 1988: 167. commonly individual peristomes completely DESCRIPTION (AMNH 1021±1023): Colo- independent of others in their row. Fascicles nies of undivided erect branches up to at 108 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 51. A±D. Annectocyma arcuata. A. General aspect (AMNH 1021; 0.5 mm). B. Brood chamber (AMNH 1022; 0.5 mm). C. Autozooidal interior with small spines (AMNH 1023; 0.05 mm). D. Au- tozooidal peristome and ori®ce (bottom) and (center) ooeciostome with oval ooeciopore (AMNH 1022; 0.1 mm). E, F. Annectocyma cf. A. major (AMNH 1024). E. General aspect (0.5 mm). F. Incompletely formed brood chamber at branch tip (0.2 mm). G±I. Annectocyma sp. G. Colony origin with divergent branches and (bottom) lateral branch (AMNH 1025; 0.5 mm). H. Oblique view of colony origin (center) and (left) lateral branch with incipient brood chamber (AMNH 1026; 0.5 mm). I. Ancestrula and early autozooids (AMNH 1025; 0.1 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 109 least 4 mm long, narrow at base and broad- divergent peristomes, with curvature increas- ening upwards, from 0.5 mm basally to 1 ing away from branch surface. Autozooidal mm width at top. Individual branches gently aperture diameters and spacing relatively arched, with distinct peristome-bearing con- large. Brood chamber widening distally, in- vex frontal and zooid-barren concave abfron- corporating bases of several autozooecial tal surfaces. Autozooidal peristomes with peristomes; ooeciostome not formed. small diameter, distributed in regular quin- REMARKS: The single specimen has the cuncical pattern over frontal surface. Brood same growth habit as Annectocyma arcuata. chambers in¯ated, located distally in region However, the autozooids and autozooidal of maximum branch width, occupying over spacing are distinctly larger and fall within half the branch width. Ooeciostomes more or the lower range of sizes of A. major as given less centrally placed in brood chamber with by Harmelin (1976), which is greater than smaller diameter basally than peristomes of that for any of the other Mediterranean spe- autozooids, then compressed into transverse- cies of Annectocyma. Having only one spec- ly elongated aperture with transverse length imen and in the absence of information on greater than peristomial diameter of auto- the ooeciostome, we are reluctant ®rmly to zooids. Several autozooidal peristomes pen- assign the specimen to A. major. etrate brood chambers. MEASUREMENTS (SKELETAL): AD 165 Ϯ 10 REMARKS: Colony morphology and zooidal ␮m, 150±180 (1, 10), AS 506 Ϯ 86, 400± measures of the Adriatic specimen corre- 640 (1, 10). spond exactly with attributes determined by Harmelin (1976). All Harmelin's specimens were individual branches, as is the Adriatic Annectocyma sp. specimen. However, the Adriatic specimen Figure 51G±I arises from a short section of a proximally DESCRIPTION (AMNH 1025, 1026): Colo- broken adnate portion, leaving open the pos- nies encrusting, consisting of one or more sibility that more than one branch arose from lobe-shaped, multiserial branches. First- a linearly extended encrusting base. formed encrusting branch extending in same DISTRIBUTION: This species is previously known from Sicily and the coast of Pro- direction as ancestrula or diverging oblique- vence. Harmelin (1976) reported the species ly. In two of the three specimens, a second at depths of 60 to 150 m, growing on shells branch budded obliquely back from ®rst of dead molluscs on the coralligenous sea asexual zooid, extending opposite growth di- ¯oor. rection of ancestrula and primary branch. MEASUREMENTS (SKELETAL): AD 141 Ϯ 7 Additional encrusting branches in these two ␮m, 133±150 (1, 10), PD 167 Ϯ 23, 137± specimens originating adventitiously about 200 (1, 10), AS 343 Ϯ 40, 283±417 (1, 10), midway along length of primary branch and GL 595 Ϯ 21, 583±620 (1, 3), GW 417 (1, extending perpendicularly from branch mar- 1). gin. No erect branches present in available specimens. Autozooids with small diameters Annectocyma cf. A. major (Johnston, 1847) but relatively widely spaced. Peristomes in Figure 51E, F available specimens relatively short. Ances- Alecto major Johnston, 1847: 281. trulae short, with the protoecial disc (233 Ϯ Stomatopora major: Hincks, 1880: 427. 15 ␮m, N ϭ 3) almost three times diameter Diaperoecia major: Harmelin, 1976: 79. Zabala, of the peristome (833 Ϯ 6 ␮m, N ϭ 3) ex- 1986: 623. tending from it. Annectocyma major: Hayward and Ryland, REMARKS: The specimens are more deli- 1985b: 1077. Zabala and Maluquer, 1988: 167. cate than either of the two other species of Entalophora proboscidea: Calvet, 1931: 33. Annectocyma found in the vicinity of Rovinj DESCRIPTION (AMNH 1024): Colony erect, and have a different growth habit, but we single, gradually widening, arched branch have not assigned them to a species because with incomplete terminal brood chamber. of a lack of information on gonozooids. Autozooids with moderately short, gradually MEASUREMENTS (SKELETAL): AD 119 Ϯ 12 110 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

␮m, 100±140 (3, 22), AS 445 Ϯ 68, 320± autozooids originating against substratum, 560 (3, 22). under distal end of ®rst autozooid, then di- vergent to right and left. Each second gen- GENUS ENTALOPHOROECIA HARMELIN, 1976 eration autozooid establishing a gently curved, adnate multizooidal runner, generally Entalophoroecia de¯exa (Couch, 1844) three autozooids wide. Each divergent runner Figure 52A±D potentially dichotomously dividing into two Tubulipora de¯exa Couch, 1844: 107. distal, divergently curved, multizooidal run- Pustulipora de¯exa: Johnston, 1847: 279. ners as in parent generation. At irregularly Pustolopora de¯exa: Heller, 1867: 125. distributed points on original or subsequent Entalophoroecia de¯exa: Harmelin, 1976: 108. encrusting lobes of the colony, distal budding Hayward and Ryland, 1985a: 113. Zabala, zone potentially dividing into two regions, 1986: 630. Zabala and Maluquer, 1988: 168. one remaining encrusting and the other be- Entalophora gallica d'Orbigny, 1853: 781. coming completely encircled by exterior DESCRIPTION (AMNH 1027, 1028; CMRR wall, establishing an erect portion of colony. 2275): Colonies light golden-brown, encrust- Peristomes of autozooids in encrusting por- ing multiserial runners to erect, with bifur- tions of colonies potentially much longer cating, radially symmetrical branches. than on erect branches. Longest peristomes Branch surfaces typically ®nely corrugated, belonging to autozooids in deep recesses, zooidal boundaries not clear. Autozooidal such as interior surfaces of ridges in pecten- peristomes bent at acute angles relative to the ids. distal growth direction of the branches; dis- Tentacles clear, 14; lophophores small, dis- tances between points of origin, and distanc- tally ¯aring, conical; at maximum protrusion, es between outer ends, of peristomes highly bases of lophophores located within peri- variable. stome, below level of aperture, in all material Brood chambers only at branch tips, very studied. long and incorporating bases of several au- OCCURRENCE: This is a common, wide- tozooids, though their width may not exceed spread, and highly variable species offshore three or four zooids at any given place. of Rovinj. It occurs on a variety of hard sub- Brood chambers progressively more swollen strata, from bivalve shells to Cellaria. Har- and wider nearer distal ends, near which their melin (1976) reported the species from shal- relatively narrow but elongate ooeciostomes lowest waters to depths of at least 300 m and are located. Ooeciostomes circular in cross growing on a huge number of substrata from section, variably tapered distally, except in marine grasses to rock surfaces of caves. some ending as a small ¯ared lip. DISTRIBUTION: E. de¯exa is common in the Exterior walls perforated by round pseu- eastern North Atlantic, from Arctic Norway dopores with centripetally directed spines ex- southward and in the Mediterranean Sea. tending from surrounding wall. Pseudopores MEASUREMENTS (SKELETAL): AD 130 Ϯ 11 densely distributed on the surface of brood ␮m, 110±160 (8, 80), AS 676 Ϯ 140, 400± chambers, intermediate on exterior walls of 960 (5, 50), BrD 553 Ϯ 74, 400±720 (5, 26), recumbent portions of autozooids, and least GL 2216 Ϯ 128, 2040±2400 (3, 5), GW 828 abundant on peristomes. Ϯ 125, 660±960 (3, 5), OsD 85 Ϯ 7, 80±95 Protoecia large (250 Ϯ 26 ␮m, N ϭ 3), (3, 5). (POLYPIDE): IH 0 Ϯ 0 ␮m (2, 12), MD hemispherical to domed button-shaped, with 28 Ϯ 3, 25±30 (2, 3), LD 609 Ϯ 108, 380± distal portion of ancestrula extending as a 720 (2, 14), TL 571 Ϯ 85, 280±680 (2, 13). uniform tube with smaller diameter (117 Ϯ 15 ␮m, N ϭ 3) than in asexually budded au- Entalophoroecia robusta Harmelin, 1976 tozooids. Distal-most portion of ancestrula Figure 52E±G elevated as a peristome, and ®rst asexual au- Entalophoroecia robusta Harmelin, 1976: 116. tozooid budded from its undersurface and Zabala, 1986: 636. Zabala and Maluquer, 1988: oriented either in the same direction as the 168. ancestrula or diverging obliquely to right or DESCRIPTION (AMNH 1029): Colony erect left. Second generation of asexually budded with bifurcating, radially symmetrical, rela- 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 111

Fig. 52. A±D. Entalophoroecia de¯exa. A. General aspect (AMNH 1027; 1 mm). B. Ooeciostome (center) (AMNH 1027; 0.1 mm). C. Pseudopores (AMNH 1027; 0.02 mm). D. Ancestrula (bottom center and early autozooids on Cellaria branch (AMNH 1028; 0.5 mm). E±G. Entalophoroecia robusta (AMNH 1029). E. General aspect (0.5 mm). F. Branch tip (0.5 mm). G. Pseudopores, more abundant on branch surface (left) than on (right) peristome (0.05 mm). 112 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

tively heavily calci®ed and robust branches. DESCRIPTION (AMNH 947, 1030±1032; Autozooidal boundaries clearly visible on CMRR 2276): Colonies white, multiserial, colony surface. Autozooidal peristomes long, discoidal, with basal wall extended as broad curved abruptly away from axis of branch lamina (up to 1 mm) beyond colony-margin but oriented relative to neighboring auto- zooidal buds. Colonies typically several mil- zooids, with apertures relatively uniformly limeters, occasionally over 1 cm, in diameter; spaced. adherent to the substratum or with elevated Gonozooids not restricted to branch ter- perimeter. One or more generations of sec- mini; brood chambers incorporating the ba- ondary, ``daughter'' colonies commonly bud- ses of several adjacent autozooids. Ooecio- ded peripherally on large colonies. pores circular in cross section, slightly small- Protoecial cone ¯ared to overlap protoe- er in diameter than autozooidal peristomes, cium early in astogeny, establishing circular contiguous with base of autozooidal peri- colony with autozooids radiating from colo- stome, but shorter. ny center. Autozooids budded only in contin- Exterior walls perforated by elongate oval uous budding zone around colony perimeter, pseudopores apparently lacking centripetally ®rst indicated by nearly parallel, radiating in- directed spines. Pseudopores least abundant terior-wall ridges on inner portion of peri- on peristomes. metrical basal wall lamina. Ontogenetically Primary zone of astogenetic change and older autozooids developing exterior wall in- encrusting portions of colonies not observed. cluding short to long, isolated peristomes REMARKS: The species is much less abun- aligned in poorly organized radiating rows. dant than Entalophoroecia de¯exa in our ma- Ontogenetically oldest autozooids (in colony terial but can be readily distinguished be- center) capped by single, ¯at, slightly sub- cause of its larger peristome diameters, the merged terminal diaphragms. relatively wider and shorter ooeciopore, larg- Gonozooids with broad, strongly in¯ated er branches, and perhaps also by greater reg- brood chambers, parallel with colony margin ularity in placement of peristomes relative to and brood chambers laterally encompassing one another. several rows of autozooecial peristomes. DISTRIBUTION: This species is at present Ooeciostome on peripheral margin of gono- known only from the Mediterranean Sea, zooid, short, recurved, usually slightly more identi®ed from the French and Spanish slender than autozooidal peristomes. Ooecio- coasts, but Harmelin (1976) considers that it pores may be circular or laterally elongate. is more widely spread, being confused with Multiple gonozooids possible, with brood other species of Entalophoroecia. chambers formed simultaneously, usually in MEASUREMENTS (SKELETAL): AD 186 Ϯ 13 single band near colony perimeter. ␮m, 160±200 (2, 20), AS 759 Ϯ 144, 600± Interior wall surfaces apparently lacking 1200 (2, 20), BrD 796 Ϯ 107, 600±1000 (2, spines but with small, uniformly spaced 20). bumps. Pseudopores in exterior wall partially occluded by centripetally converging spines. FAMILY PLAGIOECIIDAE CANU, 1918 Pseudopores more densely spaced on brood chamber surface than on autozooids. GENUS PLAGIOECIA CANU, 1918 Tentacles clear, 10; tentacles short (®g. Plagioecia patina (Lamarck, 1816) 54A) and lophophores small (®g. 54B), con- Figures 53A±F, 54 ical along colony margin, grading to cam- pylonemidan and obliquely truncate in on- Tubulipora patina Lamarck, 1816: 163. togenetically older autozooids; only marginal Diastopora patina: Hincks, 1880: 458. autozooids with functioning polypides. Short Berenicea patina: Marcus, 1940: 73. introverts (longest in ontogenetically oldest Plagioecia patina: Canu, 1918: 327. Harmelin, 1976: 129. Hayward and Ryland, 1985a: 97. autozooids) visible in some colonies (®g. Diastopora simplex Busk,1875: 28. 54A). Diastopora latomarginata: Waters, 1879: 272. REMARKS: Nielsen (1970) described em- Friedl, 1917: 277. bryology, settlement, and metamorphosis of Discosparsa annularis Heller, 1867: 123. P. patina, but either there is variation in ten- 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 113

Fig. 53. A±F. Plagioecia patina. A. Colony with budded daughter colony (AMNH 947; 0.5 mm). B. Lateral view of colony with budded daughter colony (AMNH 947; 0.5 mm). C. Brood chamber (top) near colony margin (AMNH 1030; 0.5 mm). D. Brood chamber and (top center) distal ooeciostome with circular ooeciopore (AMNH 1030; 0.1 mm). E. Marginal zone of functional autozooids (right) and (left) nonfunctional autozooids with broken peristomes and terminal diaphragms (AMNH 1031; 0.2 mm). F. Pseudopores (AMNH 1032; 0.01 mm). 114 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

found in a range of orientations of substra- tum and intensities of light. In the Mediter- ranean and the northern Adriatic, diverse hard substrata are occupied, particularly bi- valve shells (especially undersurfaces; Mc- Kinney, 2000), other calci®ed bryozoans, rocks from pebbles to cave surfaces and rock walls, and arti®cial substrata. It is also known to occur on rhizomes of Posidonia (Harmelin, 1976). P. patina is one of the more abundant bryozoan epibionts of Cel- laria spp. within Cellaria meadows offshore of Rovinj (McKinney and Jaklin, 2000), where it is attached to the Cellaria branches only by the central part of the colony and typically radiates from the point of attach- ment as relatively planar disk. DISTRIBUTION: P. patina is common in the northeastern Atlantic, from Arctic Norway southward, and in the Mediterranean Sea. It has been reported from many other parts of the world, but such reports are thought to be in error (Hayward and Ryland, 1985a). MEASUREMENTS (SKELETAL): AD 102 Ϯ 11 ␮m, 85±130 (4, 40), AS 207 Ϯ 42m 110± 280 (4, 40), GL 889 Ϯ 137, 600±1100 (7, 14), GS 2511 Ϯ 445, 1600±3100 (7, 14), OsDMN 85 Ϯ 15, 60±120 (7, 14), OsDMX: 94 Ϯ 17, 65±120 (7, 14). (POLYPIDE): IH 81 Ϯ 58 ␮m, 0±200 (1, 12), LDMn 345 Ϯ 45, Fig. 54. Plagioecia patina. A. Ontogenetic in- 260±400 (1, 11), LDMx 407 Ϯ 47, 320±500 crease in tentacle and introvert length: ®lled cir- (2, 30), MD 15 (1, 1), TLMn 289 Ϯ 50, 200± cles plot maximum tentacle length within lopho- 340 (1, 11), TLMx 355 Ϯ 77, 210±560 (2, phores (Y ϭ 198.490 ϩ 0.975X) and open circles plot lengths of exposed portions of introverts (Y 33). ϭ 2.685 ϩ 0.440X). B. Ontogenetic increase in maximum lophophore diameter (Y ϭ 268.652 ϩ Plagioecia sarniensis (Norman, 1864) 0.630X). Figure 55A±E Diastopora sarniensis Norman, 1864: 89. Hincks, 1887: 308. tacle number between the Adriatic and the Berenicea sarniensis:? Harmer, 1915: 114. Canu North Sea (Nielsen reported 8 tentacles; and Bassler, 1928: 65. specimens studied here have 10), or two dif- Microecia sarniensis: Canu, 1918: 326. ferent species were studied by Nielsen and Plagioecia sarniensis:? Canu and Bassler, 1925: by us. Embryos were reported June±August 65. Harmelin, 1976: 136. Hayward and Ryland, for specimens from the Isle of Man (Eggle- 1985a: 100. ston, 1969), and embryos were seen in our DESCRIPTION (AMNH 1033): Colony specimens during June. Large, fertile colo- white, completely encrusting, including mar- nies commonly have abundant small colonies ginal lamina, multiserial; irregularly discoi- within 2 cm of their margins, apparently off- dal with rejuvenated areas extending as spring that settled near the maternal colony. lobes. Ancestrula and early astogenetic stage OCCURRENCE: Plagioecia patina ranges be- obscured by multiple small intracolony over- tween 8 m and at least 75 m depth in the growths. Mediterranean (Harmelin, 1976), where it is Autozooids budded only in continuous 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 115

Fig. 55. A±E. Plagioecia sarniensis (AMNH 1033). A. General aspect (0.5 mm). B. Brood chamber (0.2 mm). C. Broken ooeciopore atop short ooeciostome (0.1 mm). D. Pseudopore (0.01 mm). E. Broken autozooidal peristomes transformed into secondary nanozooids by funnel terminal diaphragm (0.1 mm). F±H. Eurystrotos compacta (AMNH 931). F. General aspect (0.5 mm). G. Brood chamber (center) surrounded by peristomes of autozooids (0.2 mm). H. Distal end of ancestrula (small oblique tube, bottom center) and early autozooids (0.2 mm). 116 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 budding zone around colony perimeter. On- MEASUREMENTS (SKELETAL): ADMN 98 Ϯ togenetically older autozooids developing 7 ␮m, 80±110 (1, 10), ADMX 119 Ϯ 3, exterior wall including long, generally iso- 110±120 (1, 10), AS 244 Ϯ 36, 180±300 (1, lated peristomes aligned in poorly organized 10), GL 520 Ϯ 28, 500±540 (1, 2), GW 1215 radiating rows, locally in short connate se- Ϯ 244, 1000±1560 (1, 4), OsDMN 60±70 (1, ries. Peristomes commonly slightly tapered 2), OsDMX 80 (1, 2). toward aperture. Ontogenetically oldest au- tozooids (in colony center) capped by single, GENUS EURYSTROTOS HAYWARD AND slightly submerged terminal diaphragms per- RYLAND, 1985b forated centrally or slightly distally by small Eurystrotos compacta (Norman, 1867) pore at top of inverted, funnel-shaped exten- Figure 55F±H sion. Locally, rim of aperture extended as Alecto compacta Norman, 1867: 204. slight peak at end of small keel on distal Eurystrotos compacta: Hayward and Ryland, edge. 1985a: 94, 1985b: 1075. Zabala and Maluquer, Rare nanozooids budded against basal 1988: 171. lamina at growing edge of colony; peri- Diastopora suborbicularis Hincks, 1880: 464. stomes as short as funnel-shaped extensions Berenicea suborbicularis: Marcus, 1940: 74. of autozooidal terminal diaphragms, with di- Microecia suborbicularis: Harmelin, 1976: 122. ameter approximately one-fourth that of au- Zabala, 1986: 644. tozooids. DESCRIPTION (AMNH 931): Colonies en- Gonozooids with broad, strongly in¯ated crusting, multiserial, thin, up to 5.0 ϫ 4.6 brood chambers, parallel with colony margin mm, with irregular outline; consisting of arc- and brood chambers laterally encompassing shaped, laterally coalescent subcolonies orig- several rows of autozooidal peristomes. Ooe- inating by continued normal growth from a ciostome located near peripheral margin of small group of ancestral zooids along edge gonozooid; short, recurved, more slender of an older subcolony in which growth has than autozooidal peristomes. Ooeciopore ceased along most of arc-shaped perimeter. may be capped by terminal diaphragm form- Autozooids small, budded in quincuncial se- ing funnel, constricting diameter to one-half, ries, with clearly visible adnate proximal por- then broadly ¯ared along outer edge. tions and distal peristomes extending well Pseudopores in nonperistomial exterior above the general colony surface. (Peri- wall partially occluded by centripetally con- stomes of the available specimens are cor- verging spines. Pseudopores more densely roded, so length cannot be determined.) Col- spaced on brood chamber surface than on au- ony surface corrugated by ®ne growth lines, tozooids. Pseudopores in peristomes smaller, especially visible on peristomes. Surface of apparently without partial occlusion by con- colony abundantly perforated by circular verging spines. pseudopores, less abundant on the peristo- REMARKS: The centrally perforated termi- mial skeleton but particularly dense on brood nal diaphragms atop peristomes provide chamber surfaces. egress for a single, nonciliated tentacle of a Gonozooids abundant, with 17 fully secondary nanozooid for which no function formed brood chambers present in the larger is obvious (SileÂn and Harmelin, 1974). They specimen. Brood chambers located near the lack testes that would indicate male poly- outer edges of subcolonies, averaging just morphs, and their position and behavior do over one zooid row distance from outer edg- not indicate cleaning similar to nanozooids es (250 Ϯ 89 ␮m, N ϭ 16). Brood chamber of Diplosolen obelium. Widely scattered pri- width about twice chamber length, extending mary nanozooids in P. sarniensis (and in P. as short lobes and without engul®ng peri- patina) have been reported previously by Sil- stomes of adjacent autozooids. Ooeciostomes eÂn and Harmelin (1974). short, recurved, and are located on the distal DISTRIBUTION: The species is known in the edge of brood chambers, distinctly smaller eastern Atlantic from southern Britain to An- than autozooidal peristomes; varying from gola and as far east in the Mediterranean as circular to oval with maximum diameter the Aegean Sea. transverse. 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 117

Ancestrula small, with peristomial diame- tiserial, usually adnate with peripheral basal ter equal to diameter of ooeciostomes. A sin- lamina less than 1 mm wide. Small and most gle autozooid budded from ancestrula, suc- large colonies circular, some large colonies ceeded by a median distal autozooid and two with lobes; typically a few millimeters in di- laterally curved distolateral zooids, initiating ameter, largest colonies slightly over 10 mm. a budding zone that widens and recurves rap- Where growing on bryozoan substrata, grow- idly. The lateral ends of the marginal bud- ing edge may grow free and arch up into ding zone meet and coalesce at approximate- hemispherical blisters over zooidal ori®ces of ly the ®fth to sixth asexually budded gener- underlying substratum. ation, generating a continuous perimetrical Autozooids and nanozooids regularly al- budding zone, and leaving a small oval gap ternating in both longitudinal and lateral on either side of primary astogenetic zone of rows, distributed in highly ordered quincun- change. cial series. Autozooidal diameter small, prox- REMARKS: Only two specimens of this spe- imal portion adnate, distal portion as free cies were found, neither of which was alive peristome with height coordinated with that when observed in the laboratory. They have of neighbors; peristomes very long where many features in common with shallow-wa- colony or local portion of colony located in ter colonies of Eurystrotos compacta (Har- depression. melin, 1976; Hayward and Ryland, 1985a), Autozooidal tentacles colorless, 10, short- differing in having smaller brood chambers est at colony margins (®g. 57A); lophophores than normal for the species and that differ campylonemidan, grading from smallest from British specimens in being transversely along colony margin to largest away from elongate rather than longitudinally elongate. colony margin (®g. 57B) within peripheral They possibly belong to an undescribed spe- band of functioning autozooids; peristomes cies of Eurystrotos. of autozooids in colony centers broken off OCCURRENCE: On glass bottle, 25±30 m and autozooids closed by terminal dia- deep, west slope of Banjole. phragm. DISTRIBUTION: E. compacta is known in the Nanozooids regularly budded against basal eastern North Atlantic from Denmark to lamina, in alternation with autozooids, at southern Britain, the Mediterranean Sea, and growing edge of colony. Peristomes shorter the Adriatic Sea. than, and diameter approximately one-fourth MEASUREMENT (SKELETAL): ADMN 60 Ϯ 4 that of, autozooids. Nanozooids with one ␮m, 55±65 (1, 10), ADMX 65 Ϯ 7, 60±80 thin, colorless tentacle (devoid of cilia; Borg, (1, 10), AS 213 Ϯ 18, 180±240 (1, 10), GL 1926) approximately half the length of au- 266 Ϯ 29, 220±320 (1, 16), GW 476 Ϯ 58, tozooidal tentacles. 400±560 (1, 16), OsDMN 37 Ϯ 3, 35±40 (1, Gonozooids with nearly equidimensional, 11), OsDMX 41 Ϯ 3, 35±45 (1, 11). in¯ated brood chambers, about same size as but less in¯ated than blisters where colony GENUS DIPLOSOLEN CANU, 1918 arches over zooidal ori®ces where growing on Pentapora fascialis. Brood chamber in- Diplosolen obelium (Johnston, 1838) corporating base of several peristomes of ad- Figures 56A±D, 57 jacent autozooids, with single curved, short Tubulipora obelia Johnston, 1838: 269. ooeciostome about two-thirds the diameter of Diastopora obelia: Johnston, 1847: 277. Heller, autozooidal peristomes and located in mid- 1867: 123. Waters, 1879: 273. region of brood chamber. Diplopora obelia: Jullien and Calvet, 1903: 116, Exterior walls about 50 ␮m thick, perfo- 162. rated by circular to elongate oval pseudo- Diplosolen obelia: Canu and Bassler, 1920: 745. pores. Brood chamber wall more densely Hayward and Ryland, 1985a: 102. perforated than exterior wall of other zooids; Diplosolen obelium: Harmelin, 1969a: 1183, 1976: 145. Zabala, 1986: 641. pseudopores fewer higher on peristomes. REMARKS: The resting position of the sin- DESCRIPTION (AMNH 1034, 1035; CMRR gle tentacles of the reduced nanozooid polyp- 2277): Colonies white, encrusting, thin, mul- ides is horizontal with the colony surface, ex- 118 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 56. A±D. Diplosolen obelium. A. General aspect (AMNH 1034; 0.5 mm). B. Brood chamber with ooeciostome (near center) intermediate in size between larger autozooidal peristomes and smaller nanozooid peristomes (AMNH 1034; 0.1 mm). C. Lateral view of growing edge of colony (AMNH 1034; 0.5 mm). D. Ancestrula and early autozooids followed by alternating autozooids and nanozooids (AMNH 1035; 0.5 mm). E±G. Frondipora verrucosa. E. General aspect of frontal surface (AMNH 1036; 0.5 mm). F. Reverse surface of branch (AMNH 1037; 0.5 mm). G. Brood chamber (AMNH 1036; 0.2 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 119

position (SileÁn and Harmelin, 1974), inter- preted as a surface-cleaning behavior. OCCURRENCE: Colonies grow on a variety of hard substrata, especially bivalve shells and the erect bilaminate bryozoan Pentapora fascialis. On disarticulated shells, they occur more commonly on interior surfaces but are larger and more commonly fertile on exterior surfaces (McKinney, 2000). DISTRIBUTION: Found at inner shelf depths throughout the northeastern Atlantic from the Barents Sea to northwestern Africa and throughout the Mediterranean Sea. MEASUREMENTS (SKELETAL): PD 82 Ϯ 8 ␮m, 70±92 (3, 30), AS 277 Ϯ 41, 220±400 (3, 30), NAD 14 Ϯ 4, 10±24 (3, 21), GL 1172 Ϯ 22, 980±1560 (3, 6), GW 1192 Ϯ 27, 960±1320 (3, 6), OsD 39 Ϯ 2, 35±40 (3, 5). (POLYPIDE): IH 0 Ϯ 0 ␮m (2, 29), LD 372 Ϯ 66, 240±580 (2, 29), MD 25±30 (2, 2), TLMn 277 Ϯ 51, 140±240 (2, 14), TLMx 296 Ϯ 104, 140±620 (2, 28).

FAMILY FRONDIPORIDAE BUSK, 1859 GENUS FRONDIPORA LINK, 1807 Frondipora verrucosa (Lamouroux, 1821) Figures 56E±G, 58 MadreÂpore rameux, Marsigli, 1725: 105. Krusensterna verrucosa Lamouroux, 1821: 41. Frondipora verrucosa: Busk, 1875: 39. Waters, Fig. 57. Diplosolen obelium. A. Ontogenetic 1879: 279. Hincks, 1887: 308. Zabala, 1986: increase in maximum tentacle length (Y ϭ 654. Zabala and Maluquer, 1988: 173. 213.782 ϩ 0.135X); open circles plot maximum ?Frondipora reticulata: de Blainville, 1834: 406. tentacle lengths of anomalous lophophores adjoin- Frondipora maderensis Johnson, 1897: 64. ing excurrent chimney where colony folded sharp- Frondipora gracilis Canu and Bassler, 1930a: 87. ly over edge of shell substratum; longest tentacles Harmelin, 1969a: 1187. located on sides of lophophores adjacent to chim- ney. B. Ontogenetic increase in lophophore di- DESCRIPTION (AMNH 1036, 1037; CMRR ameter (Y ϭ 300.205 ϩ 0.118X); open circles rep- 2278): Colonies erect, highly branched, with resent the two anomalous lophophores described local lobes and branch ends occupied by fas- for A. Regression lines exclude the lophophores cicles of interior-walled autozooids surround- adjoining substratum de®ned excurrent chimney. ed by extensive exterior wall areas interrupt- ed locally by isolated or small groups of au- tozooids; usually canary yellow, some or- ange-tinted yellow. Branch widths variable, tending from the nanozooid ori®ce toward generally 1±2 mm, separated by spaces ap- the colony center, as if oriented by the co- proximately equal in width; fascicles prefer- lonial feeding current, which is centripetal entially oriented on individual branches, pro- and thin, adjacent to the colony surface (Mc- ducing a poorly to well de®ned frontal sur- Kinney, 1992). Intermittently, the tentacle face. Branches may anastomose locally. Col- will either rotate rapidly forward and back to onies increasing in width more rapidly than the normal position, or it will make a lateral, in height so that larger colonies up to several circular sweep that ends back at the normal centimeters across but usually extending no 120 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

surface. Single ooeciostome per brood cham- ber oval, slightly larger than autozooecial or- i®ces, and atop a short, slightly ¯ared tube. Tentacles clear or white except for irreg- ularly distributed brown segments, 11±12, longest and constituting largest lophophores along exterior-wall margin of fascicle (®g. 58A, B); lophophores campylonemidan, most strongly obliquely truncate along exte- rior-wall margin of fascicle. OCCURRENCE: In the vicinity of Rovinj, this species occurs on diverse bivalves and occasionally on other calci®ed bryozoans such as Cellaria and Pentapora. It was very rare during the late 1980s but had become moderately common at depths over 35 m, es- pecially in the vicinity of the islands Pelago and Sv. Ivan, by 1997 and 1998. DISTRIBUTION: Common in the Mediterra- nean, less commonly reported from the east- ern North Atlantic. MEASUREMENTS (SKELETAL): ADMN 170 Ϯ 21 ␮m, 120±200 (4, 40), ADMX 194 Ϯ 19, 160±240 (4, 40), GL 1740 Ϯ 108, 1600± 1850 (1, 5), GW 1600 Ϯ 162, 1450±1800 (1, 5), OsDMN 176 Ϯ 17, 150±200 (1, 5), OsDMX 199 Ϯ 20, 180±220 (1, 5). (POLYP- IDE): IH 0 Ϯ 0 ␮m (2, 32), LDMn 433 Ϯ 73, 300±560 (2, 10), LDMx 503 Ϯ 94, 300±660 (2, 27), MD 35 (1, 1), TLMn 387 Ϯ 82, 240± 500 (2, 14), TLMx 596 Ϯ 151, 390±860 (2, Fig. 58. Frondipora verrucosa. A. Increase in 20). maximum tentacle length toward edge of fascicles Ϫ0.123 (Y ϭ 1076.626X ). B. Increase in maximum SUBORDER ARTICULATA BUSK, 1859 lophophore diameter toward edge of fascicles (Y ϭ 585.039XϪ0.048. FAMILY CRISIIDAE JOHNSTON, 1847 GENUS CRISIA LAMOUROUX, 1812 more than 2±3 cm from substratum. Auto- Crisia ®stulosa Heller, 1867 zooids closely packed within fascicles, elon- Figure 59A, B gate tubular parallel with branch axis, more abundantly originating in interior regions of Crisia ®stulosa Heller, 1867: 118. Waters, 1879: 268. Friedl, 1917: 275. Harmelin, 1968: 427. fascicles than along the exterior-walled pe- rimeter. Each fascicle a mosaic of small, DESCRIPTION (AMNH 1038): Colonies newly budded autozooids and variably elon- bushy, branches robust, articulated, biserial, gate autozooids with fully developed diam- ¯at, white. Colony fragments at least 4.5 mm eters. Some autozooid diameters very slight- high, generally highly branched. Branch in- ly larger along the perimeter of fascicles than ternodes locally giving rise to at least one within the centers. lateral branch as well as distally-formed de- Gonozooids located between fascicles, scendant internode. Internodes ¯at, long, brood chambers spreading broadly across the straight to gently curved, composed of three colony surface, commonly enveloping sev- to six autozooids in available material. Lat- eral isolated autozooids with short peri- eral branches almost always at third zooid. stomes projecting above the brood chamber Zooids long, particularly the second in each 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 121

Fig. 59. A, B. Crisia ®stulosa (AMNH 1038). A. Branch segment (0.5 mm). B. Peristome with annular constrictions (0.1 mm). C±F. Crisia ramosa.C. Portion of fertile branch segment (AMNH 1039; 0.5 mm). D. Kenozooids at colony base (AMNH 1040; 0.5 mm). E. Pseudopores of brood chamber (AMNH 1039; 0.02 mm). F. Brood chamber with distal ooeciostome ¯ared to large circular ooeciostome (AMNH 1039; 0.1 mm). G±J. Crisia recurva. G. Fertile branch segment (AMNH 1041; 0.5 mm). H. Entire colony attached to Cellaria branch (AMNH 1042; 0.5 mm). I. Brood chamber with terminal ooeciostome and broken ooeciopore (lectotype, UIIZ 315; 0.062 mm). J. Base of attachment with radiating kenozooids (AMNH 1042; 0.1 mm). 122 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 internode, with ori®ces widely spaced along proximately centered at distal end of brood each side of branch. Circular autozooidal or- chamber. Each gonozooid generally situated i®ces alternating from side-to-side of branch- near middle of fertile internode, distal to ®rst es, at end of short to extended peristomes and commonly beyond second lateral branch, only slightly curved toward frontal sides of as previously noted by Ryland (2000). Fertile branches. colonies collected from June to November Gonozooids absent in present material, but (i.e., during all times of year when collec- reported and illustrated by Harmelin (1968) tions were made). as elongate cylindrical, slightly expanded Adnate attachment zooids (``rhizoids'') proximally, with a distally placed, small ooe- abundant, giving rise to multiple erect branch ciopore on a short ooeciostome situated just systems. Some adnate zooids with terminal distal to a small area lacking pseudopores. ori®ces giving rise to erect branch systems, DISTRIBUTION: First described from the while others are end-to-end kenozooids, as Adriatic Sea, this species has been reported described by SileÂn (1977) for Crisia cornuta. in relatively small numbers from the Adriatic Tentacles clear, 8; lophophores conical to and western Mediterranean, with a single re- ¯aring conical, radially symmetrical. Base of port (Aguirrezabalaga et al., cited in Alvarez, lophophore generally at ori®ce or within 1994a) from the Atlantic coast of Spain. peristome, but unusually may be up to 0.04 MEASUREMENTS (SKELETAL): AD 90 Ϯ 9 mm above the ori®ce. ␮m, 72±105 (1, 19), AS 615 Ϯ 108, 403± OCCURRENCE: The species occurs in the 816 (1, 23), ASW 812 Ϯ 135, 616±1210 (1, northern Adriatic on bryozoans (especially 17), BrD 274 Ϯ 42, 206±330 (1, 8), ZL (2nd Cellaria and Pentapora), shells, skeletalized in fascicle) 1230 Ϯ 120, 1030±1390 (1, 9). and ¯eshy algae, Microcosmus, and ®rm hu- man ejecta such as plastic and glass. C. ra- Crisia ramosa Harmer, 1891 mosa was probably referred to by Heller Figure 59C±F (1867: 118) as Crisia eburnea (Linnaeus), the only one of the several species of Crisia Crisia ramosa Harmer, 1891: 134. Friedl, 1917: that he noted to be abundant in the Adriatic. 275. Marcus, 1940: 45. Harmelin, 1968: 419. DISTRIBUTION: C. ramosa ranges through- Hayward and Ryland, 1985a: 52. Alvarez, out the Mediterranean and in the Atlantic 1994a: 41. ? Crisia eburnea: Heller, 1867: 118. Friedl, 1917: from northeast England to the Azores and 275. Cape Verde Islands. MEASUREMENTS (SKELETAL): AD 69 Ϯ 9 DESCRIPTION (AMNH 1039, 1040; CMRR ␮m, 60±80 (5, 52), AS 354 Ϯ 38, 260±420 2279): Colonies bushy, branches articulated, (3, 30), ASW 412 Ϯ 63, 300±500 (3, 30), biserial, ¯at, white. Colonies up to 2 cm BrD 256 Ϯ 37, 180±340 (4, 34), GL 655 Ϯ high, generally highly branched, with branch 59, 580±740 (3, 6), GW 507 Ϯ 59, 460±600 internodes commonly giving rise to one or (3, 6), OsD 111 Ϯ 14, 85±120 (3, 6). (POL- more lateral branches as well as distally- YPIDE): IH 1 Ϯ 7 ␮m, 0±40 (4, 35), LD 349 formed descendant internode. Internodes ¯at, Ϯ 51, 240±460 (4, 34), MD 22 Ϯ 5, 20±30 long, straight to gently curved, usually com- (3, 4), TL 281 Ϯ 54, 160±420 (4, 30). posed of 10 to 20 autozooids, except smaller number associated with rhizoids in basal seg- Crisia recurva Heller, 1867 ments. Lateral branches almost always at Figure 59F±J third zooid or higher, with second lateral Crisia recurva Heller, 1867: 118. branch, if present, about 5 zooids farther along. Circular autozooidal ori®ces alternat- LECTOTYPE (chosen here): UIIZ 315. ing from side-to-side of branches, at end of DESCRIPTION (AMNH 1041, 1042): Small short peristomes gently curved toward fron- bushy colonies of biserial, articulated, ¯at tal sides of branches. branches, white, less than 1 cm high. Colo- Gonozooids large, with distal in¯ated nies generally highly branched, with branch brood chambers. Ooeciopores circular, at end internodes commonly giving rise to one or of frontally directed, ¯ared ooeciostome ap- more lateral branches and, in some, a distally 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 123 formed internode. Internodes ¯at, short, Lichenopora hispida: Hincks, 1880: 473. straight to gently curved, usually of 15 to 20 Disporella hispida: Borg, 1944: 230. Hayward autozooids, except smaller number associat- and Ryland, 1985a: 128. Alvarez, 1992: 204; ed with attachment zooids in basal segments. 1994c: 493. Lateral branches almost always arising at Lichenopora mamillata Lagaaij, 1952: 181. third zooid or higher, with second lateral DESCRIPTION (AMNH 1043±1049; CMRR branch if present originating about 3 zooids 2280±2282): Colonies yellow or white, mul- farther along. Circular autozooidal ori®ces tiserial, ¯at discoidal to low moundlike, com- alternating from side-to-side of branches, at monly with slightly depressed center in in- end of long strongly frontally curved peri- fertile colonies, to at least 5.6 mm diameter; stome. usually completely adnate with peripherally Gonozooids large, with conspicuously in- extended basal wall. Margin locally elevated ¯ated brood chambers. Brood chambers typ- where nearing contact with other encrusting ically with a short, broad, median keel along organisms. Compound colonies reported distal frontal surface and circular ooeciopore elsewhere (Hayward and Ryland, 1985a) not at end of frontally directed, slightly ¯ared observed. ooeciostome approximately centered at distal Autozooids budded only in continuous end of brood chamber. Each gonozooid gen- budding zone around colony perimeter. On- erally near middle of fertile internode, at a togenetically older autozooids generally with point at which a lateral branch is formed. isolated apertures aligned in single radiating Fertile colonies collected in June, July, and rows, locally in short connate series. Younger November. zooids around perimeter of larger colonies REMARKS: Heller's specimen chosen as generally in quincuncial pattern. Outer ends lectotype is illustrated here as ®g. 59I. This of ontogenetically oldest autozooids extend- species has not been recorded since being ed as peristomes, some with granules (more named by Heller from sparse material found common on yellow colonies) or tiny spines at Lesina, Croatia. The species is distinctive, (seen only on some white colonies) on upper easily recognized by the tiny branches, small surface. Small, blunt spines line the auto- zooids, strongly curved peristomes, and zooidal chambers. Peristomes variable in abundant and uniquely shaped brood cham- length, generally short, diminishing in length bers. Colonies of C. recurva commonly are toward perimeter, absent in youngest auto- attached to basal segments of Cellaria within zooids. Rim of peristomial apertures gener- the meadow west of Banjole (McKinney and ally extended as paired spines on upper side, Jaklin, 2000), the only locality in which we commonly with additional spines. Autozooi- have found them. dal apertures circular to oval, with long axis DISTRIBUTION: Known only from the north- parallel with colony growth direction. eastern Adriatic Sea. Autozooids separated by polygonal alveoli MEASUREMENTS (SKELETAL): AD 57 Ϯ 7 of equal to slightly smaller diameter. Onto- ␮m, 45±70 (3, 39), AS 303 Ϯ 44, 230±400 genetically older alveoli progressively ®lled (3, 34), ASW 257 Ϯ 31m 200±312 (3, 31), centripetally, some closed entirely. BrD 151 Ϯ 21, 120±187 (3, 17), GL 351 Ϯ Brood chambers in central depressed area 24, 310±380 (3, 15), GW 292 Ϯ 39, 220± of colony, with domed roof of interior wall, 360 (3, 15), OsD 68 Ϯ 14, 40±80 (3, 13). extending as short lobes between proximal portions of inner portions of autozooidal SUBORDER RECTANGULATA WATERS, 1887 rows. Ooeciostomes short, cylindrical, situ- FAMILY LICHENOPORIDAE SMITT, 1866 (1867) ated near perimeter of brood chambers; ooe- GENUS DISPORELLA GRAY, 1848 ciopores nearly circular, about 100 ␮m in di- ameter. Disporella hispida (Fleming, 1828) Tentacles clear, 9±10; lophophores Figures 60A±J, 61, 62A, B obliquely truncate, broadly conical along col- Discopora hispida Fleming, 1828: 530. ony margin to campylonemidan in ontoge- Tubulipora hispida: Johnston, 1847: 268. netically older autozooids, increasing in size Discoporella hispida: Busk, 1875: 30. away from colony margin, youngest auto- 124 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 60. A±J. Disporella hispida. A±D. Yellow variety. A. General aspect (AMNH 1043; 0.5 mm). B. Autozooids (ori®ces at top) largely buried below alveoli (AMNH 1044; 0.2 mm). C. Spines within autozooid closed by inset diaphragm (AMNH 1043; 0.02 mm). D. Growing edge of colony, with minimal ori®cial spines (AMNH 1043; 0.1 mm). E±J. white variety. E. Colony with incompletely formed central brood chamber (AMNH 1046; 0.5 mm). F. Spinose ori®ce of autozooid, and alveoli (AMNH 1046; 0.1 mm). G. Spines within autozooid (AMNH 1047; 0.01 mm). H. Growing edge of colony with spinose autozooidal ori®ces (AMNH 1048; 0.1 mm). I. Fertile colony with completely formed brood chamber and ooeciostome (AMNH 1047; 0.5 mm). J. Ooeciostome (AMNH 1047; 0.05 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 125

Fig. 61. Disporella hispida. A, B. Yellow colonies with 9 tentacles. A. Ontogenetic increase in tentacle and exposed introvert length: Filled circles plot maximum tentacle length within lophophores (Y ϭ 106.231 ϩ 0.250X), open circles plot minimum tentacle lengths (Y ϭ 106.340 ϩ 0.159X), and diamonds plot exposed introvert length (Y ϭϪ27.526 ϩ 0.090X). B. Ontogenetic increase in lophophore diameter (Y ϭ 189.359 ϩ 0.110X). C, D. White colonies with 10 tentacles. C. Ontogenetic increase in tentacle and exposed introvert length: Filled circles plot maximum tentacle length within lophophores (Y ϭ 156.909 ϩ 0.514X), open circles plot minimum tentacle lengths (Y ϭ 172.051 ϩ 0.226X), and diamonds plot exposed introvert length (Y ϭϪ35.225 ϩ 0.166X). D. Ontogenetic increase in lophophore diameter (Y ϭ 192.398 ϩ 0.330X). zooids equitentacled but progressively more also addressed the taxonomic confusion in inequitentacled with age; ontogenetically the genus. Although Alvarez (1992: 209) older autozooids commonly with lopho- chose a neotype for D. hispida, Gordon and phores elevated on introverts. Taylor (2001) point out that there are two REMARKS: Several species names have problems with the neotype: it is not topotyp- been proposed for specimens of Disporella ic, and its ori®ces are neither ``expanded'' from the North Atlantic and Mediterranean, nor are they hispid, which are attributes orig- yet the differentiating characteristics of the inally indicated for the species by Fleming proposed species are not clear. Alvarez (1828). (1992) attempted to sort out the species of It is not clear whether one or two species Disporella from the region, naming four new are included here in ``Disporella hispida''. species, and Gordon and Taylor (2001) have Four of the species described by Alvarez 126 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270

Fig. 62. A, B. Disporella hispida, yellow variety. A. Ancestrula and early autozooids (AMNH 1048; 0.1 mm). B. Young colony (AMNH 1049; 0.5 mm). C±I. Patinella radiata. C. Fertile colony (AMNH 1050; 0.5 mm). D. Short ooeciostome with circular ooeciopore (AMNH 1050; 0.1 mm). E. Radiating fascicles of autozooids separated by alveoli, and growing edge of colony (AMNH 1051; 0.1 mm). F. Stellate-headed spines within autozooids (AMNH 1051; 0.02 mm). G. Broken ancestrula (AMNH 1052; 0.1 mm). H. Ancestrula and ®rst autozooids of young colony (specimen destroyed; 0.05 mm). I. An- cestrula and ®rst zooids of young colony in early stages of re¯ection of basal lamina back over ances- trula, establishing radial growth (AMNH 1048; 0.2 mm). 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 127

dition, specimens from the Arctic that are identi®ed as D. hispida by Shunatova and Ostrovsky (2001) have only 8 tentacles per lophophore, and none of the specimens have been seen with exposed introverts. More ex- tensive documentation is needed in order to determine whether the various polypide mor- phologies of ``D. hispida'' are due to species differences or are ecophenotypes. However, both types of colonies are illustrated here. Fig. 58A±D portray yellow colonies; ®g. 58E±J and 60A, B portray white colonies. Brood chambers were found in some of the smallest colonies and were absent in the largest colonies, and where found, the colo- nies were fending off encroaching competi- tors for substratum space. This brood cham- ber occurrence pattern suggests that they are initiated when lethal environmental cues are encountered, previously suggested as com- mon in encrusting cyclostomes (McKinney and Taylor, 1997). DISTRIBUTION:``D. hispida'' occurs throughout the Mediterranean Sea, the Ba- rents Sea, and on both shores of the northern Atlantic Ocean. MEASUREMENTS (SKELETAL): ADMN 78 Ϯ 10 ␮m, 60±100 (8, 100), ADMX 102 Ϯ 12, 70±130 (7, 90), AS 211 Ϯ 31, 140±300 (7, 90), AlvD 97 Ϯ 18, 60±190 (7, 90), Gap 434 Ϯ 62, 350±540 (1, 10). (POLYPIDE): IN 41 Ϯ Fig. 63. Patinella radiata. A. Ontogenetic in- 60 ␮m, 0±220 (3, 49), LD 297 Ϯ 56, 160± crease in tentacle and introvert length: Filled cir- 440 (3, 46), MD 25 Ϯ 5, 20±30 (2, 3), TLMn cles plot maximum tentacle length within lopho- phores (Y ϭ 182.581 ϩ 0.300X) and open circles 292 Ϯ 79, 120±420 (3, 31), TLMx 402 Ϯ plot lengths of exposed portions of introverts (Y 132, 140±600 (3, 32). ϭϪ3.267 ϩ 0.036X). B. Ontogenetic increase in maximum lophophore diameter (Y ϭ 201.918 ϩ GENUS PATINELLA GRAY, 1848 0.26X). Patinella radiata (Audouin, 1826) Figures 62C±I, 63 (1992) are hispid, i.e. have spines projecting Melobesia radiata Audouin, 1826: 235. from the skeletal aperture [D. hispida (Flem- Discoporella radiata: Busk, 1875: 32. Waters, ing); D. robusta Alvarez, 1992; D. zurigneae 1879: 276. Alvarez, 1992; D. alboranensis Alvarez, Lichenopora radiata: Hincks, 1880: 476. Borg, 1992]. Skeletally, the specimens from the 1944: 222. Hayward and Ryland, 1985a: 124. Alvarez, 1990: 28, 1993: 262, 1994c: 500. Rovinj area ®t only D. hispida among the six species of the genus that Alvarez (1992) de- DESCRIPTION (AMNH 1048, 1050±1052; scribed and illustrated (despite his choice of CMRR 2283): Colonies encrusting, multi- neotype, which was not illustrated). How- serial, discoidal, small (up to 5.2 mm diam- ever, living colonies in our material fall into eter in present material), with perimeter of two discrete groups: yellow colonies with calci®ed basal wall extending as a sheet be- lophophores of 9 tentacles, and white colo- yond zooidal budding zone. Colonies basi- nies with lophophores of 10 tentacles. In ad- cally white, some with purple granules below 128 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 270 outer cuticle, giving light, speckled violet ap- 963). Gordon and Taylor (1997) revised the pearance to colonies viewed without magni- concept of the genus, restricting it to species ®cation. Autozooids budded against basal which are either turbinate as is the type spe- wall in continuous zone around colony pe- cies, or have a short central pedunculate at- rimeter, rapidly diverging from basal wall at tachment with continued disk-like expansion roughly 30Њ angle, commonly increased fur- above the substratum. Both turbinate and pe- ther away from basal wall. Zooids budded in dunculate forms have large expanses of basal linear series, generating single-zooid rows wall beyond the central point of attachment separated by alveoli, many of which also of the colony. The most recent known spe- originate against the basal wall in the bud- cies of the Lichenopora, as rede®ned by Gor- ding zone. Outer ends of zooids extended as don and Taylor (1997), is Miocene. As sug- peristomes, longer in ontogenetically older gested by Gordon and Taylor (1997), the ge- zooids, organized in connate fascicles. Small, neric name Patinella Gray, 1848, with type blunt spines with stellate heads abundant in species Madrepora verrucaria Linnaeus, is autozooidal chambers. In larger colonies, available for many living species formerly peristomes of younger zooids near colony placed into Lichenopora. Patinella and Dis- perimeter possibly isolated rather than con- porella are here treated as distinct genera, tinuing the connate fascicles. Skeletal aper- based on SchaÈfer's (1991) indication that the tures of zooids slightly elongate parallel with type species of the two genera (Madrepora growth direction and extended into acute verrucaria and Disporella hispida Gray, projection on distal side. 1848) differ in skeletal microstructure and in Brood chambers located in central depres- shape and perforation of the brood chambers. sion of colony that results from the radial and DISTRIBUTION: The species occurs through- obliquely upward growth of autozooids. out the Mediterranean Sea, and ranges in the Brood chambers developing a convex roof Atlantic Ocean from the Mediterranean covered by alveoli, with ooeciostome located northwards to southern Britain; reports from centrally to peripherally within area de®ned more distant regions are suspect (Hayward by inner ends of autozooidal fascicles. Ooe- and Ryland, 1985a). ciopores about 140 ␮m in diameter, circular MEASUREMENTS (SKELETAL): ADMN 83 Ϯ to slightly oval; ooeciostomes short, and 11 ␮m, 60±120 (5, 50), ADMX 110 Ϯ 13, gently ¯ared. 80±140 (5, 50), Gap 215 Ϯ 51, 100±340 (5, Protoecia exterior-walled, low hemispher- 46), GL 140±160 (2, 2), GW 120±160 (2, 2). ical,160±180 ␮m diameter, with low margin- (POLYPIDE): IH 15 Ϯ 25 ␮m, 0±100 (3, 42), al rim. Ancestrula and initial asexually bud- LD 324 Ϯ 86, 100±450 (3, 41), MD 20 (1, ded zooecia grow as an encrusting protoecial 1), TLMn 193 Ϯ 62, 60±320 (2, 26), TLMx cone bounded by exterior wall, upper edge 340 Ϯ 133, 60±720 (3, 42). of which at some point bending back on it- self, establishing radial growth of colony. ACKNOWLEDGMENTS Tentacles clear, 8, shortest at colony mar- gin (®g. 63A); lophophores campylonemi- We thank the National Geographic Society dan, strongly obliquely truncate, grading and the US National Academy of Sciences± from small and equitentacled along colony Yugoslavia Scienti®c Exchange Program for edge to largest and most inequitentacled ad- support of ®eldwork (to FKM). Nenad jacent to chimney at colony center (®g. 63B); Smodloka, DusÏan Zavodnik, Andrej Jaklin, lophophores closer to colony center support- and the other staff of the RudÅer BosÏkovic ed on short extroverts in some colonies (®g. Institute Center for Marine Research±Rovinj 63A). for providing laboratory space and acquisi- REMARKS: Living species previously in- tion of specimens through dredging and div- cluded in Lichenopora Defrance (1823) ap- ing. Rupert Riegel and Konrad Thaler kindly parently can no longer be considered to be arranged the opportunity to examine and bor- part of this genus, which has as its type the row Heller's available types from the Uni- Eocene species Lichenopora turbinata Defr- versity of Innsbruck Institute of Zoology. ance, 1823 (designated by d'Orbigny, 1853: Mary Spencer Jones and Kevin Tilbrook ar- 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 129 ranged access to comparative specimens in fragilis sp. n. y C. labiata sp. n. Cahiers de The Natural History Museum, London. Mar- Biologie Marine 30: 143±165. jorie McKinney served willingly and capably Audouin, J. V. 1826. Explication sommaire des planches de polypes de l'E gypte et de la Syrie. as general factotum during ®eld and labora-  tory work and spent many hours in the dark- In J.C. Savigny, Description de l'Egypte, his- toire naturelle 28: 225±244. Paris: C.L.F. room processing photographs. Ruth Dewel, Panckoucke. 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APPENDIX 1 2002 HAYWARD AND MCKINNEY: ADRIATIC BRYOZOA 139

APPENDIX 1 (Continued)