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Home , Adeonidae, Alcyonidiidae, Alcyonidium (bryozoan), Alcyonidium hirsutum, Amathia vidovici, Ascophora

MARINE BRYOZOANS (ECTOPROCTA) OF THE INDIAN RIVER AREA (FLORIDA)

JUDITH E. WINSTON

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 173 : ARTICLE 2 NEW YORK : 1982

MARINE BRYOZOANS (ECTOPROCTA) OF THE INDIAN RIVER AREA (FLORIDA)

JUDITH E. WINSTON Assistant Curator, Department of American Museum of Natural History

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 173, article 2, pages 99-176, figures 1-94, tables 1-10 Issued June 28, 1982 Price: $5.30 a copy

Copyright © American Museum of Natural History 1982 ISSN 0003-0090 CONTENTS

Abstract 102 Introduction 102 Materials and Methods 103 Systematic Accounts 106 Ctenostomata 106 polyoum (Hassall), 1841 106 Alcyonidium polypylum Marcus, 1941 106 Nolella stipata Gosse, 1855 106 Anguinella palmata van Beneden, 1845 108 pavida Saville Kent, 1870 108 Sundanella sibogae (Harmer), 1915 108 alternata Lamouroux, 1816 108 Amathia distans Busk, 1886 110 (Heller), 1867 110 Bowerbankia gracilis Leidy, 1855 110 Bowerbankia imbricata (Adams), 1798 Ill Bowerbankia maxima, New Ill (Delle Chiaje), 1828 113 Valkeria atlantica (Busk), 1886 114 Aeverrillia armata (Verrill), 1873 114 Cheilostomata 114 Aetea truncata (Landsborough), 1852 114 Aetea sica (Couch), 1844 116 tenuissimum (Canu), 1908 116 IConopeum seurati (Canu), 1908 117 arborescens (Canu and Bassler), 1928 117 Membranipora savartii (Audouin), 1926 119 Membranipora tuberculata (Bosc), 1802 119 Membranipora tenella Hincks, 1880 120 Membranipora sp 120 bellula (Hincks), 1881 120 doma (d'Orbigny), 1859 122 Discoporella umbellata, subspecies depressa (Conrad), 1841 122 Aplousina gigantea Canu and Bassler, 1927 123 Parellisina latirostris Osburn, 1940 123 Antropora leucocypha (Marcus), 1937 123 Floridina antiqua (Smitt), 1873 125 Floridinella typica Canu and Bassler, 1928 125 coriacea (Johnston), 1847 125 Thalamoporella floridana Osburn, 1940 126 Thalamoporella falcifera (Hincks), 1880 126 Synnotum aegyptiacum (Audouin), 1826 127 Scrupocellaria regularis Osburn, 1940 127 neritina (Linnaeus), 1758 129 Bugula stolonifera Ryland, 1960 129 Bugula turrita (Desor), 1848 130 Bugula minima Waters, 1909 130 100 Bugula uniserialis Hincks, 1884 130 Caulibugula pearsei Maturo, 1966 131 hirtissima (Heller), 1867 131 Beania klugei Cook, 1968 131 Beania mirabilis Johnston, 1840 133 Cribrilaria radiata (Moll), 1803 133 Membraniporella aragoi (Audouin), 1826 134 Reginella floridana (Smitt), 1873 134 Bellulopora bellula (Osburn), 1950 134 Exechonella antillea (Osburn), 1927 136 Escharoides costifer (Osburn), 1914 136 Savignyella lafontii (Audouin), 1826 136 Reptadeonella costulata (Canu and Bassler), 1928 137 Tremoschizodina lata Smitt, 1873 137 Hippaliosina rostrigera (Smitt), 1873 139 (Moll), 1803 139 Watersipora subovoidea (d'Orbigny), 1852 139 Crepidacantha setigera (Smitt), 1873 140 Hippoporina verrilli Maturo and Schopf, 1968 140 Parasmittina nitida morphotype B Maturo and Schopf, 1968 142 Parasmittina spathulata (Smitt), 1873 142 Phylactellipora aviculifera (Osburn), 1914 143 Lagenicella marginata (Canu and Bassler), 1930 143 cornuta (Gabb and Horn), 1862 144 Schizoporella floridana Osburn, 1914 144 (Johnston), 1847 145 Escharina pesanseris (Smitt), 1873 145 Stylopoma spongites (Pallas), 1766 145 Cigclisula turrita (Smitt), 1873 147 Cleidochasma porcellanum (Busk), 1860 147 Cleidochasma contractum (Waters), 1899 148 Hippoporidra calcarea (Smitt), 1873 148 Microporella umbracula (Audouin), 1826 150 Hippothoa flagellum Manzoni, 1870 150 Celleporella carolinensis Ryland, 1979 151 Trypostega venusta (Norman), 1864 151 Pasythea tulipifera (Ellis and Solander), 1786 152 Vittaticella uberrima Harmer, 1957 152 Vittaticella contei (Audouin), 1826 154 Celleporina hassalli (Johnston), 1847 154 154 elongata Milne-Edwards, 1838 154 Crisia eburnea (Linnaeus), 1758 155 Plagioecia dispar Canu and Bassler, 1928 155 Distribution and Ecological Factors 156 Literature Cited 169

101 ABSTRACT The distribution and ecology of marine bryo- richer in species. Twenty-three species were found zoans of the Indian River area on the east coast at Sebastian Inlet inner breakwater, 31 at North of Florida was studied through collections made Beach breakwater, Fort Pierce, 36 at Walton at 21 stations over the course of a year. Bryozoans Rocks, and 31 at Seminole Shores. Offshore areas were identified from collections of all substrata were also sampled. Twenty-one species were (e.g., shells, hydroids, algae, rock, ) on found at Capron Shoals. Twenty-eight species which colonies might be able to grow. were identified in samples from two R/V Gosnold Eighty-four species of bryozoans have been ob- cruises. tained thus far from the waters of the Indian River In the Indian River area some bryozoan repro- region. All collections indicated that the avail- duction occurred year-round, but many species ability of a suitable substratum is the chief factor reproduced primarily from late fall to early spring, governing distribution of bryozoan species. In the in contrast to the late summer-early fall peak re- river the main substrata are the relatively short- production of bryozoan populations in temperate lived blades of seagrasses on which small, rapidly seas. Biogeographically, the species collected off- growing bryozoan species are found. Eighteen shore had generally tropical affinities, whereas species were found in the Indian River; 12 of them those collected at coastal and river stations in- in waters of salinities less than 30%o (estuarine cluded a number of tropical species, but also many conditions). Coastal stations, with more varied species with broader Western Atlantic or cosmo- substrata available for settlement and with a wider politan distributions. range of microenvironmental conditions, were

INTRODUCTION

Bryozoans are tiny marine organisms, and Puerto Rico, and Mature (1968) in- which form encrusting or arborescent colo- cluded shallow water stations in his study of nies in almost any marine environment the bryozoans from the southeast coast of the where there is a suitable surface for attach- United States. Shier (1964) collected bryo- ment. zoans from beach drift material on the north- From the number of studies (Smitt, 1872, west coast of Florida. Because no one has 1873; Osburn, 1914, 1927, 1940; Canu and collected bryozoans from intertidal and es- Bassler, 1928a; Lagaaij, 1963; Shier, 1964; tuarine localities along the Atlantic coast of Mature, 1968; Long and Rucker, 1970; Pow- Florida, it seemed important not only to ob- ell, 1971) that have been carried out in the tain data on the distribution of species, but last hundred years on the bryozoan fauna of also to make collections in various habitats the Gulf of Mexico and the Caribbean Sea throughout the course of a year, in to it might be supposed that this fauna was one learn more about substratum preference, of the best known in the world. According to temperature and salinity tolerance, repro- the checklist given by Schopf (1973) 278 ductive periods and other ecological consid- species of cheilostomes (the most diverse or- erations. der of living bryozoans) have been reported from the tropical Western Atlantic region in water of less than 125 m. depth. However, ACKNOWLEDGMENTS most of these studies were carried out by The collections and observations for the dredging material. Less is known of the present study were carried out in 1974 and species found in the shallowest coastal waters 1975 during the tenure of a Smithsonian and the subtidal and intertidal habitats of the Post-doctoral Fellowship. Systematic study, inlets and . Osburn (1914, 1940) did museum comparisons, and illustration of some intertidal collecting and shallow water specimens continued over the next few years, (less than 55 m.) dredging in the Tortugas and I am indebted to many people for their

102 1982 WINSTON: MARINE BRYOZOANS 103 aid during this time: to Dr. Mary E. Rice of enced areas (above Sebastian) ranges of en- the Smithsonian Institution, Fort Pierce Bu- vironmental conditions are wider and more reau, for arranging several stays for me there unpredictable than in the southern part of the as a Visiting Scientist and for the provision region (Young and Young, 1977). of space and equipment; to Ms. Julie Piraino Biogeographically this area lies in a tem- for her virtuosity at the control board of the perate-subtropical transition zone between Scanning Electron Microscope and to Ms. the Carolinean and Caribbean faunal prov- Cindy Hunter for help with photography. I inces. Many tropical species reach their am indebted also to many people at the Har- northern limits in the area (Briggs, 1974). bor Branch Foundation, Fort Pierce, for their Low temperature stress may be particularly assistance: to Dr. Kevin Eckelbarger, Ms. Pat severe, sometimes causing spectacular mor- Linley, Dr. Nat Eiseman, Ms. Kris Metzger, tality of tropical fishes (Gilmore, Bullock, Mr. David Mook and especially to Ms. Mary and Berry, 1978), and affecting invertebrates Ann Capone of the Indian River Coastal as well. Zone Survey for sorting and preliminary Bryozoans were collected by various means identification of bryozoans from quantitative at four stations within the Indian River, four collections. Thanks also to Dr. Frank coastal stations, and 13 offshore stations (fig. Maturo of the University of Florida for the 1). loan of bryozoan specimens from his collec- tion, to Dr. Alan Cheetham of the Smithson- INDIAN RIVER STATIONS ian Institution (NMNH) for many discus- sions of bryozoan systematics, and to Drs. The seagrasses Syringodium filiforme and Cheetham and Jeremy Jackson of the Johns Halodule wrightii form extensive beds in the Hopkins University for their critical evalu- river. There are a few beds of Thalassia tes- ation of the manuscript, and to the secretaries tudinum (which is approaching the northern at the Harbor Branch Foundation, the Johns limits of its distribution in this area) (Thomp- Hopkins University, and to Ms. Monica son, 1978). In addition to the seagrasses, free- Werner at the American Museum of Natural drifting macroalgae may contribute a large History for processing various drafts of this amount to the benthic , as well as work. providing space and food for other organisms (Benz, Eiseman, and Gallaher, 1979). Bryozoans were obtained from the Indian MATERIAL AND METHODS River from samples of seagrasses taken by quantitative methods (Young, 1975). Qual- LOCALITIES STUDIED itative collections were made at the same sta- The Indian River is a shallow bar-built tions (Haulover Canal, Titusville, Banana , separated from the Atlantic River, Eau Gallic, Sebastian Grass Flat, Se- by a continuous line of barrier islands. It ex- bastian, Link Port Grass Bed, Link Port, Fort tends along the east coast of Florida for about Pierce) in March to obtain bryozoans from 180 km., from Cape Canaveral south to all substrata (e.g., pilings, roots, Stuart. Three inlets (Sebastian, Fort Pierce, shells, submerged wood, beverage cans, and St. Lucie) cut through to the river in the buoys and floats). The stations at Haulover, southern half of the region. At its northern Sebastian and Link Port were visited also at edge, the river is connected to Mosquito La- other times of the year. Bryozoan traps, con- goon by the Haulover Canal. In the river sa- sisting of wooden slide boxes, filled with glass linity ranges from extremely brackish to hy- microscope slides and covered with plastic persaline (5-55%o), averaging about 25%o. window screening, were suspended from a The average water depth is 1.5 m., therefore, piling and from panel arrays in the canal at wind-driven tidal movements, turbulence, Link Port. Bryozoans settled and grew on the and temperature changes can affect much of undersurface of the slides, which were re- the water column. North of the inlet-influ- moved, examined, and replaced periodically. 104 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

2800-

COCOA1

ATLANTIC OCEAN

MELBOURNE 2730-

FLORIDA

VERO BEACH

2700-

FORT PIERCE

10 20 km I i I

8030 8000 1 I FIG. 1. Map of Indian River region showing collecting localities. 1. Haulover Canal. 2. Banana River. 3. Sebastian Inlet. 4. Sebastian Grass Flat. 5. Link Port. 6. Fort Pierce Breakwater. 7. Walton Rocks. 8. Seminole Shores. 9. Capron Shoals. 10. Gosnold Station 766. 11. Gosnold stations 696-698, 700-702, 734, 735, 737, 738, 757, 759; Sebastian pinnacle area.

COASTAL STATIONS consist primarily of sand and shell gravel. Both intertidally and subtidally substrata Hard substrata are provided subtidally by along the outer coast of the barrier islands coquinoid rock ledges and sabellariid worm 1982 WINSTON: MARINE BRYOZOANS 105 mounds, and intertidally by sabellariid Cruises of the R/V Gosnold in September mounds, beach rock ledges and the rocks of 1974 and February 1975, yielded specimens inlet breakwaters. of bryozoans from the Oculina pin- Four coastal stations (Sebastian Inlet nacles at 60-90 m. depth. Stations sampled Breakwater, North Beach Breakwater, Fort were all in the vicinity of the Sebastian Pin- Pierce Inlet, and Walton Rocks and Seminole nacle System (20*49.5' N and 79°58' W) a Shores, Hutchinson Island) were sampled complex of topographic prominences formed several times during the year. Each of the by oolitic limestone mounds and relict localities differed in ecological conditions reefs, with substrata ranging from rock, shell and in the types of environments available hash, sand, to Oculina rubble, other uncon- for bryozoan colonization. The locality cho- solidated coral debris, and a small amount sen for sampling within Sebastian Inlet was of living Oculina varicosa (Avent, King, and the breakwater adjacent to a muddy tidal flat Gore, 1977; Thompson and Gilliland, 1980). area. On one side of this breakwater, the On the February cruise collections were also strong current passing through the inlet ex- made at two stations immediately off Fort poses organisms living on the rocks to strong Pierce Inlet (at 10 m. depth) in a sand bottom water action. On the tidal flat side of the area. breakwater, the chief influence on distribu- tion of sessile appears to be the large EXAMINATION OF SPECIMENS amount of fine sediment which settles from the water in this calm area. The North Beach In making these collections, the different Breakwater at Fort Pierce Inlet is exposed to microenvironments such as crevices, pro- very strong wave action, but many protected tected surfaces of rocks, shells, various algae, habitats are present within caves and crevices hydroid stems, and roots were examined between the rocks. The Walton Rocks local- carefully for bryozoans. Samples of each sub- ity is on the ocean side of Hutchinson Island, stratum were taken to the laboratory and ex- immediately south of the intake canal for amined further under the dissecting scope, coolant water for the nuclear power (at so that detailed observations could be made the time of this study the power plant was on behavior and of bryozoan not yet in operation). Here the primary sub- species living on them. Examining such sub- strata for attachment are beachrock ledges strata as large masses of hydroids in this way and loose rock in sandy channels between the was a tedious process, but many tiny and ledges and the shore. The habitat is exposed non-calcified species were found that would to strong wave action, causing constant probably have been overlooked in the field. movement of sand and changes in sand level, Morphological measurements were made of and, on very low tides to the drying action the most common species and notes were of sun and wind. The sabellariid reefs at Sem- taken on the characteristic behavior and col- inole Shores are strongly developed on the oration of the species when alive and infor- coquinoid ledge base, and habitats available mation was recorded on growth stage of the to bryozoans include the worm tubes them- , presence of reproductive structures selves, as well as loose rocks and rock ledges or , and occurrence of degenerated and hydroid and algal substrata. with storage granules (common in overwintering or oversummering colonies).

OFFSHORE STATIONS ILLUSTRATION OF SPECIMENS In addition to these intertidal stations, Most of the species found in these collec- samples were taken several times during the tions have been illustrated. For species with- year at Capron Shoals, a subtidal (8-15 m. out calcification or with very delicate calci- depth) area about 7 km. south of Fort Pierce fication drawings have been made from Inlet, where substrata ranged from coarse preserved material or slides of living colo- shell debris to fine shell hash and sand. nies. Calcified species are illustrated by SEM. 106 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

The specimens were first treated in Clorox waters of both the Atlantic and Pacific. Gulf (either thoroughly or lightly depending on of Mexico. degree of calcification), then cleaned ultra- sonically in distilled water, air-dried and Alcyondium polypylum Marcus, 1941 mounted on stubs. Specimens were sputter- Figure 2 coated with gold-palladium and scanned us- Alcyonidium polypylum Marcus, 1941, p. 63. ing a Zeiss Novascan 30 Scanning Electron Microscope. Representative groups of zooids DESCRIPTION: Colonies transparent orange were photographed at 100X. Each specimen (single layer) to opaque brick red, forming a illustrated by SEM has received an AMNH uni- to multi-laminar crust from which erect catalogue number. This number is given in knobs may arise. Zooids polygonal in shape, parentheses at the beginning of the descrip- averaging .47 X .34 mm. Walls thick, with tion of that species. a smooth, glistening surface. No oral papillae. Polypides with orange and gut, and SYSTEMATIC ACCOUNTS translucent sheaths. Fifteen tenta- cles, diameter averaging .523 Eighty-four species of bryozoans were col- mm. lected from river, coastal, and offshore sta- OCCURRENCE AND ECOLOGY: Collected at tions of the Indian River area. These species three locations, Walton Rocks, Seminole are described below. The classification of the Shores (on beach rock and shell) and Capron Ctenostomata follows Prenant and Bobin, Shoals (on shell). At Walton Rocks A. poly- 1966; of the Cyclostomata, Osburn, 1953; pylum formed thin crusts on the undersur- and of the Cheilostomata, Cook, 1968a; Ry- faces of flat pieces of beach rock in the sandy land and Hayward, 1977; Hay ward and Ry- channel between the beach and the first beach land, 1979. rock ledge. According to Marcus (1941), col- onies live four years or more. However, in ORDER CTENOSTOMATA BUSK, 1852 this unstable habitat (face down on sand with SUBORDER CARNOSA GRAY, 1848 level of sand and water constantly changing) FAMILY JOHNSON, 1849 it is unlikely that colonies achieved this age, ALCYONIDIUM LAMOUROUX, 1812 though multilayer crusts occurred. Massive Alcyonidium polyoum (Hassall), 1841 colonies found in April and present through Figure 3 the summer months. In September only sin- gle-layered colonies collected at Walton and Sarcochiton polyoum Hassall, 1841, p. 484. Alcyonidium polyoum: Marcus, 1937, p. 125. Os- Seminole. Reproduction of colonies from burn, 1953, p. 727. Mature, 1957, p. 18. Shier, Capron Shoals noted in October and Novem- 1964, p. 642. Amsr and Pallares, 1965, p. 299. ber. Reproducing zooids have polypides with large intertentacular organs and yellow-or- DESCRIPTION: Colonies forming a gelati- ange eggs visible through the thin walls of nous whitish to brownish crust, never rising unilaminar parts of the colonies. into knobs or branches. Zooids hexagonal to DISTRIBUTION: Brazil, Atlantic coast of irregularly polygonal .46 X .22 mm. in size, Florida. This is the first record of the species with transparent frontal wall through which from Florida. the polypide may be distinguished. Raised oral papillae present or not depending on SUBORDER PALUDICELLEA ALLMAN, 1856 degree of retraction of polypide. Polypide FAMILY NOLELLIDAE HARMER, 1915 with 14 tentacles; lophophore averaging .331 GENUS NOLELLA GOSSE, 1855 mm. in diameter. OCCURRENCE AND ECOLOGY: Occurred only Nolella stipata Gosse, 1855 once in this study, on the panels at Link Port Figure 5 in Aprjl. Nolella stipata Gosse, 1855, p. 35. Osburn, 1953, DISTRIBUTION: Cold and warm-temperate p. 737. Mature, 1957, p. 21. 1982 WINSTON: MARINE BRYOZOANS 107

, 0.5 mm .

FIGS. 2-7. 2. Alcyonidium polypylum. 3. Alcyonidium polyoum. 4. , showing arising from base of zooids, rather than zooids budded separately from stolon as in Bowerbankia spp. 5. Nolella stipata. 6. Sundanella sibogae, group of four zooids. 7. Anguinella palmata.

Nolella gigantea Marcus, 1937, p. 131. a bryozoan. Polypide with 17 tentacles and Nolella dilatata Marcus, 1938, p. 53. Osburn, an equi-tentacled lophophore averaging .616 1940, p. 337. mm. in diameter. DESCRIPTION: Colony of widely spaced tu- OCCURRENCE AND ECOLOGY: Collected at bular zooids connected by a delicate stolon. three stations in the river, the Link Port grass Zooids tall and cylindrical (2.1 X .17 mm.), beds, Fort Pierce Breakwater and Walton their chitinous walls usually covered with a Rocks. Occurs year round. Cryptic in habit, layer of silt, thus resembling the mud tubes collected on shells, algae, seagrass, and the of amphipods or . Only careful bryozoan Zoobotryon. examination for the presence of or DISTRIBUTION: Western Atlantic: Canada reveals that the tubes belong to to Brazil. Caribbean. 108 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

GENUS ANGUINELLA VAN BENEDEN, 1845 OCCURRENCE AND ECOLOGY: Known to Anguinella palmata van Beneden, 1845 prefer brackish water (Braem, 1951). Col- Figure 7 lected only once in the Indian River, settling on the panels at Link Port in December, but Anguinella palmata van Beneden, 1845, p. 34. to be expected in brackish water throughout Osburn, 1914, p. 219; 1940, p. 338. Marcus, the area. 1937, p. 133. Mature, 1957, p. 21. Shier, 1964, DISTRIBUTION: Cosmopolitan in brackish p. 649. water. Western Atlantic: Chesapeake Bay to DESCRIPTION: Colonies limp brown tufted Brazil. Not previously reported from Florida. masses resembling dirt-coated seaweed. Zooids cylindrical (averaging .79 X . 13 mm.), GENUS SUNDANELLA BRAEM, 1939 covered with fine layer of silt making them Sundanella sibogae (Harmer), 1915 completely opaque. Polypide with 10 tenta- Figure 6 cled lophophore (averaging .205 mm. in di- ameter) which is protruded from the terminal Victorella sibogae Harmer, 1915, p. 45. Marcus, aperture of . No zooids off 1937, p. 129. Osburn, 1940, p. 336. Sundanella sibogae: Braem, 1939, p. 267. Marcus, sides of mature zooids. Only zooids at branch 1941, p. 69. Mature, 1957, p. 20. Shier, 1964, tips with functional polypides. p. 648. OCCURRENCE AND ECOLOGY: Common only at one locality, Sebastian Inlet. Most abun- DESCRIPTION: Colonies encrusting, zooids dant in April when breakwater rocks were budding distally producing a single series coated with mud and muddy masses of An- (unlike Victorella the stolon portion only oc- guinella dangled from them. Collected only casionally visible because zooids are large between January and April with the excep- and jumbled on top of each other). Lateral tion of a colony lh cm. in height which was budding occurring, but not observed in uni- found at Walton Rocks, in September. serial Indian River area specimens. Zooids DISTRIBUTION: Cosmopolitan. Western oval to teardrop-shaped, .50 X 1.45 mm. in Atlantic: Massachusetts to Brazil, Gulf of size, yellowish brown in color (partially due Mexico, and Caribbean. to adherence of silt particles). Lophophore with 31 tentacles and an average diameter of FAMILY HINCKS, 1880 .820 mm. GENUS VICTORELLA SAVILLE KENT, 1870 OCCURRENCE AND ECOLOGY: Collected both in the river (at Haulover Canal, Titusville) Victorella pavida Saville Kent, 1870 and along the coast (Walton Rocks and Sem- Figure 4 inole Shores). Probably occurs year-round. Victorella pavida Saville Kent, 1870, p. 34. Mar- As colonies consist of very small encrusta- cus, 1940, p. 329; 1953, p. 312. Osburn, 1944, tions on Syringodium, algae, and hydroid p. 17. roots it is found only by careful examination DESCRIPTION: Colonies consisting of creep- of these substrata. Reproductive season un- ing chains of zooids, often developing into known. fuzzy branching masses. Superficially resem- DISTRIBUTION: Western Atlantic: Beaufort bling Bowerbankia, but while Bowerbankia to Brazil. Gulf of Mexico. zooids bud from a stolon the stolon arises from the base of each zooid in Victorella (fig. SUBORDER VESICULARINA JOHNSTON, 1847 4). Zooid chains simple in young colonies as FAMILY JOHNSTON, 1838 colonies develop zooids elongate and bud GENUS .4M4 77/7.4 LAMOUROUX, 1816 additional stolons or zooids forming a mas- Amathia alternata Lamouroux, 1816 sive network. Zooids cylindrical, averaging Figure 8 .15 X .60 mm. (reported to reach 2-3 mm. Amathia alternata Lamouroux, 1816, p. 160. in old colonies). Polypides with eight tenta- Shier, 1964, p. 651. cled campylonemidan lophophore, averaging Amathia convoluta (in part) Mature, 1957, p. 22. .369 mm. in diameter. Figure 11. 1982 WINSTON: MARINE BRYOZOANS 109

i 2mm

FIGS. 8-16. 8. Amathia alternata. 9. Amathia vidovici. 10. Amathia distans. 11. Bowerbankia max- ima. 12. Bowerbankia imbricata. 13. Zoobotryon verticillatum. 14. Aeverrillia armata. 15. Bowerbankia gracilis. 16. Bowerbankia maxima showing size of zooids and expanded lophophores in contrast with those of B. gracilis (white pigment appears black in transmitted light). 110 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

DESCRIPTION: Colonies erect, bushy and node. Often growing ends of stolons with no branching, springy to the touch, yellow- zooids for the first two internodes, or with brown in color. Zooids tubular, connate to a clump of just a few zooids. the tips, averaging .56 mm. in length and . 15 OCCURRENCE AND ECOLOGY: Occurred mm. in diameter, arranged in neatly paired year-round at Fort Pierce Breakwater, Wal- double rows along a stolon, each double row ton Rocks and Seminole Shores. Most abun- covering about four-fifths of the length along dant in December at Fort Pierce Breakwater. an intemode. Lines of zooids straight or Also collected associated with algae on the twisting as much as 90 degrees around the continental shelf of Florida (Winston and stolon. Zooids of successive internodes ori- Eiseman, 1980). ented at 180-degree angles to each other. DISTRIBUTION: Circumtropical. Cosmo- Polypide with eight equal tentacles and an politan in warm seas. Western Atlantic: average lophophore diameter of .302 mm. Beaufort to Brazil. Caribbean. OCCURRENCE AND ECOLOGY: Collected at Capron Shoals and at Sebastian Inlet, Walton Amathia vidovici (Heller), 1867 Rocks and Seminole Shores. Occurred year- Figure 9 round, but most abundant in the winter. Col- Valkeria vidovici Heller, 1867, p. 128. onies attached to shell, to beach rock (espe- Vescicularia dichotoma Verrill, 1873, p. 709. cially to rock overhangs), attached to and Amathia dichotoma Osburn, 1912, p. 254. entangled in red alga Solieria tenera (at the Amathia vidovici: Osburn, 1940, p. 340; 1953, p. Sebastian Grass Flats). No colonies reached 741. Prenant and Bobin, 1956, p. 283. the large sizes (up to 30 cm.) Shier (1964) DESCRIPTION: Colonies branching, semi- observed on the Gulf Coast; many were quite erect, whitish to tan in color, very lightly chi- tiny. tinized. Stolons wider (.13-20 mm.) and DISTRIBUTION: Cape Hatteras to Florida stiffer than those of Amathia distans. Zooids and Gulf of Mexico. May also occur in the only connected for part of their length of Caribbean where its identity has been con- about .4 mm. Arranged in small groups, four fused with Amathia convoluta. to eight pairs making a short spiral round the stolon in the distal half of the intemode, leav- Amathia distans Busk, 1886 Figure 10 ing the proximal half of the intemode bare. OCCURRENCE AND ECOLOGY: Present year- Amathia rfwtans Busk, 1886, p. 33. Marcus, 1937, round in the Indian River area: Sebastian p. 134. Osburn, 1940, p. 339. Mature, 1957, p. Inlet, Fort Pierce Breakwater and Seminole 23. Cook, 1968a, p. 228. Shores on the coast, and found attached to Amathia brasiliensis Busk, 1886, p. 34. Osburn, Solieria tenera at the Sebastian Grass Flat in 1940, p. 339. lAmathia goodei Osburn, 1914, p. 219. March. DISTRIBUTION: Western Atlantic: Massa- DESCRIPTION: Colonies delicate and trans- chusetts to Gulf of Mexico. Caribbean. Also, lucent, consisting of a thin (.11-.12 mm.) Pacific from Southern California to the Ga- dichotomously branching stolon with clumps lapagos, Mediterranean, East Atlantic and of zooids at intervals. Walls of zooids and Indian Ocean. stolons of Florida specimens a chitinous tan color, with bright yellow spots of pigment all Bowerbankia gracilis Leidy, 1855 over, causing living colonies to appear yel- Figure 15 low. Zooids connate and a little shorter than Bowerbankia gracilis Leidy, 1855, p. 142. Osburn, those of A. alternata, about .4 mm. in height. 1914, p. 218; 1940, p. 34 (in part; see B. max- Arranged in double series at the distal end ima). Marcus, 1938, p. 56. Prenant and Bobin, of the internodes, each series completing a 1956, p. 303. Mature, 1957, p. 25. Shier, 1964, full or partial spiral around stolon, usually p. 652 (in part; see B. maxima). Ryland, 1965, only filling about half the space along inter- p. 78. Amor and Pallares, 1965, p. 298. 1982 WINSTON: MARINE BRYOZOANS 111

DESCRIPTION: Colonies straw-colored, stolon, remaining creeping or growing up creeping along hydroids, algae, or other sub- from the surface as bushy tufts. Zooids vase- strata, often forming entangled masses, con- shaped to subcylindrical. For functional sisting of tubular flask-shaped zooids ar- zooids, mean length (N = 5) averaging 0.38 ranged singly or in pairs or clumps along a mm., mean width 0.17 mm. For degenerated stolon which is about 0.05 mm. in diameter zooids mean length .38 mm., mean width and divided into internodes by septa. Zooids 0.19 mm. Stolon diameter averaging 0.23 attached to stolon basally and separately bud- mm. Small star-shaped black pigment spots ded from it. Zooid walls lightly chitinized, in zooids and stolons of living colonies, but internal structures visible. With polypides polypides unpigmented. Polypides with a giz- expanded zooids erect and long, in live In- zard and ten tentacles. Lophophore diameter dian River area specimens 0.38-0.80 mm. averaging .246 mm. in length (mean .60 mm.) and .09-14 mm. TAXONOMIC DISCUSSION: The zooids of this in width. With polypides retracted or non- species in the Indian River are smaller than functional, zooids shorter and commonly those described elsewhere. The zooid clusters flattened against the substratum. Caudate of Bowerbankia imbricata are supposed to be process absent from zooid base in all speci- non-helicoidal; in these specimens the zooids mens collected. Polypide with lophophore did form a partial spiral around the stolon. unpigmented, mean lophophore diameter Colony morphology and dimensions thus .315 mm., eight tentacles. Eggs brooded in- more closely resemble those described for ternally in zooids in which polypides have Bowerbankia pustulosa. However, polypides degenerated. Embryos peach colored. have ten tentacles and those of B. pustulosa OCCURRENCE AND ECOLOGY: Found both have only eight. in the Indian River and at coastal stations OCCURRENCE AND ECOLOGY: Found from (Sebastian Inlet, Walton Rocks and Fort March to September. Commonly associated Pierce Breakwater). Most common cteno- with algae. At the grass flat at Sebastian Inlet stome in the Indian River, both in these col- colonies occurred on Solieria tenera, but at lections and in those of the Indian River Sur- Fort Pierce Breakwater and Sebastian colo- vey. In the river, primarily on seagrasses, but nies were found attached to the undersurfaces also on wood, dead shells, and aluminum of rocks and ledges and in holes and channels cans. At coastal stations found on algae, dead in the worm reef mounds. shell (in the holes left by boring ), DISTRIBUTION: In the Western Atlantic, hydroids, and other bryozoans. Collected previously reported only from cooler water, year-round at coastal stations. Settled on but known from warm waters in the Medi- panels at Link Port from December through terranean and Red Sea. June. Colonies with embryos collected in the Banana River in March. Bowerbankia maxima, new species DISTRIBUTION: Western Atlantic: Green- Figures 11, 16-18 land to Brazil. Probably cosmopolitan in shallow water. DIAGNOSIS: Bowerbankia maxima is sim- ilar to Bowerbankia gracilis, but distinguish- Bowerbankia imbricata (Adams), 1798 able by its white pigmentation when living. Figure 12 No white coloration is mentioned in any species of Bowerbankia described previously. Sertularia imbricata Adams, 1798, p. 11. It can also be distinguished from B. gracilis Bowerbankia imbricata: Rogick and Croasdale, on the basis of its greater dimensions. 1949, p. 47. Prenant and Bobin, 1956, p. 293. HOLOTYPE: AMNH 572. Ryland, 1965, p. 79. Amor and Pallares, 1965, p. 296. PARATYPES: AMNH 573, 574. ETYMOLOGY: The species name is from the DESCRIPTION: Colony composed of clusters Latin maximus (greatest). of zooids, clumped along a relatively thick DESCRIPTION: Colonies, like those of B. 112 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

FIGS. 17-18. Bowerbankia maxima, new species. 17. Group of zooids encrusting alga Solieria, show- ing white pigmentation. 18. Lophophore of Bowerbankia maxima, showing distribution of white pigment in tentacles and lophophore base. 1982 WINSTON: MARINE BRYOZOANS 113 gracilis, found creeping along grasses, hy- (1964) material from the northwest Gulf of droids, algae and other substrata; the fuzzy Mexico (0.5-1.5 mm. X 0.15-0.20 mm.) and grayish white network of zooids and stolons that of Osburn (1940) from Puerto Rico often developing into partially erect branches (0.65-1.20 X 0.16-0.20 mm.) probably in- and clusters or pendulous masses. Zooids cludes both B. gracilis and B. maxima. The arranged in irregularly sized clumps along the species which Marcus (1937) called B. cau- stolons. Functional zooids subcylindrical, data is in the same size range as B. maxima becoming more elongated when polypide is (1.0-1.5 mm. X 0.16-0.20 mm.) and, as the expanded. presence or absence of caudate processes has In living specimens length of functional been shown to be a variable character, per- zooids (N = 25) averaging 1.14 mm. (range haps might have affinities with it. 0.91-1.46 mm.), width averaging .230 mm. OCCURRENCE AND ECOLOGY: Collected (range .200-.273 mm.); length of degenerated from March to November in the Indian River zooids (N = 21) averaging .575 mm. (range on grass and the alga Solieria. At coastal sta- .419-737 mm.), width averaging .248 mm. tions on shells, algae, hydroids, other bryo- (range .200-300 mm.); stolons (N = 20) av- zoans and rock. Young colonies found in late eraging .134 mm. (range .091-182 mm.). March and larger colonies in succeeding Orifice round when polypide expanded and months. Most abundant in September, es- square when polypide retracted. When living, pecially at Walton and Sebastian where col- immediately distinguishable from other onies formed large masses directly on the Bowerbankia species because of brilliant beach rock ledge or breakwater surface, and white pigmentation of zooids and polypides, overgrew hydroids and other organisms. Re- the white color being derived from star- productive season unknown. shaped white cells in zooids and stolons (fig. DISTRIBUTION: Atlantic coast of Florida. 16) and from the diadem of white pigment Also found at Discovery Bay, and Port Royal, at the base of the lophophore and the white Jamaica. As the taxonomic discussion indi- pigmentation on the tentacles (fig. 18). Pol- cates, distribution may be more widespread. ypide with a large well-developed averaging .119 X .139 mm. in size (N = 20), a long introvert region, and a lophophore GENUS ZOOBOTRYON EHRENBERG, 1831 averaging .751 mm. in diameter, with eight equal tentacles curved strongly outward at Zoobotryon verticillatum (Delle Chiaje), 1828 the tips. Eggs and brooding not observed. Figure 13 DISCUSSION: Maturo (1957) described Hydra verticillata Delle Chiaje, 1828, p. 203. Bowerbankia gracilis from the Beaufort area Zoobotryon pellucidum Ehrenberg, 1831. Marcus, as having zooids about .70 mm. in length by 1937, p. 139. Osburn, 1940, p. 341. .15 mm. in width. This fits B. gracilis as Zoobotryon verticillatum Osburn, 1953, p. 742. understood here also. In the Indian River Maturo, 1957, p. 25. Cook, 1968a, p. 229. area specimens there was almost no overlap Zoobothryon verticillatum Prenant and Bobin, 1956, p. 288. between Bowerbankia gracilis and Bower- bankia maxima with respect to sizes of func- DESCRIPTION: Colonies made up of irreg- tional zooids (as measured in live specimens). ularly or trifurcately branching masses of sto- The length of degenerated zooids of B. max- lons and zooids. Zooids ovoid, .38 X .58 ima was within the range of zooids of B. gra- mm. in size, occurring in clusters on either cilis, but these degenerated zooids were very side of the thick transparent stolon. Individ- much wider (.273 mm. vs. .106 mm.) than ual polypides with a large gizzard. Lopho- B. gracilis zooids. Some of the variability of phore with eight short equal tentacles and a "Bowerbankia gracilis'"'' as described from the diameter averaging .320 mm. Western Atlantic and Caribbean by various OCCURRENCE AND ECOLOGY: Collected both authors may actually be due to the presence in the river (Haulover Canal, Sebastian Grass of both species in their samples. Shier's Flats) and along the coast. Most abundant at 114 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

Walton Rocks. Overwintering colonies ini- GENUS AEVERRILLIA MARCUS, 1941 tiated rapid growth about the first of April, Aeverrillia armata (Verrill), 1873 producing colonies the size of a fist in two Figure 14 weeks time. In early summer (June) large masses of Zoobotryon, intermingling with Vesicularia armata Verrill, 1873, p. 710. algae and hydroids, dangled from beach rock Aeverrillia armata: Marcus, 1941, p. 76. Mature-, ledges. Sexual reproduction began then, and 1957, p. 26. Shier, 1964, p. 652. colonies were abundant throughout summer DESCRIPTION: Colony composed of a nar- months, though, as the summer passed, the row creeping stolon (about .04 mm. in di- transparent new growth became encrusted ameter) divided into internodes by irregu- with red and green filamentous algae, dia- larly spaced septa. Zooids arising (usually in toms, and silt. The muddy masses of mature pairs) from kenozooids placed on either side colonies provided attachment for other bryo- of stolons. Zooids ovoid, with a rounded zoans such as Savignyella, Amathia, Bower- base, tapering slightly toward the distal end bankia, Nolella, and Aetea. By October most which is provided with four movable spines. of these masses, along with most macro-algae Campylonemidan lophophore with eight ten- found on the ledges, had sloughed off or been tacles. When the polypide is expanded the ripped away. lophophore is surrounded by a setigerous col- DISTRIBUTION: A circumtropical species. lar equal in length to the tentacles. This collar Western Atlantic: Beaufort to Brazil. Gulf of may or may not project from the aperture Mexico and the Caribbean. when the polypide is retracted, depending on the degree of retraction. Gizzard present. OCCURRENCE AND ECOLOGY: Collected at SUBORDER STOLONIFERA EHLERS, 1876 three coastal stations: Sebastian Inlet, Fort FAMILY VALKERIIDAE HINCKS, 1877 Pierce Inlet and Seminole Shores. Although GENUS VALKERIA FLEMING, 1823 previously recorded from waters of lowered Valkeria atlantica (Busk), 1886 salinity (to 12% in Chesapeake Bay according to Osburn, 1944) it was not taken within the Farrella atlantica Busk, 1886, p. 37. river. Most common from August to January, Valkeria atlantica: Osburn, 1940, p. 342. but probably occurring year-round. Found on DESCRIPTION: Colonies of this inconspic- stems of the hydroids Thyroscyphus ramosus uous and delicate species found creeping over and Eudendrium carneum and on Amathia algae, hydroids, and shells. Zooids tall, slen- spp. No evidence of reproduction seen. der (.60-80 X .10 mm. when living), usually DISTRIBUTION: Western Atlantic: Maine to paired, arising from the distal end of a very Brazil and Gulf of Mexico. Distribution not short joint or vestigial intemode. Polypides well known; it is another species which is with a campylonemidan lophophore with usually only collected accidentally. eight tentacles. OCCURRENCE AND ECOLOGY: One colony ORDER CHEILOSTOMATA found on a hydroid stem at the North Beach SUBORDER ANASCA LEVINSEN, 1909 breakwater in August. Because of its small FAMILY SMITT, 1867 size and creeping habit this species is easy to GENUS AETEA LAMOUROUX, 1818 overlook, but as a careful search of hydroid Aetea truncata (Landsborough), 1852 stems, algae, and other likely substrata was Figures 19, 21 made at each collection in this study, Valk- Anguinaria truncata Landsborough, 1852, p. 288. eria atlantica seems to be truly rare in this Aetea truncata: Canu and Sassier, 1928, p. 51. area. Hastings, 1930, p. 702. Marcus, 1938, p. 11. DISTRIBUTION: Brazil, Puerto Rico, Flor- Osburn, 1940, p. 346; 1950, p. 12. Cook, 1968, ida. Knowledge of distribution limited, prob- p. 137. ably circumtropical. lAetea truncata: Shier, 1964, p. 606. 1982 WINSTON: MARINE BRYOZOANS 115

FIGS. 19-24. 19. Aetea truncata, group of zooids. 20. Aetea sica, two zooids and stolon. 21. Aetea truncata, portion of basal region of zooids, showing lack of annulation. 22. Aetea sica, portion of basal region of zooid, showing fine annulation. 23. Cupuladria doma. 24. Discoporella umbellata, subspecies depressa. 116 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

DESCRIPTION: (AMNH 575) Colonies con- September and October. At Fort Pierce sisting of creeping stolonate portions from Breakwater, Walton Rocks, and Seminole which tubular portions arise. Tubular por- Shores on drift Sargassum and on attached tions are divided into a basal "stalk"-like re- algae and bushy bryozoans: Amathia, Zoo- gion and a terminal apertural region with the botryon, and Bugula. In the river on sea- operculum at the upper end of its frontal sur- grasses and the red alga Solieria tenera. Re- face. Polypide extending down into the basal productive season unknown. portion. Both stem and terminal regions DISTRIBUTION: Cosmopolitan except in po- spotted with minute tubercles, but not an- lar regions. Western Atlantic: Cape Hatteras nulated (fig. 21). Tubular portions straight, to Brazil. Caribbean. about 0.60-0.80 mm. in height and about .06 mm. at the widest part of the apertural FAMILY BUSK, 1854 region. Terminal region about one-half to GENUS CONOPEUM GRAY, 1848 one-third height of tube. No ovicells. Em- bryos brooded in membranous ovisacs which Conopeum tenuissimum (Canu), 1908 are only present at the time of reproduction. Figures 32, 44 OCCURRENCE AND ECOLOGY: Collected in Membranipora tenuissima Canu, 1908, p. 253. November at Sebastian Inlet, colony creeping Electra laciniosa Shier, 1964, p. 612. over the stems of Thyroscyphus ramosus. Not Conopeum laciniosum Cook and Hayward, 1966, reproducing at the time of collection. p. 442. DISTRIBUTION: Worldwide except in polar Conopeum tenuissimum: Dudley, 1973b, p. 284. seas. Western Atlantic: Cape Hatteras to Bra- zil. Gulf of Mexico and Caribbean. DESCRIPTION: (AMNH 584) Colony form- ing a delicate whitish crust on seagrass, shells and other hard substrata. Zooids oval, av- eraging .53 X .25 mm. Each zooid with a Aetea ska (Couch), 1844 short pair of distal spines and with lateral Figures 20, 22 spines running along the edge of the gym- Hippothoa ska Couch, 1844, p. 102. nocyst; number of lateral spines varies, usu- Aetea azorensis Jullien and Calvet, 1903, p. 123. ally from five to seven pairs. No proximal Aetea ska Osburn, 1914, p. 186; 1927, p. 124; spine. Operculum chitinous. Budding pattern 1940, p. 364; 1950, p. 12. Ryland, 1965, p. 18. distal-proximal, primary bud from ances- Prenant and Bobin, 1966, p. 83. Amor and Pal- trula usually distal, and next proximal. lares, 1965, p. 308. Growth of colony proceeding in both direc- Not Aetea ska Canu and Bassler, 1928a, p. 51, pi. tions, with intermediate budding giving rise 1, fig. 2. to a double fan-shaped colony (fig. 44), which Aetea recta Calvet, 1931, p. 71. eventually becomes rounded in form. Polyp- DESCRIPTION: (AMNH 576) Like Aetea ide with 11-13 tentacles (averaging 12) and truncata, colonies consisting of stolons di- lophophore averaging .475 mm. in diameter. lated into zooid bases from which protrude No ovicells, reproduction by means of many erect tubes. Unlike Aetea truncata basal por- small eggs released into sea water through tions marked by many fine annulations (fig. intertentacular organs of polypides and de- 22). Tubular portions of zooids with very veloping into planktonic larvae. slightly expanded apertural regions. Propor- OCCURRENCE AND ECOLOGY: Collected at tion of "head" to "stalk" varying but usually all six stations in the Indian River, at salin- one to three (rather than about one to two as ities varying between 18 and 40 %. Present in A. anguina). Erect portion about 0.1-1.8 throughout the year. Reproduction chiefly in mm. in height. Embryos brooded in external spring and fall. Large old fouled colonies col- ovisacs. lected in late March at the Banana River sta- OCCURRENCE AND ECOLOGY: Occurred tion had areas in which zooids were filled from February to October, most abundant in with storage products and the frontal partially 1982 WINSTON: MARINE BRYOZOANS 117

"closed" or calcified, and regenerating areas spines are fewer than those of C. tenuissimum having new zooids with functional polypides. in water of the same salinity, e.g., one to three The species is able to grow to reproductive for C. seurati vs. five for C. tenuissimum at size in less than a month (Dudley, 1973a), Link Port, and some colonies with no lateral enabling it to live successfully on small and/ spines at all. The budding pattern is not ex- or unstable substrata. Even the narrow blades actly as described for European specimens of of the seagrass Syringodium provided enough C. seurati by Cook and Hayward (1966), but substratum for colonies to grow to a repro- it appears quite similar. Until further infor- ductive stage, as colonies with intertentacular mation on early colony astogeny of American organs were seen on them. specimens can be obtained, it seems best to DISTRIBUTION: Estuaries on the Atlantic identify them with C. seurati. and Gulf coasts of the United States. Also in OCCURRENCE AND ECOLOGY: Collected in some Pacific coast localities, apparently in- the Indian River only from the panels at Link troduced along with from the Gulf Port, but probably widespread within the Coast (Lagaaij and Cook, 1973). river. A winter species in Florida, occurring from December through May. In December IConopeum seurati (Canu), 1908 (when the largest numbers of C. tenuissimum Figures 31, 43 were settling) the ratio of C. tenuissimum to Nitscheina seurati Canu, 1928, p. 26. C. seurati colonies was greater than 99-1, but : Prenant and Bobin, 1966, p. in January the numbers were more equal and 127. Ryland and Hayward 1977, p. 62. by May only seurati was settling. Thus, while the two species occurred in the same habitat DESCRIPTION: (AMNH 583) Colonies during the part of the year their seasons of forming white crusts on seagrasses and other reproduction were offset—the largest peak substrata. Zooids ovoid, averaging .55 X .33 for C. tenuissimum being in late fall—that mm. in size, having pair of long-pointed dis- for C. seurati in late winter-spring. Cono- tal spines. Lateral spines variable in number peum seurati also seemed to be a better space or entirely lacking. No proximal spine. Prox- competitor than C. tenuissimum (its colonies imal budding pattern, unlike other members overgrew those of C. tenuissimum). The of the genus. In colonies from the Indian species occurred here in salinities ranging River the ancestrula first buds proximally from 18 to 40%o. In European waters it occurs and that zooid gives rise to one distal and in salinities down to less than \%, and is a two distqlateral zooids; the distolateral zooids true brackish water bryozoan (Winston, then bud proximally, as well as distally and 1977). laterally, thus forming a circle around the DISTRIBUTION: East coast of Florida (es- ancestrula and ensuring growth in both distal tuarine habitats). Estuarine habitats in Great and proximal directions from the ancestrular Britain, Northern Europe, and Mediterra- region (fig. 43). Polypides with mean tentacle nean. Not previously reported from the number of 15 and mean lophophore diameter Western Hemisphere. of .621 mm. No ovicells. Reproduction by means of non-brooded eggs and planktonic GENUS MEMBRANIPORA BLAINVILLE, 1830 larvae (Cook, 1966). Membranipora arborescens DISCUSSION: Colonies are not easily distin- (Canu and Bassler), 1928 guishable from those of Conopeum tenuis- Figures 25, 26 simum, but because of the slightly larger zooid size and the difference in budding pat- Bijlustra savartii Audouin, Smitt, 1873, p. 20 pi. 4, figs. 92-95 Not savartii Audouin. tern, the lacelike pattern appears slightly Acanthodesia arborescens Canu and Bassler, 1928c, more regular than in C. tenuissimum. Lateral p. 15. walls of zooids are somewhat more calcified Conopeum commensale Marcus, 1937, p. 35, Not than those of C. tenuissimum and the distal Conopeum commensale Kirkpatrick and Metz- spines are longer and more pointed. Lateral elaar. Marcus, 1938, p. 16; 1939, p. 126; 1941, 118 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

FIGS. 25-30. 25. Membranipora arborescens, showing chitinous spinules in frontal membrane. 26. Membranipora arborescens, only. 27. Membranipora tuberculata. 28. Membranipora savartii. 29. Membranipora tenella. 30. Membranipora sp. 1982 WINSTON: MARINE BRYOZOANS 119

p. 16; 1950, p. 30; 1957, p. 37. Lagaaij, 1963, Membranipora savartii (Audouin), 1926 p. 166. Shier, 1964, p. 610. Figure 28 Membranipora arborescens: Cook, 1968a, p. 138; 1968c, p. 121. Flustra savartii Audouin, 1826, p. 240. savartii: Smitt, 1873, p. 20. DESCRIPTION: (AMNH 577, 578) Colonies Acanthodesia savartii: Canu and Bassler, 1928a, white, encrusting, multilaminar, to erect, fo- p. 14. Marcus, 1937, p. 40. Osburn, 1940, p. liaceous, or bilaminar branching. Zooids reg- 352. ularly shaped, subrectangular (distal and Membranipora savartii: Osburn, 1950, p. 27. Ma- proximal walls straight, lateral walls curving), ture, 1957, p. 35. Shier, 1964, p. 670. Long and Rucker, 1970, p. 19. in encrusting colonies about .30 X .25 mm. in size. Opesia oval, cryptocyst granular, nar- DESCRIPTION: (AMNH 579) Colony whit- row, slightly wider proximally than laterally, ish, encrusting, one or more layers in thick- with denticles projecting into the opesia giv- ness. Zooids elongate, rectangular or quad- ing it a serrated edge. Paired gymnocystal tu- rangular, averaging .27 X .55 mm., with a bercles (occasionally very large and heavy) denticulated shelf formed by cryptocyst un- may occur. Zooids outlined by a dark brown der the proximal half of the membranous line. Frontal membrane armed with delicate frontal surface. Denticulation of edge of cryp- chitinous spinules, oriented into the frontal tocyst variable even within colonies. A large membrane from its edge and scattered over proximal often present, sometimes its surface (see fig. 25). Their development having a number of teeth (as in specimen variable, some colonies or areas of a colony illustrated) or teeth almost completely lack- lacking them entirely, in other colonies or ing. Cryptocyst heavily calcified and covered areas of colonies, strongly developed. with fine tubercles, mural rim beaded. No DISCUSSION: This species has long been ovicells or avicularia. confused with Conopeum commensale. Ac- OCCURRENCE AND ECOLOGY: Collected at cording to Cook (1968c) Conopeum com- five stations: Sebastian Grass Flats, Sebastian mensale occurs only in association with gas- Inlet, Walton Rocks, Seminole Shores, and tropod shells inhabited by pagurid crabs, Capron Shoals. Occurred year round, most whereas Membranipora arborescens occurs abundant in April and October. Found on on various substrata, shells, other bryozoans, dead shell and beach rock substrata at all etc., occasionally including shells occupied localities. by pagurids, but not limited to them. Con- DISTRIBUTION: Cosmopolitan in warm opeum commensale is not known from the water. Western Atlantic: Beaufort to Brazil. Western Hemisphere. The presence of a dark Caribbean and Gulf of Mexico. brown line outlining zooids and chitinous spinules on the frontal membrane is char- Membranipora tuberculata (Bosc), 1802 acteristic of encrusting phases of M. arbores- Figure 27 cens. Flustra tuberculata Bosc, 1802, p. 143. OCCURRENCE AND ECOLOGY: Collected in Membranipora tehuelcha Osburn, 1914, p. 193. April at Walton Rocks, encrusting an empty Nitscheina tuberculata: Canu and Bassler, 1928a, gastropod shell, and at Capron Shoals on a p. 18. shell fragment, also in February on a piece Membranipora tuberculata: Marcus, 1973, p. 33. of wire in beach drift on the North Beach, Osburn, 1940, p. 349. Maturo, 1957, p. 33. Fort Pierce. Probably common subtidally Shier, 1964, p. 609. throughout the area. DESCRIPTION: (AMNH 580) Colonies DISTRIBUTION: Western Atlantic: Cape forming white crust which may almost cover Hatteras to Brazil. Gulf of Mexico. Also algae (primarily the gulfweed, Sargassum). known from West Africa and from the Pacific Zooids rectangular, about .24 X .54 mm. in coast of America, Mexico to Equador. size with large membranous frontal area and 120 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173 two tall rounded tubercles at the distal cor- Stream, but M. tenella was never found on ners. In young colonies twinned ancestrula the Sargassum itself. visible, in older colonies origins usually over- DISTRIBUTION: East coast of Florida and grown. Polypide with 15 tentacles and an Brazil. To be expected also wherever Gulf average lophophore diameter of .478 mm. Stream flotsam is cast up. No ovicells. Eggs released through interten- tacular organs into the sea where they de- Membranipora sp. velop into cyphonautes larvae. Figure 30 OCCURRENCE AND ECOLOGY: Found year- round whenever drifting Sargassum (S. na- DESCRIPTION: (AMNH 582) Colonies yel- tans and S. fluviatilis) collected. Also oc- lowish white, encrusting hard substrata cast curred occasionally on attached brown and ashore as beach drift. Zooids rectangular, at Fort Pierce Inlet, Walton Rocks about .50 X .20 mm., distal wall strongly and Seminole Shores. Reproduction not arched, separating lines between zooids. noted in colonies from drifting algae, but a Gymnocyst variable, covering about one- colony from an attached red alga at North fourth of the zooid length, the opesia irreg- Beach breakwater contained many polypides ularly oval. Specimens collected character- with intertentacular organs. No information ized by the presence of broad heavy spines, on reproductive season available, but cy- occurring chiefly on the proximal half of the phonautes larvae have been collected in off- zooid, broad, and unpointed in shape and shore tows throughout the year (M. projecting forward at an angle over the frontal E. Rice, personal commun.). membrane. The ends may be bent, broad- DISTRIBUTION: Circumtropical. Atlantic ened or bifid and some times large and cer- coast, Gulf of Mexico, and Caribbean wher- vicorn. No ovicells or avicularia. ever Sargassum drifts ashore. DISCUSSION: Florida specimens are very similar in appearance to Electra biscuta Membranipora tenella Hincks, 1880 which was described by Osburn (1950) from Figure 29 Mazatlan, Mexico and the Secas Islands, Membranipora tenella Hincks, 1880, p. 376. Panama. The specimens described by Osburn Electra tenella: Marcus, 1937, p. 38. had shieldlike spines bending across the dis- tal end of the operculum. These specimens DESCRIPTION: (AMNH 581) Colonies yel- have some of the shieldlike spines, but they lowish white, encrusting plastic and wood project over the opesia from the sides or objects deposited in beach drift. Zooids rect- proximal end. Osburn's specimens show two angular, about .45-.50 mm. in length, vary- conical spines opposite the distal end of the ing in width between about .18 and .25 mm. operculum. In Florida specimens the distal Opesia elongate-oval, taking up almost all the margin of the opesia has no spines. frontal surface, slightly narrower at the distal OCCURRENCE AND ECOLOGY: Like Mem- end, the rim finely beaded. Gymnocyst very branipora tenella, Membranipora sp. was narrow, with a smooth surface in which the found only on wood and plastic objects in intercalary cuticle is only faintly visible and beach drift. It did not occur on algae. with one or two small (.02-.03 mm.) tuber- DISTRIBUTION: East coast of Florida. cles developed proximally. No spines pres- ent, and specimens lacking the chitinous spi- nules on the frontal membrane and the FAMILY STACH, 1937 cryptocystal denticles characteristic of Mem- GENUS ELECTRA LAMOUROUX, 1816 branipora arborescens. Electra bellula (Hincks), 1881 OCCURRENCE AND ECOLOGY: Found on Figure 33 various wood and plastic substrata in beach Membranipora bellula Hincks, 1881, p. 149. drift deposited by storms. The presence in Electra bellula: Marcus, 1937, p. 37. Osburn, the same drift of pelagic Sargassum spp. and 1940, p. 355.Lagaaij, 1963, p. 170. Shier, 1964, Physalia indicated an origin in the Gulf p. 611. 1982 WINSTON: MARINE BRYOZOANS 121

FIGS. 31-36. 31. Conopeum seurati. 32. Conopeum tenuissimum. 33. Electra bellula. 34. Aplousina gigantea. 35. Parellisina latirostris. 36. Antropora leneocypha, zooids of young colony.

DESCRIPTION: (AMNH 585) Colony en- transparent appearance, uniserial, biserial to crusting, delicately calcined, with a glassy sheetlike. Zooids about. 18 X .34 mm. in size 122 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173 with an oval opesial region and a generally than one series. Polypide with 13-16 tenta- large narrow calcined proximal gymnocystal cles (according to Cook, 1965b). No ovicells. region. A large flattened branching spine pro- OCCURRENCE AND ECOLOGY: Living speci- jecting over the proximal end of the opesia mens collected in 10 m. on a coarse shell- from the calcified region. In addition, usually sand bottom off Ft. Pierce Inlet February two distal spines and often several spines 1975. The tiny of this species also projecting from the proximal gymnocyst. found among the shell fragments that make Polypides with ten tentacles; mean lopho- up the beach sand in the area. Most common phore diameter .225 mm. lunulitiform species in the area, probably OCCURRENCE AND ECOLOGY: Occurred occurring everywhere bottom and depth con- throughout the year in the Indian River. Col- ditions are suitable. lected on seagrass (primarily Syringodium) DISTRIBUTION: Tropical and subtropical at Link Port and at St. Lucie Inlet. Also col- waters of the Atlantic. Western Atlantic: lected in April and July at Walton Rocks on Cape Hatteras to Florida. Gulf of Mexico and brown algae. Reproductive season unknown. Caribbean. DISTRIBUTION: Western Atlantic: Florida to Brazil. Caribbean and Gulf coast of Flor- Discoporella umbellata, subspecies ida. depressa (Conrad), 1841 Figure 24 FAMILY LAGAAU, 1952 Lunulites depressa Conrad, 1841, p. 348. GENUS CUPULADRIA CANU AND Cupularia umbellata Smitt, 1873, p. 14. Canu and BASSLER, 1919 Bassler, 1928a, p. 64. Hastings, 1930, p. 718. Cupuladria doma (d'Orbigny), 1859 Cupularia lowei Osburn, 1914, p. 194. Discoporella umbellata Osburn, 1940, p. 374; Figure 23 1950, p. 113. Maturo, 1957, p. 41. Shier, 1964, p. 621. Cheetham and Sandberg, 1964, p. 1022. Discoflustrellaria doma d'Orbigny, 1851, p. 561. Discoporella umbellata subsp. depressa Cook, Cupularia doma: Smitt, 1873, p. 15. Canu and 1965, p. 180. Bassler, 1923, p. 77; 1928a, p. 64. Discoporella doma: Osburn, 1940, p. 374. Maturo, DESCRIPTION: (AMNH 587) Colony free- 1957, p. 41. Cheetham and Sandberg, 1964, p. living, reddish brown, larger and flatter than 1022. Shier, 1964, p. 621. that of C. doma, cup- or bowl-shaped. Basal Cupuladria doma: Gautier, 1962, p. 54. Cook, surface concave, divided by radial ridges 1965b, p. 216; 1968, p. 145. Prenant and Bobin, bearing tubercles. Zooids rhomboidal, aver- 1966, p. 314. aging .34 X .23 mm., regularly arranged, with DESCRIPTION: (AMNH 586) Colonies cup- a granular cryptocyst formed by fused spi- shaped, pink to reddish brown when living, nous processes dividing up the frontal sur- 4-5 mm. in size, free-living on sandy bot- face into a distal semicircular opesia and a toms, basal surface usually flat, showing ra- proximal area punctured by about six or eight dial ridges with tubercles. Zooids rhomboi- small opesiules. Interzooecial vibraculum at dal, small, about .28 X .18 mm. in size, the distal end of each zooid. Polypides av- regularly arranged. Opesia subtrifoliate, eraging 13 tentacles; mean lophophore di- spade-shaped distally, with lacy proximal ameter .453 mm. No ovicells. portion formed by inward projections of OCCURRENCE AND ECOLOGY: Living colo- granular cryptocyst as two to three pairs of nies collected in 10 m. of water off Fort Pierce denticulate spinules. An interzooecial vibrac- Inlet (in association with Cupuladria doma) ulum distal to each zooid. In mature colonies in February and at Capron Shoals in July. the central and peripheral zooids closed by Common in the same sand bottom environ- continuous granular calcification. Peripheral ments as C. doma, although, according to vibracula large, giving the "cup" a scalloped Maturo (1968), it is less abundant than C. edge, and occasionally occurring in more doma in these environments. 1982 WINSTON: MARINE BRYOZOANS 123

DISTRIBUTION: Primarily a tropical and side of the orifice (see ovicelled zooids, fig. subtropical western Atlantic subspecies: Cape 35). Ovicell hyperstomial with a beaded sur- Hatteras to Florida. Gulf of Mexico and the face. Characterized by the presence of vicar- Caribbean. Also Pacific coast from Point ious avicularia with a short spatulate man- Conception, California to Ecuador. dible, placed at proximal end of an autozooid sized kenozooid. OCCURRENCE AND ECOLOGY: Encrusting FAMILY ALDERINIDAE CANU AND BASSLER, dead shells and Oculina from offshore sta- 1927 tions. Colonies with ovicells present (Sep- GENUS APLOUSINA CANU AND BASSLER, 1927 tember). DISTRIBUTION: Florida, Caribbean, Gulf of Aplousina gigantea Canu and Bassler, 1927 Mexico. Figure 34 Aplousina gigantea Canu and Bassler, 1927, p. 3; GENUS ANTROPORA NORMAN, 1903 1928a, p. 20. Osburn, 1940, p. 357. Maturo, 1957, p. 38. Shier, 1964, p. 612. Antropora leucocypha (Marcus), 1937 Figures 36, 37 DESCRIPTION: (AMNH 588) Colonies en- crusting on shell or Oculina rubble. Zooids Crassimarginatella leucocypha Marcus, 1937, p. large, measuring about .80X1.00 mm., 46; 1938, p. 20. Cheetham and Sandberg, 1964, p. 1017. ovoid to hexagonal in shape, frontal surface Conopeum reticulum (in part) Osburn, 1940, p. membranous, edged by a narrow low-beaded 351. cryptocyst and an occasional narrow proxi- Antropora leucocypha: Shier, 1964, p. 613. mal gymnocyst. Ovicell endozooidal, a small hood-shaped structure formed by the exten- DESCRIPTION: (AMNH 590, 591) Colony sion of the distal zooid wall under the gym- encrusting, grayish white to pink or magenta nocyst and cryptocyst of the distal zooid. in color, uni- to multilaminar, encrusting OCCURRENCE AND ECOLOGY: Collected only various substrata, most commonly gastropod at offshore stations. One of most abundant shells. Zooids irregularly oval, narrowing a species on dead Oculina and shell at 80-90 bit distally, about .30-35 mm. in length by m. depth. Colonies with ovicells were col- .20-.25 mm. width. Gymnocyst hardly de- lected in February. veloped, cryptocyst very narrow distally and DISTRIBUTION: Western Atlantic: Beaufort laterally, slightly wider proximally, closely to Florida. Caribbean. Gulf of Mexico. beaded and crenulate. Opesia also narrow distally, wider proximally. Triangular spaces between autozooids, filled by kenozooids, FAMILY ALDERINIDAE CANU AND BASSLER, deposits with central openings, or 1927 solid tubercles, some transformed into avic- GENUS PARELLISINA OSBURN, 1940 ularia. Zooids illustrated (fig. 36) surrounded Parellisina latirostris Osburn, 1940 by many small kenozooids, as is often the Figure 35 case, but kenozooids and avicularia also often absent from large areas of colonies. Parellisina latirostris Osburn, 1940, p. 361. La- Zooids quite variable in shape, and in sec- gaaij, 1963, p. 175. Long and Rucker, 1970, p. 19. ondary layers irregularly oriented. Polypides transparent to whitish in color; mean tentacle DESCRIPTION: (AMNH 589) Colony en- number 12, mean lophophore diameter .326 crusting. Zooids oval to irregular in shape, mm. Ovicell endozooidal. Opercula variable .45 to .55 mm. in length and .30 to .45 mm. in size in both sterile and fertile zooids, and in width. Opesia filling most of frontal area, fertile zooids thus difficult to detect. a beaded mural rim marking the cryptocyst. DISCUSSION: Antropora leucocypha is very Small spines occasionally occurring on either similar to A. tincta and, according to Cook 124 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

FIGS. 37-42. 37. Antropora leucocypha, zooids of an old multilamellar colony. 38. Floridina antiqua. 39. Micropora coriacea. 40. Floridinella typica. 41. Thalamoporella floridana. 42. Thalamoporella fal- cifera. (1968), can only be distinguished from it by the size of the avicularia, and the shape of the frequency of occurrence of kenozooids, their mandibles. Variation observed in Flor- 1982 WINSTON: MARINE BRYOZOANS 125 ida specimens included these differences as straight or only slightly curved in comparison well as differences in texture of the crypto- with those of autozooids. cyst. Smaller (younger ?) unilaminar colonies OCCURRENCE AND ECOLOGY: One of the (generally found on gastropod shells inhab- most common species from shell and Oculina ited by gastropods) resembled the description rubble in Gosnold collections in September of Antropora leucocypha while large multi- and February. None of these colonies showed laminar colonies on gastropod shells inhab- zooids with elongated opesiae. ited by hermit crabs approached the descrip- DISTRIBUTION: A warm water species tion of Antropora tincta (fig. 37). known from both the Atlantic and the Pa- OCCURRENCE AND ECOLOGY: Common in- cific. Western Atlantic: Cape Hatteras to tertidally at all coastal stations throughout Florida. Caribbean and Gulf of Mexico. the year. Although occasionally found on dead shells, colonies usually encrusted the GENUS FLORIDINELLA CANU AND shells of living Thais haemastoma floridana BASSLER, 1917 living on and under breakwater rocks, beach rock ledges, and sabellariid mounds, or dead Floridinella typica Canu and Bassler, 1928 Thais shells inhabited by hermit crabs. Figure 40 DISTRIBUTION: Western Atlantic: Cape Floridinella typica Canu and Bassler, 1928a, p. Hatteras to Brazil. Caribbean and Gulf of 372. Mexico. DESCRIPTION: (AMNH 594) Colonies en- crusting. Zooids irregularly oval with proxi- FAMILY ONYCHOCHELLIDAE JULLIEN, 1882 mal end elongated, about .55-60 mm. in GENUS FLORIDINA JULLIEN, 1881 length by .30-. 3 5 mm. in width, separated Floridina antiqua (Smitt), 1873 from each other by a distinct groove. Opesia Figure 38 triangular, proximal edge serrate, surrounded by a fiat, oval, or diamond-shaped, coarsely Mollia antiqua Smitt, 1873, p. 12. beaded cryptocyst area. Edge of cryptocyst Floridina antiqua: Canu and Bassler, 1928a, p. 60. finely beaded, meeting the rim of the zooids Hastings, 1930, p. 715. Osburn, 1940, p. 372; laterally, but having below its proximal bor- 1950, p. 102. Shier, 1964, p. 617. Cook, 1964a, der an area of gymnocyst often raised into a p. 70; 1968a, p. 151. large tuberosity deceptively like an ovicell. Ovicells endozooidal. No avicularia. DESCRIPTION: (AMNH 582) Colonies en- OCCURRENCE AND ECOLOGY: Collected in crusting, usually unilaminar. Zooids subhex- September and February encrusting shell and agonal (distal ends rounded, proximal cor- Oculina rubble at continental shelf stations. ners sharp) about .40 mm. in length by .40 Also encrusting a dead shell fragment at Cap- to .35 mm. in width. Opesia trifoliate with ron Shoals in October. No information on laterally directed opesiular indentations and reproduction. a curved proximal edge, rest of frontal mem- DISTRIBUTION: East and West coast of Flor- brane covered by a granulated cryptocyst, ida. depressed in the center, and with a raised and more closely granulated margin. Distinct FAMILY GRAY, 1848 grooves between zooids. Vicarious avicularia GENUS MICROPORA GRAY, 1848 scattered among the zooids. These are hex- agonal structures almost as long as auto- Micropora coriacea (Johnston), 1847 zooids but narrower and very pointed at the Figure 39 distal end with dark brown winged mandibles Flustra coriacea Johnston, 1847, p. 349. which are hooked at the end (specimen il- Micropora coriacea: Smitt, 1973, p. 13. Canu and lustrated shows skeleton only, mandibles not Bassler, 1928a, p. 62. Hastings, 1930, p. 719. present). Ovicell endozooidal, opesiae offer- Osburn, 1940, p. 373; 1950, p. 105. Marcus, tile zooids elongated with proximal edge 1949, p. 16; 1953, p. 285. Prenant and Bobin, 126 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

1966, p. 332 (part). Ryland and Hayward, 1977, perforate and raised into adoral tubercles. p. 113. One of opesiules usually larger than other and DESCRIPTION: (AMNH 593) Colonies en- bounded by a descending tubular portion of crusting. Zooids regularly arranged, very cryptocyst continuing to the basal wall of the wide, rhombic in shape, about .50 mm. long zooid. Interzooecial avicularia shaped like by .40-50 mm. wide, separated by a shallow gothic arches; the point of the mandible usu- groove. Frontal membrane flat proximally ally bent. Spicules in the form of large and and concave distally where lateral walls rise small calipers occurring in the . to a smooth mural rim. Curving epaulets on (See specimen illustrated.) No ovicells. either side of semicircular orifice. Cryptocyst DISCUSSION: Osburn (1940) described finely beaded and penetrated by numerous Thalamoporella floridana as a subspecies of small round perforations. Ovoid opesiules Thalamoporella gothica. Thalamoporella placed proximal and lateral to the orifice just gothica was originally described by Busk below "shoulders" of zooid. Ovicell subim- from the Pacific coast of Mexico. This species mersed, with a beaded imperforate roof has a large bilobed ovicell which may brood formed from cryptocyst and triangular center several embryos at a time. Ovicells have from gymnocyst of the distal zooid, and a never been reported from Atlantic speci- smooth surfaced inner layer surrounding the mens. There are other differences: Atlantic zooid operculum which closes the ovicell. specimens seem always to have adoral tu- Embryos orange-red. bercles, whereas west coast material has OCCURRENCE AND ECOLOGY: Found only wider opesiae and no adoral area. Spicules at offshore stations encrusting shell and Ocu- are variable. Spicules in the form of open lina rubble. Colonies with ovicells containing compasses occur in T. gothica. Only calipers embryos in February. have been described in Florida and Puerto DISTRIBUTION: Widespread in temperate Rico specimens, but a few spicules in the seas. Western Atlantic: Cape Hatteras to Bra- form of open compasses were found in Beau- zil. Caribbean and Gulf of Mexico. fort area specimens. Until more is known about this species it seems preferable to sep- arate it from T. gothica. FAMILY OCCURRENCE AND ECOLOGY: Occurred LEVINSEN, 1909 year-round at Fort Pierce Breakwater. Most GENUS THALAMOPORELLA HINCKS, 1887 abundant species encrusting the stems of the Thalamoporella floridana Osburn, 1940 hydroid Thyroscyphus ramosus in winter Figure 41 months; colonies less abundant in summer. Those collected in October had inner sections Thalamoporella rozierii Osburn, 1914, p. 196. Thalamoporella gothica new var. floridana Os- filled with brown bodies, but a newly budded burn, 1940, p. 378. outer zone. Also collected encrusting red al- Thalamoporella gothica var. floridana: Mature, gae at Walton Rocks, and at Sebastian Inlet 1957, p. 40. where large flower-like masses were found Thalamoporella gothica: Shier, 1964, p. 620. attached to breakwater rocks in January. Pol- ypides of a colony collected in January had DESCRIPTION: (AMNH 595) Colonies en- large bulbous intertentacular organs. crusting sheets, sometimes rising into erect DISTRIBUTION: Cape Hatteras to Florida. bilaminar frills or rosettes, grayish white in Gulf of Mexico and the Caribbean. color. Zooids quadrangular in shape, variable in size, about .45-.50 mm. long X .20-25 Thalamoporella falcifera (Hincks), 1880 mm. wide, separated by distinct grooves. Figure 42 Frontal membrane covered by light perforate Steganoporella rozieri forma falcifera Hincks, calcification except for the distal opesia and 1880, p. 380. two large rounded opesiules proximal to it. Thalamoporella falcifera: Osburn, 1914, p. 197. Opesia hoof-shaped. Area on either side im- Marcus, 1937, p. 52. Mature, 1957, p. 40. 1982 WINSTON: MARINE BRYOZOANS 127

DESCRIPTION: (AMNH 596) Glassy white substrata. Zooids in pairs, back to back along colonies encrusting benthic or drifting algae. the branches, with a joint between each pair. Zooids elongate, quadrangular, averaging Zooids scoop-shaped, averaging .24 X .10 .61 X .26 mm., with distinct grooves between mm. in size, thinly calcified, with a large fron- zooids and a delicately tuberculate and per- tal area. Two types of avicularia: box-shaped forate cryptocyst covering all but the area of sessile avicularia in one or both distal corners opesiules and opesia. Like Thalamoporella of zooids, and sometimes also bulbous pe- floridana this species has a hoof-shaped ope- dunculate avicularia on the dorsal surface sia with rounded tubercles on either side. (shown in colony illustrated). Attachment Avicularia interzooidal, about two-thirds the radicles may have ends elaborated into hook- size of zooids, with elongate, curved man- like processes to grapple the substratum. No dibles. Spicules in the form of compasses. ovicells. Embryos brooded in modified (en- Ovicells very large and prominent, consisting larged) individuals (gonozooids). Polypides of two hemispherical valves joined at a me- with ten tentacles, and a lophophore aver- dian suture. Polypides with a mean tentacle aging .231 mm. in diameter. number of 16, and a lophophore averaging OCCURRENCE AND ECOLOGY: Occurred .460 mm. in diameter. year-round at all coastal stations. Also col- OCCURRENCE AND ECOLOGY: Reported only lected in April and October at Capron Shoals from algae. In the Indian River area collected (10 m.). Found primarily on hydroid stems on Sargassum fluitans only, in deep water (both Thyroscyphus and Eudendrium) but algae collections (Winston and Eiseman, also occurred on shell fragments and beach 1980) also found on three species of benthic rock. Zooids containing present in algae. Occurred year-round, but most abun- July. Colonies largest and most abundant dant during the fall when almost every plant from October to December. Synnotum is a of Sargassum fluviatilis cast ashore was en- warm water species, and while it can be col- crusted. Membranipora tuberculata some- lected all year-round in the Indian River area, times also encrusted 51. fluviatilis but T. fal- colonies may overwinter in a dormant con- cifera was not found on S. natans, the most dition. Those collected at Sebastian Inlet in common species of drift Sargassum and the January after a cold spell had lowered water one usually encrusted by M. tuberculata. temperatures, contained only brown bodies Ovicells very rare. Some found in collections in distal zooids, and starchy-looking granular from continental shelf waters. material in proximal zooids. DISTRIBUTION: Western Atlantic: Beaufort DISTRIBUTION: Distributed around the to Brazil. Caribbean. Also reported from world in warm water. Western Atlantic: Cape Ceylon, Java, and Australia. Hatteras to Brazil. Caribbean and Gulf of Mexico. FAMILY GREGORY, 1893 GENUS SYNNOTUM PIEPER, 1881 FAMILY SCRUPOCELLARIIDAE LEVINSEN, 1909 Synnotum aegyptiacum (Audouin), 1826 GENUS SCRUPOCELLARIA VAN BENEDEN, 1845 Figure 53 Scrupocellaria regularis Loricaria aegyptica Audouin, 1826, p. 243. Osburn, 1940 Synnotum aviculare Osburn, 1914, p. 191. Figure 54 Synnotum aegyptiacum: Osburn, 1927, p. 126; Cellularia cervicornis Smitt, 1872, p. 14. Osburn, 1940, p. 42; 1950, p. 151. Marcus, 1937, p. 58; 1914, p. 192 (not Scrupocellaria cervicornis 1938, p. 26. Mature, 1957, p. 42. Shier, 1964, Busk, 1852). p. 622. Scrupocellaria regularis Osburn, 1940, p. 384; 1950, p. 144. Cheetham and Sandberg, 1964, DESCRIPTION: (AMNH 597) Delicate erect p. 1025. Shier, 1964, p. 622. branching and stolonate colonies resembling tiny strings of glass beads, entangled among DESCRIPTION: (AMNH 598) Colony erect, hydroid stems and algae or attached to other jointed, branching, forming short pinkish tan 128 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

10.5 mm |

FIGS. 43-52. 43. Conopeum seurati, initial growth pattern. 44. Conopeum tenuissimum, initial growth pattern. 45. Beania mirabilis. 46. Beania klugei. 47. Entire colony of Bugula minima attached to algal surface. 48. Type 3, branching pattern of Bugula sp. 49. Type 4, Branching pattern, Bugula stolonifera. 50. Bugula turrita. 51. Bugula stolonifera. 52. . (figs. 48 and 49, after Maturo, 1966).

tufts on various substrata. Zooids narrow frontal area. Scutum large, almost covering proximally and wider distally, opesia ovoid, the opesia, and paddle-shaped with pointed occupying about two-thirds of frontal area. edges and symmetrical alcicorn decoration Zooids averaging .51 mm. in length by .20 (not visible in SEM). Polypides with obliquely mm. in width. Two to seven simple or truncate lophophores, averaging .464 mm. in branched distolateral spines around mural diameter, with 13 translucent pink tentacles. rim. Innermost proximal spine specialized as Avicularia in two locations: on inner sides a shield or scutum bent over membranous of gymnocysts and on outer sides of zooids 1982 WINSTON: MARINE BRYOZOANS 129 at the level of the opesia. Vibracula delicate, ter and early spring (Mook, 1976). In more relatively short, placed on the proximal part temperate waters Bugula neritina reproduces of the back surface of each zooid. Ovicells in summer and fall, and old colonies may hyperstomial, flat at the top, with a few large overwinter. In the Indian River area repro- pores. Embryos pinkish red. duction occurs in the cooler months and old OCCURRENCE AND ECOLOGY: Occurred colonies appear to "over summer." Colonies year-round at coastal stations: Sebastian In- collected at Walton Rocks in June and July let, Fort Pierce Breakwater, Walton Rocks, were rusty brown in color and almost com- and Capron Shoals. Also found in March at pletely degenerated, with many zooids dead the Sebastian Grass Flats on Solieria. Young and filled with , but with functional colonies found from January to March. Col- polypides still present at the most distal ends onies containing embryos and larvae col- of branches. Cold can kill Bugula neritina, lected in March and April. also, especially exposure to cold air. Large DISTRIBUTION: Western Atlantic: Ber- intertidal colonies collected at Sebastian Inlet muda, Cape Hatteras to Florida. Gulf of after a cold spell were dead, whereas subtidal Mexico and the Caribbean. Also, Gulf of colonies were unaffected. California. DISTRIBUTION: One of the most abundant and troublesome fouling organisms, widely FAMILY GRAY, 1848 distributed throughout warmer waters of the GENUS BUGULA OKEN, 1815 world, especially in harbor and ports. Bugula neritina (Linnaeus), 1758 Figure 52 Bugula stolonifera Ryland, 1960 Figures 49 and 51 Sertularia neritina Linnaeus, 1758, p. 815. Bugula neritina: Osburn, 1914, p. 186; 1927, p. 1'Bugula californica Marcus, 1937, p. 71. 126; 1940, p. 389. Marcus, 1937, p. 67. Shier, Bugula californica Maturo, 1957, p. 45. 1964, p. 624. Ryland, 1965, p. 45. Maturo, Wugula avicularia Shier, 1964, p. 624. 1966, p. 560. Cook, 1968a, p. 157. Bugula stolonifera Ryland, 1960, p. 78; 1965, p. 50. Maturo, 1966, p. 568. Cook, 1968a, p. 160. DESCRIPTION: Erect branching colonies forming brownish or reddish purple tufts on DESCRIPTION: Colonies grayish tan, erect any possible substratum. Zooids alternating and branching, forming a fan or funnel biserially up the branches, large, averaging (young colonies) or a dense tuft (older colo- .97 X .28 mm. in size, tapering proximally, nies). Smaller than the colonies of Bugula and with frontal membrane taking up whole neritina, usually 3-4 cm. in height. Zooids frontal surface. No avicularia and no spines, long and slender, tapering proximally, aver- although outer distal corners of the zooids aging .78 X .19 mm. in size, and with sharply pointed. Polypides also large; mean U-shaped frontal membrane taking up three- tentacle number 23, and mean lophophore quarters or more of frontal surface. Branch- diameter .764 mm. Ovicells large and glob- ing pattern predominantly type four (see fig. ular, attached to distal corners of zooids; 49). Outer distal corner of zooid elongated oriented at an angle to the branch axis. Em- into a spine, occasionally very large, with one bryos dark brown, about .25 mm. in diam- or two spines below this, and one spine at the eter. inner distal corner. Pedunculate bird's head OCCURRENCE AND ECOLOGY: In the Indian avicularia from one-quarter to one-half the River collected on seagrasses, shells, way down lateral edge of frontal membrane. docks and canal walls, floats, rotting wood, Avicularia with rounded heads, decurved and algae. Also found at coastal stations (Se- beaks, and according to Maturo (1966) oc- bastian Inlet and Walton Rocks) and at Cap- curring in three size classes, large, medium, ron Shoals. Settlement on panels and bryo- or small, depending on their position with zoan traps at Link Port from January through respect to branch bifurcations. Polypides mid-June with peak of settlement in late win- with an obliquely truncate lophophore av- 130 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173 eraging .441 mm. in size and 14 tentacles. tached to undersides of beach rock ledge. Ovicells subglobular; embryos yellow-brown. Maturo (1966) noted that specimens from the OCCURRENCE AND ECOLOGY: Also a com- south coast were smaller and less robust than mon fouling species. Collected in December those from New England. The specimens at Sebastian Inlet and January at Sebastian from Walton Rocks were also small (less than Inlet and Capron Shoals. Settled on panels 2.5 cm. in height). Colonies collected in April at Link Port from December through April contained ovicells with embryos. (Mook, 1976). Colonies commonly occurred DISTRIBUTION: Massachusetts to Florida. in association with Bugula neritina, attached Also reported from Brazil (Marcus, 1937), to the proximal portions of the B. neritina but according to Maturo (1966) that record colonies. It is less tolerant of warm temper- may refer to closely related species. atures than B. neritina. No young colonies were seen in the river after early May, and Bugula minima Waters, 1909 no oversummering colonies were found. Figure 47 DISTRIBUTION: Eastern and Western Atlan- Bugula neritina var. minima Waters, 1909, p. 136. tic and Mediterranean. Western Atlantic: Osburn, 1914, p. 187. Hastings, 1930, p. 704. Massachusetts to Florida, Gulf of Mexico, Bugula minima: Osburn, 1940, p. 390; 1950, p. Brazil. 155. DESCRIPTION: Colony brownish red, re- Bugula turrita (Desor), 1848 sembling a miniature Bugula neritina colony. Figure 50 First two zooids budded uniserially, rest of Cellularia turrita Desor, 1848, p. 66. colony consisting of biserial branches. Zooids IBugula turrita: Marcus, 1937, p. 68. with frontal membrane covering most of Bugula turrita: Mature, 1957, p. 123; 1966, p. frontal surface, outer distal end a sharp angle. 563. Avicularia rather elongate, with a long beak, DESCRIPTION: Erect, branching colonies, attached at proximal end of zooid. with biserially arranged zooids, main OCCURRENCE AND ECOLOGY: Collected at branches having secondary branches ar- two stations: Fort Pierce Breakwater and ranged in a spiral pattern about the long axis. Seminole Shores, from January until June. Branch bifurcations type three (fig. 48). Found on algae, Thyroscyphus, sponges, and Zooids averaging .58 X .17 mm. tapering Amathia distans. proximally, frontal membrane taking up DISTRIBUTION: Probably circumtropical. about two-thirds of their frontal length. Dis- Western Atlantic: East coast of Florida, Tor- tal spine formula 2:1, the frontal outer distal tugas, Brazil, and Caribbean. spine usually very short, the outer dorsal one may be very long. Inner distal spines also Bugula uniserialis Hincks, 1884 sometimes well developed, although those of Indian River specimens quite short. Polyp- Bugula uniserialis Hincks, 1884, p. 367. Hastings, 1930, p. 705. Marcus, 1937, p. 72. ides with an obliquely truncate lophophore averaging .393 mm. in diameter and 14 ten- DESCRIPTION: A tiny delicate, straw-col- tacles. Avicularia located about halfway ored species, similar to Bugula minima, but down the frontal margin, about .15 mm. in lacking its red coloration. Zooids alternating, length, with a decurved beak. Ovicells posi- proximal portion very narrow, frontal mem- tioned slightly to one side of the center of the brane taking about half zooid length, outer zooid and tilted toward the axis of the branch. distal edge sharply angular but not developed Ovicells containing embryos globose. Devel- into a spine. Bird's beak avicularia attached oping or senescent ovicells shallow caplike by a peduncle to the most proximal portion structures. of the zooid. Polypides with 16 tentacles. OCCURRENCE AND ECOLOGY: Collected in Ovicells described but not occurring in our April and November at Walton Rocks, at- specimen. 1982 WINSTON: MARINE BRYOZOANS 131

OCCURRENCE AND ECOLOGY: Collected at FAMILY CANU AND BASSLER, 1927 Seminole Shores in October on a red alga GENUS BEANIA JOHNSTON, 1840 (Laurencia sp.) attached to worm reef. Pol- Beania hirtissima (Heller), 1867 ypides with Bugula-type behavior, though Figure 56 they scan more actively than those of species with large colonies. Hastings (1930) de- Diachoris hirtissima Heller, 1867, p. 94. scribed zooids containing eggs and sperm. Beania hirtissima: Marcus, 1937, p. 62. Osburn, The colony collected here was not reproduc- 1940, p. 397; 1950, p. 579. Lagaaij, 1963, p. ing. 180. Maturo, 1966, p. 579. DISTRIBUTION: Western Atlantic: East coast of Florida and Brazil. Also Galapagos, South- DESCRIPTION: (AMNH 600) Colonies ern California, and West Australia. A very forming spiny brownish mats attached to inconspicuous species generally associated hydroid stems or encrusting flat surfaces. with algae. Its distribution may be more Zooids shaped like split peanut kernels, av- widespread. eraging .64 X .29 mm. in size. Frontal surface flat, slightly constricted below the operculum GENUS CAULIBUGULA VERRILL, 1900 and bristling with spines around its margins. Eight to 10 spines radiating out around the Caulibugula pearsei Maturo, 1966 orifice, eight to 14 curving over the rest of Figure 55 the frontal membrane, and about 10 more Caulibugula sp. Osburn, 1951, p. 137. projecting outward from lateral walls below Caulibugula pearsei Maturo, 1966, p. 575. frontal membrane. Zooids closely joined to each other, six tubular processes connecting DESCRIPTION: (AMNH 599) Colonies, up each one with the six adjacent zooids. Polyp- to 6 cm. in height, composed of fan-shaped ides with 18 tentacles and lophophores av- branches of zooids arising from a stalk of eraging .520 mm. in diameter. No ovicells kenozooids. Kenozooids forming stalks or avicularia. 1.5-3.6 mm. long and .19-40 mm. wide, OCCURRENCE AND ECOLOGY: Occurred with annulations at proximal end. Branch year-round at Fort Pierce Breakwater, Wal- zooids biserial, turned so that alternating ton Rocks, and Seminole Shores. An impor- zooids face each other, scoop-shaped, aver- tant component of the Thyroscyphus stem- aging .40 mm. in length by .21 mm. in width, and-root-mat fauna. Most plentiful in August with a U-shaped frontal membrane taking up when many other species were absent from about three-quarters of the frontal surface. the actively growing Thyroscyphus. Also The semicircular distal area around the ori- found at Walton Rock encrusting beach rocks fice bearing from one to four spines, depend- partially buried in sand. Surfaces of these ing on the position of the zooid on the branch. matlike loosely adhering colonies almost Spines very long (those in the specimen il- completely coated with sand grains, making lustrated are broken off short), curving over them difficult to recognize as living organ- the frontal surface of the branch. Small pe- isms. Reproductive season unknown. dunculate avicularia sometimes present near DISTRIBUTION: Widely distributed in warm proximal edge of frontal membrane. Polyp- waters. Western Atlantic: Cape Hatteras to ides with 12 tentacles and an obliquely trun- Brazil. Caribbean and Gulf of Mexico. cate lophophore averaging .277 mm. in di- ameter. No ovicells observed. OCCURRENCE AND ECOLOGY: Collected at Beania klugei Cook, 1968 Walton Rocks in May attached to undersur- Figure 46 face of beach rock ledge and entangled with Beania intermedia Osburn, 1914, p. 189; 1940, p. Bowerbankia imbricata and Sundanella si- 398. Hastings, 1930, p. 705. Shier, 1964, p. 624. bogae. Maturo, 1966, p. 579. DISTRIBUTION: Cape Hatteras to Florida. Beania klugei Cook, 1968a, p. 164. 132 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

E'

FIGS. 53-58. 53. Synnotum aegyptiacum. 54. Scrupocellaria regularis. 55. Caulibugula pearsei. 56. Beania hirtissima. 57. Membraniporella aragoi. 58. Cribrilaria radiata.

DESCRIPTION: Colonies uniserial, vinelike, icles. Zooids boat-shaped, thinly calcified, or semi-erect. Attached to substratum by rad- averaging .73 X .30 mm. in size, with flat 1982 WINSTON: MARINE BRYOZOANS 133 frontal membrane and rounded abfrontal DESCRIPTION: Colony uniserial, vinelike, surface, proximal end a narrow tube by which irregularly branching, adhering by radicles to the zooid was budded from the zooid prox- hydroids, algae, and other bryozoans. Zooids imal to it. Two minute distal projections at boat-shaped, lenticular in outline, about .6 distal end above operculum. On either side by 15 mm., frontal membrane flat. One erect of operculum usually a stalked "bird's beak" spine on either side of the operculum, plus avicularium. Polypides with average lopho- one or two spines on the distal margin, and phore diameter of .657 mm., and 26 tenta- five to 11 pairs of lateral spines overarching cles. No spines. No ovicells. the frontal membrane. Zooids suberect; each TAXONOMIC DISCUSSION: Cook (1968a) has connected to the next by a stolon-like elon- differentiated this species from Beania inter- gation of the basal portion of the zooid. No media which has smaller zooids, distal and ovicells. No avicularia. Attachment radicles lateral spines, larger avicularia, and lateral on abfrontal side of proximal end of zooid. tubules originating at the mid-zooid level. All OCCURRENCE AND ECOLOGY: Collected at the specimens listed in the synonymy above Sebastian Inlet, Fort Pierce Breakwater, and were found to be referable to B. klugei. NOTE: Seminole Shores among the hydroid roots. Indian River specimens of B. klugei have Also collected at Capron Shoals (10 m.) on been listed as B. intermedia in Winston a dead shell. One degenerating colony was (1978) and Winston and Eiseman (1980). found in August. It probably occurs primarily OCCURRENCE AND ECOLOGY: Common at in fall and winter months, but is a very in- both river and coastal stations, where the tiny conspicuous species and might often be over- colonies occurred primarily on hydroid roots looked. Embryos probably develop inside the and algae. An important component of the zooids; there is no information available on Thyroscyphus community at the Fort Pierce its reproduction. Breakwater and found entangled with hy- DISTRIBUTION: May be worldwide in warm droids and algae at the other coastal stations and temperate waters. Not often recorded. as well. Also common on the drift alga Soliera Western Atlantic: Florida to Brazil. Gulf of tenera in Indian River seagrass beds, and col- Mexico and the Caribbean. lected once (in October) at Capron Shoals (10 m.) on dead shell. B. klugei occurred year- FAMILY HINCKS, 1880 round. Most abundant in the winter and GENUS CRIBRILARIA CANU AND spring. Degenerating colonies found at Se- BASSLER, 1928 bastian Inlet in January, however, after a cold spell brought the water temperature below Cribrilaria radiata (Moll), 1803 15° C. Many colonies found between the end Figure 58 of August and December lacked the paired Eschara radiata Moll 1803, p. 63. avicularia. The significance of this is not Cribrilina radiata: Smitt, 1873, p. 22. known. Puellina radiata: Canu and Bassler, 1928a, p. 73. DISTRIBUTION: Probably widely distrib- Osburn, 1940, p. 406. uted in tropical and subtropical waters. West- Colletosia radiata: Marcus, 1937, p. 73. Maturo, 1957, p. 48. Shier, 1964, p. 625. ern Atlantic: Cape Hatteras to Florida. Ca- Cribrilaria radiata: Cheetham and Sandberg, ribbean and Gulf of Mexico. 1964, p. 1026. Cook, 1967, p. 333; 1968a, p. 172. Long and Rucker, 1970, p. 19. Beania mirabilis Johnston, 1840 DESCRIPTION: (AMNH 602) Colonies en- Figure 45 crusting, inconspicuous, pink to white in Beania mirabilis Johnston, 1840, p. 272. Osburn, color, but living colonies often fouled green 1914, p. 189; 1940, p. 398; 1950, p. 170. Has- by algae. Zooids irregularly ovoid in shape, tings, 1930, p. 705. Marcus, 1937, p. 60. Pren- separated by deep grooves, about .45 X .35 ant and Bobin. 1966, p. 552. Cook, 1968a, p. mm. in size. Frontal surface made of five to 163. Ryland and Hayward, 1978, p. 150. 12 pairs of radiating costae separated by rows 134 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173 of evenly spaced lacunae. Usually a some- OCCURRENCE AND ECOLOGY: Collected only what larger pore just below the orifice. There once at Gosnold station 696 encrusting dead may be tubercles on the outer ends of the shell. Ovicells present at time of collection costulae. Orifice semicircular, surrounded by (September). five to seven distal spines and may have a DISTRIBUTION: Western Atlantic: Cape small proximal umbo. Ovicell helmet-shaped, Hatteras to Florida. Also reported from the imperforate, with several bumps or a keel on Red Sea and the East Indies. its frontal surface. Embryos rose-pink in color. Interzooecial avicularia with long pointed mandibles in a channeled rostrum GENUS REGINELLA JULLIEN, 1886 may be present. Reginella floridana (Smitt), 1873 OCCURRENCE AND ECOLOGY: Collected in Figure 59 September and February at Gosnold stations Cribrilina figularis var. floridana Smitt, p. 1873. where it was common encrusting shell and Cribrilina floridana Osburn, 1914, p. 195. Oculina rubble. Ovicells contained embryos Puellina floridana: Canu and Bassler, 1928a, p. at the time of collection in February. Ovi- 74. celled specimens also collected at Capron Pelmatopora {sensu latd) apsata Shier, 1964, p. Shoals in October. 626. DISTRIBUTION: Cosmopolitan. Western Reginella floridana: Cheetham and Sandberg, Atlantic: Cape Hatteras to Brazil. Caribbean 1964, p. 1026. and Gulf of Mexico. DESCRIPTION: (AMNH 603) Colony en- crusting. Zooids irregularly oval in shape, GENUS MEMBRANIPORELLA SMITT, 1873 about .30 X .45 mm. in size, frontal shield Membraniporella aragoi (Audouin), 1826 made up of five to seven pairs of radially Figure 57 arranged costae with rows of small lumen pores separating them, the outer pores es- Flustra aragoi Audouin 1826, p. 240. pecially may develop into hollow tubules. Membraniporella aragoi Harmer, 1926, p. 473. Not Membraniporella aragoi Marcus, 1938, p. 30. Orifice semicircular with two erect bifid or trifid distal spines and two flattened lateral DESCRIPTION: (AMNH 601) Colonies en- spines which arch over the orifice, joining at crusting. Zooids about .35 X .50 mm. in size, its center. Ovicell unknown. No avicularia. with a small cryptocyst and gymnocyst and DISCUSSION: Shier (1964) described a new a large oval frontal membrane. Spines (usu- species Pelmatopora (sensu lato) apsata, ally three or four per side) arising outside based on material from the Gulf coast of mural rim, curving over frontal membrane, Florida, but his figure shows quite clearly its branching once or twice at the tips which identity with Reginella floridana. interlock like clasped hands over the center OCCURRENCE AND ECOLOGY: Collected at of the zooid. Non-ovicelled zooids with two Gosnold station 696 and 702 encrusting dead to four hollow straight or branching spines shell and Oculina. at distal end. Only one pair of spines showing DISTRIBUTION: Cape Hatteras to the Tor- in ovicelled zooids. Ovicells hemispherical tugas. Northwest gulf coast of Florida. and heavily calcified, with lunate inner area and raised lip. No avicularia. GENUS BELLULOPORA LAGAAIJ, 1963 TAXONOMIC NOTE: Cook (1967) has sepa- Bellulopora bellula (Osburn), 1950 rated M. aragoi from Membraniporella mar- cusi on the basis of number and form of the Colletosia bellula Osburn, 1950, p. 188. Bellulopora bellula: Lagaaij, 1963, p. 183. spines. Spines of M. marcusi (=M. aragoi of Marcus 1938, p. 30) show third or fourth DESCRIPTION: Colonies encrusting, only a order branching, those of M. aragoi have few millimeters in size. Zooids ovoid, sepa- usually only a single branch. rated by deep grooves, about .35 to .50 mm. 1982 WINSTON: MARINE BRYOZOANS 135

...... * * • » • jm ^k * # . * * ' • * !• V ','••«•

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FIGS. 59-64. 59. Reginella floridana. 60. Exechonella antillea. 61. Escharoides costifer. 62. S'awg- nyella lafontii. 63. Reptadeonella costulata. 64. Tremoschizodina lata. in size, frontal surface covered by six or seven lacunae between them. Orifice keyhole- radiating pairs of costae with rows of slitlike shaped, with a small pedicellate avicularium 136 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173 on either side. Ovicell small, hemispherical, DESCRIPTION: (AMNH 605) Zooids ovoid, and carinate. inflated, glassy, about .36 X .27 mm. Row of OCCURRENCE AND ECOLOGY: Collected in large pores around margin, with ribs between September at Gosnold station 697 encrusting them, converging toward the center and fad- a shell fragment. Ovicelled at time of collec- ing out as they approach. Peristome a short tion. tube with six to eight thick, jointed spines. DISTRIBUTION: Western Atlantic: Cape A pointed mucro with an open area in the Hatteras to Florida. Gulf of Mexico. Origi- center projecting over orifice. Large avicu- nally described from the Gulf of California laria on margin of mucronate process. Usu- and the Galapagos. ally one on each side, occasional zooids with only one. Avicularia with tiny elongate-tri- FAMILY HARMER, 1957 angular mandible oriented laterally and pro- GENUS EXECHONELLA CANU AND jecting upward. Ovicells hyperstomial, round BASSLER, 1927 and prominent, also with marginal distal and Exeehonella antillea (Osburn), 1927 lateral pores in more than one row, and a raised rim with strong ribs converging on the Figure 60 center. Only four spines showing on ovicelled Lepralia antillea Osburn, 1927, p. 128. zooids. Ancestrula about .30 mm. in size, Exeehonella pumicosa Canu and Sassier, 1928a, differing in shape from subsequent zooids, p. 70. with a circle of 12 jointed spines surrounding Exeehonella antillea: Osburn, 1940, p. 366; 1950, the orifice. Embryos bright orange. p. 95. Shier, 1964, p. 616. Cook, 1967, p. 337; 1968a, p. 97. OCCURRENCE AND ECOLOGY: Found only on algae. Occurred on drift Sargassum col- DESCRIPTION: (AMNH 604) Colony en- lected at Seminole Shores in February and crusting, unilaminar, grayish yellow in color, also on four species of red, green, and brown attached by dorsal processes. Zooids oval, algae in collections from the continental shelf large, about .80 X .50 mm. in size, separated (Winston and Eiseman, 1980). All specimens by a deep groove. Frontal surface covered by in Florida collections very small, forming large pores, surrounded by a collar, giving the circular colonies only a few millimeters in surface of the zooids the texture of terrazzo size. Zooids only three generations out from pavement. Marginal pores smaller and un- the ancestrula may have ovicells, and almost collared. Orifice large, shaped like two at- every zooid was ovicelled in all but the small- tached semicircles, the distal one larger than est colonies. Polypide with 15 tentacles; loph- the proximal. Peristome wide-rimmed, rather ophore diameter about .350 mm. flat proximally, becoming raised into a collar DISTRIBUTION: Florida Atlantic coast and around distal margin of orifice. Tiny avicu- continental shelf, Tortugas, Florida, Bahia de laria sometimes occurring near edge of fron- Santos, Brazil. tal surface. No ovicells. FAMILY LEVINSEN, 1909 OCCURRENCE AND ECOLOGY: Encrusting a GENUS SAVIGNYELLA LEVINSEN, 1909 Thyroscyphus stem at Fort Pierce Breakwater in March. Savignyella lafontii (Audouin), 1826 DISTRIBUTION: Western Atlantic: Cape Figure 62 Hatteras to Florida. Caribbean and Gulf of Eucratea lafontii Audouin, 1826, p. 242. Mexico. Also known from the Gulf of Cali- Savignyella lafontii: Osburn, 1914, p. 197. Mar- fornia and from West Africa. cus, 1937, p. 78. Cook, 1968, p. 178. Savignyella lafonti: Osburn, 1927, p. 126; 1952, Escharoides costifer (Osburn), 1914 p. 288. Figure 61 Catenaria lafontii: Hastings, 1930, p. 732. Escharella costifera Osburn, 1914, p. 203. DESCRIPTION: (AMNH 606) Colony com- Escharoides costifera: Marcus, 1938, p. 38. posed of uniserial jointed branches, brick red 1982 WINSTON: MARINE BRYOZOANS 137

or brownish when living due to coloration of to center of zooid. Orifice semi-elliptical, polypides and/or embryos within the zooids. proximal margin straight, distal margin Zooids trumpet-shaped with tubelike proxi- rounded, with a low peristome. Avicularium, mal portion and inflated distal portion, vary- with a rounded base and a pointed mandible, ing in length between 0.75-1.5 mm. Distal located between ascopore and orifice and portion with numerous pores penetrating oriented obliquely distally and outwardly frontal surfaces. Orifice semicircular, sur- from the zooid center, touching the outer rounded by a raised peristome with four stout margin of the orifice. No ovicells. Embryos spines and a large pointed suboral avicular- brooded in gonozooids, in this species similar ium with a triangular mandible. Ovicells in size to the autozooids and differentiated globular, also perforated by pores. Polypide from them only in the more hoodlike devel- with 17-19 tentacles (Marcus, 1937). opment of the peristome. OCCURRENCE AND ECOLOGY: Found at all DISCUSSION: Cheetham and Sandberg coastal stations: Sebastian Inlet, Fort Pierce (1964) were unaware of Canu and Bassler's Breakwater, Walton Rocks, and Seminole introduction of this species when they de- Shores, and in the Sebastian grassbed in the scribed it as new (A. H. Cheetham, personal Indian River. Most commonly occurred on commun.). the roots of Thyroscyphus and other hydroids OCCURRENCE AND ECOLOGY: Thick encrus- where colonies often intermingled with those tations on Oculina and shell rubble in con- of Vittaticella. The delicate branching colo- tinental shelf collections (30-90 m.). Not col- nies are not usually obvious until substratum lected at coastal stations. Colonies have dark material is examined under the microscope. brown coloration when live and keep a brown Also found on algae, sponges, and Zoobot- and white speckled appearance even when ryon. Occurred year-round, most abundant preserved in alcohol. from September to December, and especially DISTRIBUTION: East and west coasts of abundant at Fort Pierce Inlet. Reproduction Florida, Gulf of Mexico, north coast of Flor- may be year-round in this area. Osburn found ida, Straits of Florida. This species has been colonies reproducing in June in the Tortugas. confused with R. violacea and R. plagiopora, Colonies collected in the Indian River in Jan- but its identity was clarified by Cheetham uary were reproducing. and Sandberg, 1964. DISTRIBUTION: Worldwide in warm waters. Western Atlantic: Bermuda, east coast of FAMILY BASSLER, 1953 Florida to Brazil. Caribbean. Gulf of Mexico. GENUS TREMOSCHIZODINA DUVERGIER, 1921 FAMILY JULLIEN, 1903 Tremoschizodina lata Smitt, 1873 GENUS REPTADEONELLA BUSK, 1884 Figure 64 Reptadeonella costulata Gemellipora lata Smitt, 1873, p. 36. (Canu and Bassler), 1928 Tremoschizodina lata: Canu and Bassler, 1928a, Figure 63 p. 131. Cook, 1968, p. 184.

Adeona costulata Canu and Bassler, 1928b, p. 94. DESCRIPTION: (AMNH 608) Colony en- Reptadeonella hastingsae Cheetham and Sand- crusting. Zooids relatively large, about berg, 1964, p. 1039. .65 X .55 mm. in size, surfaces covered with DESCRIPTION: (AMNH 607) Colony en- shallow regularly shaped pores. Orifice horse- crusting. Zooids approximately .50 X .50 shoe-shaped distally, shallowly V-shaped mm. in size, irregularly rhomboidal in shape, proximally, with very slight lateral indenta- separated from each other by a distinct tions. Ovicells endozooidal. Ovicelled zooids threadlike ridge. Characterized by row of cir- distinguished by the larger, more transversely cular marginal pores, granular secondary cal- elliptical orifice. Very rare frontal avicularia, cification, and crescentic ascopore proximal not occurring in the colonies examined. 138 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

FIGS. 65-70. 65. Hippaliosina rostrigera. 66. Watersipora subovoidea. 67. Cryptosula pallasiana. 68. Crepidacantha setigera. 69. Hippoporina verrilli. 70. Parasmittina spathulata.

OCCURRENCE AND ECOLOGY: Only in con- in September and February did not have tinental shelf collections. Colonies collected ovicelled zooids. 1982 WINSTON: MARINE BRYOZOANS 139

DISTRIBUTION: A tropical species, known agonal in shape, the front surface covered by from the east coast of Florida, Florida Straits, large pores sunk in heavy calcification. Very Gulf of Mexico, and West Africa. often zooids with a suboral umbo giving col- ony surface a beaded appearance to unaided eye. Orifice very large, bell-shaped, anterior GENUS HIPPALIOSINA CANU, 1918 part deeper and rounded, posterior a shallow Hippaliosina wstrigera (Smitt), 1873 arc, sides parallel, cardelles small. Narrow Figure 65 peristome forming a neat frame around the Escharella wstrigera Smitt, 1873, p. 57. orifice. No ovicells. Embryos brooded in Lepralia wstrigera: Osburn, 1914, p. 211. ovisacs within zooids. No orificial dimor- Hippaliosina wstrigera: Canu and Sassier, 1928a, phism. Embryos detected in living specimens p. 130. Hastings, 1930, p. 729. Osburn, 1940, by their orange color. Rare small suboral p. 448; 1952, p. 475. Cheetham and Sandberg, avicularia. None observed on specimens col- 1964, p. 1044. Shier, 1964, p. 641. lected in this study. OCCURRENCE AND ECOLOGY: DESCRIPTION: (AMNH 609) Colonies en- Occurs on al- crusting, orange when live. Zooids variable most any hard substratum: wood, glass, and in shape and size (.35-60 mm. in length by beachrock. A common and cosmopolitan .25-30 mm. in width). Frontal surface gran- fouling species, tolerant of reduced salinities. ular with one or two rows of marginal pores, Not found in the Indian River but did occur a distinct threadlike rim between zooids. at coastal stations between April and Decem- Orifice horseshoe-shaped distally, arc-shaped ber. Embryos noted in colonies only in April, proximally, narrow peristome raised distally. but colonies active at all times they were ob- Avicularia single or paired, with a rounded served. base and pointed mandible directed obliquely DISTRIBUTION: Cosmopolitan, but disjunct inward from edge of zooid toward distal mar- in many cases. Distribution may be related gin of orifice. Ovicell endozooidal. Fertile to proximity to shipping lanes (Marcus, zooids distinguished by wide short orifices. 1942). OCCURRENCE AND ECOLOGY: On Oculina and shell rubble in continental shelf collec- GENUS WATERSIPORA NEVIANI, 1895 tions. Also found at Seminole Shores in Oc- Watersipora subovoidea (d'Orbigny), 1852 tober in depressions on loose subtidal beach Figure 66 rock. No colonies in this material had fertile zooids. Cellepora subovoidea d'Orbigny, 1852, p. 402. Lepralia cucullata Osburn, 1914, p. 211. DISTRIBUTION: Warm waters of Western Watersipora cucullata Hastings, 1930, p. 729. Atlantic and east Pacific. Cape Hatteras, Gulf Marcus, 1937, p. 118; 1938, p. 46. Osburn, of Mexico, Caribbean, Galapagos, Panama, 1940, p. 449. and Gulf of California. Dakaria subovoidea: Harmer, 1957, p. 1022. Watersipora subovoidea: Gautier, 1962, p. 183. GENUS CRYPTOSULA CANU AND Ryland, 1965, p. 68. Cook, 1968a, p. 184. BASSLER, 1925 DESCRIPTION: (AMNH 610) Colonies uni- Cryptosula pallasiana (Moll), 1803 laminar encrusting to bilaminar frilled, Figure 67 brownish orange to black when living. Zooids large, averaging .82 mm. in length by .38 mm. Eschara pallasiana Moll, 1803, p. 57. in width, elongate but irregular in shape. Cryptosula pallasiana: Canu and Sassier, 1925, p. Frontal surface slightly curved and evenly 33. Marcus, 1942, p. 58. Osburn, 1952, p. 47. Maturo, 1957, p. 58. perforated all over by large pores. Orifice mushroom-shaped, ovoid anteriorly, with a DESCRIPTION: (AMNH 611) Colonies en- small semicircular sinus. Operculum dark crusting, or sometimes rising into frills, pink, brown to black. No avicularia. Polypides beige, or white when live. Zooids about bright orange-red, average lophophore di- .60 X .40 mm. in size, usually regularly hex- ameter .66 mm., average tentacle number 21. 140 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

No ovicells. Embryos brooded in internal fice. In living specimens setose spines occur- ovisacs. ring between marginal pores, on bleached OCCURRENCE AND ECOLOGY: Most abun- specimen illustrated only basal joints of these dant encrusting bryozoan at all coastal sta- spines remain. Ovicell hyperstomial, closed tions. A well-known fouling organism, its by operculum, may have a porous central success attributed to its rapid rate growth area. (Osburn, 1914) and its ability to grow on al- OCCURRENCE AND ECOLOGY: Found only most any surface including copper antifoul- in Gosnold collections, on shell fragments, ing paint (Wisely, 1958). Most abundant en- ovicelled in September. crusting the rocks of the breakwaters at about DISTRIBUTION: Circumtropical. Tropical the mean low water level. Only encrusting east Pacific, West Africa, Florida, and Carib- bryozoan growing on cemented sand tubes bean. of the reef-building sabellariid Phragmato- poma lapidosa, attaching both on outer sur- FAMILY HIPPOPORINIDAE BASSLER, 1953 faces of the worm reef and in caves and cav- GENUS HIPPOPORINA NEVIANI, 1895 ities within it. These colonies were subject to Hippoporina verrilli Maturo and the cycle of growth and decay of the worm Schopf, 1968 tubes over the year; by late summer when old Figure 69 worms died and most tubes crumbled, most Watersipora died, though colonies on other Escharella pertusa (Esper) Verrill, 1875, p. 41 substrata remained healthy. Present year- (part). round, reproducing from November to April, Lepralia pertusa (Esper) Osburn, 1912, p. 241 (part). with maximum production of larvae in No- Hippodiplosia americana (Verrill) Hastings, 1930, vember and December. Where the water was p. 725. Marcus, 1937, p. 101. Osburn, 1952, p. calm and/or competition for space with hy- 339. Maturo, 1957, p. 51. Lagaaij, 1963, p. 192. droids intense (as at Sebastian), Watersipora Shier, 1964, p. 631 (Not of Verrill, 1875). grew out in bilaminar frills. Also found once Hippoporina americana Cook, 1964b, p. 6; 1968a, attached to algae at the Sebastian Grass Flats. p. 188. DISTRIBUTION: Synonymy of this species Hippoporina verrilli Maturo and Schopf, 1968, p. is very confused, but it appears to be world- 56. Powell, 1971, p. 771. wide in warm seas. Western Atlantic: east DESCRIPTION: (AMNH 613) Colonies en- coast of Florida to Brazil. Caribbean. Gulf crusting shell, wood and other hard substrata. of Mexico. Zooids regular in shape, in young colonies measured in this study usually quadrangular, CREPIDACANTHIDAE LEVINSEN, 1909 averaging .40 X .29 mm., up to .47 X .39 in CREPIDACANTHA LEVINSEN, 1909 old colonies (Maturo and Schopf, 1968). Crepidacantha setigera (Smitt), 1873 Frontal surface of zooids perforated by about Figure 68 10-20 medium-sized evenly spaced pores. Zooids separated from each other by slightly Escharella setigera Smitt, 1873, p. 58. raised rims. Orifice rounded distally, with Crepidacantha setigera: Canu and Bassler, 1928a, p. 135. Cook, 1968a, p. 186. sides almost straight and slightly curved proximal portion behind delicate condyles. DESCRIPTION: (AMNH 612) Colonies en- Often a prominent umbo just below orifice. crusting, pinkish when alive. Zooids pyri- Polypide with 12 tentacles. Lophophore di- form, about .40 X .60 mm. in size, surface ameter .428 mm. Avicularia variable, many inflated, faintly granular, separating grooves colonies lacking them. When present may deep, row of small marginal pores present. occur on one or both sides of the orifice and Orifice keyhole-shaped, distal portion cir- may be narrow, ovoid or have round base cular and proximal border broadly arcuate and triangular mandible. Narrow ones di- between stout condyles. Avicularia with long rected proximolaterally, the other two types setose mandibles located on each side of ori- may be oriented proximolaterally, laterally, 1982 WINSTON: MARINE BRYOZOANS 141

l*V '*•**> ^f"*% • ' * FIGS. 71-76. 71. Parasmittina nitida morphotype B. 72. Phylactellipora aviculifera. 73. Lagenicella marginata. 74. Schizoporella comuta. 75. Schizoporella floridana. 76. Schizoporella unicornis.

proximomedially, laterally or distomedially. granular calcification around the outside and Ovicells globular, hyperstomial, with rough a smoother porous central area which may 142 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173 also become covered by granular secondary and an orificial collar raised into lappets on calcification. Embryos reddish orange in each side. Avicularia variable, one to three color. per zooid. If two or more are present, they OCCURRENCE AND ECOLOGY: When living, are of different types. Avicularia with pointed colonies from the Indian River are ivory to mandibles placed lateral to the orifice (usu- beige in color, but in other areas color may ally extending up onto the collar), and on the vary from golden yellow to red. Collected at distal or lateral side of the orifice. Oval three stations: Capron Shoals, Fort Pierce avicularia, variously oriented, occurring lat- Breakwater, and on the panels at Link Port eral or proximal to orifice (opposite acute in the Indian River. One of the most abun- avicularium if both occur). Giant (interzooi- dant bryozoans recruited on panels in the dal) avicularia may occasionally occur. Ovi- river in 1974 and 1975 (Mook, 1976). Eu- cell hyperstomial and prominent, upper rim rythermal, but reproduction heaviest in and sides usually covered by granular calci- cooler months. Settlement was heaviest in fication, front penetrated by two rows of large fall (October to January) though colonies set- pores. Embryos pinkish orange. Polypides tled in bryozoan traps through June. with 13 tentacles and an average lophophore DISTRIBUTION: This species was confused diameter of .419 mm. with H. americana until quite recently, but DISCUSSION: Traditionally a whole com- is apparently widely distributed in warm- plex of species have been lumped together temperate to tropical waters, especially in under Smittina trispinosa. Mature and Schopf seasonally brackish lagoons, harbors, or es- (1968) separated two of these varieties into tuaries. Western Atlantic: Cape Cod to Bra- Parasmittina nitida morphotype A and Para- zil. Gulf of Mexico. Pacific Coast: Gulf of smittina nitida morphotype B. Morpho- California to the Galapagos. West Africa. type A is the form figured by Verrill as Dis- copora nitida. Morphotype B has been shown FAMILY LEVINSEN, 1909 by Mature (1973) and Humphries (1975) to GENUS PARASMITTINA OSBURN, 1952 be genetically distinct and should be recog- nized as a new species. Parasmittina nitida morphotype B OCCURRENCE AND ECOLOGY: Occurred Mature and Schopf, 1968 from April to December at Walton Rocks as Figure 71 one of the most common bryozoans on the Smittina trispinosa (Johnston) Osburn, 1912, p. subtidal beachrock stones. Also found on 246 (part), pi. 27, fig. 65a. Canu and Sassier, dead shells at Capron Shoals in April and 1923, p. 143. Rogick and Croasdale, 1949, p. October. Appeared to reproduce year-round 67, pi. 10, figs. 68-71. Osburn, 1952, p. 412 in the Indian River area, and also reported (part), pi. 49, fig. 7. Maturo, 1957, p. 55 (part), by Humphries (1975) to reproduce year- fig. 62. round in the Gulf of Mexico. Smittina trispinosa var. nitida (Verrill) Osburn, DISTRIBUTION: Distribution not well known 1912, p. 246. Marcus, 1937, p. 104. due to the problems with , but Parasmittina nitida morphotype B Maturo and Schopf, 1968, p. 41, fig. 120. Humphries, 1975, probably occurs in the Western Atlantic from p. 20, pi. 4. New England to Brazil, in the Gulf of Mexico, and the Caribbean. DESCRIPTION: (AMNH 614) Colonies en- crusting, one or two layers thick, color vari- Parasmittina spathulata (Smitt), 1873 able, whitish buff to yellow. Zooids quadrate, Figure 70 averaging .45 X .29 mm. in size, frontal wall solid and coarsely granular in the center with Escharella jacotini var. spathulata Smitt, 1873, p. 60. a row of large marginal pores. Primary orifice Smittina trispinosa var. spathulata Osburn, 1914, round with short toothed condyles and a ly- p. 208; 1927, p. 29. rula one-third to one-fourth width of orifice. Smittina trispinosa spathulata Canu and Bassler, Two spines on distal edge of primary orifice 1928a, p. 114. 1982 WINSTON: MARINE BRYOZOANS 143

Smittina trispinosa var. spathulosa [sic] Osburn, ber. This chamber, with a membranous or 1940, p. 435. chitinous covering but no hinge or mandible, Parasmittina spathulata: Osburn, 1952, p. 415. resembling an aborted avicularium, thus giv- Cheetham and Sandberg, 1964, p. 1038. Long ing the species its name. Ovicell prominent, and Rucker, 1970, p. 20. helmet-shaped, finely granular in texture DESCRIPTION: (AMNH 613) Colonies en- with a curving distal flange meeting the lat- crusting, often plurilaminar. Zooids about eral processes of the orificial collar of the fer- .60 X .40 mm. in size, variable in shape, ir- tile zooid. Ovicell not closed by the opercu- regular, but often rounded distally and with lum, its opening a low arch (barely visible in sharp proximal edges. Frontal surface flat, upper right zooid of specimen illustrated). coarsely granular with a row of large marginal Embryos yellowish brown in color. pores. Orifice subcircular with short con- OCCURRENCE AND ECOLOGY: Collected on dyles, large truncate lyrula and low peristome dead shells at Capron Shoals in April and raised more laterally and usually hiding the October and on the continental shelf pinna- condyles. Two to four spines on orifice (usu- cles in February. Colonies were reproducing ally covered by secondary calcification). in October. Avicularia variable in size, position, and DISTRIBUTION: East coast of Florida, Tor- number but usually one with long pointed tugas, northwest Florida. mandible in the vicinity of the orifice and directed proximally. Ovicells prominent, hy- GENUS LAGENICELLA CHEETHAM AND perstomial, top and sides often becoming SANDBERG, 1964 covered in granular calcification, frontal sur- Lagenicella marginata (Canu and Bassler), face with numerous irregularly spaced col- 1930 lared pores. Figure 73 OCCURRENCE AND ECOLOGY: Found only in Gosnold collections. One specimen ovi- Lagenipora marginata Canu and Bassler, 1930, p. celled when collected (September). 36. DISTRIBUTION: Western Atlantic: Cape Lekythopora longicollis Lagaaij, 1963, p. 199. Hatteras-Brazil. Gulf of Mexico and Carib- Lagenicella marginata: Cheetham and Sandberg, 1964, p. 1041. bean. Galapagos. DESCRIPTION: (AMNH 616) Colonies en- FAMILY PHYLACTELLIPORIDAE BASSLER, 1953 crusting, made up of clusters of small zooids. GENUS PHYLACTELLIPORA, BASSLER, 1953 Zooids flask-shaped, about .40 X .25 mm. in size, including peristome. Frontal wall in- Phylactellipora aviculifera (Osburn), 1914 flated, surface perforated by small pores Figure 72 which are almost lost beneath thick wavy lon- Phylactella collaris var. aviculifera Osburn, 1914, gitudinally ribbed calcification. Peristome p. 213. long, smoother, imperforate, zooid con- Smittina (sensu latd) thrincota Shier, 1964, p. 637. stricted at base of peristome and then just slightly flaring. Pair of very small (0.010 DESCRIPTION: (AMNH 615) Colony en- mm.) acute avicularia occurring at the top of crusting, white in color. Zooids ovoid to hex- a tubular chamber extending from the lateral agonal, about .45 X .30 mm. in size with a rim of the peristome. Ovicells distal to peri- granular, evenly porous, slightly inflated stome, opening high up its distal side, with frontal wall, and a depression between a crescentic densely porous frontal area and zooids. Orifice subcircular, round distally, smoothly calcified top and sides. with a pair of strong cardelles near the prox- color pinkish red. imal end. Distal rim of orifice with slightly OCCURRENCE AND ECOLOGY: Collected once bowed proximal margin, orificial collar high at Seminole Shores encrusting drift Sargas- and thin laterally and jutting out proximally sum sp. Colonies occurred in the crotches to form a triangular shelf with a central cham- where leaves attached to the main stem of 144 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173 the alga. Colonies had ovicells with embryos ondary cover in what looks like a row of in April. Lagaaij (1963) reported L. margi- pores. Embryos orange-red to cherry red. nata to be a "fairly common species in the OCCURRENCE AND ECOLOGY: Found at all Gulf of Mexico" and "presumably encrusting three coastal stations and at Capron Shoals. a perishable substratum." It probably is com- Most abundant on beachrock and dead shell mon on drift and rooted Sargassum and in subtidal pools at Walton Rocks and Sem- other algae. Found on the deep-water alga inole Shores. Found from April to December Rhodymenia pseudopalmata off the Florida (may be present year-round). Most abundant coast (Winston and Eiseman, 1980). and reproducing October to December. Col- DISTRIBUTION: East coast of Florida, Gulf onies collected at Walton Rocks in December of Mexico, Galapagos. had most zooids ovicelled and filled with em- bryos. JULLIEN (1903) DISTRIBUTION: Range incompletely known SCHIZOPORELLA HINCKS, 1877 as the species has been confused with Schi- zoporella biaperta as well as several other Schizoporella cornuta (Gabb and Horn), species. Probably occurs from Woods Hole, 1862 Massachusetts to Florida, the Gulf of Mex- Figure 74 ico, and the Caribbean. Also along the Pacific Reptescharellina cornuta Gabb and Horn, 1862, coast of North America and reported from p. 147. West Africa. Schizoporella incrassata Canu and Bassler, 1928a, p. 93. (Not Hincks, 1882). Schizoporella floridana Osburn, 1914 Schizoporella canui Osburn, 1940, p. 422. Schizoporella cornuta: Osburn, 1952, p. 320. Ma- Figure 75 ture, 1957, p. 50. Shier, 1964, p. 628. Cheetham Schizoporella floridana Osburn, 1914, p. 205; and Sandberg, 1964, p. 1030. Cook, 1968a, p. 1927, p. 126; 1940, p. 29. Canu and Bassler, 192. 1928a, p. 93. Cheetham and Sandberg, 1964, p. 1030. DESCRIPTION: Colonies encrusting, pink to Schizoporella unicornis Shier, 1964, p. 629. salmon in color. Zooids oval to hexagonal in Schizoporella unicornis var. floridana Shier, 1964, shape, about .35 X .25 mm. in size. Bound- p. 630. aries between zooids often not distinct due to heavy calcification. Frontal surface per- DESCRIPTION: (AMNH 617) Colony en- forated by numerous pores (.015-.02 mm. in crusting, unilaminar or multilaminar to fo- diameter) and becoming increasingly irreg- liaceous or tubular. Zooids rectangular, av- ular to age due to secondary calcification. eraging .49 X .36 mm. in Indian River Orifice slightly more than semicircular, with specimens, heavily calcified, with a granular two cardelles and a shallow V-shaped sinus texture and numerous large (.02 mm. or proximally. Elliptical avicularia, paired, or, more) pores sunk in depressions in the cal- in these specimens, more commonly single, cification. A heavy suborificial umbo often raised on mammillated processes just below present. Orifice often situated somewhat at and beside the orifice with mandibles di- an angle at one side of semicircular distally, rected up at a 4 5-degree angle and outward with a broad U-shaped sinus; the zooid mid- toward the sides of the zooid. Ovicell hyper- line. Avicularia with rounded bases and stomial, globular and prominent; differing pointed mandibles occurring singly, or in from the other two described species of pairs, beside and just below the orifice, di- Schizoporella in not having pores all over rected upward at about a 30-degree angle on ovicell surface, but having an outer rim of a mammillate process. Polypides pink in heavy calcification and a circular central area color, mean tentacle number 17, mean lo- radially grooved to a greater or lesser degree, phophore diameter .509 mm. Ovicells large, the grooves ending under the rim of the sec- hyperstomial, slightly wider than long, cov- 1982 WINSTON: MARINE BRYOZOANS 145 ered with pores. Embryos pale orange in ture (1968) lists it as ranging extensively color. north and south of Cape Hatteras. Figure 3, OCCURRENCE AND ECOLOGY: Found en- no. 3 of Long and Rucker (1970) is definitely crusting Thalassia in the Link Port grassbeds. this species. It is one of the principal fouling Colonies on Thalassia blades did not grow organisms in many parts of the world. large or become multiserial. Ovicelled zooids occurred only a few generations out from the GENUS ESCHARINA ancestrula and colonies only a few millime- MILNE-EDWARDS, 1838 ters in size were producing embryos. Colonies Escharina pesanseris (Smitt), 1873 recruited on panels at Link Port from January Figure 77 to August, peak settlement (according to Hippothoa pesanseris Smitt, 1873, p. 43. Mook, 1976) in late spring to early summer Escharina pesanseris: Osburn, 1914, p. 207. (chiefly April and May). Not collected at any Mastigophora pesanseris: Osburn, 1927, p. 130; coastal stations and massive coral-like colony 1940, p. 452; 1952, p. 479. Canu and Bassler, forms described from the Gulf coast of Flor- 1928a, p. 133. Hastings, 1930, p. 722. Marcus, ida and from other areas were not seen. 1939, p. 142. DISTRIBUTION: Western Atlantic, Beaufort, Escharina pesanseris: Cook, 1968a, p. 195. Long N.C., Bermuda to Florida Gulf of Mexico, and Rucker, 1970, p. 19. Caribbean. This "species" may actually be DESCRIPTION: (AMNH 618) Colonies en- a variety of Schizoporella errata, a common crusting and small, less than 1 cm. in di- fouling species in ports and harbors of the ameter. Zooids polygonal, usually five-sided Mediterranean and Europe (Ryland, 1965). about .60 X .55 mm. in size, frontal surface covered with fine pores which may become Schizoporella unicornis (Johnston), 1847 filled in by secondary calcification, and with Figure 76 distinct grooves between them. Orifice semi- Lepralia unicornis Johnston, 1847, p. 320. circular, proximal border straight with a nar- Schizoporella unicornis: Mature, 1957, p. 49. Ry- row, deep, notched sinus. Peristome forming land, 1965, p. 65. Cook, 1968a, p. 101. Long a thick rim around the orifice and bearing six and Rucker, 1970, p. 19. to eight spines. Avicularia in pairs on each side of the orifice, directed distally, with a DESCRIPTION: Colony encrusting, unilam- inar to plurilaminar, white to pink in color. fin-shaped "duck-foot" mandible (this chi- Zooids regularly rectangular in shape, about tinous mandible lacking in Clorox-treated .45 X .35 mm. in size, frontal surface per- specimen illustrated). Ovicells (not illus- forated by a large number of small pores (.01 trated) thin-walled, globular, small and im- mm. or less in diameter). Zooids separated perforate. from each other by a sharp depression, fron- OCCURRENCE AND ECOLOGY: Collected only tal wall often raised into umbo below the at offshore stations, February 1975, on dead orifice. Orifice central, slightly more than shells. The small colonies, often fouled and semicircular with a deep V-shaped sinus. eroded, are easy to overlook, but once noted Ovicells hyperstomial, large, globular, po- the fan-shaped avicularia are distinctive. rous, with a radial pattern making edge ap- DISTRIBUTION: Circumtropical. Western pear fluted and also often with a pointed Atlantic: Cape Hatteras to Brazil. Caribbean. umbo at the top. Gulf of Mexico. OCCURRENCE AND ECOLOGY: Collected on GENUS STYLOPOMA LEVINSEN, 1909 beachrock stones at Walton Rocks in May. Reproducing at that time. Stylopoma spongites (Pallas), 1766 DISTRIBUTION: The true range of this Figure 78 species in the Western Atlantic is unknown Eschara spongites Pallas, 1766, p. 45 (part). because of its confusion with £. errata. Ma- Hippothoa spongites: Smitt, 1873, p. 42. 146 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

FIGS. 77-82. 77. Escharina pesanseris. 78. Stylopoma spongites. 79. Cigclisula turrita. 80. Cleido- chasma porcellanum. 81. Cleidochasma contractum. 82. Hippoporidra calcarea. 1982 WINSTON: MARINE BRYOZOANS 147

Schizoporella spongites: Osburn, 1914, p. 207. FAMILY CANU AND Stylopoma spongites: Osburn, 1927, p. 128. Canu BASSLER, 1917 and Bassler, 1928a, p. 91. Hastings, 1930, p. GENUS CIGCLISULA CANU AND BASSLER, 1929 721. Stylopoma informata Marcus, 1937, p. 91. (Not Cigclisula turrita (Smitt), 1873 Lonsdale, 1845); 1955, p. 296. Osburn, 1940, Figure 79 p. 424; 1952, p. 336. Shier, 1964, p. 631. Lepralia turrita Smitt, 1873, p. 65. "Stylopoma spongites" (Levinsen): Cheetham and Holoporella turrita: Osburn, 1914, p. 217; 1927, Sandberg, 1964, p. 1031. p. 131. Canu and Bassler, 1928a, p. 145. Has- Stylopoma spongites (Levinsen): Long and Rucker, tings, 1930, p. 732. 1970, p. 19. Cigclisula turrita: Harmer, 1957, p. 1059. Stylopoma spongites (Pallas): Thomas and Has- Trematooecia turrita: Powell, 1971, p. 773. tings, 1967, p. 316. DESCRIPTION: (AMNH 620) Colony en- DESCRIPTION: (AMNH 619) Colony en- crusting, plurilaminar, massive peach col- crusting to tubular, usually plurilaminar, ored live. Zooids erect, about .50 X .50 mm. white to orange in color. Zooids usually reg- in size, orifice central, surrounded by several ularly quadrangular in shape, about .50 X .35 thick tubercles or projections, thickly calci- mm. in size, frontal wall perforated by nu- fied, with walls penetrated by small scattered merous small pores, with intervening calci- pores. Orifice subcircular proximally, semi- fication becoming increasingly thick and circular distally. Tiny ovate avicularia oc- granular with age, sometimes with a low curring in various positions on the frontal umbo below the orifice (which tends to be- surface and on the tips of the tubercles. Large come buried with increasing calcification). spatulate avicularia sometimes occurring. Orifice semicircular, with straight proximal Ovicells (according to Harmer, 1957 and border and deep V-shaped to linear sinus. Osburn, 1940) spherical, inconspicuous, Peristome low, usually not forming a notice- opening into the distal end of the peristome, able ridge except on zooids at the growing with a rounded membranous area when edge. Avicularia varying in size and shape. young which becomes heavily calcified and Small avicularia with short triangular man- porous with age. dibles occurring beside the orifice or on the OCCURRENCE AND ECOLOGY: Occurred only frontal surface of zooids or ovicells. Occa- in continental shelf collections. Specimens sional large spatulate vicarious avicularia collected in September and February had no also occurring. Ovicell distinctive, huge in ovicells. comparison with zooids, about .55 mm. in DISTRIBUTION: East coast of Florida, Gulf diameter, globular, completely covering the of Mexico, Caribbean. Panama. Also re- orifice of the mother zooid, covered with ported from the Mediterranean and the Pa- pores and usually bearing several small avic- cific. ularia on its surface. OCCURRENCE AND ECOLOGY: Found in con- tinental shelf collections in September en- GENUS CLEIDOCHASMA HARMER, 1957 crusting dead Oculina. Collected twice on Cleidochasma porcellanum (Busk), 1860 beachrock stones at Seminole Shores, in Figure 80 April and October. None of these specimens had ovicells. Lepralia porcellana Busk, 1860, p. 283. Lepralia cleidostoma Smitt, 1873, p. 62. DISTRIBUTION: Western Atlantic: Cape Hippoporina cleidostoma Canu and Bassler, 1928a, Hatteras, Bermuda to Brazil. Gulf of Mexico p. 104. and Caribbean. Galapagos. A warm water Hippoporina porcellana: Hastings, 1930, p. 721. species abundant in the Caribbean and Gulf Marcus, 1937, p. 96. Osburn, 1940, p. 428; of Mexico. 1952, p. 344. Shier, 1964, p. 633. 148 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

Cheidochasma porcellanum: Cheetham and Sand- ginal pores. Edges of zooids indistinct except berg, 1964, p. 1032. Cook, 1964b, p. 11; 1968a, for pores. Orifice rounded distally and beaded p. 198. Long and Rucker, 1970, p. 19. Powell, at edge, with strong cardelles and rounded 1971, p. 771. sinus proximally. Four to eight hollow spines DESCRIPTION: (AMNH 621) Colonies en- around orifice. Avicularia variable, some- crusting, white to beige or peach in color, times entirely lacking near edge. Small ovoid unilaminar to plurilaminar. Zooids hexago- avicularia may be present on either side of nal in shape, .60 X .45 mm. in size, slightly orifice, having serrated mandibles, raised at inflated, calcification thick and sparsely gran- an angle, and proximolaterally directed. ular, with three to five marginal pores, a faint Spatulate distolaterally directed avicularia line of calcification marking edges. Orifice may also occur beside orifice. Ovicell hyper- keyhole-shaped, round proximally, with car- stomial, usually broader than long, imper- delles directed backward and a deep and forate, with an outer heavily calcified front broad sinus below that. Avicularia varying having faint radial striations, and a less cal- in size and position, in these specimens rare, cified central area. Ovicells covering all but when present located below the aperture, di- two spines of fertile zooids. According to rected laterally, and with a long pointed man- Marcus (1937) polypides have 12 tentacles dible. Ovicell hyperstomial, imperforate, and are dark yellow in color. granular. Near the growing edge ovicells OCCURRENCE AND ECOLOGY: One of the prominent and showing longitudinal stria- most common species in the Gosnold collec- tions. In older areas of the colony ovicells tions, often forming massive encrustations embedded in calcification of distal zooids and on dead Oculina. Specimens collected in Sep- almost invisible except for slight swelling of tember had ovicells. the distal zooids and squared-off lines above DISTRIBUTION: Western Atlantic: Cape the aperture of the fertile zooid marking their Cod to Brazil. Caribbean and Gulf of Mexico. openings. Embryos reddish pink. Eastern Pacific, Madeira, and West Africa. OCCURRENCE AND ECOLOGY: Colonies with ovicells containing embryos present encrust- GENUS HIPPOPORIDRA CANU AND ing Oculina in Gosnold collections of Feb- BASSLER, 1927 ruary 1975. Hippoporidra calcarea (Smitt), 1873 DISTRIBUTION: Probably circumtropical. Figure 82 Western Atlantic: Cape Hatteras to Brazil. Gulf of Mexico and Caribbean. Lepralia edax (forma calcarea) Smitt, 1873, p. 63. Lepralia edax Osburn, 1914, p. 212. Cleidochasma contractum (Waters), 1899 Hippoporidra calcarea: Canu and Sassier, 1928a, Figure 81 p. 140.

Lepralia contracta Waters, 1899, p. 11. DESCRIPTION: (AMNH 623) Colonies en- Perigastrella contracta: Hastings, 1930, p. 722. crusting gastropod shells occupied by hermit Marcus, 1937, p. 98. crabs. Massive, often developing nodules, Hippoporina contracta: Osburn, 1940, p. 428; long prominences or cylindrical branches, 1952, p. 346. Marcus, 1955, p. 298. Shier, 1964, p. 632. commonly one on either side of the shell. Cleidochasma contractum: Cook, 1964b, p. 14; Two basic kinds of zooids. Small autozooids 1968a, p. 198. Cheetham and Sandberg, 1964, forming the bases of branches and promi- p. 1032. nences (only ones illustrated in fig. 82) and large cortical zooids with reduced orifices. DESCRIPTION: (AMNH 622) Colonies en- Autozooids about .30 X .30-40 mm. in size, crusting, unilaminar to plurilaminar, white with a suborificial umbo, their frontal surface in color. Zooids about .43 X .36 mm. in size, very granular. There with a row of marginal oval to hexagonal in shape, with rough, thick pores and more irregular inner row of pores. and uneven calcification and a row of mar- Depressions between zooids; in some areas 1982 WINSTON: MARINE BRYOZOANS 149

FIGS. 83-88. 83. Microporella umbracula. 84. Hippothoaflagellum. 85. Celleporella carolinensis. 86. Trypostega venusta. 87. Pasythea tulipifera. 88. Celleporina hassalli.

zooids set off from each other by a thin cal- gether so closely and at such differing levels careous rim, in other areas zooids backed to- and angles that boundaries become hard to 150 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173 distinguish. Small avicularia with blunt man- broken off. Avicularia with very pointed dibles present on the frontal surface of many mandibles on one or both sides of ascopore, zooids, and large avicularia with long pointed directed distolaterally. Ovicell large, globu- mandibles found between zooids. Ovicell lar, porous, closed by zooidal operculum. hyperstomial and hooded with a raised Embryos reddish pink in color. smooth rim and an inverted U-shaped fron- OCCURRENCE AND ECOLOGY: Occurs year- tal area that may be membranous or become round in the Indian River area, reproductive partly covered by calcification. in fall and winter. Colonies with ovicells and OCCURRENCE AND ECOLOGY: Collected in embryos occurred in late October at Capron 7 m. of water on the subtidal worm reef off Shoals, in December at Walton Rocks and Pepper Park, Fort Pierce in April. Also ovicelled specimens taken in the Gosnold col- dredged in 67-70 m. offshore in September. lection in September. Buge (1957) and Cook (1964b) studied the DISTRIBUTION: Circumtropical. Western association of Hippoporidra spp. with gastro- Atlantic: Florida. pod shells inhabited by hermit crabs and con- cluded that it was primarily between the GENUS HIPPOTHOA LAMOUROUX, 1821 bryozoan and the , as they are not found on shells that contain living gastro- Hippothoa flagellum Manzoni, 1870 pods. Usually the colony is formed by a Figure 84 which settles near the orifice of the shell (out Hippothoa flagellum Manzoni, 1870, p. 6. Hincks, of range of the chelae of the crab). Colonies 1880, p. 293. Osburn, 1952, p. 278. Gautier, in this genus appear to be sexually dimorphic. 1962, p. 118. Cook, 1968a, p. 176. Ovicelled zooids are female. According to Cook's studies (1968b), cortical zooids have DESCRIPTION: (AMNH 625) Colony uni- very small polypides with only six non-cil- serial, composed of straggling networks of iated tentacles and may be male zooids. zooids. Autozooids smooth-walled, ovoid or DISTRIBUTION: Florida, Gulf of Mexico. teardrop-shaped, the proximal parts elon- gated into narrow tubular connections. Au- FAMILY HINCKS, 1880 tozooid orifices rounded distally, with prox- GENUS MICROPORELLA HINCKS, 1877 imal sinus. Female zooids shorter than autozooids, to which they are usually at- Microporella umbracula (Audouin), 1826 tached by a short tubular process, and having Figure 83 a smooth and helmet-shaped ovicell with a Flustra umbracula Audouin, 1826, p. 239. single central pore. The orifice which closes Microporella ciliata var. coronata Hastings, 1930, the ovicell, is semicircular, slightly curved p. 727. proximally, but without a sinus. Microporella coronata Osburn, 1952, p. 386. DISCUSSION: I follow Cook (1968a) in keep- Microporella umbracula: Harmer, 1957, p. 964. Powell, 1971, p. 772. ing H. flagellum separate from H. distans as described by Harmer (1957), as his material DESCRIPTION: (AMNH 624) Colony en- from the East Indian region differs with re- crusting, unilaminar, pink to orange when spect to the shape of the orifice of ovicelled alive, showing a great deal of variation in size zooids. and shape of zooids, position of avicularia, OCCURRENCE AND ECOLOGY: Found only etc. Zooids about .60 X .60 mm. in size, sur- in continental shelf collections. One of the face inflated with smooth or roughened cal- most common species on dead shell and cification perforated by small pores. Zooids Oculina rubble in 40-90 m. depth. Ovicelled separated by a depression. Orifice semicir- zooids observed at the time of collection in cular. Below it a crescent-shaped ascopore September. edged with a row of fine teeth, usually four DISTRIBUTION: Probably cosmopolitan in spines, and often dark green in color, occur- warm seas. Western Atlantic: Cape Hatteras ring around proximal rim of orifice but often to Florida (Maturo, 1968, p. 278). If Osburn's 1982 WINSTON: MARINE BRYOZOANS 151

Hippothoa distans is really this species, then mm. in length in Atlantic material), ovicells it has also been recorded from the Tortugas. with many evenly sized regularly spaced pores, and a colony which is whitish and cal- GENUS CELLEPORELLA GRAY, 1848 cified except at the edges and able to form several layers of zooids. Celleporella carolinensis Ryland, 1979 OCCURRENCE AND ECOLOGY: Found en- Figure 85 crusting Sargassum in beach drift at Semi- Hippothoa hyalina Smitt: Verrill, 1878, p. 305. nole Shores in February and July. Colony (?) Hippothoa hyalina (Linnaeus) Osburn, 1932, collected in February had white embryos. p. 443. DISTRIBUTION: The previous identification (Probably) Hippothoa hyalina (L.) Marcus, 1937, of this species with Celleporella hyalina p. 79, pi. 17, fig. 42. Hippothoa hyalina (L.) (in part) Marcus, 1938, p. makes its exact range difficult to determine, 74-77, 119, pi. 20, fig. 54 (not 56), pi. 21, fig. but Celleporella carolinensis probably occurs 55. from Cape Cod to Brazil. Ryland collected (?) Hippothoa hyalina (L.) Osburn, 1944, p. 42, it at Woods Hole and at Beaufort, N.C. It is fig. 26. here reported from Florida. Its normal hab- Hippothoa hyalina (Linnaeus) Maturo 1957, p. 48, itat appears to be the brown alga Sargassum fig. 51. filipendulum, on which the colonies may be Celleporella carolinensis Ryland, 1979, p. 612, abundant and yet unnoticed because their fig. 1. transparency renders them almost invisible.

DESCRIPTION: (AMNH 626) Colony uni- GENUS TRYPOSTEGA LEVINSEN, 1909 laminar, a transparent glistening layer of en- crusting zooids. Zooids polymorphic, of three Trypostega venusta (Norman), 1864 types: autozooids, very small male zooids, Figure 86 and female zooids bearing ovicells. Auto- Lepralia venusta Norman, 1864, p. 84. zooids oblong-elongate, averaging .36 X .18 Gemellipora glabra forma striatula Smitt, 1873, mm. in size and having polypides with 11 p. 37. tentacles, mean lophophore diameter .253 Trypostega venusta: Osburn, 1914, p. 198; 1940, mm. Male zooids about one-quarter the size p. 409; 1952, p. 280. Canu and Bassler, 1928a, of autozooids, female zooids (with ovicells) p. 77. Marcus, 1938, p. 35. Shier, 1964, p. 627. Cook, 1968a, p. 177. about half autozooid size. All zooids smooth surfaced with faint transverse growth ridges. DESCRIPTION: (AMNH 627) Colony en- Opecula of male zooids about one-third the crusting, surface smooth and glassy when size of that of the autozooid, but similar in young, becoming whitish with age. Zooids shape, both are horseshoe-shaped distally, rhomboidal, numerous pores evenly distrib- with a rounded proximal sinus. Orifice of fe- uted over the surface. Faint longitudinal male zooid (which closes the ovicell) lunate striations and transverse growth lines, some- and with a barely apparent sinus. Ovicell times with a small umbo proximal to the globular and prominent, with only a few scat- orifice. Zooids measuring about .45 X .30 tered pores and a ridge along the orifice mar- mm. Orifice pyriform: rounded distally, two gin that develops into a slight keel. Early cardelles and a V-shaped sinus proximally. stages of embryos yellow, late stages white. Rectangular zooeciules or dwarf zooids lo- Larva whitish, subspherical, without pigment cated at the distal end of each autozooid. spots (Ryland, 1979). Ovicells deeply embedded, porous, and cov- DISCUSSION: This species was recently de- ering the proximal part of the zooeciule, scribed by Ryland (1979) who separated it which then appears to be part of the ovicell. from Celleporella hyalina on the basis of OCCURRENCE AND ECOLOGY: Abundant in zooid size, early astogeny, unilaminar growth shelf waters on shell and Oculina rubble. Also form, and ovicell characters. Celleporella hy- occurred subtidally in 10 m. of water at Cap- alina has larger zooids (autozooids .45-0.5 ron Shoals. Ovicells present on specimens 152 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

collected in September. The zooeciules of this by.10 mm. wide. Polypides with 10 tentacles, species have been found to contain a polyp- average lophophore diameter .167 mm. No ide, but no and thus seem not to be ovicells or avicularia. male zooids (as in Hippothoa) but they do OCCURRENCE AND ECOLOGY: Found at only have a large opercular muscle and so, ac- one station: Fort Pierce Breakwater, but cording to Marcus (1938), should be consid- sometimes quite abundant there. Its distri- ered to be avicularia. Although zooeciules bution is tropical and colonies found in win- ordinarily are located at the distal ends of ter months (December and January) were zooids they are occasionally found inside the fouled with and appeared senescent, opercula of autozooids, apparently replacing although still having some functional polyp- dead autozooids. Trypostega colonies when ides. Most common in summer on Thyro- alive are often so fouled by algae that the scyphus roots and stems and on the break- colonies themselves resemble greenish and water rocks just below the Watersipora zone. yellowish patches of algae. The pores become Young colonies noted in October. darkened by this accumulation of algae, giv- DISTRIBUTION: Florida, Caribbean, Brazil, ing the colony surface a speckled appearance. West Africa. DISTRIBUTION: Atlantic, Pacific, and In- dian but with greatest distribution in VITTATICELLIDAE HARMER, 1957 warm water. Not known from the Mediter- GENUS VITTATICELLA MAPLESTONE, 1901 ranean. Western Atlantic: Cape Hatteras to Vittaticella uberrima Harmer, 1957 Brazil. Caribbean and Gulf of Mexico. Figure 90 PASYTHEIDAE DAVIS, 1934 ? Vittaticella elegans (Busk) Osburn, 1940, p. 464 PASYTHEA LAMOUROUX, 1812 (Not elegans Busk, 1852, p. 361). Vittaticella uberrima Harmer, 1957, p. 772. La- Pasythea tulipifera gaaij, 1963, p. 202. Cook, 1968a, p. 179. (Ellis and Solander), 1786 Figure 87 DESCRIPTION: (AMNH 629) Colony, erect, jointed, branching, up to 5-8 cm. in length. tulipifera Ellis and Solander, 1786, p. 27. Zooids arranged in a single series, all facing Pasythea tulipifera: Marcus, 1938, p. 37. Osburn, the same way, tubular at the base and broad- 1940, p. 462. Cook, 1968a, p. 186. ening toward the orifice, characterized by DESCRIPTION: (AMNH 628) Colony com- vittae (longitudinal grooves with pores) ex- posed of erect jointed branches rising from tending down either side of the frontal surface a creeping stolon. Axis of branch consisting and by a triangular lateral process on either of tubular kenozooids separated by chitinous side of the orifice, bearing an avicularium joints, triads of zooids arising from the distal with a triangular mandible. Zooids about ends of these on opposite sides. Middle zooid .20 X .50 mm. in size. Ovicells embedded in of triad connected by another joint to the base of distal zooid, often several in sequence kenozooid and facing away from the stalk. on an internode. Frontal surface of ovicell Other two zooids budded off the central flat, indented, surrounded by a beaded bor- zooid and facing toward the stalk. Joints der. Embryos brownish, larvae dark brown movable. When colony is alive, triads can be in color. drawn up parallel to the kenozooid stalk or OCCURRENCE AND ECOLOGY: Large colo- held out almost perpendicular to it. Distal nies collected at Walton Rocks in April and tips of zooids sometimes elongated into Fort Pierce Breakwater in December. Ovi- spines. Both zooids and kenozooids with cells with embryos present in both instances. scattered pores on their surfaces. Orifice with Osburn noted reproduction occurring in an orbicular top portion and a sinuate bottom Puerto Rican specimens in February, but not portion. Zooids averaging .40 mm. in length in mid-summer. 1982 WINSTON: MARINE BRYOZOANS 153

FIGS. 89-94. 89. Vittaticella contei. 90. Vittaticella uberrima. 91. Craw eburnea, branch with gono- zooid, showing position of oeciopore. 92. Crisia elongata, gonozooid in side view. 93. Crisia elongata zooids. 94. Plagioecia dispar, view of colony, showing arc-shaped brood chamber. 154 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

DISTRIBUTION: Cosmopolitan in warm Celleporina hassalli: Hayward and Ryland, 1979, water. Western Atlantic: East coast of Flor- p. 276. ida, Bermuda. Caribbean and Gulf of Mex- DESCRIPTION: (AMNH 631) Colony pluri- ico. laminar, forming small nodular masses around hydroid stems, etc. Zooids erect, av- Vittaticella contei (Audouin), 1826 eraging .40 X .30 mm. across exposed frontal Figure 89 surfaces, piled together irregularly so orifices Eucratea contei Audouin, 1826, p. 242. are at several levels and zooids at various Catenicella contei: Marcus, 1937, p. 76; 1938, p. angles to each other. Surface inflated, with a 31. row of marginal pores and often some pores Vittaticella contei: Osburn, 1940, p. 465. toward the orifice as well. Orifice round with a shallow V-shaped sinus. Peristome high DESCRIPTION: (AMNH 630) Small, deli- with a transversely elongated secondary ori- cate, translucent white colonies, with branches fice. Polypides with mean tentacle number only a few millimeters in length. Zooids of 13, mean lophophore diameter .373 mm. short, wide distally and smoothly tapering Peristomial avicularia, often paired, directed proximally, averaging .33 X .14 mm. in size, distally, and usually tilted at a 4 5-degree an- fertile zooids often shorter. Vittae usually only down the proximal half of the zooids. gle to the orifice. Large spatulate avicularia sometimes occurring. Ovicells prominent, Avicularia lacking, replaced by a long pointed with a circular membranous or lightly cal- process on either side of the orifice (which cified frontal area edged by radiating ridges may have a central membranous area). Ori- of calcification. Embryo pinkish red, larvae, fice semicircular. Polypides with 12 tentacles; pink. lophophore diameter averaging .253 mm. OCCURRENCE AND ECOLOGY: Living colo- Ovicells large, embedded in the distal zooid, nies pink in color, due to pale pink or polyp- outlined by a beaded border, and with a lon- ides and pinkish red of brooded embryos gitudinal groove down the front, having a showing through translucent calcification. transversely elongate orifice separate from and just above the orifice of the fertile zooid. Encrusted Thyroscyphus at the North Beach Breakwater, abundant from October-April, Embryos pinkish red. but disappeared in the summer months. Re- OCCURRENCE AND ECOLOGY: Found year- producing from November-February. round at all coastal stations, but more abun- DISTRIBUTION: Western Atlantic: Cape dant at Fort Pierce Inlet and Seminole Shores. Colonies, like those of Savignyella, Hatteras to Florida. May be cosmopolitan small in size and generally cryptic in habit. but identity has been confused, especially Occurred chiefly in the roots of Thyroscyphus with Celleporina costazii. and Endendrium, but also found on sponges, beach rock or breakwater rock surfaces, and ORDER CYCLOSTOMATA Sargassum. DIVISION ARTICULATA BUSK, 1852 DISTRIBUTION: Western Atlantic: Cape FAMILY JOHNSTON, 1838 Hatteras to Brazil. Gulf of Mexico and Ca- GENUS CRISIA LAMOUROUX, 1812 ribbean. Widely distributed in warm waters, Crisia elongata Milne Edwards, 1838 but noted only rarely due to small size and Figures 92,93 cryptic habit. Crisia elongata Milne-Edwards, 1838, p. 203. Osburn, 1940, p. 328; 1953, p. 684. Shier, 1964, FAMILY CELLEPORINIDAE HARMER, 1957 p. 647. GENUS CELLEPORINA GRAY, 1848 Crisia eburnea form denticulata Smitt, 1872, Celleporina hassalli (Johnston), 1847 p. 4. Figure 88 ICrisia denticulata Osburn, 1914, p. 185. Crisia denticulata Canu and Bassler, 1928a, p. Lepralia hassalli Johnston, 1847, p. 30. 156. 1982 WINSTON: MARINE BRYOZOANS 155

DESCRIPTION: (AMNH 633) Colonies bushy Crisia eburnea: Mature, 1957, p. 30. Cook, 1968a, white tufts found attached to undersides of p. 235. rock overhangs, hydroids, and algae, or DESCRIPTION: (AMNH 632) Colonies embedded in sponges. Zooids tubular, aver- forming delicate white tufts on stems of algae. aging .07 X .60 mm, and punctate, some- Branches curving slightly inward; joints yel- times with faint annular striations. Short lowish in color. Zooids tubular, punctate, peristome turned outward and sometimes about .10 X .25 mm. in size, orifices usually with a small denticle on its distal border. In- pointed distally. Internodes short, but vari- temodes of branches formed by alternating able in number of zooids, usually with five series of zooids, about 14-16 per internode to seven. Gonozooid usually occurring be- (though variable), connected by brown or tween the fourth and fifth zooid of a fertile black chitinous joints. Polypides with eight internode, elongate, club-shaped and rounded tentacles and a campylonemidan lophophore distally. Oeciopore a short tube with a trans- averaging .266 mm. in diameter. Gonozooid versely ovoid opening (fig. 91). short, broadly inflated, widest at distal end, generally with a faintly annulated surface. OCCURRENCE AND ECOLOGY: Found only Oeciopore a transverse slit located back of once in these collections, at the Sebastian the distal end of the gonozooid where it rests Grassbed on the red alga Solieria tenera. Col- against branch zooids. Mature larvae ovoid, onies small, but with gonozooids containing solid, and ciliated, greenish white in color, ripe embryos and larvae. Larvae which with no pigment spots. Gonozooids contain- emerged from the oeciopore while the colony ing embryos and developing larvae yellowish was being examined under the microscope brown in color. remained for a moment or two on the colony OCCURRENCE AND ECOLOGY: Collected and then took off for the water surface like year-round at three coastal stations, Sebas- tiny bees. tian Inlet, Fort Pierce Breakwater, and Sem- DISTRIBUTION: Common in the cold waters inole Shores. A dominant species in the lower of Europe and America. Western Atlantic: intertidal at Fort Pierce Breakwater during Labrador to Florida. Also found in West winter and early spring on Thyroscyphus and Africa. Eudendrium, as well as in association with sponges. Colonies grew vegetatively during DIVISION HAGENOW, 1851 the fall and gonozooids started developing in FAMILY GREGORY, 1899 early December. Gonozooids with embryos GENUS PLAGIOECIA CANU, 1918 were present in January and February. Al- Plagioecia dispar Canu and Bassler, 1928 though these large mature colonies were often Figure 94 fouled by fungi and diatoms they contained Plagioecia dispar Canu and Bassler, 1928a, p. 159. functional and active polypides. The species was less abundant in summer, but young col- DESCRIPTION: (AMNH 634) Colonies en- onies did occur. However, these colonies did crusting on dead shell and coral, fan-shaped not appear to contain functional polypides. when young, becoming subcircular with Many zooids had peristomes extended and growth. Zooids tubular, embedded for most openings closed off by diaphragms and ap- of their length, and separated from each other peared to be dormant or oversummering at by furrows. Peristomes short (about. 10 mm. the time. or less) and semi-erect with oval orifices, not DISTRIBUTION: Circumtropical. Western arranged in series. Brood chamber a promi- Atlantic: Florida. Gulf of Mexico and the nent swelling, making an arcuate band well Caribbean. inside the growing edge of the colony, and normally surrounding a number of peri- Crisia eburnea (Linnaeus), 1758 stomes (12 in the specimen illustrated). Figure 91 DISCUSSION: This species closely resembles Sertularia eburnea Linnaeus, 1758, p. 810. Plagioecia patina and may, in fact be syn- 156 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

TABLE 1 tea (Calvert, 1903) in the lack of concentric Indian River wrinkles on the colony and in slightly smaller (All Collections) micrometric measurements. Faint concentric HAULOVER CANAL wrinkles can be seen in the colony illustrated 1. Bowerbankia gracilis here, however. 2. Bowerbankia maxima OCCURRENCE AND ECOLOGY: Found at 3. Sundanella sibogae Gosnold stations encrusting shell and Ocu- 4. Zoobotryon verticillatum lina rubble. Brood chambers present on col- 5. Bugula neritina onies collected in February. 6. Bugula stolonifera DISTRIBUTION: Cape Hatteras to Straits of 1. Conopeum tenuissimum Florida. BANANA RIVER 1. Bowerbankia gracilis DISTRIBUTION AND ECOLOGICAL 2. Bugula neritina FACTORS 3. Conopeum tenuissimum The factors influencing bryozoan distri- SEBASTIAN GRASSBED bution have been discussed by many authors. 1. Crisia eburnea Most factors are interrelated and difficult to 2. Amathia alternata assess separately, as, for example, the rela- 3. Amathia vidovici tionships between turbulence, rate of depo- 4. Bowerbankia maxima sition, temperature, and water depth (see 5. Bowerbankia gracilis 6. Bowerbankia imbricata Cook, 1968a). 7. Zoobotryon verticillatum SALINITY 8. Beania klugei 9. Bugula turrita The factor known to most critically restrict 10. Bugula neritina the distribution of marine bryozoans is sa- 11. Conopeum tenuissimum linity. Only about 6 percent of living species 12. Membranipora savartii are able to survive in waters of reduced sa- 13. Scrupocellaria regularis 14. Savignyella lafontii linity (under 30%.) (Winston, 1977). 15. Watersipora subovoidea At coastal and offshore stations (Walton Rocks, Seminole Shores, Capron Shoals, and LINK PORT GRASSBED Gosnold stations) in the Indian River area 1. Amathia distans salinities remained at normal marine levels 2. Bowerbankia gracilis (35 to 37%o) throughout the survey. Within 3. Bowerbankia maxima 4. Nolella stipata the Indian River lagoon salinities varied both 5. Beania klugei with location and with time. The total range 6. Bugula neritina measured in 1974-1975 by the Indian River 7. Conopeum tenuissimum Coastal Zone Survey was from 14 to 34%o, 8. Electra bellula whereas at the inlets (Fort Pierce, Sebastian, 9. Schizoporellafloridana St. Lucie) salinity ranges between 30 and ST. LUCIE INLET 35%o were recorded (Young, 1975). 1. Bowerbankia gracilis Eighteen species of bryozoans were found 2. Zoobotryon verticillatum in the Indian River (table 1). Twelve species 3. Bugula stolonifera (three ctenostomes): Bowerbankia gracilis, 4. Bugula neritina Bowerbankia maxima, Zoobotryon verticil- 5. Conopeum tenuissimum latum: and (nine cheilostomes): Conopeum 6. Electra bellula seurati, Conopeum tenuissimum, Electra bellula, Beania klugei, Bugula neritina, Bug- ula stolonifera, Hippoporina verrilli, Schizo- onymous with it. According to Canu and floridana, and Watersipora subovo- Bassler (1928) it differs from Diastopora lac- idea) occurred in waters of less than 30%o 1982 WINSTON: MARINE BRYOZOANS 157 salinity. Other species, e.g., Nolella stipata, TABLE 2 Amathia alternata, Membranipora savartii, Link Port—Barge and Piling Bryozoan Traps were collected at stations within the river, (Glass Slides) but not at times when the salinity was under SPECIES 30%,. CTENOSTOMES Of the species that occurred in brackish 1. Alcyonidium polyoum (30%o) water only three: Conopeum tenuis- 2. Bowerbankia gracilis simum, Conopeum seurati, and Bowerbankia 3. Victorella pavida gracilis are truly brackish water species (hav- ing their chief abundances in brackish water ANASCAN CHEILOSTOMES and only occurring occasionally in fully ma- 1. Bugula neritina rine water). The other species are euryhaline 2. Bugula stolonifera 3. Conopeum seurati species characteristic of areas (like harbors) 4. Conopeum tenuissimum in which sudden strong variations in salinity 5. Scrupocellaria regularis and temperature may occur, but on a short- term basis, and in which conditions are gen- ASCOPHORAN CHEILOSTOMES erally marine (Gautier, 1962; Winston, 1977). 1. Hippoporina verrilli 2. Schizoporellafloridana 3. Watersipora subovoidea SUBSTRATUM The presence of a suitable surface for larval settlement and colony growth and the asso- zoan species most successful on this type of ciated category of variety of microhabitats substratum were those with short -spans, available (on or including different substrata) small zooid and colony size, and high repro- at any locality are among the most important ductive rates such as the Bowerbankia spp., environmental controls of bryozoan distri- and two species of Conopeum, of which Co- bution, as students of both living and fossil nopeum tenuissimum (able to reproduce in populations have pointed out (e.g., Chee- three weeks or less [Dudley, 1973a]) appeared tham, 1963; Cook, 1968a; Voigt, 1973; to be dominant. These species were present Thomsen, 1977). Many studies have shown during most of the year and could apparently the preference of bryozoan larvae of various survive considerable fluctuations in temper- species for a certain substratum (Ryland, ature and salinity. Of the other common 1959, 1962, 1976; Ryland and Stebbing, species, two, Bugula neritina and Bugula sto- 1971; Stebbing, 1972), or the association of lonifera, are common fouling species. Their colonies of a species with a substratum (Hay- distribution appears to be limited to tem- ward, 1973; Hayward and Ryland, 1975; perature rather than by substratum prefer- Moore, 1973; Rogick and Croasdale, 1949; ence as they were abundant only during the Stebbing 1971; Winston and Eiseman, 1980) winter months (see also Mook, 1976). or with a particular microhabitat or portion Another species, Schizoporella floridana, is of a substratum (Hayward and Harvey, known to form massive encrustations around 1974). As the station descriptions show, lo- Thalassia in localities farther south in Flor- calities sampled varied considerably with re- ida Bay (Multer, 1975). In the Indian River spect to type and number of substrata avail- area the colonies of S. floridana on Thalassia able. never reach more than a few millimeters in In the Indian River (table 1) the chief sub- size, yet these tiny colonies were often com- strata sampled were three species of sea- pletely mature, containing many ovicells. grasses, Halodule wrightii, Syringodium fili- The bryozoan traps set out at Link Port (table forme, and Thalassia testudinum. All are 2) supported a fauna similar to that found on relatively short-lived. In addition, seagrasses seagrasses, with the major exception of Hip- offer minimal surface area for somatic growth poporina verrilli. This encrusting cheilo- of colonies. As would be expected the bryo- stome, common in tropical estuarine envi- 158 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

TABLE 3 flat side of the breakwater, fine-grained sed- Sebastian Inlet - -Inner Breakwater—Sebastian, iment smothered colonies of encrusting Florida forms growing directly on the surface of the SPECIES rocks. Two normally sheetlike encrusting CYCLOSTOMES forms, Watersipora subovoidea and Thala- moporella floridana, there assumed an es- 1. Crisia elongata chariform growth habit by producing ruffled CTENOSTOMES bilaminate from an encrusting base. 1. Aeverrillia armata Many of the bryozoans found on the Sebas- 2. Amathia alternata tian Breakwater were arborescent types which 3. Amathia vidovici required very little attachment surface. One 4. Anguinella palmata 5. Bowerbankia maxima of the dominant species was the ctenostome 6. Bowerbankia gracilis Anguinella palmata, whose pendant colonies 7. Nolella stipata resembled mud-covered seaweed. Zoobo- 8. Zoobotryon verticillatum tryon, another ctenostome, was also common hanging from the undersides of the jetty ANASCAN CHEILOSTOMES rocks. The other species found at Sebastian 1. Aetea ska 2. Aetea truncata were mostly the delicate erect or encrusting 3. Antropora leucocypha forms that were characteristically associated 4. Beania klugei with the bases and stems of hydroids. 5. Bugula neritina Thirty-one species were collected from the 6. Bugula stolonifera North Beach Breakwater, Fort Pierce Inlet 1. Membranipora savartii (table 4), including one cyclostome, nine 8. Synnotum aegyptiacum ctenostomes and 21 cheilostomes. More col- 9. Thalamoporella floridana lections were made at the North Beach Break- ASCOPHORAN CHEILOSTOMES water than at any other locality, as it was the 1. Savignyella lafontii closest site to gather material for morpho- 2. Watersipora subovoidea logical and behavioral studies (Winston, 3. Vittaticella contei 1978), but the high number of species from 4. Schizoporella cornuta this locality probably reflected chiefly the 5. Cryptosula pallasiana number of available microhabitats. Var- iously oriented rock surfaces and the surfaces of caves and crevices between the rocks, pieces of wood wedged between the rocks and ronments was collected on oyster shells in the holes, and hollows in the sabellariid mounds Indian River, as well as on panels and slides, that encrusted many rocks, all provided sub- but was not found on seagrasses. strata. The North Beach Breakwater is ex- The four coastal localities studied were posed to strong wave and tidal action, but alike in containing a large number of micro- very few bryozoans lived in the most exposed habitats, but at each locality the environ- situations. One species that did is Watersi- mental conditions were somewhat different, pora subovoidea which formed a blackish and these differences were reflected in the crust on the rocks at the upper intertidal level. bryozoan fauna each locality supported. At The channels receiving the strongest wave the Sebastian Inlet inner breakwater 23 surges from the ocean to the inlet were com- species of bryozoans were collected (table 3) monly lined with colonial and including one cyclostome, eight ctenostomes sponges; among these were found the bushy and 14 cheilostomes. The breakwater, which white colonies of Crisia elongata, which offered sheltered conditions and a range of reached maximum development in associa- hard substrata, supported a great many ses- tion with certain sponges. Below the upper sile organisms: sponges, hydroids, soft , intertidal level the rocks (especially in chan- tunicates, as well as bryozoans. On the tide- nels and crevices) were covered by various 1982 WINSTON: MARINE BRYOZOANS 159

TABLE 4 TABLE 5 North Beach Breakwater—Fort Pierce Inlet - Walton Rocks - -Hutchinson Island, Florida Fort Pierce, Florida SPECIES SPECIES CYCLOSTOMES CYCLOSTOMES 1. Crisia elongata 1. Crisia elongata CTENOSTOMES CTENOSTOMES 1. Alcyonidium polypylum 1. Aeverrillia armata 2. Amathia alternata 2. Amathia distans 3. Amathia distans 3. Amathia vidovici 4. Anguinella palmata 4. Anguinella palmata 5. Bowerbankia maxima 5. Bowerbankia maxima 6. Bowerbankia gracilis ' 6. Bowerbankia imbricata 7. Bowerbankia imbricata 7. Nolella stipata 8. Nolella stipata 8. Valkeria atlantica 9. Sundanella sibogae 9. Zoobotryon verticillatum 10. Zoobotryon verticillatum ANASCAN CHEILOSTOMES ANASCAN CHEILOSTOMES 1. Aeteasica 1. Aetea ska 2. Antropora leucocypha 2. Antropora leucocypha 3. Beania hirtissima 3. Beania hirtissima 4. Beania klugei 4. Beania klugei 5. Beania mirabilis 5. Bugula neritina 6. Bugula turrita 6. Bugula stolonifera 7. Bugula minima 7. Bugula turrita 8. Membranipora tuberculata 8. Bugula uniserialis 9. Scrupocellaria regularis 9. Caulibugula pearsei 10. Synnotum aegyptiacum 10. Electra bellula 11. Thalamoporella floridana 11. Membranipora savartii ASCOPHORAN CHEILOSTOMES 12. Membranipora tuberculata 13. Scrupocellaria regularis 1. Celleporina hassalli 14. Synnotum aegyptiacum 2. Cryptosula pallasiana 15. Thalamoporella floridana 3. Exechonella antillea 4. Hippoporina verrilli ASCOPHORAN CHEILOSTOMES 5. Pasythea tulipifera 1. Cryptosula pallasiana 6. Savignyella lafontii 2. Celleporella carolinensis 7. Schizoporella cornuta 3. Microporella umbracula 8. Vittaticella contei 4. Parasmittina nitida morphotype B 9. Vittaticella uberrima 5. Savignyella lafontii 10. Watersipora subovoidea 6. Schizoporella cornuta 7. Schizoporella unicornis 8. Vittaticella contei 9. Vittaticella uberrima hydroids, of which the largest and most abun- 10. Watersipora subovoidea dant was Thyroscyphus ramosus. It was among the stems and roots of T. ramosus that most of the bryozoan species occurred. were Beania hirtissima, Beania klugei, Pa- Thalamoporella floridana did not form folia- sythea tulipifera, Synnotum aegyptiacum, ceous colonies as at Sebastian but instead Vittaticella contei, and Savignyella lafontii all grew in tubular fashion around the bases of of which occurred year-round. Thalamopo- the hydroid stems, the colonies only occa- rella floridana, Crisia elongata, and Celle- sionally rose into a bilaminate frill. The char- hassalli flourished during the winter acteristically hydroid-associated bryozoans months, whereas the stoloniferous cteno- 160 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

TABLE 6 sandy trough between the ledge and the Seminole Shores—Hutchinson Island, Florida beach. These subhabitats differed consider- ably with respect to physical stability. The SPECIES coquinoid ledge substratum was relatively CYCLOSTOMES stable, though environmental conditions var- 1. Crisia elongata ied seasonally (see section on temporal vari- CTENOSTOMES ation). Erect bryozoans attached directly to 1. Aeverrillia armata the undersurfaces of beachrock ledges, while 2. Alcyonidium polypylum other forms encrusted or attached to co-oc- 3. Amathia alternata curring hydroids and algae. 4. Amathia distans The other subhabitat, that of smaller 5. Amathia vidovici stones, shells, and boulders lying in the sand 6. Anguinella palmata channel between the ledge and the beach was 7. Bowerbankia maxima 8. Bowerbankia gracilis a most unstable one as the substrata were 9. Sundanella sibogae constantly being covered and uncovered by 10. Zoobotryon verticillatum shifting sand. During a one-month period, the level of sand in this area could change by ANASCAN CHEILOSTOMES 30 cm. or more. Colonies on stones that were 1. Aetea sica 2. Antropora leucocypha completely buried were destroyed by being 3. Beania hirtissima abraded off. Yet the stones, when only par- 4. Beania klugei tially covered with sand or other stones, sup- 5. Beania mirabilis ported a diverse fauna of encrusting bryo- 6. Bugula minima zoans, notably the ctenostome Alcyonidium 7. Bugula uniserialis polypylum, the cheilostomes Parasmittina 8. Membranipora savartii nitida morphotype B, Schizoporella cornuta, 9. Membranipora tuberculata Schizoporella unicornis, Cryptosula palla- 10. Scrupocellaria regularis siana, Membranipora savartii, and the ubiq- 11. Synnotum aegyptiacum uitous Watersipora subovoidea. Often colo- ASCOPHORAN CHEILOSTOMES nies on the underside of the stones were lying 1. Celleporina hassalli directly on the sand, but apparently the sand 2. Cryptosula pallasiana was porous enough and the water action suf- 3. Hippaliosina rostrigera ficient so that feeding and respiration could 4. Savignyella lafontii occur, and only long-term burial killed the 5. Schizoporella cornuta colonies. 6. Parasmittina nitida morphotype B 7. Stylopoma spongites At Seminole Shores (table 6) 31 bryozoans 8. Vittaticella contei were collected including one cyclostome, 10 9. Watersipora subovoidea ctenostomes, and 20 cheilostomes. The areas exposed at low water at Seminole Shores con- sisted mostly of sabellariid reef on coquinoid rock base. stomes were more common in summer and The dominant substrata for attachment, early fall. were the worm tubes themselves, the beach- At Walton Rocks 36 species including one rock stones and ledges, and the algae and cyclostome, 10 ctenostomes and 25 cheilo- hydroids (especially Eudendrium carneum) stomes were found, making it the richest lo- attached to the rocks. Watersipora subovo- cality in terms of total number of species idea was the only bryozoan commonly found (table 5). on worm tubes. As the beachrock at Seminole Two distinct subhabitats were available for Shores is more scoured than at Walton Rocks colonization by bryozoans at Walton Rocks; and contains fewer crevices for the attach- the coquinoid ledge and the loose beachrock ment of algae and hydroids, the total number stones and boulders resting on and in the of bryozoan species was smaller than at Wal- 1982 WINSTON: MARINE BRYOZOANS 161

TABLE 7 TABLE 8 Capron Shoals—Buoy 10A off Hutchinson Gosnold Collections (Oculina pinnacles and sand Island, Florida bottoms) Species Found on Oculina and on Gastropod Shells, Oyster or other Bivalve Shells SPECIES Stations 696-698, 700-702; 734, 735, 737, 738, CTENOSTOMES 757, 758, 759 1. Atcyonidium polypylum 2. Bowerbankia gracilis SPECIES CYCLOSTOMES ANASCAN CHEILOSTOMES 1. Plagioecia dispar 1. Aetea ska 2. Beania intermedia ANASCAN CHEILOSTOMES 3. Beania mirabilis 1. Antropora leucocypha 4. Bugula neritina 2. Aplousina gigantea 5. Bugula stolonifera 3. Bellulopora bellula 6. Cribrilaria radiata 4. Cleidochasma contractum 7. Discoporella umbellata depressa 5. Cleidochasma porcellanum 8. Fioridinella typica 6. Crepidacantha setigera 9. Membranipora arborescens 7. Cribrilaria radiata 10. Membranipora savartii 8. Cigclisula turrita 11. Scrupocellaria regularis 9. Floridina antiqua 12. Synnotum aegyptiacum 10. Fioridinella typica ASCOPHORAN CHEILOSTOMES ASCOPHORAN CHEILOSTOMES 1. Hippoporina verrilli 1. Escharina pesanseris 2. Microporella umbracula 2. Hippaliosina rostrigera 3. Parasmittina nitida morphotype B 3. Hippoporidra calcarea 4. Phylactellipora aviculifera 4. Hippothoa flagellum 5. Schizoporella cornuta 5. Membraniporella aragoi 6. Trypostega venusta 6. Micropora coriacea 1. Watersipora subovoidea 7. Microporella umbracula 8. Parasmittina spathulata 9. Parellisina latirostris 10. Phylactellipora aviculifera ton Rocks. Arborescent ctenostomes, Ama- 11. Reginella floridana thia spp., Bowerbankia spp., and Zoobotryon 12. Reptadeonella costulata verticillatum, and the cheilostomes, Bugula 13. Stylopoma spongites spp., and Scrupocellaria regularis were abun- 14. Tremoschizodina lata dant. The bottom surfaces of loose beach- 15. Trypostega venusta rocks in pools in the ledge were often com- Species found in sand from Station 766 pletely covered by encrusting cheilostomes. ANASCAN CHEILOSTOMES Two ctenostomes and 19 cheilostomes oc- 1. Discoporella umbellata depressa curred at Capron Shoals (table 7) where 2. Cupuladria doma patches of broken shells alternated with areas of sand bottom. Discoporella umbellata de- pressa, a free-living form, with a tiny conical colony that maintains itself on the shifting surface of the sand by means of long move- shell fragments, hence the dominant forms able spinelike vibracula, was present in sam- were encrusting. Most common were Alcyo- ples taken in July. This was the only living nidium polypylum, Microporella umbracula, species taken from the sand substratum Trypostega venusta, Cribrilaria radiata, and (though dead colonies of another lunuliti- Phylactellipora aviculifera. Panels which had form species, Cupuladria doma, were col- been exposed at Capron Shoals for six lected there). The main substrata available months were examined in January. Four to bryozoan settlement were dead shells and species (all common fouling species) oc- 162 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

TABLE 9 TABLE 9-(Continued) Association of Bryozoan Species with Various Substrata Anguinella palmata Beania hirtissima WOOD (Sticks and Boards) Beania klugei Bowerbankia gracilis Beania mirabilis Crisia elongata Bowerbankia maxima Cryptosula pallasiana Bugula stolonifera Hippoporina verrilli Bugula minima Membranipora tenella Bugula uniserialis Membranipora sp. Celleporina hassalli Schizoporella cornuta Crisia elongata Schizoporella floridana Exechonella antillea Scrupocellaria regularis Pasythea tulipifera Thalamoporella floridana Savignyella lafontii Scrupocellaria regularis LIVE GASTROPOD SHELLS Sundanella sibogae Antropora leucocypha Synnotum aegyptiacum GASTROPOD SHELLS OCCUPIED BY HERMIT CRABS Thalamoporella floridana Valkeria atlantica Antropora leucocypha Hippoporidra calcarea Vittaticella contei Watersipora subovoidea PLASTIC AND METAL Conopeum tenuissimum ALGAE Membranipora arborescens Amathia alternata Membranipora tenella Amathia distans Membranipora sp. Amathia vidovici OTHER BRYOZOANS {Zoobotryon, Bugula neritina, Aetea sica Amathia spp.) Beania hirtissima Aetea ska Beania klugei Aeverrillia armata Bowerbankia gracilis Bowerbankia gracilis Bowerbankia imbricata Bugula minima Bowerbankia maxima Nolella stipata Bugula neritina Savignyella lafontii Bugula minima Vittaticella contei Bugula stolonifera Bugula uniserialis SPONGES Celleporella carolinensis Bugula minima Crisia eburnea Crisia elongata Electra bellula Savignyella lafontii Escharoides costifer SABELLARIID TUBES Lagenicella marginata Membranipora tuberculata Amathia distans Microporella umbracula Bowerbankia imbricata Nolella stipata Crisia elongata Pasythea tulipifera Watersipora subovoidea Savignyella lafontii FREE LIVING Scrupocellaria regularis Cupuladria doma Synnotum aegyptiacum Discoporella umbellata depressa Thalamoporella falcifera Thalamoporella gothica HYDROIDS (Thyroscyphus and Eudendrium) Vittaticella contei Aetea truncata Watersipora subovoidea Aeverrillia armata Zoobotryon verticillatum Amathia distans 1982 WINSTON: MARINE BRYOZOANS 163

TABLE 9-(Continued)

SEAGRASSES (Syringodium, Halodule, Thalassia) Nolella stipata Aetea Parasmittina nitida morphotype B Amathia distans Parasmittina spathulata Beania klugei Parellisina latirostris Bowerbankia gracilis Pasythea tulipifera Bowerbankia maxima Plagioecia dispar Bugula neritina Phylactellipora aviculifera Bugula stolonifera Reginella floridana Conopeum tenuissimum Reptadeonella costulata Electra bellula Scrupocellaria regularis Membranipora savartii Schizoporella cornuta Nolella stipata Schizoporella unicornis Schizoporella floridana Stylopoma spongites Sundanella sibogae Synnotum aegyptiacum Watersipora subovoidea Thalamoporella floridana Tremoschizodina lata ROCK, SHELL AND CORAL Trypostega venusta Vittaticella uberrima Aetea sica Alcyonidium polypylum Amathia alternata Amathia distans Antropora leucocypha curred: Bugula neritina, Bugula stolonifera, Aplousina gigantea Hippoporina verrilli, and Watersipora subo- Beania hirtissima voidea. There are similarities between the Beania klugei Capron Shoals fauna and those of the two Beania mirabilis Bellulopora bellula intertidal stations, Walton Rocks and Sem- Bowerbankia gracilis inole Shores, which have the most similar Bowerbankia imbricata substrata. Only two ctenostomes were found Bowerbankia maxima at Capron Shoals, but one of these, Alcyo- Bugula neritina nidium polypylum was a dominant form at Bugula stolonifera Seminole and Walton rocks. Only four Bugula turrita species of the encrusting cheilostomes, Mem- Caulibugula pearsei branipora arborescens, Floridinella typica, Cleidochasma contractum Phylactellipora aviculifera, and Trypostega Cleidochasma porcellanum venusta, were not found farther inshore. The Crepidacantha setigera Crisia elongata last three, however, were collected from the Cribrilaria radiata deeper water Gosnold stations. An indication Cryptosula pallasiana of the primary importance of substratum Cigclisula turrita over other factors is that of the cheilostomes Escharina pesanseris collected during the Gosnold cruise one-third Floridina antiqua (nine species) occurred also in very shallow Floridinella typica subtidal to intertidal inshore stations. Hippaliosina rostrigera Both lunulitiform species, Cupuladria Hippoporidra calcarea doma and Discoporella umbellata depressa Hippoporina verrilli were also taken from sand bottoms of Fort Hippothoa flagellum Membranipora savartii Pierce Inlet and farther offshore. Twenty- Membraniporella aragoi eight species of cheilostomes were obtained Micropora coriacea from samples taken on the fossil Oculina Microporella umbracula reefs (table 8). Table 9 summarizes the association of 164 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173 bryozoan species found in Indian River area probably related to seasonal changes in other collections with various substrata: algae, sea- physical conditions, or to appearance or dis- grasses, wood, hydroids, other bryozoans, appearance of ephemeral substrata (e.g., algae rock and dead shell, gastropod shells, sponges, and certain hydroids). All habitats studied in and sabellariid tubes. detail showed such changes. At Sebastian Inlet January collections were TEMPORAL VARIATION marked by the occurrence of Watersipora subovoidea in foliaceous sheets at upper low- Even with collections made over the course water level, and by the presence of brittle, of a year it is only for the more abundant flower-like clusters of Thalamoporella flori- species that any reasonable idea of season- dana on the undersides of rocks in protected ally of occurrence and reproduction can be crevices. A cold spell in January in which the gained, but since few studies of bryozoans water temperature dropped below 15° C. from warm water regions have covered even killed some species (e.g., Bugula neritina, but this length of time all the information ob- did not affect others e.g., Bugula stolonifera). tained has been included in table 10. Offshore In April breakwater rocks were noticeably collections (Gosnold stations) were made coated with mud, and the most abundant only in September and February so not much bryozoan was the ctenostome Anguinella information can be gained from species palmata, whose colonies became coated with which were present only at those stations. sediment. September collections showed the What is obvious for coastal and river dwell- area dominated by hydroids (including Pen- ing species is that more than half of them naria tiarella, Thyroscyphus ramosus, and were present year-round (34 out of 64 species Eudendrium carneum), while colonies of soft collected at inshore stations). This number corals (Telesto sp.) covered the bases of the might be expected to rise with more intensive rocks along with masses of anemones and collection, as so many of these species are sabellid worms. Ruffled colonies of Water- small and cryptic. Abundance and condition sipora were again present, but ctenostomes during the course of the year varied from were still dominant, in the form of large species to species, however. Certain species mud-covered masses of Zoobotryon and appeared to be chiefly restricted by summer Bowerbankia maxima hanging from rock temperatures. Bugula neritina, Thalamopo- undersurfaces below the Watersipora level. rella floridana, Crisia elongata, and Amathia Bryozoans here grew more actively in the fall alternata achieved their maximum distri- than in the spring and summer months; the bution and abundance in winter. Colonies fauna was most diverse in the month of No- collected in summer had a senescent "over- vember when 16 species were collected. Al- summering" appearance, e.g., the loss of pig- though substrata were still muddy, more en- ment and degeneration of polypides in Bug- crusting species (Schizoporella cornuta, ula neritina, the fouled appearance and Antropora leucocypha, Cryptosula palla- calcification of diaphragms in Crisia elon- siana, Membranipora savartii, and Watersi- gata. Other species: Pasythea tulipifera, Syn- pora subovoidea) were present on shells, glass, notum aegyptiacum, Beania hirtissima, were and stones in crevices among the breakwater most abundant in summer months and col- rocks than at other times. Masses of Amathia onies were few or had a degenerated appear- spp. were most common a month later in ance in winter collections. A third group: December. Alcyonidium polypylum, Bowerbankia max- At the North Beach Breakwater, Fort ima, and Thalamoporella falcifera had peak Pierce Inlet, the hydroid Thyroscyphus ra- abundances in fall. This last category, and the mosus was abundant year-round. In January patterns of occurrence of remaining species this hydroid was dormant (with no feeding cannot be related to times or minimum or hydranths), but its perennial roots, stalks, maximum water temperatures. Variation in and branches provided an important sub- their occurrence and abundance over time is stratum for bryozoans, especially Crisia elon- 1982 WINSTON: MARINE BRYOZOANS 165

gata, Thalamoporella floridana, and Water- pifera was very common both on Thyroscy- sipora subovoidea. In this high energy phus and on the undersurfaces of rock crev^ environment the Watersipora and Thala- ices just below the Watersipora level. moporella colonies on hydroid roots re- Bowerbankia maxima was abundant. By Au- mained chiefly encrusting, only occasionally gust the worm mounds were worn and bro- forming bilaminar frills. Vittaticella contei ken. There were fewer sponges and tunicates and Savignyella lafontii were common on and few anemones, but Thyroscyphus and Thyroscyphus and among massive Eudendrium were still thriving; on them the colonies. Antropora leucocypha was common most abundant bryozoan was Beania hirtis- on the shells of snails (mainly Thais hae- sima. Very little Watersipora was present in mostoma floridana) living on the breakwater comparison with winter months. Some col- rocks. In February Crisia elongata was most onies encrusting the rocks were alive; most abundant and reproducing actively. Thala- of the colonies on the worm reef were dead. moporella and Watersipora were still flour- Fall (October) brought a return of the cooler ishing and were joined on the hydroid stems water fauna. At this time Watersipora colo- by Celleporina hassalli which was also re- nies began growing rapidly and on the Thy- producing. March was the month showing roscyphus (which was still active) Pasythea, greatest diversity of the Thyroscyphus fauna: Synnotum, and particularly Savignella were Thalamoporella, Crisia, Celleporina, Syn- abundant. Young colonies of Crisia (without notum aegyptiacum, Vittaticella contei, Sa- gonozooids) occurred, and Thalamoporella vignyella, Pasythea tulipifera, Bugula min- colonies were actively budding again. By ima, Beania hirtissima, and Beania klugei December when the water temperature were all present Scrupocellaria regularis was reached 20° C. crevices under rocks were common both on hydroid stems and break- crowded with masses of bushy bryozoans: water rocks. In April five days of very rough Crisia (with gonozooids developing), Ama- seas resulted in many of the sabellariid thia distans and Vittaticella uberrima (colo- masses being smashed and overturned; the nies five to eight cm. in size with gonozooids holes inside these structures were exposed containing embryos). On the Thyroscyphus, and the worms themselves were killed. The Thalamoporella, Amathia distans, Synno- storm also caused shifts in levels of sand in tum, and Crisia were dominant; Pasythea the crevices, burying many of the hydroids still occurred, but colonies were fouled and and scouring rock surfaces. On Thyroscyphus senescent in appearance. in the low to mid-intertidal level Thalamo- High tides and strong surf made it almost porella, Crisia, Beania hirtissima, and Cel- impossible to collect at Walton Rocks in win- leporina were the most common species. ter months as the coquinoid ledge remained Watersipora was abundant on worm tubes underwater. Watersipora occurred on the and in holes in the worm reef, whereas ledge itself. Species of Bugula were attached Amathia vidovici formed clumps on under- to the undersides of the ledge, whereas Vit- surfaces of rocks and in the holes in the worm taticella contei, Amathia convoluta, Bower- reef exposed by the storms. April marked the bankia gracilis, Beania klugei, Synnotum ae- first new growth of Thyroscyphus. By June gyptiacum, and Sundanella sibogae occurred and July Thyroscyphus colonies were flour- on hydroids attached to the ledge. By April ishing and the hydroid Eudendrium carneum environmental conditions had ameliorated. was also plentiful, whereas the sabellariid There was rapid growth of algae on the ledges mounds were collapsing. Noticeably fewer and also of Zoobotryon verticillatum. Bow- bryozoans occurred on Thyroscyphus in sum- erbankia gracilis occurred on Zoobotryon mer months when the hydroid was growing and algae, Electra bellula on the algae, actively and producing stinging nematocysts. whereas Scrupocellaria regularis, Bugula tur- Thalamoporella and Celleporina were lack- rita, and Vittaticella hung from the ledges. ing and Crisia was less abundant, though The sandy channel between the surf line and young colonies were present. Pasythea tuli- the ledges was deeply scoured. Loose beach- 166 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173 rocks lying on the sand had undersurfaces ledges making it impossible to collect there. covered by encrusting bryozoans: Alcyonid- The channel area was still scoured out and ium polypylum, Membranipora savartii, beachrock stones were abundant. Encrusting Beania hirtissima, Parasmittina nitida mor- species had large colonies. Parasmittina, S. photype B, Schizoporella cornuta, and Schi- cornuta, and Microporella were reproducing; zoporella unicornis. In June the beach profile A. polypylum, S. unicornis, and Cryptosula had flattened, the channel had filled in (to 30 occurred as large colonies, but were not re- to 60 cm. above its winter depth) and sand producing. had buried most beachrocks, boulders, and At Seminole Shores in January the worm dead shells. Zoobotryon, Amathia distans, reef was vigorously growing. Watersipora and Bugula turrita were attached to the occurred directly on worm tubes. Other bryo- ledges. Bugula neritina was present in the zoans were found only on hydroid stems and form of faded brown senescent colonies in algae attached to worm reef or rock surfaces. which only the distalmost zooids had func- Although the number of species was similar tional polypides. Bowerbankia maxima, to that at the other coastal stations colony Thalamoporella, and Watersipora occurred numbers were low and colonies found were on red algae, and A. polypylum, Schizoporella small in size and very cryptic, adhering cornuta, and Parasmittina had survived on closely to the rare beachrock stones or almost a few beachrocks (on most beachrocks there invisible among the hydroid roots. In Janu- were only dead and abraded colonies). At the ary species from hydroid-algal habitats in- July collection the sand level in the channel cluded Beania hirtissima, Beania mirabilis, had returned to the level of winter months. Synnotum, Sundanella, Crisia, Amathia vi- Many dead Watersipora colonies were noted dovici, and Bugula minima. As at Walton on the ledges (where they had been killed by strong surf and high seas hampered collec- the previous high sand level), but live colo- tion. In February the reef was completely nies were abundant in the pockets in the covered and only uprooted Sargassum fili- worm reef mounds caused by their seasonal pendulum and drift Sargassum spp. could be degeneration. Cryposula and Microporella collected. June found much of the worm reef umbracula encrusted dead shell. Zoobotryon eroded and dying, and sand had covered most was less luxuriant than in June. In September loose beachrock and shell material in ledge sand had again filled in around the rocks so pools, the quantities of hydroids and algae that only the upper surface of the ledges was had diminished. Zoobotryon specimens col- exposed. Zoobotryon persisted on the ledges, lected contained sperm. Aetea sica, Bower- though colonies were fouled with filamentous bankia gracilis, Beania klugei, Crisia, Ama- algae. Large branching masses of Bowerban- thia distans, Bugula minima, and Vittaticella kia maxima were attached directly to the contei occurred on hydroids or algae. Only rock surface. Alcyonidium polypylum, Wa- two encrusting species were found in June: tersipora, Parasmittina, and S. cornuta were Membranipora savartii and Schizoporella present as small colonies (young or regener- cornuta, but in July six encrusting species ated) on beachrocks. Fall (October) brought were collected from beachrock stones in high tides, surf, and winds. Most algae, hy- back-ledge pools: A. polypylum, Parasmit- droids, and bryozoans had disappeared from tina, Cryptosula, S. cornuta, and Stylopoma undersides of ledges, the channel area was spongites. In September young colonies of A. scoured out again and full of loose stones on polypylum, Watersipora, Beania klugei, and which encrusting species, especially Schizo- Amathia alternata were found on beach rock. porella unicornis, Schizoporella cornuta, and Masses of Zoobotryon and Bowerbankia Membranipora savartii, were abundant. Bug- maxima were tangled among the ula turrita, Bugula neritina, and Scrupocel- attached to the worm reef. In October the laria regularis were collected from the ledges worm reef mounds were still degenerated, in November. In December surf covered the though there had been some new settlement 1982 WINSTON: MARINE BRYOZOANS 167

TABLE 10 Seasonal Occurrence and Breeding in Bryozoans of the Indian River Area

COLONIES WITH EMBRYOS AND/OR SPECIES SPECIES COLLECTED (MOST ABUNDANT) YOUNG SETTLED COLONIES OBSERVED Bowerbankia gracilis Year-round December-June Bowerbankia imbricata March-September — Bowerbankia maxima March-November (September) March (young colonies) Zoobotryon verticillatum April to October June Amathia alternata Year-round (Winter) — Amathia distans Year-round (December) — Amathia vidovici Year-round — Valkeria atlantica August — Aeverrillia armata Year-round (Fall) — Alcyonidium polyoum April April Alcyonidium polypylum Year-round September-November Victorella pavida December December Sundanella sibogae Year-round — Nolella stipata Year-round — Anguinella palmata January-April — Aetea truncata November — Aetea sica February-October (Sept.-Oct.?) — Membranipora arborescens February-April (+?) — Membranipora savartii Year-round (April & October) — Membranipora tuberculata Year-round — Membranipora tenella Year-round (?) — Membranipora sp. Year-round (?) — Conopeum seurati December-May Dec-May (Late Winter-Spring) Conopeum tenuissimum Year-round Spring and Fall (December) Electra bellula Year-round — Cupuladria doma Year-round (?) — Discoporella umbellata depressa Year-round (?) — Aplousina gigantea February February Parellisina latirostris February February Antropora leucocypha Year-round — Floridina antiqua February — Micropora coriacea February February (?)" Floridinella typica October, February — Thalamoporella floridana Year-round (Winter) January Thalamoporella falcifera Year-round (Fall) — Synnotum aegyptiacum Year-round (Summer) — Scrupocellaria regularis Year-round January-April Bugula neritina Year-round (Winter) Jan .-June (late winter/early spring) Bugula stolonifera December-May December-April Bugula turrita November-April April Bugula minima January-June — Bugula uniserialis October — Caulibugula pearsei May — Beania hirtissima Year-round (August) — Beania klugei Year-round (Winter & Spring) — Beania mirabilis Fall and Winter — Membraniporella aragoi February February (?) Cribrilaria radiata October, February October & February Bellulopora bellula February (?) 168 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 173

TABLE 10 -(Continued) COLONIES WITH EMBRYOS AND/OR SPECIES SPECIES COLLECTED (MOST ABUNDANT) YOUNG SETTLED COLONIES OBSERVED Reginella floridana February Exochonella antillea March Escharoides costifer February February Savignyella lafontii Year-round (September-December) Year-round (?) Reptadeonella costulata February Tremoschizodina lata February Hippaliosina rostrigera October & February Plagioecia dispar February February (?) Parasmittina spathulata February February (?) Parasmittina nitida morphotype B Year-round Year-round Phylactellipora aviculifera Year-round (?) October Lagenicella marginata April April Schizoporella cornuta Year-round (October-December) October-December Schizoporella floridana January-August January-August (April-May) Schizoporella unicornis May and October May Escharina pesanseris February Stylopoma spongites February, April, October Cigclisula turrita February Cleidochasma porcellanum February February (?) Cleidochasma contractum February February (?) Hippoporidra calcarea April, September Microporella umbracula Year-round Late Fall and Winter Hippothoa flagellum February February (?) Celleporella carolinensis February, July February Trypostega venusta April, October, February Pasythea tulipifera Year-round (Summer) October Vittaticella uberrima April, December April, December Vittaticella contei Year-round Year-round (?) Celleporina hassalli October-April November-February Crisia eburnea March March Crisia elongata Year-round (Winter) January-February " (?) = Colonies ovicelled only.

of worms. There was very little algae on the spongites and Hippaliosina rostrigera also ledges except for one red alga, Lawrencia sp.; found. No collections could be made in No- the hydroid Eudendrium was thriving. On vember or December as the combination of Lawrencia were Bugula uniserialis, Vittati- poor tides, strong winds and high surf con- cella contei, Aetea sica, and Synnotum, ditions again prevailed. whereas Beania klugei, Vittaticella, Sunda- nella, B. uniserialis, Synnotum, and Scru- REPRODUCTION pocellaria occurred on Eudendrium. Beach- Information on reproduction (the presence rock stones were well scoured, and most of colonies with embryos) has been included bryozoan colonies were abraded and dead. in table 10. In temperate regions a few bryo- Membranipora savartii was fairly abundant zoan species (chiefly those with boreal dis- in depressions in the rocks, living (though tributions) reproduce in winter, but repro- partially abraded) colonies of Stylopoma duction and settlement of the majority of 1982 WINSTON: MARINE BRYOZOANS 169 species occurs in late summer and early fall I have also collected it), the range of mea- (Ryland, 1970). In tropical regions like West surements given for Bowerbankia gracilis Africa (Cook, 1968a) on the other hand, set- from subtropical and tropical localities by tlement occurs from November through other authors suggests that some of their April with the period of heaviest settlement material might be referable to B. maxima. from late January through April. In the In- One species, Conopeum seurati, has not dian River area some bryozoan reproduction previously been recorded from the Western occurs year-round, but peaks of reproductive Atlantic. Seven species: Victorella pavida, activity occurred in winter-spring and Alcyonidium polypylum, Crisia eburnea, fall-winter. Bugula neritina reproduces in Bugula uniserialis, Celleporella carolinensis, summer in Swansea, Wales, and from April Pasythea tulipifera, and Lagenicella margi- to December in Beaufort, N.C. (Ryland, nata have not previously been recorded from 1970). This species in Florida reproduced Florida. Reptadeonella costulata and Regi- from January to June with peak settlement nella jloridana have not been recorded from in late winter and early spring. Conopeum the east coast of Florida, although both occur tenuissimum reproduces between April and in the Gulf of Mexico. Phylactellipora avic- November in Chesapeake Bay (Dudley, ulifera has not been recorded north of the 1973a), between March and October in Beau- Tortugas. One species, Membranipora te- fort (Mature, 1959). Peak reproduction in nella, was recorded only once before from Chesapeake Bay occurred in summer. Indian Florida (the original description by Hincks, River populations reproduced from fall to 1880) and once from Brazil (Marcus) 1937, spring with peak reproduction in December. but may actually be quite common on float- Conopeum seurati which co-occurred with C. ing objects. Another species found on floating tenuissimum, started reproducing later and substrata, Membranipora sp., may also be had its peak of reproduction later in late win- common. ter and early spring. In general, most species found in the river and at coastal stations (intertidal to subtidal) BlOGEOGRAPHY are cosmopolitan or circumtropical in their It is difficult to assess the affinities of this distribution, although Bugula turrita, Bugula fauna as many species are probably more stolonifera, Aeverrillia armata, Membranip- widespread than has been recognized, but ora savartii, Conopeum tenuissimum, Cau- would not be identified or reported in general libugula pearsei, and Plagioeocia dispar have surveys because of the small size of their col- Western Atlantic or Western Atlantic plus onies and the cryptic nature of their habitats. Caribbean and Gulf of Mexico distributions. Only one new species Bowerbankia maxima The affinities of the species collected from is described here. Although its distribution the continental shelf Oculina reefs are gen- is here listed as Florida and Jamaica (where erally tropical.

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