Bryozoa of the Southeast Coast of the United States: Bugulidae and Beaniidae (Cheilostomata: Anasca) 1

BRYOZOA OF THE SOUTHEAST COAST OF THE UNITED STATES: BUGULIDAE AND BEANIIDAE (CHEILOSTOMATA: ANASCA) 1

FRANK J. S. MATURO, JR. Department of Zoology, University of Florida, Gainesville

ABSTRACT The species of Bugula, Caulibugula, and Beania which occur from Cape Hatteras to Florida are described and figured. The genus Bugula includes B. Iwritina, B. tun"ita, B. fulva, B. stolonifera, B. microoecia, B. rylandi sp. n., B. marcusi sp. n., and B. grayi sp. n. Records of BURlIla avicularia and B. cali/ornica from the western Atlantic are erroneous or dubious. Bugula fulva and B. stoloni/era are recognized for the first time in the American fauna. The northern range limit of Bugula microoecia is extended from the Gulf of Mexico to North Carolina. The genus Caulibugula is represented by C. dendrograpta (Waters) and C. pearsei sp. n. Caulibugula armata Verrill is redefined. Both Beania hirtissima and B. intermedia are now known to occur as far north as Cape Lookout, N. C.

INTRODUCTION AND ACKNOWLEDGMENTS While the author was engaged in a study of the Bryozoa of Beaufort, North Carolina, and vicinity (Maturo, 1957), it became apparent that a rich bryozoan fauna is present in the offshore waters of the Carolina coast. Furthermore, except for a few limited localities, this fauna has not been examined in any detail from Cape Hatteras to southern Florida. The present report is the first of a series which will consider the Bryozoa of the offshore waters of this large area. The material upon which this study is based was obtained from a number of sources. The author collected in the areas off Beaufort, Cape Lookout, and New River, North Carolina; Sapelo Island, Georgia; and Fernandina Beach and Matanzas Inlet, Florida. Very valuable collections from off Oregon Inlet, Cape Hatteras, and Cape Lookout were generously donated by Dr. Maximo Cerame-Vivas and Dr. 1. E. Gray of Duke University, obtained under the auspices of the latter's grants NSF G-13952 and G-25128. The writer was especially fortunate to have the opportunity to examine a very extensive collection of bryozoans from several hundred stations along and off the Georgia coast collected by Mr. Milton Gray, sponsored by the Trustees of the Sapelo Research Foundation. Various individuals have also contributed specimens and are acknowledged where appropriate throughout the text.

'This study was supported by National Science Foundation Grant NSF G-24309. Part of the manuscript was prepared during tenure of a National Academy of Sciences-National Research Council Research Associateship at the Smithsonian Institution. 1966] Maturo: Bryozoa of Southeast United States 557 Special thanks are expressed to Dr. Cerame- Vivas and Dr. Gray for their cooperation on many collecting trips; and to Miss Patricia L. Cook, British Museum (Natural History), for aid in comparing my specimens with appropriate collections in the Zoology Department. I am indebted to Dr. H. K. Brooks, Mr. Herbert Eppert, and Mr. Reginald Scolaro of the Geology Department, University of Florida, who rendered valuable services as a SCUBA diving team in localities off North Carolina and Florida; to the Duke University Marine Laboratory and the University of Georgia Marine Institute for providing generous use of their facilities; to the staff of the United States National Museum for making available its collection; to Dr. Willard Hartman for the loan of specimens from the Yale Peabody Museum; to Mr. Thomas Hopkins, Mr. Arnold Ross, and especially, Mr. Scolaro, for invaluable aid as research assistants. III the interest of saving space I have omitted from the following accounts detailed lists of station sites from which the species were collected and have replaced them with general locality names. These names used in conjunction with the depth information are usually sufficient to establish the general habitats and ranges on U. S. Coast and Geodetic Survey charts. Where pertinent, compass bearings and other data are given to help define northern or southern range limits along the southeast coast. The format of this report has been simplified as much as possible to provide only the essential systematic information; the synonymy is given only at the species level and this is limited to the original designation, followed by pertinent American citations. Keys to the genera and species included in the report are provided to aid the non-specialist in making identifications. Also, the confusing and complex terminology prevalent in bryozoan literature has been avoided as far as possible. All terms used in the descriptions are those adopted and defined by Hyman (1959). Particular specimens referred to in the descriptions are identified by catalog numbers: numbers preceded by USNM designate specimens in the collections of the U. S. National Museum; BMNH means British Museum (Natural History); YPM is Yale Peabody Museum. Holotypes and figured specimens have been deposited in the U. S. National Museum. All figures were prepared with the aid of a Wild M5 drawing apparatus.

FAMILY BUGULlDAE GRAY, 1848 Key to Genera Considered Below Colony with segmented stalk composed of higWy modified zooids (kenozooi ds ) Caul ibug u la Colony without stalk, or if stalk present, it is composed of unsegmented regular zooids and radicles (radicular fibers or rhizoids) Bugula 558 Bulletin of Marine Science [16(3) Genus Bugula Oken, 1815 Genotype.-Sertularia neritina Linnaeus, 1758. Bugula was automatically rejected as a generic name when the Inter- national Commission on Zoological Nomenclature (1956) rejected Oken's names. However, in the interest of stability, this most familiar of ecto- proct generic names continues in use pending the time when it is officially reinstated. Description.-Colonies are unilaminar, erect and branching; zooids arranged in two or more series, alternate, facing same direction, usually truncate distally and narrowed proximally with a forked proximal end, the distal end of each zooid extending above the base of the succeeding one. The frontal membrane occupies nearly all of the frontal surface; the operculum is indistinct or absent. Spines may be present, especially at the distal angles of the zooids. Pedunculate avicularia are usually present. Ovicells are hyperstomial, typically globular, occasionally reduced, and usually attached by a short stalk to the distal end of the zooid. Discussion.-Ryland (1960) has prepared an excellent review of the generic characters and a valuable discussion of useful features for the separation of species. Most of these will become obvious in the following key and descriptions; however, two characters need some explanation here. These are spine formula and bifurcation type. The number and position of spines on the distal angles of zooids can be expressed as a formula, such that 3: 1 means three spines on the outer distal angle (always given first) and one spine on the inner distal angle. In the description of species with a 2: 1 formula, two terms are introduced to identify the outer spines: frontal outer distal spine and dorsal outer distal spine; the former, often being merely a short conical or denticle-like projection, is placed to the front and at the base of the latter. The modes of branching in biserial species fall into three types as classified by Harmer (1923, 1926). These are designated as types 3, 4, and 5. The type is usually constant for a given species if one ignores the most proximal bifurcations. For purposes of classification the zooids involved in a bifurcation are designated by letters: the first zooid from which two are budded in preparation for a bifurcation is designated as A, its alternate (which also buds two) beside it is called B; the outer zooids above A and Bare C and D, respectively, the inner zooids being E and F. Zooid G is above E; zooid H is above F. The inner angle of the bifurcation is formed by zooids E and F in type 3 (Fig. 1), by zooids F and G in type 4 (Fig. 2), and by G and H in type 5 (Fig. 3). The bifurcation patterns show most clearly on the dorsal (basal) side of the colony. The measurements included in the descriptions are made as suggested by Ryland, namely, they were obtained from the dorsal side, zooid length 1966] Maturo: Bryozoa of Southeast United States 559

FIGURES 1-3. Basal (dorsal) views of some species of Bugula showing bifurcation types (see text for explanation). Scale indicated; avicularia not shown. 1, Type 3 (Bugula rylandi, part of holotype, USNM 11895); 2, Type 4 (Bugula stolonifera, USNM 1] 899); 3, Type 5 (Bugula fulva, USNM 11~01). being the distance between two zooid apices and zooid width being the maximum width. Zooids involved in a bifurcation or comprising the stem axis in spiral colonies were excluded.

KEY TO THE SOUTHEASTERN SPECIES OF Bugula 1. Avicularia absent, colony reddish-purple or reddish-brown ______.. .. neritina Avicularia present, branches of colony whitish, cream, buff, or yell owi sh -ora nge .______2 2. (1) Inner distal angle of zooid always unarmed, rounded or acute, outer angle with one, rarely two, spines or denticle-like pro jec ti 0ns ...... ______3 Both distal angles usually bearing at least one spine or finger- like projection ...... __ __ .___ _ .______4 3. (2) Frontal membrane occupying three-fourths or more of length of frontal surface, avicularia exceed zooid width and are situated near top or not more than one-third way down lateral margin of frontal membrane _.. .. microoecia Frontal membrane occupying two-thirds of length of frontal surface, avicularia less than zooid width and are situated half way down lateral margin of frontal membrane turrita 4. (2) Outer distal angle of zooid with one or two spines or projections, inner distal angle bearing one spine 5 Both distal angles bearing at least two spines; may have some branches m uItiser j al ..__. ._. . . . fulva 560 Bulletin of Marine Science [16(3) 5. (4) Outer distal angle of zooid with one spine or finger-like projection ._. ..__ __ 6 Outer distal angle of zooid with two spines or projections, the frontal-most one may be reduced to a denticle-like projection or knob .______7 6. (5) Lateral margin of frontal membrane inrolled to form an angle between outer distal spine and avicularium, frontal membrane occupying about two-thirds of frontal surface avicularia less than zooid width (about 0.14 mm long) and placed about halfway down margin of frontal membrane ____ turrita No angular projection of margin between distal spine and avicularium, frontal membrane occupying three-fourths or more of frontal surface, avicularium equal to zooid width (about 0.18 mm long) and placed one-fourth to one-third way down margin of frontal membrane .. .______rylandi 7. (5) Bifurcations of type 3 only . .______.___ __ 8 Bifurcations mostly of type 4, occasionally with some proximal ones of type 3 and distal-most ones of type 5 stolonifera 8. (7) Length of avicularia less than zooid width (about 0.14 mm) ______. . .__...... turrita Length of avicularia equal to or exceeds zooid width (might be smaller in most proximal part of colony), avicularium somewhat elongate rectangular ._.______9 9. (8) Avicularium conspicuously swaybacked (with a prominently concave dorsal margin), not exceeding 0.18 mm in length ___. . ._.. .marcus i Avicularium with a slightly concave dorsal margin, exceeding 0.18 mm in length . . . . . grayi

Bugula neritina (Linnaeus) Fig. 4

Sertularia neritina Linnaeus, 1758: 815. Acamarchis neritina: Verrill, 1878:304. Bugula neritina: Verrill, 1900:593. Bugula neritina: Osburn, 1914:186; 1927:126; 1940:389; 1947:22. Bugula neritina: Marcus, 1937:67. Bugula neritina: Maturo, 1957:42; 1959: 117. Bugula neritina: Shier, 1964:624.

Description.-Colonies consist of erect bushy tufts which may be 8 em or more tall; bifurcations of type 4 and 5. The species is easily recognized by its reddish-brown or reddish-purple color and its algal-like appearance. Zooids are biserial, alternate, tapering proximally, the frontal membrane extending the entire length of the frontal surface. There are no avicularia 1966J Maturo: Bryozoa ot Southeast United States 561

O.5mm

FIGURES 4-6. All figures drawn to scale indicated. 4, Bugula nerama (Linnaeus), showing three ovicells, USNM 11902; 5, Bugula microoecia Osburn, showing ovicells on the three zooids of left side, USNM 11906; 6, Bugula turrita (Desor), USNM 11908.

or spines; however, the distal corners are sharply angular. Ovicells are large, globular, lateral facing, and arise from the median distal corner of the zooids so as to form a linear series, one above the other, in mature colonies. Zooids are about 0.63 mm long and 0.27 mm wide. Distribution.-The species has a worldwide distribution in warmer waters; in the western Atlantic it ranges from North Carolina to Brazil. 562 Bulletin of Marine Science [16(3 ) Bugula microoecia Osburn Fig. 5 Bugula microoecia Osburn, 1914:187; not 1940:391 (see below). Description.-Colonies are phytoid, composed of a central stalk and stems from which very delicate, feathery, branches arise; stalk orange, branches white; up to 20 em or more tall; all branches jointed at the base; bifurcations of type 3. The stalk is composed of greatly elongated zooids, whose internal organs have degenerated to brown bodies, and of tangled masses of radic1es. Zooids are biserial, alternate, tapering slightly toward the base, the frontal membranes turned toward the median axis of the branch and extending practically the entire length of the frontal surface. The joints occur just above the basal forks of the zooids starting new branches. The spine formula is 1:0, the spine being essentially an elongation of the outer distal corner. Avicularia are placed high on the lateral margin (about one-third the way down), robust, length exceeding the width of the zooids, beak short and decurved. The ovicells appear to be very reduced, shallow arches, scarcely elevated beyond the distal margin of the zooids, the exposed frontal surface flattened as an extension of the frontal membrane of the zooid. The ovicell chamber is actually well developed, occupying a deep pocket in the dorsal side of the zooid. Regular zooids are 0.45-0.63 mm long and 0.14-0.18 mm wide; some stem zooids are more than 2 mm long. The avicularia are usually 0.18- 0.23 mm long, but those on the stem zooids may reach 0.27 mm. Remarks.-Specimens from off the Georgia coast collected in June possessed ovicells with embryos. Many of the zooids in the material collected possess a frontal membrane modified in the region of the orifice to suggest an operculum. Osburn's type material from the Tortugas islands consists of two slides collected from different substrates. Both are labeled in Osburn's hand as "type." The original description and figures agree with the material on slide USNM 11843, bearing the label "Bugula microoecia, type. Tortugas, Fla. 18 fath. on bottom of coral sand. R. C. Osburn," which is hereby designated the lectotype. Slide USNM 11882, "Bugula microoecia, type. Tortugas, Fla. 18 fath. on the leg of a crab, Hyas sp. R. C. Osburn," is designated the paralectotype. The avicularia of the type series are conspicuously smaller (about 0.14 mm) than those in the present material. But for generally larger dimensions, my specimens agree with Osburn's. The ovicells figured by Osburn (1914:Fig. 1) need interpretation. The somewhat wavy line shown running horizontally below the distal margin of the two upper ovicells is not the frontal border of the oviccll as it suggests but is actually the margin of the distorted valve formed by the 1966] Maturo: Bryozoa of Southeast United States 563 distal portion of the frontal membrane which closes the ovicell aperture. The area between these lines and the distal border (actually the frontal border) of these ovicells is in fact the exposed ovicell aperture. The corresponding line in the lower right zooid represents the distal border of the zooid showing through the transparent frontal membrane. In Figure 5 in the present study the three zooids on the left have ovicells, the middle ovicell having its aperture partially opened. Distribution.-The species is rather common on shelly bottoms offshore from Sapelo Island, Georgia, at depths of 54-78 ft (16.3-23.6 m). A single specimen of what appears to be the stalk and stems of a large colony was collected from the washings of a shrimp trawl at Fort Macon, Beaufort Inlet, North Carolina. Osburn (1940) reported it from Puerto Rico in 71/2 fathoms (13.7 m). However, his specimen (USNM 11889) has become badly distorted by decalcification. It cannot be assigned to Bugula microoecia because it has a 2: 1 spine formula, type 4 bifurcations, and no discernible joints. Bugula turrita (Desor) Fig. 6 Cellularia turrita Desor, 1848: 66. Cel/ularia fastigiata: Leidy, 1855:142. Cellularia turrita: Perkins, 1869: 161. Bugula turrita: Verrill & Smith, 1874:712. Bugula turrita: Verrill, 1878:304; 1879a:52; 1879b:189; 1879c:29. Bllgllla gracilis var. uncinata: Osburn, 1912: 224. Bugula turrita: Osburn, 1912:225; 1944:40; (?) 1947:23. ? Bugula tun'ita: Marcus, 1937:68. Bllgula turrita: Hutchins, 1945: 541. Bugllla tun'ita: Rogick & Croasdale, 1949:51. Bllgula tllrrita: Maturo, 1957:43; 1959: 123. Description.-Colony consists of erect branches with secondary branches arranged in a whorled or spiral pattern about the long axis; height up to about 8 cm; bifurcations of type 3. Zooids biserial, alternate, tapering toward the base, the frontal membranes occupying about two-thirds (sometimes three-fourths) of the length of the frontal surface, orificial area facing distally and oblique to the frontal plane. The distal spine formula is basically 2: 1 although the degree of development of these spines is quite variable. The frontal outer distal spine is usually very short (rarely, lost so that only a frontal marginal angle persists in its place) while the dorsal outer distal spine and the inner distal spine may be well developed; occasionally, all three may be reduced to very short conical projections. Zooids are 0.45-0.54 mm long and about 0.15 mm wide, exclusive of zooids involved in bifurcations or comprising the main branch axes. There are 14 tentacles. The avicularia are small, slightly less than the width of 564 Bulletin of Marine Science [16(3)

E E oIf)

FIGURES 7-11. All figures drawn to scale indicated. 7, Bugula lulva Ryland, showing ovicells, USNM 11900; 8, Bugula lulva Ryland, showing proximal spines, USNM 11901; 9, Bugula rylandi new species, holotype, USNM 11895; 10, Bugula rylandi new species, paratype showing ovicells, USNM 11896; 11, Bugula marcusi new species, holotype, USNM 11897. 1966] Maturo: Bryozoa of Southeast United States 565 the zooids (average 0.14 mm), possess a decurved beak, and are located about halfway down the lateral margin of the frontal membrane. The ovicells are elongate globose, placed slightly to the median side of the zooid axis and tilted toward the axis of the branch. The globose appearance is evident only during the time an embryo is present in the ovicell, otherwise, the maturing or spent ovicell appears to be a shallow, cap-like hood. The ancestrula and ancestrula-like primary zooids of secondary colonies are inverted cones with the frontal membrane reduced to a circular area at the distal end which is perpendicular to the zooid axis. In a previous paper (Maturo, 1957: 45) the ancestrulas were described on the basis of figures and descriptions given by Marcus (1938: 67-72 ) . Remarks.-None of the material at hand shows the circle of spines surrounding the frontal area of the ancestrula as figured by Marcus, but instead agrees with the description by Ryland (1960:98). Very probably a closer study of Marcus's species will indicate that it is not Bugula turrita but a closely related species. No specimens collected during this investigation showed the vigorous growth exhibited by New England material collected at approximately the same time of the year (late June-early July). Most were 3 cm or less in height, although one lot from off Fernandina Beach, Florida, contained rather scrawny colonies up to 5 cm high. The species was collected on rocky outcroppings in depths up to 17 fathoms (31.1 m). Reproductively mature colonies were collected in June. Distribution.-The species is distributed from Massachusetts to Florida; records for the Caribbean and Brazil should be reexamined for confirmation. Bugula fulva Ryland Figs. 3, 7, & 8 Bugula ditrupae: Marcus, 1937:69. (Not Busk, 1858). BlIgu/a fu/va Ryland, 1960:86; 1962:28. BlIgu/a aqlli/irostris Ryland, 1960: 89. Description.-Older colonies consist of erect bushy tufts while younger ones are f\abellate to cup-shaped; color yellow-brown or buff; 2-3 cm tall; bifurcations of types 4 and 5 in basal portions, but most branches arise from a simple dichotomy of multiserial rows of zooids. Zooids are biserial in basal portions of colonies, becoming multiserial distally, with branches up to seven zooids wide in material examined; marginal zooids tapering slightly toward the base with frontal membranes extending three-fourths or more of the length, internal ones rectangular or slightly hexagonal with frontal membranes extending the entire length. The distal spine formula is typically 3:3, although 4:3, 3:2, and 2:2 566 Bulletin of Marine Science [16(3) formulas have been observed in the same colonies, the latter condition typical of inner zooids bearing ovicells. There may also be one or two spines placed proximally on the sides of the frontal membrane. Avicularia with decurved beaks are present on both inner and outer series of zooids, the outer avicularia being larger (equal to or exceeding zooid width) and placed about one-third the way down the lateral margin, the inner ones being smaller (about three-fourths of the zooid width) and usually placed halfway down or lower on the margin. Ovicells are raised and globular, and aligned with the principal axis of the branch. The ancestrula is an inverted cone with a short frontal membrane surrounded by spines: one median proximal, one lateral proximal on each side, and three at each distal corner. Zooids are 0.43-0.90 mm long and 0.14-0.21 mm wide, the marginal zooids usually the longest. Marginal avicularia are 0.18-0.25 mm long; inner avicularia are 0.08-0.11 mm long. Remarks and Dislribution.-Embryos were present in ovicells of specimens of each lot collected, namely, mid-March, early May, mid-June, and late July. The species has been collected from depths of 72 feet (21.8 m) on reefs 12 miles south of Cape Lookout, North Carolina, and depths up to 175 feet (53.1 m) 85 miles ESE of Sapelo Island, Georgia. On March 18, 1962, specimens were found washed ashore in abundance at Matanzas Inlet, Florida, occurring in windrows along with B. neritina upon which they had been growing. This is the first report of the occurrence of this species from the western North Atlantic, Ryland having recorded it from the British Isles and the Mediterranean Sea. Marcus has collected it from Brazil. Two slides in the Hutchins Collection (USNM 11888: Bugula avicularia. Casco Bay. YPM-Lot 68) from Maine appear referable to this species, as are Verrill's specimens from "Ram Island Ledge" (presumably, in Great Harbor, Woods Hole) (USNM 4981). Miss Cook has kindly compared the Matanzas Inlet specimens with the types of B. fulva Ryland (BMNH 1959.3.18.5) and B. aquilirostris Ryland (BMNH 1959.1.20.1) and reports them to be in the middle of the range of variation between the types which Ryland (1962) has combined into a single species.

Bugula rylandi, sp. n. Figs. 1,9, & 10 Material.-Two slides, part of same colony, from off Sapelo Island, Georgia, 5 miles from Sapelo whistle buoy (back bearing 340°), 72 feet (21.8 m), drag 3, F. J. Maturo, Jr., call., June 6, 1962: HOLOTYPE, USNM 11895. Other specimens from off Sapelo Island, Georgia, 19 miles from sea buoy (bearing 35°), drag 12, M. Gray, coIl., January 31, 1961: PARA- 1966] Maturo: Bryozoa oj Southeast United States 567

TYPES, USNM 11896; author's collection nos. 132, 970-979, 1830 A-C, 1850 A-C, 1852-1857, from various localities off Sapelo Island. Descri[Jtion.-Colonies consist of erect branches with secondary branches arranged about them in a spiral pattern; height up to 4 em; bifurcations of type 3. Outer branches are whitish in color, whereas, the main stems which are masses of fused radicles are a rich cream in alcohol preparations. Zooids are biserial, alternate, tapering toward the base, the frontal membranes occupying three-fourths or more of the length of the frontal surface. Frontal membranes of zooids just above bifurcations may extend only half the zooid length. The distal spine formula is 1: I, both spines being prominent. The outer spine is broad-based and conical, increasing in size toward the distal end of the branches. The avicularia are placed high on the lateral margin, in a position comparable with that of the frontal outer distal spine of the 2: I formula species or slightly lower (between one-fourth to one- third of the way down the frontal membrane). Their length is about equal to the width of the zooids; the beak is decurved. Ovicells are vestigial, elevated, shallow, cap-like, poorly calcified, with the frontal surface entirely membranous; they are tilted toward the median axis of the branch. The ancestrula is an inverted cone with the frontal membrane circular and facing distally, its border devoid of spines. Zooids are 0.45-0.62 mm long and 0.15-0.23 mm wide. The ratio of length to width is quite variable, both short and long zooids may attain the maximum widths. The short and very wide zooids look markedly different from the usual form because of a tendency to develop a "shouldered" appearance at the outer distal angle where the spine originates. Avicularia are 0.14-0.20 mm long. Remarks.-Bugula rylandi can be distinguished from the other members of the genus along the southeastern coast by its 1: 1 spine formula. Bugula carvalhoi Marcus from the Brazilian coast also has a 1: 1 spine formula but it differs from B. rylandi in having fan-shaped colonies, type 4 bifurcations, longer frontal membranes, lower placed avicularia, shorter ovicells, and in being occasionally triserial. Bugula rylandi has an ancestrula like that of B. plumosa (Pallas) but it differs from that species in spine formula, shape and position of avicularia, and ovicell character. Bugula rylandi bears some resemblances to B. aperta (Hincks) but may be distinguished from it by the 1: 1 spine formula, avicularia with more strongly decurved beaks, and a tendency towards stouter zooids. Mature ovicells with embryos occurred on specimens collected in late January and early February. I take pleasure in naming this species in honor of Dr. J. S. Ryland, who 568 Bulletin of Marine Science [16(3) has done much to clarify the systematics of the genus Bugula in his studies on marine ectoprocts. Distribution.-The species has been collected to date only offshore from Sapelo Island, Georgia, in depths of about 70 feet (21.2 m). Bugula stalanifera Ryland Figs. 2 & 12 ? Bugula californica: Marcus, 1937:71. Bugula avicularia: McDougall, 1943:339. Bugula californica: Maturo, 1957:45. ? Bugula avicularia: Shier, 1964:624. Bugula stolonifera Ryland, 1960:78. Descriptian.-Colonies consist of erect branches which project out in all directions from the short basal portion giving a cup or funnel shape to the younger colonies; older colonies with additional branches forming rather dense tufts which have a somewhat whorled branching pattern. All of the specimens at hand were taken from test panel surfaces which were sub- merged for 2-week intervals; the largest specimen is 1.5 em tall. The bifurcation pattern is variable, predominantly type 4, with the type 3 condition being found in the most proximal bifurcations and type 5 generally occurring in the most distal parts. Zooids are biserial, alternate, tapered slightly toward the base, with the frontal membrane occupying about three-fourths or more of the length of the frontal surface. The distal spine formula is 2: 1 with some variation in occasional zooids. The spines are well-calcified, tubular, and usually well developed (most of the spines shown in Fig. 12 have been damaged). The "E" zooid of a bifurcation frequently may have a 1: 1 spine pattern and, in rare instances, some individuals involved in the bifurcations may possess a 2: 2 formula. The position of the avicularia varies from about one-fourth to one-half the way down the lateral margin of the frontal membrane from the base of the frontal spine. The avicularia have prominent heads with rather full, rounded profiles and short decurved beaks. They characteristically fall into three size classes: (1) very small avicularia, occasionally absent, which occur on the "E" zooids of type 4 bifurcations, 0.07-0.11 mm long; (2) large avicularia which exceed the width of the zooids, 0.18-0.23 mm long; and (3) intermediate sized avicularia, 0.14-0.17 mm long, which are usually located on zooids close to and composing the inner angle of a bifurcation. The ovicells are subglobular, high, extending above the lower margin of the frontal membrane of the next upper zooid. The zooids, exclusive of those involved in bifurcations, are 0.54-0.63 mm long and 0.16-0.18 mm wide. The ancestrula possesses a short slanting frontal membrane and a 4-2: 2 distal spine pattern plus a median proximal spine (see Maturo, 1957:Fig. 47). 1966] Maturo: Bryozoa of Southeast United States 569

Remarks.-The description given above emends to some extent the original description by Ryland. Ryland did not find the minute avicularia at the bifurcations, large avicularia exceeding the width of the zooids, or type 5 bifurcations in either British specimens or material from the Bay of Naples (1962). In fact it appeared that the American material belonged to a new species until I examined specimens in the Osburn Collection (USNM 11892: Bugula spicata Hincks. Naples. R.C.O.) which have the characters mentioned above, occur within the range of B. stolonifera, and seem clearly referable to it. I am grateful to Miss Cook for comparing some of my material with the type of B. stolonifera, BMNH 1959.3.18.1, Swansea Docks. This species bears strong resemblances to B. californica Robertson, but the latter species differs in having a strongly spiral growth form, bifurcations which superficially resemble several types in Harmer's classification but which actually are merely the splitting of at least quadriserial rows, and a tendency for wider, multiserial branches in the distal parts of the colony. Soule (1959) gives dimensions for two size classes of avicularia for B. californica; it apparently does not possess the tiny avicularium associated with the bifurcations of B. stolonifera. I am unaware of any description of the ancestrula of Pacific coast material. At least part of the material described by Marcus (1937) as B. californica appears to be the present species. I can find no appreciable difference between his figures (Plate XV, Figs. 37A and 37B) and my material. The records of Bugula avicularia (Linnaeus) from along our coast are presumed to be B. stolonifera. The possible exceptions are Verrill's records from north of Cape Cod. But specimens identified by Verrill as B. avicularia from "Ram Island Ledge" (presumably, in Great Harbor, Woods Hole) (USNM 4981) are B. fulva Ryland. Osburn (1912) did not find B. avicularia at Woods Hole but he listed it because of Verrill's report; his figure and descriptions were based on an English specimen. Maturo (1957) did not collect it at Beaufort, but included it because of McDougall's report (identified by Osburn). Through a lapsus Maturo failed to state that his description and figures were based upon a specimen given him by Osburn which was found to be British material. Specimens collected by McDougall were found in the Hutchins Collection (USNM 11886). These are labeled Bugula avicularia but are B. stolonijera. Bugula stolonijera only superficially resembles B. avicularia, i.e., in having a prominent 2: 1 spine pattern and avicularia mounted in approximately the same positions. However, the sizes and appearance of the short-beaked avicularia of B. stolonifera immediately distinguish it. It is my opinion that Osburn referred all biserial "white" bugulas from the Atlantic coast 570 Bulletin of Marine Science [16(3) which had prominent spines in a 2: 1 pattern to B. avicularia if he could not assign them to B. turrita. The record of B. avicularia from the Gulf of Mexico by Canu & Bassler (1928) is erroneous. The specimens (USNM 7457:ALBATROSSSta. D 2372) are actually the B. dentata of Osburn (1940). This seems to agree with Harmer's description (1926) of Lamouroux's species, but Osburn's, and Canu & Bassler's specimens have type 4 bifurcations rather than the type 5 of Pacific material. Distribution.-Bugula stolonifera has been collected on the Atlantic coast from Massachusetts (Hutchins, USNM 11885; Osburn, USNM 11891), Bermuda (Hutchins, USNM 11887), North Carolina, and Brazil.

Bugula marcusi, sp. n. Fig. 11 Material.-One small colony, % to 1 mile offshore, east of Bogue Pier, Bogue Bank, Beaufort, North Carolina, about 40 feet (ca. 12 m), F. J. Maturo, Jr., colI., June 13, 1963: HOLOTYPE,USNM 11897. Description.- The single colony collected is small (0.6 cm), however, it shows a spiral growth pattern developing, with type 3 bifurcations. Zooids are biserial, alternate, narrowed slightly toward the base; the frontal membranes occupying about two-thirds of the frontal surface. The orificial area of the frontal membrane turns rather abruptly from thc frontal plane to face distally. The distal border bears three spines: a short, denticle-like frontal outer distal spine; a prominent, conical, dorsal, outer distal spine; and a short, tubular, inner distal spine. The avicularia are attached halfway down or lower on the outer frontal margin; they have a relatively elongate, decurved beak and a slender, distinctly "sway-back" profile resulting from a prominent concavity along the dorsal margin where the head and beak join. The ancestrula and ancestruloid zooids have an inverted cone shape with the frontal membrane circular and facing distally, its border bearing at least four spines. These spines may be prominent or reduct:d to short denticle-like projections. There are no ovicells present on the specimen. The zooids, exclusive of those involved in a bifurcation, are 0.41-0.45 mm long, 0.14-0.16 mm wide, averages for ten zooids being 0.44 mm and 0.16 mm respectively. The avicularia exhibit a gradual increase in size from the proximal to the distal end of the colony. The smallest ones at the basal portions are 0.13 mm long while those at the distal ends are 0.17-0.18 mm long. Remarks.-Bugula marcusi shares many features with B. turrita (Desor) but differs from it in possessing larger avicularia which have a different 1966] Maturo: Bryozoa of Southeast United States 571 shape and which are generally placed lower along the frontal margin, and an ancestrula with distal spines. The avicularia strongly resemble those of B. graciLis Busk as figured by Ryland (1963: Fig. 1D) but B. marcusi differs from that species in having shorter and stouter zooids, a short frontal outer distal spine, an ancestrula with spines, and avicularia of graded sizes. It may be distinguished from B. grayi by its smaller dimensions, the shape and size of avicularia, and differences in the ancestruloid zooids. The material described by Marcus (1937, 1938) as B. turrita from Bahia de Santos, Brazil, differs from Desor's species by having an ancestrula and ancestruloid zooids with spines and avicularia of larger size and different shape. After a comparison of my specimen with the description and figures given by Marcus, I can find no important differences between them (except for a better developed frontal outer distal spine in his figures). It is my opinion that Marcus's B. turrita is B. marcusi. This species is named in honor of Prof. Ernst Marcus who has made many outstanding contributions to the study of the Ectoprocta.

Bugula grayi, sp. n. Fig. 13 Material.-One slide and one vial (alcohol) from off Sapelo Island, Georgia, 17% miles (bearing 74°) from Doboy Sound sea buoy, 65 feet (19.7 m), station 189, M. Gray, coIL, March 5, 1963: HOLOTYPE, USNM 11898. Other material from off Sapelo Island, Georgia, 5 miles from Sapelo whistle buoy (buoy bearing 340°), 73 feet (22.2 m), drag 135, M. Gray, coIL, Aprilll, 1962: PARATYPES, author's collection nos. 1847, 1849. Off Sapelo whistle buoy, 4% miles (buoy bearing 345°), 67 feet (20.1 m), drag 107, M. Gray, coiL, February 27, 1962: PARATYPE, author's collection no. 1858. Description.-Colony consists of erect branches with secondary branches arranged in a spiral or whorled pattern; height of type specimen is 6.5 cm; bifurcations of type 3. The color of the secondary branches in 70 per cent alcohol is pale cream, the main stems, which are much thickened by entwined radicles, are a dark cream. Zooids are biserial, alternate, tapering slightly toward the base, with frontal membranes occupying slightly more than three-fourths of the length of the frontal surface. The distal spine formula is typically 2: 1, with the spine at the frontal outer distal corner quite small and denticle-like (sometimes missing in the proximal parts of branches), the dorsal outer distal spine bluntly elongate and conical, and the inner distal spine narrow and tending to curve slightly across the 572 Bulletin of Marine Science [16(3)

13 O.5mm

FIGURES 12-14. All figures drawn to scale indicated. 12, Bugula stolonifera Ryland, showing ovicells and three sizes of avicularia. USNM 11899; 13, Bugula grayi new species, part of holotype, USNM 11898; 14, Cau/ibugula dendrograpta (Waters), showing part of stalk kenozooid and proximal portion of a branch, USNM 11903. 1966] Maturo: Bryozoa of Southeast United States 573 frontal membrane. The large outer spine is usually very broad-based, giving a shouldered appearance to the disto-lateral margin of the zooid, but it may be very reduced in the most proximal zooids of a branch. Avicularia are attached to the outer margin about two-thirds of the way down the frontal membrane border from the base of the frontal distal spine; this position is somewhat variable, ranging from just below halfway to three-fourths of the way down. The length of the avicularia is equal to or slightly exceeds the width of the zooids; the beak is decurved. Mature ovicells were not observed, but the developing ovicells which appear in March and spent ones found in April are shallow cap-like hoods placed slightly to the median side of the orificial rim and tilted toward the axis of the branch. The ancestrula has not been found but ancestruloid zooids of secondary colonies are inverted cones with the frontal membrane in two planes, i.e., having an orificial area perpendicular to the zooid axis and a short frontal portion which is less than one-fourth of the length of the zooid. There are three minute denticle-like spines on the orificial rim which correspond in position with the spines of more distal zooids. An avicularium is placed to one side of the base of the frontal membrane (on the same side as the pair of distal spines). Zooids, exclusive of those involved in bifurcations, are 0.46-0.63 mm long (0.53 mm average) and 0.19-0.23 mm wide (0.21 mm average). There are about 14 tentacles. Avicularia are 0.19-0.27 mm long (0.22 mm average). Remarks.-Bugula grayi is most likely to be confused with B. turrita (Desor) which it resembles in its spiral growth pattern, spine formula, bifurcation type, and ovicell form. It differs from B. turrita in having stouter zooids, proportionately longer frontal membranes, much larger avicularia (nearly twice as large) with a different shape and a more proximal position, and a different form to the ancestruloid zooids of secondary colonies. It differs from B. stolonifera Ryland in having only type 3 bifurcations, generally larger avicularia with different shape and position, conical outer distal spines rather than tubular ones, and different ancestruloid zooids. Bugula marcusi Maturo bears considerable resemblance to B. grayi but the former is a smaller species with distinctly swaybacked avicularia. The species is named in honor of Dr. 1. E. Gray of Duke University. Distribution.-Bugula grayi was obtained on shell debris from an area approximately 25 miles east of Sapelo Island, Georgia (31°24'N., 80049'W.) at depths of 65-73 feet (19.7-22.2 m). 574 Bulletin of Marine Science [16(3) Genus Caulibugula Verrill, 1900 Genotype.-Bugula (Caulibugula) armata Verrill, 1900:593. Description.-Colonies possess jointed stalks composed of elongate, cylindrical kenozooids; branches are usually biserial and fan-like, the proximal zooids of the fan more or less different from its successors, frequently being ancestruloid. Zooids sometimes divided into three sections by two joints or creases, turbinate, with a short frontal membrane and long spines; or Bugula-like, with an elongate frontal membrane and with vestigial spines or none. Avicularia pedunculate, with an acute beak and mandible, variously situated, or wanting. Ovicells hyperstomial but often reduced in size, or wanting. (After Harmer, 1926.)

KEY TO THE SOUTHEASTERN SPECIES OF Caulibugula Bifurcations of type 4; proximal zooids of branch with 6-8 long, jointed spines surrounding a frontal membrane which has a slit-like proximal prolongation . dendrograpta Bifurcations of type 5; proximal zooid of branch generally without distinct frontal membrane or spines (rarely, 3 spines present), with proximal annulations pearsei (See discussion of this species to distinguish C. armata Verrill)

Caulibugula dendrograpta (Waters) Fig. 14 Not Bugula (Caulibugula) armata Verrill, 1900:593. Stirparia dendrograpta Waters, 1913: 470. Bugula armata: Osburn, 1914: 188. Caulibugula armata: Marcus, 1938:29. Caulibugula armata: Osburn, 1940:393. Description.-Colonies consist of very slender, jointed stalks (kenozooids) with delicate flabellate branches of zooids arising just below the joints; color whitish in alcohol specimens; up to several centimeters in height; bifurcations of type 4. The stalk segments are 0.07-0.09 mm wide and up to 2 mm long in specimens examined, are slightly enlarged at the ends (nodes), and possess a pair of lateral calcareous thickenings; the proximal ends may give rise to radic1es, whereas the distal ends produce stem segments and the branch zooids. Branch zooids are biserial (following the first one which is ancestrula-like), alternate, rather strongly curved out- wards, tapering strongly toward the base, apparently divided transversely into three parts by two dorsal joints (?): one just below the level of the proximal end of the frontal membrane and the other just above the proximal fork. The frontal membrane is elongated, generally exceeding half the zooid length; operculum appears to be without a bordering sc1erite. 1966] Maturo: Bryozoa of Southeast United States 575 The frontal membrane of the primary zooid of each branch is surrounded by six to eight long, curved, jointed spines and it usually possesses a proximal slit-like prolongation. Ordinary zooids bear on their dorsal side near the distal border one to four slender, elongated spines, jointed at the base, and exceeding, in some cases, twice the length of the zooids. The full number of spines is developed only on the external zooids of a fan (whole branch), the internal ones generally possessing only one. Avicularia are small, situated on all but the primary zooids; the position variable, being attached near the proximal end in the lower zooids of a fan, while progressively more distal on the next few zooids, then on the disto-lateral corner of all succeeding zooids. Ovicells were not found on the material collected in this study, but those on specimens from Tortugas (Osburn Collection, USNM 11893) are globular, attached by a short stalk to the inner corner of the distal border of the zooids, with the inconspicuous aperture appearing to face laterally, although it is difficult to tell with certainty. Zooids are 0.40-0.50 mm long and 0.14 mm wide. Avicularia average 0.07 mm in length. Remarks.-Osburn (1914), believing he had found Verrill's C. armata at Tortugas, figured and redescribed it, calling attention to the distinctive distribution of the avicularia. In 1940, he placed Stirparia dendrograpta Waters, 1913, in the synonymy of C. armata, a course accepted by Marcus (1941). I also agree with Osburn that his Tortugas species and Water's Zanzibar species are identical. However, Osburn misinterpreted Verrill's poor description which lacked an illustration. I have examined Verrill's type specimens of Bugula (Caulibugula) armata (YPM 2891, Bermudas, 1898) and find them to be identical with what Levinsen (1909: 104-107; PI. 3, Figs. 2a-2n) described and figured as a new species, Bugula caraibica. Therefore, Bugula (= Caulibugula) caraibica Levinsen becomes a junior synonym of Caulibugula armata Verrill, with Levinsen's figures and description serving to characterize C. armata. Caulibugula dendrograpta (Waters), being the oldest available name, now becomes the valid name for the C. armata of authors. Distribution.-The species is widely distributed in warmer waters, occurring on the Atlantic Coast as far north as Cape Lookout, North Carolina.

Caulibugula pearsei, sp. n. Figs. 15 & 16 Caulibugula sp. Osburn, 1951: 137. Material.-Off Sapelo Island, Georgia, 5-6 miles from Sapelo whistle buoy (back bearing 340°), 72 feet (21.8 m), drag 1, F. J. Maturo, Jr., colI., June 6, 1962: HOLOTYPE,USNM 11904. 576 Bulletin of Marine Science [16(3)

/ I I 18

O.5mm 1966J Maturo: Bryozoa of Southeast United States 577 Same data as Holotype: PARATYPE,author's collection no. 1724. Three slides, Black Rocks off New River, North Carolina, A. S. Pearse, coIl., R. C. Osburn Collection, June 29, 1949: PARATYPE,USNM 11894. Description.-Colonies consist of conspicuously jointed stalks of elongated, uniserial kenozooids from which ftabellate branches of zooids arise just below the joints; height of fragments at hand up to 1.6 em; bifurcations of type 5. Stalk kenozooids are cylindrical to quadrangular, 1.52-3.61 mm long, 0.19-0.30 mm wide, and possess one to three proximal annulations. The quadrangular appearance of most kenozooids is the result of four longitudinal thickenings. The primary zooid of the fan-shaped branches is an abortive zooid without internal organs or spines, having only a faint suggestion of a frontal membrane, and possessing one to four proximal annulations. Branch zooids are biserial, alternate, tapering gently toward the base, occasionally divided transversely into two parts by a constriction just above the forked proximal end, with a buguloid frontal membrane occupying about three-fourths of the length of the frontal surface; zooids turned strongly toward the median axis of the branch so that essentially a right angle (or even an acute angle) is formed between the frontal membranes of two neighboring zooids. The orificial area of the frontal membrane is abruptly turned distally, its margin bearing one to four very long, curved, jointed spines. The number of spines per zooid is related to position within the fan: outer zooids of fan margin possess four spines; lateral zooids of inner bifurcations bear two to three spines; inner zooids of bifurcations bear one to two spines. Occasionally, "E" and "P" zooids of bifurcations are spineless. Avicularia are small and placed beside the proximal end of the frontal membrane; the beak is abruptly hooked. Ovicells have not been observed. Zooids are 0.36-0.45 mm long and 0.14-0.16 mm wide. Avicularia are about 0.12 mm long. Zooid fans may arise opposite one another on the same stalk segment. In one instance a fan originates terminally on a stalk; in this case the primary zooid differs from the usual condition by having three spines on one side of an indistinct frontal membrane.

~ FIGURES 15-18. All figures drawn to scale indicated. 15, Caulibugula pearsei new species, showing stem kenozooids and branch primary zooids, paratype, author's collection no. 1724; 16, Caulibugula pearsei new species, showing spine pattern and basal zooid, holotype, USNM 11904 (ovicell-like structure on righthand side is egg shell or case of some invertebrate); 17, Beania hirtissima (Heller), USNM 11905; 18, Beania intermedia (Hincks), showing avicularia and mode of branching. 578 Bulletin of Marine Science [16(3) Remarks.-The stalk calcification is somewhat variable so that some segments are like those described by Levinsen (1909) for Caulibugula caraibica (Levinsen) [= C. armata Verrill, see discussion of preceding species]. As far as I have been able to determine, only three other species of Caulibugula have type 5 bifurcations: C. exilis (MacGillivray), C. tuberosa Hastings, and C. armata Verrill [C. caraibica (Levinsen)). Among other features, C. exi/is and C. tuberosa differ from C. pearsei by possessing primary zooids with numerous jointed spines surrounding a distinct frontal membrane and by possessing stalk kenozooids lacking longitudinal calcareous thickenings. Caulibugula exilis has longer avicularia with abruptly hooked beaks, while C. tuberosa has no avicularia and possesses short circular frontal membranes. Caulibugula pearsei seems most closely related to C. armata Verrill (not Osburn) but differs from it in having a greater number of jointed spines on the zooids, a different spine pattern, shorter zooids, and avicularia present on most zooids. The primary zooids and stalk kenozooids appear to be very much alike. However, the proximal zooids of C. pearsei are much shorter, flared distally, and bear four strong spines. The species is named in honor of the late Dr. A. S. Pearse, founder of the Duke University Marine Laboratory, and first collector of the species. Distribution.-The range of the species is North Carolina to Florida.

FAMILY BEANIIDAE CANU & BASSLER, 1927 Genus Beania Johnston, 1840 Genotype.-Beania mirabilis Johnston, 1840. Description.-Zooids with a narrow, tubular, proximal end, expanded distally, generally united with six or four other zooids by connecting tubes separated by open spaces; sometimes uniserial and branching; their proximal ends not equitant on the distal portions of their predecessors. Frontal membranes occupying the entire front of the expanded portion, more or less surrounded by spines, or without these structures. Pedun- culate avicularia usually present, in some species wanting. Ovicells present, vestigial, or wanting. (After Harmer, 1926.)

KEY TO THE SOUTHEASTERN SPECIES OF Beania Zooids, multiserial; frontal membrane surrounded by numerous spines, avicularia absent hirtissima Zooids uniserial, branching; frontal membrane not surrounded by spines, avicularia usually present intermedia 1966] Maturo: Bryozoa of Southeast United States 579 Beania hirtissima (Heller) Fig. 17 Diachoris hirtissima Heller, 1867: 94. Beania hirtissima: Marcus, 1937:62. Beania hirtissima: Osburn, 1940:397; 1947:24. Beania hirtissima: Lagaaij, 1963:180. Description.-Colonies are loosely attached or partially growing free, composed of unilaminar, spinous mats of rnultiserial zooids which are separated from one another but which are joined by tubular connecting processes. The zooids are roughly oval in outline, with an indentation or constriction on either side just below the operculum suggesting a neck. Each zooid is located with the distal end elevated above the level of the next succeeding zooid. The frontal surface is occupied entirely by the frontal membrane which terminates distally in a semicircular operculum. The entire margin of the frontal surface is beset with long, slender spines: 8 to 10 erect spines surround the orificial area, 8 to 14 spines arise on each side from the border of the frontal surface and curve gently over the frontal membrane, and about 10 spines, which project outward, arise on the distal half just below (dorsal to) the level of the others. Long spines and an occasional tubular radicle also arise from the dorsal surface. Each zooid is connected by tubular processes to the six zooids which surround it. A septum is present in each tube. Zooids average 0.63 mm in length (proximal septum to distal septum) and 0.29 mm in width. There are no avicularia or ovicells. Remarks and Distribution.-The species was collected in depths of 17 fathoms (31.1 m) offshore from Cape Lookout, North Carolina, and in depths to 30 fathoms (54.9 m) off both Sapelo Island, Georgia, and Fernandina Beach, Florida. These are new range extensions for the species; the Cape Lookout record represents the northernmost report of Beania hirtissima along the Atlantic Coast. Although widely distributed in warmer waters, it has previously been reported in the western North Atlantic from Bermuda and Puerto Rico (Osburn). Lagaaij (1963) has recently found it in the Gulf of Mexico. Beania intermedia (Hincks) Fig. 18 Diachoris intermedia Hincks, 1881: 133. Beania intermedia: Osburn, 1914:189; 1940:398; 1947:25. Beania intermedia: Marcus, 1937:61. Beania intermedia: Wells et al., 1964:568. Beania intermedia: Shier, 1964:624. Description.-Colonies consist of semi-erect or loosely incrusting, uniserial zooids attached to the substrate by radic1es. The zooids are thinly 580 Bulletin of Marine Science [16(3) calcified, broadly club-shaped or boat-shaped, the proximal end beginning as a narrow tube which quickly enlarges and becomes flattened frontally by the extensive frontal membrane occupying the remaining frontal surface, the distal end rounded and bearing two minute denticle-like distal projections. Two additional denticles may be situated at the level of the attachment of the operculum but are commonly replaced by (bear ?) a pair of pedunculate avicularia. Spines are absent from the remaining margin of the frontal membrane. Daughter zooids may arise as buds in three places on the dorsal surface of a zooid: from a very short, tubular elevation in the midline at the level of the base of the operculum, or from two lateral bud areas near the proximal end of the frontal membrane. Radicles arise from a bud zone in the middorsal line between the lateral buds. Ovicells are absent. Zooids are 0.95-1.14 mm long and 0.29-0.34 mm wide. The avicularia are 0.11-0.14 mm long. Distribution.-The species is widely distributed in warmer waters. The specimens for the present study were collected by Dr. Harry Wells from scallop shells taken east of Cape Lookout, North Carolina, and represent the northernmost records for this species along the Atlantic Coast.

SUMARIO BRYOZOADE LA COSTASURESTEDE LOSESTADOSUNlDOS:BUGULIDAEY BEANIIDAE(Cheilostomata: Anasca)

Se describen e ilustran doce especies pertenecientes a las familias Bugulidae y Beaniidae que se presentan frente a la costa desde North Carolina hasta la Florida; se proveen claves para facilitar su identificaci6n. Ademas de Bugula neritina, B. turrita, B. fulva, B. stolonifera, B. microoecia, Caulibugula dendrograpta, Beania hirtissima y B. intermedia, se describen cuatro nuevas especies. Estas son: Bugula rylandi, B. marcusi, B. grayi, y Caulibugula pearsei. La distribuci6n de Bugula microoecia se extiende desde el Golfo de Mexico hasta North Carolina; el reporte de su presencia en Puerto Rico hecho por Osburn es dudoso. Bugula turrita se extiende por 10 menos hasta la Florida, pero reportes mucho mas al sur deben ser re-examinados. Bugula fulva y B. stolonifera son reconocidas por primera vez en aguas americanas, habiendo sido confundidas ambas con otras especies. La presencia de Bugula avicularia y B. californica en el Atlantico Occi- dental es dudosa. Los reportes american os de los cuales habia ejemplares (disponibles) han sido re-examinados y se encontr6 que eran err6neos. Ellos son generalmente B. stolonifera. Caulibugula armata Verrill, 1900, no es tal como 10 describe Osburn y 1966] Maturo: Bryozoa of Southeast United States 581 otros autores sino que es equivalente a Caulibugula caraibica (Levinsen), 1909, su sin6nimo joven. La Caulibugula armata de otros autores resulta ser C. dendrograpta (Waters), 1913.

LITERATURE CITED BUSK, G. 1858. Zoophytology: on some Madeiran Polyzoa. Quart. J. microscop. Sci., 6: 261-263. CANU, F. AND R. S. BASSLER 1927. Classification of the cheilostomatous Bryozoa. Proc. U. S. nat. Mus., 69 (14): 1-42. 1928. Fossil and Recent Bryozoa of the Gulf of Mexico region. Proc. U. S. nat. Mus., 72 (14): 1-199. DESOR, E. 1848. Ascidioidian polyps or Bryozoa (from Nantucket). Proc. Boston Soc. nat. Hist., 3: 66-67. GRAY, J. E. 1848. List of the specimens of British animals in the collection of the British Museum. Part 1, Centroniae or radiated animals. (Bryozoa, p. 91-151.) London, 173 p. HARMER, S. F. 1923. On cellularinc and other Polyzoa. J. Linn. Soc., Zoo\., 35: 293-361. 1926. The Polyzoa of the Siboga expedition. Part 2, Cheilostomata, Anasca. Siboga Exped., 28b: 183-501. HELLER, C. 1867. Die Bryozoen des adriatischen meeres. Verhandl. der K.-K. Zoo\.- Bot. GeseUschaft in Wien, 17: 77-136. HINCKS, T. 1881. Contributions toward a general history of the marine Polyzoa. Ann. Mag. nat. Hist., Ser. 5, 8: 122-136. HYMAN, LIBBIE H. 1959. The invertebrates: smaller coelomate groups. McGraw-Hill Book Co., Inc., New York. 783 p. HUTCHINS, L. W. 1945. An annotated check-list of the salt water Bryozoa of Long Island Sound. Trans. Conn. Acad. Arts ScL, 36: 533-551. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE 1956. Opinion 417. Rejection for nomencIatorial purposes of volume 3 (Zoologie) of the work by Lorenz Oken entitled Okens Lehrbuch der Naturgeschichte published in 1815-1816. Opin. DecIar. Inter- national Comm. Zoo\. Nomenc\., 14(1): 1-42. JOHNSTON, G. 1840. Description of a new genus of British zoophyte. Ann. Mag. nat. Hist., Ser. I, 5: 272-274. LAGAAIJ, R. 1963. New additions to the bryozoan fauna of the Gulf of Mexico. Pub!. Inst. Mar. Sci. Univ. Tex., 9: 162-236. LEIDY, J. 1855. Contributions toward a knowledge of the marine invertebrate fauna of Rhode Island and New Jersey. J. Acad. nat. Sci. Philad., ser. 2, 3: 135-152. 582 Bulletin of Marine Science [16(3)

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