Norntatesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

NorntatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2847, pp. 1-49, figs. 1-106, 1 table June 1 1, 1986

Bryozoans from Bali, Lombok, and Komodo

JUDITH A. WINSTON1 AND BEVERLY F. HEIMBERG2

CONTENTS

Abstract ...... 2 Introduction ...... 2 Study Areas ...... 2 Methods and Materials ...... 3 Systematic Accounts ...... 6 Cranosina coronata ...... 6 Parellisina curvirostris ...... 6 Caulibugula mortenseni ...... 7 Scrupocellaria spatulata ...... 7 Thalamoporella stapifera ...... 8 Thalamoporella komodoensis, New Species ...... 10 Steginoporella buskii ...... 11 Smittipora harmeriana ...... 11 Cribrilaria flabellifera ...... 14 Exechonella brasiliensis ...... 15 Coleopora verrucosa ...... 15 Hippopodina feegeensis ...... 16 Petraliella chuakensis ...... 16 " Watersipora subovoidea"...... 17 Watersipora edmondsoni ...... 17 Calyptotheca wasinensis ...... 18 Schizomavella inclusa ...... 18 Stylopoma duboisii ...... 20 Stylopoma viride ...... 21 Parasmittina signata ...... 21

1 Associate Curator, Department of Invertebrates, American Museum of Natural History. 2 Curatorial Assistant, Department of Invertebrates, American Museum of Natural History.

Parasmittina parsevalii ...... 24 Parasmittina hastingsae ...... 26 Crepidacantha carsioseta, New Species ...... 27 Fenestrulina harmeri, New Species ...... 28 Margaretta triplex ...... 30 Celleporaria sibogae, New Species ...... 30 Celleporaria aperta ...... 33 Cleidochasma porcellanum ...... 34 Cleidochasma peristomarium ...... 36 Rhynchozoon verruculatum ...... 37 Rhynchozoon rostratum ...... 38 Rhynchozoon detectum ...... 39 Drepanophora verrucosa, New Species ...... 40 Discussion ...... 41 Acknowledgments ...... 43 Literature Cited ...... 43 Glossary ...... 47

ABSTRACT Collections from three very shallow-water (3 m are reported for the first time from the Indonesian or less) Indonesian reeflocalities yielded 33 species region. Many of the species found have wide dis- of cheilostome bryozoans. Five species: Thala- tributions: 79 percent are recorded from elsewhere moporella komodoensis, Crepidacantha carsiose- in the Indo-West Pacific, 30 percent from the Red ta, Fenestrulina harmeri, Celleporaria sibogae, and Sea, and 28 percent from the Caribbean. This work Drepanophora verrucosa are described as new. Five provides further documentation of the diversity species: Exechonella brasiliensis, Watersipora ed- and broad ecological tolerances of reef-associated mondsoni, Parasmittina hastingsae, Rhyncho- cheilostome bryozoans. zoon rostratum, and Rhynchozoon verruculatum

INTRODUCTION From 1899 to 1900, the Dutch, under the study of the fine structure of bryozoan skel- direction of Dr. Max Weber, carried out the etons is becoming increasingly important to great collections ofthe Siboga expedition. The bryozoan taxonomy, and we expect the pres- bryozoans ofthis expedition were studied by ent paper to contribute to this cataloging pro- Sidney F. Harmer in a series of four mono- cess. graphs over the course ofhis career. The first part, on the Entoprocta, Ctenostomata, and STUDY AREAS Cyclostomata, was published in 1915. The last part, on the Ascophora, was compiled Figure 1 shows the location of the three from his uncompleted drafts and notes by areas where bryozoans were collected. Lo- Anna B. Hastings and published posthu- cality 1 was at Sanur on the southeast coast mously in 1957. In view ofthis opus on bryo- of the island of Bali, facing Badung Strait. zoans of the Indonesian region one might Bryozoans were collected from coral rubble question the need for further studies, but, in in 2-3 m depths. A strong current scoured fact, very little of the Siboga material was the reef in the area collected, the corals had taken from reefenvironments. Thus, the col- flattened or encrusting shapes, and the pieces lections described below, though small, do ofdead coral that remained in scour holes in add to our knowledge ofthe systematics and the reef rock were large and heavy. It ap- distribution of reef-associated bryozoans. In peared to be a relatively high-energy envi- addition, many of these species have never ronment. been studied and illustrated by SEM. The Locality 2 was 90 km east of Sanur on the 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 3

,el FIG. 1. Map ofportion of the Indonesian archipelago showing locations from which bryozoans were collected. (1) Sanur, Bali. (2) Lambuan, Lombok. (3) Komodo. island of Lombok in the Lesser Sunda Is- laboratory, the bryozoan specimens were lands. The reef was located just outside the identified and measured with a dissecting mi- harbor at Lembar, on the southwest coast of croscope. Measurements included standard Lombok. The bryozoans were collected in a features: zooid length (Lz), zooid width (Wz), back-reefarea, on coral rubble from the com- orifice length (Lo), orifice width (Wo), ovicell pacted platform of rubble that formed the length (Lov), ovicell width (Wov), avicular- back part of the reef flat (depths 0.5-1.5 m). ium length (Lav), avicularium width (Wav), This mass of rubble bordered a back-reef la- as well as other measurements necessary for goon, with sand-bottom and scattered sea- a particular species. Figure 2 shows how each grass beds. It constituted a sheltered back- of these measurements was taken. reef environment. At least one colony of each species was The third locality was approximately 380 examined under the scanning electron mi- km east of Lembar on the east coast of the croscope. Specimens were prepared for the island of Komodo, in the bay adjacent to the SEM by soaking in bleach to remove chitin- Komodo Dragon refuge. Here the reefbegan ous parts and tissue. The specimens were directly off the beach with leather corals, rinsed in water, cleaned ultrasonically, and nephteids, and other soft corals in the shal- dried before they were sputter-coated with lowest water, and scleractinians appearing in gold or platinum. slightly deeper water. Bryozoans were col- It seemed immediately apparent that the lected from coral rubble in depths of 1.5-2.0 scanning electron microscope offered a better m. This locality, though open to the water of way to illustrate bryozoan specimens. Bryo- the bay, was somewhat sheltered by the bay zoan skeletal (and more rarely chitinous) parts itself. had traditionally been illustrated by pen-and- METHODS AND MATERIALS ink drawings or, in the 20th century, by photographs. An argument can be made that well- Encrusted coral rubble was rinsed in fresh executed drawings (such as those of Peter water, trimmed to remove excess substra- Hayward) are still the best way to illustrate tum, and air-dried in the field. At the AMNH species, for they can survive photocopying, 4 AMERICAN MUSEUM NOVITATES NO. 2847

FIG. 2. Diagrams ofbryozoan zooids to show way in which each measurement was taken. Lz = zooid length, Wz = zooid width, Lo = orifice length, Wo = orifice width, Lov = ovicell length, Wov = ovicell width, Lav = avicularium length, Wav = avicularium width, Lop = opesia length, Wop = opesia width. thus extending the useful life of the mono- at skeletal structure (and the structure ofchi- graph in which they are included past the tinous parts as well) at a new level of reso- expiration ofreprints, editions, and even au- lution. In this regard we are at the point thors! However, those taxonomists not tal- reached some years ago by entomologists and ented enough to do their own illustration and protozoologists, for example. With the SEM not wealthy enough to pay an illustrator, but we are able to see the characters traditionally who wanted to objectively portray the colo- considered most important in bryozoan sys- nies they studied, turned as soon as possible tematics-orifice, frontal wall structure, pore to photography. Unfortunately, the small size patterns, avicularia-in detail never before ofbryozoan zooids means that pictures taken possible. Multiple views are necessary to il- at the high magnifications (50-60 x) neces- lustrate different polymorphs and ontogenet- sary for identification come close to the limits ic and astogenetic stages which can also fur- of resolution of the dissecting microscope. nish taxonomic characters. In addition, we Even the best photomicrographs are not en- are finding new characters, both internal tirely in focus due to the limited depth of (muscle scars, pores, spines, hooks) and ex- field at those magnifications, and many ternal (fine structure of frontal wall texture, monographs have been made less useful by pores, joints between zooids, orificial con- the poor quality of their illustration. This is dyles and sinuses, avicularian mandibles, and especially true of the early photographs like opercula) which we believe will be important those in papers by Canu and Bassler (e.g., in revealing the still tangled phylogenetic re- 1924, 1928a, 1928b, 1929), wherethe details lationships in the group. oforifices and avicularia were blacked in with In our species descriptions we have at- ink (and where the structure drawn in was tempted to include as much fine structural sometimes what the authors wished to see detail as we could obtain through SEM study. rather than what was on the specimen). While We have also attempted to illustrate as many SEM also has limits to depth of focus, with of these characters as possible. This means SEM "portraits" ofgroups ofzooids taken at that we have included several illustrations 60-100 x all details of zooids are in focus, per species, but we believe that this is nec- even when the colony surface is (as is gen- essary ifwe are to take advantage ofall char- erally the case) not perfectly even. acters available to us. Once we have achieved More and more, bryozoologists have come a better understanding ofbryozoan structure to realize the importance ofSEM as a tool in at this level, it may once again be sufficient the study of specimens. It permits us to look to illustrate a species less extensively, but it 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 5

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FIGS. 3-8. 3. Cranosina coronata, Bali (AMNH 583), growing edge ofcolony showing marginal pore chambers and transverse avicularia. Scale = 400 ,um. 4. C. coronata, zooids and avicularia. Scale = 200 ,um. 5. C. coronata, avicularian mandible. Scale = 40 ,m. 6. C. coronata, scalloped edge of avicularian mandible. Scale = 10 ,um. 7. Parellisina curvirostris, Komodo (AMNH 584), zooids and ovicells. Scale = 200 Mum. 8. P. curvirostris, close-up of zooids to show marginal spine bases. Scale = 200 ,m. will take much more study before that time We have included a glossary of bryozoan comes. terminology used in this paper. We hope this 6 AMERICAN MUSEUM NOVITATES NO. 2847 will make our use of terms clear to bryozoan OCCURRENCE: One colony was collected at workers and that it will make our descriptions Sanur. more accessible to nonspecialists interested DISTRIBUTION: Torres Strait, Indonesia, in identifying reef faunas. W. Australia, Ceylon, Indian Ocean, Mac- clesfield Bank, China Sea, Venezuela.

SYSTEMATIC ACCOUNTS Genus Parellisina Osburn, 1940 ORDER CHEILOSTOMATA BUSK, 1852 Parellisina curvirostris (Hincks), 1862 SUBORDER ANASCA LEVINSEN, 1909 Figures 7, 8 SUPERFAMILY MEMBRANIPOROIDEA Membranipora curvirostris Hincks, 1862, p. 29. BUSK, 1854 Ellisina curvirostris Harmer, 1926, p. 228. Has- tings, 1930, p. 711. FAMILY CALLOPORIDAE NORMAN, 1903 Callopora curvirostris Canu and Bassler, 1928b, p. Genus Cranosina Canu and Bassler, 1933 32. Parellisina curvirostris Osburn, 1940, p. 361. Cranosina coronata (Hincks), 1881 Cheetham and Sandberg, 1964, p. 1020. Cook, Figures 3-6 1968, p. 156. Ryland and Hayward, 1977, p. 102. Winston, 1984, p. 7. Membranipora coronata Hincks, 1881, p. 147; 1882, p. 118. Thornely, 1907, p. 186. DESCRIPTION: The colony is encrusting on Membranipora hastilis Kirkpatrick, 1890a, p. 18. coral. Zooids vary from ovoid to subtrian- Thornely, 1905, p. 110. gular in shape and have a beaded cryptocyst Biflustra coronata MacGillivray, 1891, p. 79. and a smooth gymnocyst which is widest Setosellina coronata Harmer, 1926, p. 265. Cranosina coronata Osburn, 1940, p. 363. proximally and very narrow laterally, with the opesia taking up most of the surface be- DESCRIPTION: The colony is encrusting on neath the frontal membrane. Occasional spine coral rubble. Zooids are irregularly ovoid to bases can be seen on the inner edges of the subhexagonal; a narrow densely beaded cryp- gymnocyst of zooids. A groove marks the tocyst surrounds the oval to triangular ope- mural rims ofadjacent zooids. The ovicell in sia, with beads tending to be aligned radially. this species is globular with a beaded texture Deep grooves separate zooids. At the distal like that ofthe cryptocyst. The ovicells ofthe end of-each zooid, a smooth semicircular specimen found (fig. 5) were broken. No gymnocystal hood surrounds the operculum. avicularia were present in this colony. No ovicells are present in our specimens. Measurements Harmer (1926) suggests that they are probably lacking. Located at the distal edge of Range Mean N LZ 0.360-0.468 0.425 5 most zooids are transversely directed avic- wz 0.324-0.378 0.349 5 ularia. These have a spoutlike rostrum with Wo 0.090-0.126 0.108 5 two large pointed condyles to support the se- Lov 0.144 0.144 2 Wov 0.180-0.198 0.189 2 tiform mandible. The mandible (fig. 3) is long, Lop 0.216-0.324 0.292 5 slightly curved (some ofthose illustrated have Wop 0.216-0.270 0.230 5 been distorted by the drying process), and DISCUSSION: The condition of the colony edged with finely scalloped serrations (fig. 4). and the lack ofavicularia make positive iden- Measurements tification difficult, but zooid morphology, es- Range Mean N pecially the occurrence of distal and lateral Lz 0.504-0.558 0.536 5 spine bases, indicates that it belongs to Par- Wz 0.324-0.414 0.364 5 ellisina curvirostris. Zooid measurements are Lo 0.072-0.090 0.083 5 Wo 0.144-0.180 0.171 5 in the range for the species. Lop 0.342-0.432 0.382 5 OCCURRENCE: Komodo-one worn dead wop 0.216-0.270 0.241 5 colony was found. Lav 0.126-0.180 0.155 5 Wav 0.054-0.090 0.076 5 DISTRIBUTION: Britain, East and West At- Lm 0.342-0.486 0.434 10 lantic, Mediterranean, Caribbean, Indian 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 7

Ocean, W. Pacific, Galapagos Islands, Sin- FAMILY SCRUPOCELLARIIDAE gapore. LEVINSEN, 1909 Genus Scrupocellaria Van Beneden, 1845 SUPERFAMILY BUGULOIDEA GRAY, 1848 Scrupocellaria spatulata (D'Orbigny), 1851 FAMILY BUGULIDAE GRAY, 1848 Figures 11, 12 Genus Caulibugula Verrill, 1900 Cellularia spatulata D'Orbigny, 1851. Crisia pilosa Audouin, 1826, p. 241; 1828, p. 71. Caulibugula mortenseni (Marcus), 1925 Scrupocellaria pilosa Waters, 1913, p. 478. Figures 9, 10 Scrupocellaria cervicornis Marcus, 1922, p. 425. Scrupocellaria spatulata Harmer, 1926, pp. 382- Stirpariella mortenseni Marcus, 1925a, p. 153; 384. 1925b, p. 37. Caulibugula mortenseni, Harmer, 1926, p. 465. DESCRIPTION: Colonies are erect, biserially Cook, 1968, p. 163. branching, and delicate. Zooids are elongate, DESCRIPTION: The colony is erect, with a with the oval opesia occupying only one-half jointed stalk and biserial branches. Zooids to one-third of the frontal wall; the rest is a are moderately large, narrowing proximally, smooth proximal gymnocyst. Four to five with sharply pointed outer distal corners and spines are located at the distal end of each a membranous frontal surface (fig. 9). About zooid. The scutum, on the proximal inner two-thirds ofthe way down the outer edge of side of the opesia, is narrow, covering only each zooid is a short, stout peduncle bearing part of the frontal membranous area. It is a bird's head avicularium. The avicularium slightly flattened and unbranched. Small vi- is slightly longer than the width ofthe zooids. bracula with delicate, short, curved setae oc- It has an elongated head and a strongly curved cur on the basal surface of the colony. Mar- beak (fig. 10). No ovicells were present on ginal avicularia are located distally on the our specimen. In zooids of mature colonies, sides ofzooids. They are triangular in shape. a pedunculate ovicell occurs attached to the Zooids are never more than two across, ex- distal corner of the opesia. cept at a bifurcation. No ovicells were present on this colony. Measurements Range Mean N Measurements Lz 0.540-0.630 0.598 5 N Wz 0.234-0.306 0.266 5 Range Mean Lop 0.378-0.450 0.421 5 LZ 0.306-0.360 0.346 5 Wop 0.198 0.198 5 Wz 0.144-0.180 0.162 5 Lav 0.360-0.378 0.369 2 Lop 0.180-0.234 0.202 5 WaV 0.180 0.180 2 Wop 0.126-0.144 0.137 5 DISCUSSION: Our specimen appears to be a DISCUSSION: Harmer (1926) placed Scru- very young colony of Caulibugula morten- pocellaria pusilla (Smitt, 1872) in the syn- seni, previously described from the Sunda onymy of this species. However, Osburn Strait and Java Sea. The colony has only one (1940) gave reasons, based chiefly on avic- branch bifurcation (type 5 of Harmer, 1923) ularian characteristics, for keeping the two and lacks the elongate kenozooid stalk, but species separate. On the basis of our exami- zooid and avicularian morphology corre- nation ofCaribbean specimens, we agree with spond extremely well with those of the spec- Osburn. The specimen of Scrupocellaria imen illustrated by Harmer (1926, pl. spatulata in our collection was apparently a XXXIII, figs. 19-22). In this species of Cau- young colony. It lacks ovicells and does not libugula, spines are lacking or vestigial and possess the giant spatulate marginal avicu- zooids have a shape very similar to those of laria which sometimes occur in this species Bugula neritina. (e.g., pl. XXVI, fig. 1, in Harmer, 1926). OCCURRENCE: Komodo-one very small OCCURRENCE: Komodo. colony was found. DISTRIBUTION: Red Sea, Zanzibar, Indo- DISTRIBUTION: Indonesia, West Africa. nesia. 8 AMERICAN MUSEUM NOVITATES NO. 2847

FIGS. 9-12. 9. Caulibugula mortenseni Komodo (AMNH 585), portion of branch. Scale = 400 ,um. 10. C. mortenseni, close-up of avicularium. Scale = 100 Aim. 11. Scrupocellaria spatulata, Komodo (AMNH 586), zooids. Scale = 200 ,m. 12. S. spatulata, portion of branch. Scale = 400 ,m.

SUPERFAMILY MICROPOROIDEA GRAY, 1848 Thalamoporella granulata var. stapifera Levin- sen, 1909, p. 188. FAMILY THALAMOPORELLIDAE Thalamoporella stapifera Harmer, 1926, p. 297. LEVINSEN, 1909 Soule and Soule, 1970, p. 10. Genus Thalamoporella Hincks, 1887 DESCRIPrION: Colonies are large and encrust on coral. Zooids are rectangular to sub- Thalamoporella stapifera Levinsen, 1909 hexagonal in shape. From the raised beaded Figures 13-16 border the granular cryptocyst, perforated by 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 9

FIGS. 13-18. 13. Thalamoporella stapifera, Lombok (AMNH 587), zooids and ovicell. Scale = 200 ,Am. 14. T. stapifera, Bali (AMNH 588), showing vicarious avicularium. Scale = 200 gm. 15. T. stapifera, compass. Scale = 40 um. 16. T. stapifera, calipers. Scale = 40 um. 17. T. komodoensis, Komodo (holotype, AMNH 589), zooids and ovicell. Scale = 200 um. 18. T. komodoensis, calipers. Scale = 40 Mum. evenly spaced small pores, slopes inward to- polypide tube to the orifice. The orifice-ope- ward the two large opesiules and rises as a sia is shaped like a semicircle distally and a 10 AMERICAN MUSEUM NOVITATES NO. 2847 smaller semicircle proximally . A tiny pair of foramen. Spicules include calipers in the shape condyles separates the parts. 4Calcification of ofa very broad wishbone with the ends point- the distal end of the zooid i's smooth, and ing inward. forms a narrow, slightly raised rim around HOLOTYPE: AMNH 589. the distal portion of the orifiIce. There may PARATYPE: AMNH 590. be an occasional small adoralI tubercle (e.g., ETYMOLOGY: This species was named after fig. 13). The ovicell is quite large, smooth, the island of Komodo where it was found. and oval, with a median sutu:re; it opens via DESCRIPTION: The colony is encrusting. a wedge cut out over the oriifice. Ovicelled Zooids are subhexagonal in shape, with a zooids are shorter than autoz(coids and have beaded border surrounding the granular larger opesiules and a more Suinken polypide cryptocyst, which in this species is flatter and tube. The avicularium has a long, narrow ros- less depressed centrally than in many Thal- trum with a pair of stout, triarigular condyles amoporella species. Opesiules (slightly un- at the base ofthe rostrum (fig. 16). The prox- equal in size) are found one on each side of imal end of the avicularium iis rounded and the polypide tube. The orifice-opesia is semi- has a narrow cryptocyst. A vicularia were circular distally and has a broad slightly flat- present but rare in our specime-ns. Compasses tened proximal sinus bordering the polypide and calipers were found. TheD compass (fig. tube. Calcification ofthe distal rim is smooth 15) has an angle of approxim,ately 1280. The and flat with no trace ofadoral tubercles. The calipers (fig. 16) are wishbo)ne or stirrup- proximal rim of the orifice and the polypide shaped. tube area show a more strongly granular cal- Measurements cification than the proximal part ofthe cryp- Range Mean N tocyst; this region is imperforate, while the Lz 0.576-0.846 0.738 15 cryptocyst underlying the proximal zooid Wz 0.360-0.504 0.438 15 surface is perforated by evenly spaced pores. Lo 0.180-0.306 0.227 15 Ovicelled zooids are equal in size to auto- Wo 0.198-0.234 0.215 15 zooids; they have larger opesiules, a narrower Lov 0.414-0.432 0.423 4 Wov 0.540-0.594 0.572 4 polypide tube, and a larger orifice. The ovicell Lavz 0.540-0.666 0.597 7 is globular, round in outline, with a deep cen- Wavz 0.270-0.396 0.319 7 Lav 0.414-0.486 0.460 7 tral wedge-shaped opening above the zooidal Wav 0.162-0.234 0.206 7 orifice-opesia. Its calcification is imperforate and smooth except for faint radiating lines ADDMONAL SPECIMENS EXANiSNED:t(tin27) and median suture. The vicarious avicularia 28.3.6.106, 290 B4, Siboga 279 have a smoothly rounded rostrum. The man- Roma Island, N.E. of Timor,oStation36 m. dible and the rostral foramen into which it 28.3.6.104, 290 B3 (whole mc from same fits are mushroom-shaped, semicircular dis- station). 28.9.13.46, co type.,unton Tridacna, tally, with a narrow "stem" attaching to the locality unknown, G. M. R. I_~evmnsen. hinge. The opesia is rounded proximally, OCCURRENCE: Lombok, Sarnur. margined by a granular and imperforate cryp- DISTRIBUTION: Hawaii, IndLonesia. tocyst. The proximal end of the avicularium is pointed. Spicules include very broad cali-

.AThalamonnrellaf*&ff&%PVFVff QP&&w knmndoensniVV4>&13Y new11%, t sreeciesMV\V1 pers (fig. 18). Figures 17, 18 Measurements Range Mean N DIAGNOSIS: Encrusting Thalamoporella Lz 0.360-0.810 0.566 15 with medium-size zooids (smaller than those Wz 0.270-0.468 0.377 15 of T. Zooids with Lo 0.144-0.216 0.179 15 stapifera). slightly unequal Wo 0.144-0.216 0.173 15 opesiules, a flatter, less depressed cryptocyst Lov 0.414-0.468 0.446 4 than many Thalamoporella species, and with Wov 0.468-0.522 0.513 4 Lavz 0.378-0.540 0.457 15 no adoral tubercles ornamenting the smooth, Wavz 0.198-0.288 0.233 15 flat distal rim. Ovicells round in outline, with Lav 0.180-0.324 0.268 15 a deep V-shaped central opening. Avicularia Wav 0.144-0.216 0.184 15 with mushroom-shaped mandible and rostral OCCURRENCE: Komodo. 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI I1I

FAMILY STEGINOPORELLIDAE HINCKS, FAMILY ONYCHOCELLIDAE JULLIEN, 1882 1884 emend. BASSLER, 1953 Genus Smittipora Jullien, 1882 Genus Steginoporella Smitt, 1873 Smittipora harmeriana (Canu and Bassler), 1929 Steginoporella buskii (Harmer), 1900 Figures 23, 24 Figures 19-22 Smittipora abyssicola (Smitt) Harmer, 1926, p. 259 (not Vincularia abyssicola Smitt, 1873, p. 6). Steganoporella buskii Harmer, 1900, p. 272. Velumella harmeriana Canu and Bassler, 1929, p. Thornely, 1905, p. 112. Levinsen, 1909, p. 168. 128. Hastings, 1932, p. 413. Marcus, 1938, p. 22; 1955, p. 285. Cook, 1964a, p. 46; 1968, p. 152. DESCRIPTION: The colony is flat and en- Steganoporella transversalis Canu and Bassler, crusts on coral and shell. Zooids are hexag- 1928a, p. 68. Marcus, 1949, p. 11. onal to subrectangular in shape. The granular Steginoporella buskii Pouyet and David, 1979, p. cryptocyst surface is sunken slightly in the 771. center and raised at the edge. A thin groove divides the zooids. The orifice-opesia is sub- DESCRIPTION: The greenish-colored colo- triangular. Nonbrooding zooids have slight nies are encrusting on pieces of coral. The opesiular indentations. The avicularia are in- large A zooids are square to rectangular in terzooecial, with a granular cryptocyst rela- shape and lined up in neat bricklike rows (fig. tively narrower than that of autozooids. A 19). The frontal surface of the zooid is un- pair of small hinge teeth are located on the derlain by a granular cryptocyst marked by sides of the avicularium. The ovicells are scattered pores and a slightly granular tex- thickenings of the cryptocyst of the distal ture. This cryptocyst has a narrow marginal zooid. Their surface is also granular. Opesia descending portion, a flat central portion, and of fertile zooids are enlarged, perhaps by an ascending polypide tube. The semicircular cryptocystal degeneration as in Steginoporel- operculum (fig. 21), its crescentic shelfvisible la buskii, and lack opesiular indentations. in illustrated specimens, covers the distal third ofthe zooid. A beaded ridge edges each zooid. Measurements B zooids are larger and have opercula with Range Mean N marginal teeth (fig. 22). Embryos are brooded LZ 0.486-0.612 0.567 10 inside A zooids; during this period, the cryp- wz 0.360-0.450 0.405 10 Lo 0.162-0.216 0.191 10 tocyst undergoes degeneration (Cook, 1 964a). Wo 0.162-0.216 0.193 10 Lov 0.054-0.090 0.066 3 wov 0.108-0.144 0.126 3 Measurements Lavz 0.378-0.540 0.457 10 Wavz 0.216-0.360 0.281 10 Range Mean N Lav 0.162-0.252 0.203 10 A zooids WaV 0.090-0.126 0.110 10 Lz 0.630-0.810 0.714 15 Wz 0.504-0.702 0.602 15 DIscuSSION: Canu and Bassler (1929) gave Lo 0.216-0.288 0.256 15 Wo 0.414-0.522 0.466 15 a new name to Harmer's Smittipora abyssi- Loperc 0.198-0.288 0.245 10 cola from Indonesia, but did not discuss their Woperc 0.396-0.540 0.479 10 reasons for doing so. Harmer's species is dis- B zooids tinct from the other species he named in the Lz 0.810-1.008 0.864 10 Wz 0.576-0.756 0.650 10 Siboga report, Smittipora cordiformis, but it Loperc 0.324-0.396 0.351 10 is not identical with the species now known Woperc 0.486-0.594 0.518 10 as Smittipora levinseni which Smitt (1873) described from Florida. Among other differ- OCCURRENCE: Found at Sanur. ences, Smittipora levinseni is considerably DISTRIBUTION: Gulf of Guinea (Sierra larger in zooid size. The Caribbean species Leone to Ghana), Guinea, Ceylon, Indonesia, Smittipora acutirostris is very similar in zooid Torres Strait, Arabia, South Africa, Austra- size. In comparison with acutirostris, S. har- lia, Caribbean, Brazil. meriana has slightly smaller avicularia, and 12 AMERICAN MUSEUM NOVITATES NO. 2847

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FIGS. 19-24. 19. Steginoporella buskii, Bali (AMNH 591). Scale = 400 ,um. 20. S. buskii, showing B (left) and A (right) zooids. Scale = 200 ,um. 21. S. buskii, A zooid operculum. Scale = 200 ,um. 22. S. buskii, B zooid operculum with marginal teeth. Scale = 200 ,um. 23. Smittipora harmeriana, Komodo (AMNH 592), zooids and avicularia. Scale = 200 ,um. 24. S. harmeriana, close-up of avicularium and ovicelled zooid. Scale = 100 ,um. larger orifices, with relatively greater enlarge- cryptocyst of both avicularia and autozooids ment of the orifices of fertile zooids. The appears more sharply depressed from zooid 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 13

FIGS. 25-30. 25. Cribrilariaflabellifera, Lombok (AMNH 593), zooids, ovicell, and avicularia. Scale = 200 ,um. 26. Exechonella brasiliensis, Komodo (AMNH 594), zooids near growing edge. Scale = 200 ,um. 27. E. brasiliensis, close-up of zooid, showing rimmed pores. Scale = 200 ,um. 28. Hippopodina feegeensis, Bali (AMNH 595), ancestrular region. Scale = 400 ,um. 29. H. feegeensis, Komodo (AMNH 596), zooids near growing edge of colony. Scale = 400 ,um. 30. H. feegeensis, Lombok (AMNH 597), ovicelled zooid. Scale = 200 ,um.

margins and avicularian condyles may be ADDmONAL SPECIENS EXAMINED: (BMNH) more pronounced. Smittipora harmeriana: 28.3.6.71, 323D, 14 AMERICAN MUSEUM NOVITATES NO. 2847

- lmrv.'% Im* p '. I 401- .3 i. I :1., . J...0 :t 2 . 16*

FIGS. 31-34. 31. Petraliella chuakensis, Komodo (AMNH 598). Scale = 400 um. 32. P. chuakensis, close-up of ovicell and avicularia. Scale = 200 ,um. 33. Coleopora verrucosa, Komodo (AMNH 599). Scale = 200 Mm. 34. C. verrucosa, close-up showing slitlike pores and hollow papillae. Scale = 20 ,um.

Badjo Bay, W. Flores, Station 50. Cribrilaria radiata var. flabellifera Kirkpatrick, 82.2.23.485-489, Torres Strait, Prince of 1888, p. 75. Wales Channel. 1975.7.16.41, l1/2 km E. of Colletosia radiata Harmer, 1926, p. 475, in part. Pulau Putri Ketjil. Smittipora Puellina radiata var. flabellifera Thomely, 1912, cordiformis: p. 144. 28.3.6.74, 364C Makassar, S.W. Celebes, Colletosia radiataflabellifera Soule, 1959, p. 48. Station 71. Part oftype specimen, 85.12.29.3, Cribrilariaflabellifera Harnelin, 1970, p. 94; 1978, Mergui, Burma. p. 186. Banta and Carson, 1977, p. 392. Win- OCCURRENCE: Two colonies were collected ston, 1984, p. 13. at Komodo. DISTRIBUTION: Indonesia. DESCRIPTION: Colonies are small and encrusting. Zooids are ovoid to rhomboid in shape, with six to nine pairs of costae. Sep- SUPERFAMILY CRIBRILINOIDEA HINCKS, 1879 arating the costae is a line of evenly spaced FAMILY CRIBRILINIDAE HINCKS, 1880 pores. The first pair of costae (or apertural bar) is more extensively fused and tends to Genus Cribrilaria Canu and Bassler, 1928 be thicker than the other costae and slightly Cribrilaria flabellifera (Kirkpatrick), 1888 raised. Above the apertural bar, but under Figure 25 the orifice, is a larger pore. The orifice is semi- 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 15 circular and is bordered distally by six spines. Measurements The ovicell is shaped like a helmet with a Range Mean N small keel. Avicularia are interzooecial. They Lz 0.774-0.936 0.850 5 are rhomboid in shape with;a smooth Wz 0.450-0.738 0.590 5 calci- Lo 0.180-0.216 0.198 5 fied proximal gymnocyst like that of an au- Wo 0.180-0.234 0.212 5 tozooid. The palate is roun(ded proximally Lav 0.090 0.090 and the rostrum is hourglass--shaped, flaring Wav 0.036 0.036 out at the distal end. The aviicularium man- DISCUSSION: The Indonesian species ofEx- dible is broad proximally and c:omes to a point echonella discussed by Harmer (1957) are E. distally, resembling a poplar Ileaf (not shown magna and E. tuberculata. Unlike E. brasi- in fig. 25, but see Winston, 1 984, fig. 25). liensis, both species have tubular peristomes. Measurements In addition, in E. tuberculata, the rims ofthe Range Mean N large pores form spinelike projections, very Lz 0.342-0.414 0.378 5 different from the flattened pore rims of E. Wz 0.234-0.324 0.274 S brasiliensis, while zooids of E. magna are Lo 0.054-0.072 0.061 S characterized by relatively large frontal avic- Wo 0.072-0.090 0.083 5 Lov 0.126 0.126 2 Uularia with curved triangular mandibles. Such Wov 0.180 0.180 2 avicularia are lacking in E. brasiliensis. Lavz 0.180-0.270 0.230 5 Wavz 0.126-0.180 0.148 5 In its flared peristome, orifice shape, and Lav 0.126-0.144 0.130 5 overall zooid size, E. brasiliensis most closely Wav 0.090-0.108 0.094 5 resembles the Caribbean and West African OCCURRENCE: Lombok-oine colony was species, E. antillea. However, the peristome found. ofE. brasiliensis is more flared and lacks the DISTRIBUTION: Philippines, Indian Ocean, spinous projections found in many Carib- New Guinea, Mediterranean , Baja Califor- bean colonies of E. antillea, while the pri- nia, Belize, Costa Rica. mary orifice has a more shallow proximal portion. The most striking difference, though, FAMILY EXECHONELLIDAE IiARMER, 1957 is in the number of frontal pores, which is Genus Exechonei/a less than 25 in E. brasiliensis but approxi- Canu and Bassler, 1927 mately 50-60 in E. antillea colonies from the Exechonella brasilifensis Caribbean. Canu and Bassler, 1928 OCCURRENCE: One colony was collected at Figures 26, 27 Komodo. ,I Brazil. F-riAnnl n hrvifirno;ov C-ninlni Rqcl ?r I 92Rq DISTRIBUTION: Indonesia, p. 72. DESCRIPTION: The colony is small and en- Genus Coleopora Canu and Bassler, 1927 crusting on coral. The zooids are rhomboid Coleopora verrucosa in shape. The frontal surface is convex and Canu and Bassler, 1924 perforated with large pores that have a smooth Figures 33, 34 flat collar surrounding them. The zooids are separated by a deep valley. The orifice is al- Coleopora verrucosa nom. nud. Canu and Bassler, most square in shape and surrounded by a 1924, p. 67; 1927, p. 6; 1929, p. 267. smooth flared peristome. The inner rim of Teuchopora verrucosa Harmer, 1957, p. 898. the primary orifice can be seen within the DESCRIPTION: The colony is small and en- peristome. A pair of stout, triangular con- crusting on coral. Zooids are rhomboidal in dyles are present about two-thirds ofthe way shape. The frontal surface (fig. 34) is perfo- down from the top ofthe orifice. The adven- rated with slitlike to triangular pores and is titious avicularia are oval and have a cross- marked with occasional hollow papillae. bar. They are rare and very small. When pres- These papillae are smooth-textured and nar- ent they are located on the side of the zooid row toward the distal end. -The orifice is sur- and directed transversely. No ovicells were rounded by a high peristome which flares out present on our specimen. slightly. The orifice is almost square and has 16 AMERICAN MUSEUM NOVITATES NO. 2847 a pair of condyles pointing down into the younger zooids the surface is covered with opening. There are no aviculairia. No ovicells tiny pores and is smoother than in older were found on the one specinnen collected. zooids though the pores have raised edges. Measurements The surfaces ofolder zooids are also covered Range Mean N with pores but, with increasing secondary cal- Lz 0.702-1.170 0.868 5 cification, their surfaces become more gran- Wz 0.450-0.720 0.601 5 ular in texture. The orifice has a slightly raised Lo 0.162-0.198 0.184 5 peristome and two condyles pointing down Wo 0.180-0.216 0.198 5 and in. Its distal portion is circular, its prox- DISCUSSION: Harmer (1957) included Re- imal portion slightly concave. Most zooids cent and fossil species in Teuchopora (Ne- have one or two avicularia located above the viani, 1895). However, PoluiZZi (1977) has orifice and directed more or less transversely shown that Teuchopora casttrocarensis, the inward. The avicularia have slightly curved type ofthe genus, differs from Recent species triangular mandibles. The ovicells are large in frontal wall structure, type (ofbudding, and and helmet-shaped with a perforated, heavily other general characters. We btave placed Co- calcified surface. As shown in figure 30, a leopora verrucosa in the Exechonellidae on triangular avicularium, shorter than those of the basis of frontal wall and[ orificial char- autozooids, may be located on the side ofthe acters. However, the specie4s mresentlv in- ovicell. cluded in the genus Coleoporia may not con- Measurements stitute a monothetic group. F urther study is Range Mean N needed. Lz 0.684-1.080 0.860 15 OCCURRENCE: One colony was found at Wz 0.450-0.864 0.652 15 Komodo. Lo 0.180-0.270 0.247 15 Wo 0.216-0.270 0.247 15 DISTRIBUTION: Indonesial, Philippines, Lov 0.432 0.432 Ceylon. Wov 0.522 0.522 Lav 0.180-0.270 0.220 13 Wav 0.090-0.108 0.095 13 SUBORDER ASCOPHORA LE,VINSEN, 1909 OCCURRENCE: Komodo, Sanur, and Lom- SUPERFAMILY SCHIZOPOIRELLOIDEA bok. JULLIEN, 1883 DISTRIBUTION: Circumtropical. FAMILY HIPPOPODINIDAE LEVINSEN, 1909 FAMILY PETRALIELLIDAE HARMER, 1957 Genus Hippopodina Levinsen, 1909 Genus Petraliella Canu and Bassler, 1927 Hippopodina feegeensis (Busk), 1884 Figures 28-30 Petraliella chuakensis (Waters), 1913 Figures 31, 32 Lepraliafeegeensis Busk, 1884, p. 144. Lepralia pulcherrima Canu and Bassler, 1928a, p. Petraliella chuakensis Waters, 1913, p. 518. Liv- 82. ingstone, 1926, p. 99. Canu and Bassler, 1929, Hippopodina feegeensis Levinsen, 1909, p. 353. p. 251. Hastings, 1932, p. 436. Harmer, 1957, Hastings, 1930, p. 729. Marcus, 1939, p. 116. p. 698. Osburn, 1940, p. 412. Harmer, 1957, p. 974. Hippopetraliella (Serripetraliella) chuakensis has- Powell, 1971, p. 771. Banta and Carson, 1977, tingsae, Stach, 1936, p. 371. p. 413. Winston, 1984, p. 19. DESCRIPTION: Colonies are encrusting. Cosciniopsisfallax Canu and Bassler, 1929, p. 296. Zooids are irregular in shape. Their surface DESCRIPTION: Colonies are encrusting on is covered with many large pores and is pieces of shell and coral. Zooids are rectan- heavily calcified. A raised line ofcalcification gular to polygonal in shape. The thin line of marking the meeting of the frontal edge of calcification marking the intercalary cuticle lateral walls and intercalary cuticle sharply is noticeable in younger zooids (fig. 29) but delimits zooid boundaries. Generally there becomes obscured by secondary calcification are two avicularia on a zooid, one on each in the older regions ofthe colony (fig. 30). In side ofthe orifice. When an ovicell is present 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 17 there may be only one aviculairium. The avic- Measurements ularia are curved and directed distally. A Range Mean N complete cross-bar is present on some but Lz 0.720-1.116 0.886 15 not all avicularia. The orifice is curved dis- Wz 0.396-0.684 0.486 15 Lo 0.216-0.252 0.238 15 tally, with parallel sides; its piroximal edge is Wo 0.252-0.306 0.275 15 serrated. Below the orifice is;a smooth shelf; Ws 0.144-0.198 0.161 15 on the bottom edge of the stieii is a row oI DISCUSSION: Zooids of "Watersipora sub- 7-9 bumps. Ovicells are round to elliptical ovoidea" are longer than those of Watersi- in shape. The surface is covered with many pora edmondsoni. The other main differences small perforations and has a]bumpy texture. are the shape of the orifice and sinus. The Older ovicells become calcified over and bur- orifice of W. edmondsoni is not as large and ied in the secondary calcificaltion of the sur- the sinus is not as broad as those of "W. rounding zooids. subovoidea." The condyles are more pro- Measurementss nounced in W. edmondsoni, extending be- Range Mean N yond the rim of the sinus and pointing down Lz 0.360-0.558 0.475 5 into it. Wz 0.306-0.486 0.367 5 We have followed Soule and Soule (1975) Lo 0.144-0.162 0.151 5 in calling this species the "Watersipora sub- Wo 0.126-0.198 0.154 5 ovoidea ofHarmer." It may turn out, as they Lov 0.198-0.216 0.205 5 Wov 0.198-0.288 0.227 5 suggest, that these specimens will be referable Lav 0.162-0.198 0.169 5 to Watersipora aterrima (Ortmann, 1890), Wav 0.054-0.090 0.068 5 but as we do not have any Japanese material OCCURRENCE: Komodo-c ne colony was for comparison, we have not used that name collected. here. Our specimen fits Harmer's illustration DISTIUBUTION: Zanzibar, Torres Strait, (pl. LXIX, fig. 12) in the shape ofthe aperture Philippines, Borneo. and sinus and the presence of a thin collar. We agree with the Soules that peristomial (presence of collar or oral lappets) and ori- FAMILY WATERSIPORIDAE VI[GNEAUX, 1949 ficial characters (particularly the size and shape of the sinus, the sinus ledge, and the Genus Watersipora Neviani, 1895 condyles), are important in defining species "Watersipora subovcgidea" and species groups in Watersipora. The pres- fide Harmer, 19:57 ence or absence ofthe two distolateral areolae Figures 35-37 may also be useful.

1957 ADDmONAL SPECIMENS ExAMmED: (BMNH) Dakaria subovoidea (D'Orbigny' 1957, 367.A.6, Siboga Station 181, Ambon, an- p. 1022, in part. Harmer, "Watersipora subovoidea" fide Harmer, Soule and chorage. 367.A.3, cleaned specimen from Soule, 1975, p. 308. same station, specimen illustrated in Harmer (1957), pl. LXIX, fig. 12. DESCRIPTION: Colonies are encrusting on OCCURRENCE: Komodo. pieces of coral. Dried colonie,s are an irides- DISTRIBUTION: Indonesia and probably cent black. Zooids are ovoid the surface is elsewhere. The extent of distribution uncer- covered with evenly spaced, large pores. A tain until the species of this genus have re- thin calcified line separates each zooid. The ceived revisionary study. orifice is semielliptical, openiing into a broad sinus. The sinus ledge projects distally into Watersipora edmondsoni the orifice (fig. 37). A short cc ndyle does not Soule and Soule, 1968 quite reach its edge. There is a slight peri- Figures 38-40 stome around the orifice, and 1the suboral area is granular but imperforate. TNwo large areolae Watersipora edmondsoni Soule and Soule, 1968, occur in the oral region of each zooid, one p. 215. on each side of the sinus. Th ere are no ovi- DESCRIPTION: Dried colonies are an irides- cells. Embryos are brooded iInternally. cent black and encrust on pieces of coral. 18 AMERICAN MUSEUM NOVITATES NO. 2847

Zooids are ovoid to diamorad shaped. The Schizoporella nivea var. wasinensis Waters, 1913, zooid surface is covered by:many pores ex- pp. 504, 535. cept for a small region near tiie orifice. There Emballothecafurcata Levinsen, 1909, p. 334. are two prominent areolae, oine on each side Calyptotheca wasinensis Harmer, 1957, p. 1017. of the orificial sinus, close to the edge of the DESCRIPTION: Colonies are encrusting. zooid. A thin calcified line marks the bound- Zooids are rectangular and slightly convex, ary of each zooid. The orifice is oval proxi- the frontal surface perforated and heavily cal- mally, with a pair of sharp-pointed condyles cified, the pores sunken in the calcification. marking the beginning of the narrow semi- A calcified ridge separates the zooids. A small circular sinus. There is a slight peristome umbo is present under the orifice. The orifice around the orifice. There are no avicularia is wider than long, with large, blunt lateral and no ovicells. Embryos are brooded inter- condyles and opens into a broad, shallow si- nally. nus (fig. 42). No ovicells were present in our Measurementss specimen. According to Harmer, they are glo- Range Mean N bose, often with triradiate sutural lines, and Lz 0.684-0.954 0.812 15 the fertile zooid has a wider orifice than au- Wz 0.414-0.702 0.559 15 tozooids. Avicularia are located in the distal Lo 0.162-0.216 0.202 15 left and right corners of the zooid. There are Wo 0.198-0.234 0.216 15 Ws 0.090 0.090 15 usually two avicularia per zooid. The avicularia are small and point proximally. DISCUSSION: Our specimens have narrower sinuses than those of the Watersipora ed- Measurements mondsoni colonies in our collections from Range Mean N Hawaii. They are similar to Lz 0.486-0.630 0.554 5 the specimens Wz 0.270-0.414 0.320 5 illustrated by Soule and Soule (1975, pl. 3, Lo 0.126-0.162 0.140 S fig. 5 and pl. 3, fig. 6). It may be true, as Wo 0.144-0.162 0.148 5 suggested by Soule and Soule (1975), that a Lav 0.054 0.054 5 larger degree of speciation has occurred in Wav 0.036 0.036 5 Indo-Pacific watersiporids than had been re- OCCURRENCE: One colony was found at alized. However, until more material is avail- Sanur. able for study it seems best to identify our DISTRIBUTION: Ceylon, Red Sea, Sey- specimens with edmondsoni. chelles, Amirante Is., Providence (Indian ADDMONAL SPECIMENS EXAMINED: Schizo- Ocean), Zanzibar, E. Africa, Indonesia. porella aterrima (Harmer list, no. 898), Torres Strait, Haddon Collection, no. 122, reg. Feb. Genus Schizomavella 24, 1898; specimen illustrated in Soule and Canu and Bassler, 1917 Soule (1975), pl. 3, fig. 6. Watersipora subovoidea, 1981.4.1.3 pt. (Harmer list, no. Schizomavella inclusa (Thornely), 1906 1252), Ceylon, 1905; specimen illustrated in Figures 43, 44 Soule and Soule (1975), pl. 3, fig. 5. Schizomavella subimmersa var. Waters, 1889, p. OCCURRENCE: Sanur. 11. DISTRIBUTION: Hawaii, Indonesia. Schizoporella triangula Thornely, 1905, p. 115. Schizoporella avicularis Thornely, 1905, p. 116. FAMILY SCHIZOPORELLIDAE JULLIEN, 1882 Schizoporella inclusa Thornely, 1906, p. 450. Schizomavella (Metroperiella) lepraliodes Canu Genus Calyptotheca Harmer, 1957 and Bassler, 1929, p. 307. Calyptotheca wasinensis (Waters), 1913 Schizomavella inclusa Harmer, 1957, p. 1028. Figures 41, 42 DESCRIPTION: Colonies are encrusting on Schizoporella nivea (not S. nivea Busk, 1884, p. coral rubble. Zooids are polygonal and slight- 163) Thornely, 1905, p. 114; 1907, p. 189. ly convex. The zooid frontal surface is heavi- Schizoporella nivea Waters, 1909, p. 168, var. mil- ly calcified and covered with pores except for lanensis (name suggested but retracted). the area around the orifice. There is a row of Thornely, 1912, p. 148. Waters, 1913, p. 502. narrow marginal areolae. A thin line of cal- 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 19

i IX-Jito

FIGS. 35-40. 35. Watersipora "subovoidea," Komodo (AMNH 600). Scale = 400 ,um. 36. W. "subovoidea." Scale = 200 ,m. 37. W. "subovoidea," close-up of orifice. Scale = 40 ,um. 38. W. edmondsoni (AMNH 601), close-up oforifice. Scale = 40 ,m. 39. W. edmondsoni. Scale = 400 ,um. 40. W. edmondsoni. Scale = 200,gm.

cification divides zooids. The orifice is An adventitious avicularium is located in the rounded distally, tapering toward the large middle of each zooid below the sinus of the blunt condyles and small semicircular sinus. orifice. Avicularia have a rounded opesia and 20 AMERICAN MUSEUM NOVITATES NO. 2847

FIGS. 41-44. 41. Calyptotheca wasinensis, Bali (AMNH 602). Scale = 200 ,um. 42. C. wasinensis, close-up of orifice. Scale = 100,um. 43. Schizomavella inclusa, Komodo (AMNH 603). Scale = 400,gm. 44. S. inclusa, close-up of orifice. Scale = 40 ,um. a subtriangular rostrum with a complete cross- 1030 for justification of the synonymy for bar. They are directed proximally. Large, vi- this species. carious avicularia occur rarely in this species. OCCURRENCE: One colony was found at The mandible of such an avicularium is il- Komodo. lustrated in Harmer, 1957, pl. LXVI, fig. 7. DISTRIBUTION: Ceylon, Indonesia, Austra- No ovicells were present on our specimens. lia. According to Harmer (1957) they are immersed, porous, and sometimes edged by a Genus Stylopoma Levinsen, 1909 semicircular ridge of calcification (Harmer, 1957, pl. LXVI, fig. 4). Stylopoma duboisii (Audouin), 1826 Figures 45-47 Measurements Mean N Flustra ?duboisii Audouin, 1826, p. 239; 1828, p. Range 66. Savigny, pl. VIII, figs. 41, 42. Lz 0.414-0.468 0.443 5 Wz 0.216-0.378 0.313 5 Lepralia schizostoma MacGillivray, 1869, p. 135; Lo 0.090-0.126 0.108 5 1879, Dec. 4, p. 33. Wo 0.090-0.108 0.097 5 Lepralia (Schizoporella) assimilis Haswell, 1880, Lav 0.054-0.090 0.072 5 p. 39 (not Lepralia assimilis Hassall, 1842, p. Wav 0.036-0.054 0.047 5 412). DISCUSSION: See Harmer, 1957, pp. 1029- Schizoporella spongites (in part) Thornely, 1905, 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 21

p. 14(nodescr.); 1907, p. 189 (nodescr.); 1912, DESCRIPTION: Colonies are encrusting and p. 148 (no descr.). green in color. Zooids are rhomboidal in the Stylopoma duboisii Harmer, 1957, p. 1033. Pow- initial layer ofgrowth, becoming irregular in ell, 1967b, p. 168. Cook, 1968, p. 196. frontally budded layers. The zooid surface is DESCRIPrION: Colonies are encrusting on thick, granular, and covered with pores. In coral. Zooids are rectangular in shape. The older sections of the colony the calcification zooid surface is covered with pores, except becomes so rough and heavy that it may form for the area in the immediate vicinity of the spiny ridges between the pores. The orifice is orifice, which has a granular texture. In older round distally, opening into a broadly sections ofthe colony the area between pores V-shaped sinus. The beginning of the sinus is thick and granular; the pores are sunken is marked by a pair of blunt-pointed, proxi- in polygonal pits. In younger sections it is mally directed condyles. In younger zooids only slightly roughened. A thin irregular line the area around the orifice is smooth except of calcification marks the division between for a few tubercles. In older zooids this area zooids. The orifice is semicircular distally, becomes quite granular (fig. 50). There are with serrated condyles (fig. 47) that may come two kinds of adventitious avicularia. Both right up to the narrow, slitlike central sinus. have a triangular base which then narrows to There are two types of avicularia: small, tri- a point. They differ in size and location: the angular, and adventitious or large, spatulate, larger avicularia are located on one side of and vicarious. The small avicularia occur sin- the orifice and are directed distolaterally; the gly or in pairs on each zooid, beside or just small avicularia are located under the orifice below the orifice. They are directed distolat- and point in the same direction. No vicarious erally or transversely outward. The large vi- avicularia were present in these colonies. They carious avicularia are spatulate (fig. 46) and are illustrated in Harmer, 1957, pl. LXXIV, less frequent. No ovicells were present in fig. 12. No ovicells were present on colonies either colony collected. When present they collected, but according to Harmer's (1957) are very large, porous, and cover the fertile illustration (pl. LXXIV, fig. 8), they are very zooid orifice (Harmer, 1957). large, covered with the same granular, porous calcification, and occlude the fertile zooid Measurements orifice. They have an independent circular, Range Mean N sometimes tubular, opening. Lz 0.378-0.630 0.478 15 wz 0.252-0.450 0.331 15 Measurements Lo 0.090-0.126 0.101 15 Range Mean N Wo 0.108-0.126 0.118 15 LZ 0.540-0.990 0.718 15 Adventitious avicularia wz 0.378-0.810 0.541 15 Lav 0.036-0.072 0.053 15 Lo 0.162-0.216 0.191 15 Wav 0.018-0.036 0.035 15 wo 0.162-0.216 0.180 15 Spatulate avicularifa Lav, 0.072-0.144 0.100 7 Wav, 0.054-0.090 0.062 7 Lav 0.324-0.378 0.351 2 LaV2 0.126-0.234 0.168 9 Wav 0.162-0.180 0.171 2 WaV2 0.072-0.108 0.088 9 Lavz 0.486-0.540 0.513 2 Wavz 0.234-0.288 0.261 2 OCCURRENCE: Lombok and Sanur. OCCURRENCE: Komodo and Sanur. DISTRIBUTION: Philippines, Ceylon, Indian DISTRIBUTION: W. Africa, Red Sea, Indian Ocean, Indonesia, Great Barrier Reef, north- Ocean, Ceylon, Indonesia, Australia. east Australia, Red Sea. Stylopoma viride (Thomely), 1905 Figures 48-50 FAMILY SMITTINIDAE LEVINSEN, 1909 Schizoporella viridis Thornely, 1905, p. 116; 1912, Genus Parasmittina Osbum, 1952 p. 148. Waters, 1909, p. 147. Marcus, 1921b, p. Parasmittina signata (Waters), 1889 17. Livingstone, 1926, p. 84. Figures 51-54 Stylopoma grandis Canu and Bassler, 1929, p. 316. Stylopoma viride Hastings, 1932, p. 425. Harmer, Smittina signata Waters, 1889, p. 17. 1957, pp. 1036-1038. Smittina marmorea Waters, 1909, p. 157. 22 AMERICAN MUSEUM NOVITATES NO. 2847

-,W 10 0.1 . . L. III Jr .4 -*O(# '- A-

.71, i 11 -.: % 1, .-AI...-1 t,

I . t I i K"4- 0- %, *1A 00 -, It '411R,I

Z~*q I'.*~ ~*OA**1~ ! FIGS. 45-50. 45. Stylopoma duboisii, Bali (AMNH 604). Scale = 200 ,um. 46. S. duboisii, large interzooecial avicularium. Scale = 200,gm. 47. S. duboisii, close-up of orifice. Arrow points to serrated condyle. Scale = 20 ,um. 48. S. viride, Lombok (AMNH 605), close-up of orifice. Scale = 200 um. 49. S. viride. Scale = 400 ,m. 50. S. viride. Scale = 200 ,m.

Smittia signata Thomely, 1912, p. 151 (no descr.). Smittina signata Hastings, 1932, p. 429. Harmer, Schizoporella horsti Osburn, 1927, p. 127. 1957, p. 928. Lacerna signata Canu and Bassler, 1929, p. 308. Parasmittina signata Lagaaij, 1963, p. 197. 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 23

FIGS. 51-54. 51. Parasmittina signata, Bali (AMNH 606), zooids and ovicells. Scale = 200 ,um. 52. P. signata, Komodo (AMNH 607), showing giant spatulate avicularium. Scale = 200 ,um. 53. P. signata, showing single ephemeral oral spine. Scale = 100 ,um. 54. P. signata, close-up ofovicelled zooids. Scale = 100 Jim. Rimulostoma? signata Cheetham and Sandberg, as the smaller ones. Both point proximally. 1964, p. 1028. The smaller avicularia are more abundant DESCRIPTION: Colonies are encrusting. than the larger type. The ovicell is round, Zooids are rectangular to polygonal. The tex- with scattered pores on the surface and a ridge ture of the calcification of the frontal wall is of calcification around the edges. rough, especially in the older portions of the Measurements colony, but imperforate except for a row of Range Mean N large areolae around the zooid margin and Lz 0.342-0.576 0.443 15 an occasional small pore, usually near the Wz 0.270-0.414 0.307 15 Lo 0.090-0.144 0.118 15 avicularium. The orifice is round distally. Wo 0.108-0.144 0.122 15 Proximally, a deep narrow sinus occurs be- Lov 0.162-0.216 0.180 8 Wov 0.234-0.288 0.248 8 tween two square-edged condyles. There is Lav 0.054-0.108 0.082 12 no lyrula. Three oral spines are present on Wav 0.036-0.054 0.045 12 young zooids. Two kinds of adventitious Spatulate avicularia avicularia are found. Small avicularia with Lav 0.270 0.270 2 subtriangular rostra are located below and to Wav 0.090 0.090 2 one side of the orifice. Large spatulate avic- DISCUSSION: The generic placement of this ularia may be located in the same position distinctive species, the only smittinid lacking 24 AMERICAN MUSEUM NOVITATES NO. 2847 a lyrula except for members ofthe genus Co- lia? Challenger. 32.4.20.51 pt., Australia, donellina, is difficult. Harmer (1957) placed Great Barrier Reef Exp., Station xxii. it in Smittina, but that genus is now generally 32.4.20.119, Australia, Great Barrier Reef reserved for those species with a porous fron- Exp., off N.E. Low Island. 99.7.1.2014, Busk tal wall (Hayward and Ryland, 1979, p. 98). Collection. The imperforate frontal wall and the row of OCCURRENCE: Sanur and Komodo. marginal pores, as well as the finely porous DISTRIBUTION: Australia, Red Sea, Indian ovicell, are characteristics found in Smittoi- Ocean, Philippines, Tahiti, Indonesia, Gulf dea and Parasmittina. But Smittoidea is lim- of Mexico, Cura9ao, Mexico. ited to those species with a median adven- titipus avicularium, whereas the small and Parasmittina parsevalii (Audouin), 1826 large avicularia of signata are located to one Figures 55-58 side ofthe orifice. Apparently on the basis of Cellepora parsevalii (Flustra?) Audouin, 1826, p. the location of avicularia and the structure 238; 1828, p. 64 (as Persevalii). Savigny, pl. VII, ofthe frontal wall and ovicells, Lagaaij (1963) figs. 91-94. placed the species in Parasmittina. Smittia trispinosa var. i Hincks, 1884, p. 361. Within the Smittinidae the absence of a Smittia obstructa Waters, 1889, p. 18. lyrula is almost certainly a derived character. Smittina tripora Canu and Bassler, 1929. For that reason it might seem desirable to Smittina trispinosa var. applicata Canu and Bas- place the species in a genus of its own. How- sler, 1929, p. 345. ever, to use the absence of a character to Smittina obstructa Hastings, 1932, p. 431. monothetically define a group is usually con- Parasmittina parsevalii Harmer, 1957, p. 941. sidered bad taxonomic practice. Based on the Powell, 1967b, p. 172. Cook, 1968, p. 215. presence of derived characters-variety and DESCRIPTION: Colonies are large and en- positioning of avicularia, frontal wall mor- crusting on corals. Zooids are rectangular in phology, and ovicell morphology-the species shape, heavily calcified, and the rough-tex- appears to fit within the definition of Para- tured frontal surfaces are imperforate except smittina and is probably appropriately left in along the margins where there is a row of that genus. areolae. The orifice is roundish to square. At Though this species apparently has a cir- the proximal end is a square lyrula, flanked cumtropical distribution, Harmer (1957) by condyles. Two spines are located at the noted that zooid size varied considerably in distal end ofyoung zooids (fig. 55). The short colonies from different locations and depths; peristome is raised laterally into two lappets in fact, examination of specimens to which which flare out slightly at the top. On zooids hehadaccess shows that some specimens have with ovicells, the peristome becomes more zooids almost twice as long as others; this circular (fig. 57). There are three kinds ofad- amount ofvariation seems suspiciously large. ventitious avicularia, two small and one large. In addition, it should be noted that Carib- The small avicularia are found oriented in bean and West African populations are not almost every direction. Small oval avicularia only somewhat smaller in size than those from near the proximal edge ofthe orifice are usu- the Indo-Pacific, but also lack the large avic- ally directed proximally. Small triangular ularia (P. L. Cook, personal commun.). Only avicularia occur beside the orifice on either further study will show whether we are deal- the lateral or proximal margins of the peri- ing with one extremely variable species or a stome. They point either distally or distolat- species complex. erally. The small avicularia in the ancestrula ADDrIToNAL SPECIMENS ExAMINED: (BMNH) area are scattered with no apparent pattern. 81A, Siboga Station 105, Sulu Archipelago, The large avicularia are spatulate; the spoon- 275 m. 472D, Siboga Station 99, N. Ubian, shaped rostra have a scalloped edge. They Sulu Archipelago, 16-23 m. Japan, Owston take up almost halfofthe zooid bearing them, Collection, SD, reg. June 23, 1902. Japan, which is distorted in shape, with orifices dis- Mitsukuri Collection, C, reg. Sept. 23, 1896. placed laterally and sometimes partially oc- 1961.11.2.17, Mudlump no. SP5, A. H. cluded. The large avicularia are directed Cheetham. 1973.3.22.84 pt., Gold Coast, proximolaterally. Ovicells are round. The Bassindale Collection. 34.11.12.65, Austra- frontal surface is covered with small pores 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 25

FIGS. 55-58. 55. Parasmittina parsevalli, Bali (AMNH 608), growing edge of colony. Scale = 1 mm. 56. P. parsevalii, Sanur (AMNH 609), close-up of giant spatulate avicularium. Scale = 200 ,um. 57. P. parsevalii, Komodo (AMNH 6 10), ovicelled zooids showing various types and orientations ofavicularia. Scale = 200 ,tm. 58. P. parsevalii, Sanur, ovicelled region of same colony seen in figures 55 and 56. Scale = 200,m. and slits, and there is a peripheral layer of DISCUSSION: Most of the specimens ex- secondary calcification. amined by Harmer (1957) including the specimen from Ghardaqa, Red Sea, appear to be- Measurement,'s long to one species. There are two exceptions. Range Mean N The specimen from Torres Strait (Haddon Lz 0.360-0.612 0.444 15 Collection, reg. Feb. 24, 1898) does not be- Wz 0.270-0.468 0.362 15 long to that species. In the specimen from Lo 0.108-0.198 0.138 15 Wo 0.108-0.162 0.131 15 Skeat, Malay Peninsula, which Harmer illus- Lov 0.162-0.198 0.183 6 trated as a variation because of its reversed Wov 0.216-0.270 0.237 6 avicularia (pl. LXV, fig. 6), the avicularia are Small avicularia not just reversed, but have a very bulbous Lav 0.054-0.108 0.077 8 rostral surface; that specimen probably rep- Wav 0.054 0.054 8 resents a third species. Avicularia near orifice Lav 0.108-0.144 0.117 8 ADDmONAL SPEcvIMENs EXAMiNED: (BMNH) Wav 0.054-0.072 0.065 8 38I, Siboga Station 77, Borneo Bank, Strait Spatulate avicularia of Makassar, 59 m; specimen illustrated in Lav 0.288-0.342 0.313 5 Harmer (1957), pl. LXV, fig. 5. 45B, same Wav 0.090-0.162 0.122 5 station. 388C, 388C-1, Siboga Station 81, 26 AMERICAN MUSEUM NOVITATES NO. 2847

,,~6

~ ~ ~ 4

P, 1tj; --

FIGS. 59-63. 59. Parasmittina hastingsae, Komodo (AMNH 61 1), close-up ofzooid at growing edge to show broad lyrula and two oral spines. Scale = 100 ,um. 60. P. hastingsae, close-up ofovicell. Scale = 100 ,um. 61. P. hastingsae, close-up of giant triangular avicularium, showing serrated edge of rostrum. Scale = 100 ,um. 62. P. hastingsae, showing giant avicularium. Scale = 100 um. 63. P. hastingsae, showing small oval avicularium. Scale = 40 ,um.

Borneo Bank, Strait of Makassar, 0-34 m. no. 1379, Ghardaqa, Red Sea, Crossland, 4d. 563A, Siboga Station 37, Sailus Ketjil, Pa- Harmer list, no. 1409, Torres Strait, Had- ternoster Is., 0-27 m. 289A, Siboga Station don, reg. Feb. 24, 1898, 175. 47b, Sumbawa, N., 296 m. 23J, 325G 2, OCCURRENCE: Sanur and Komodo. One of 325N-1, Siboga Station 50, Badjo Bay, W. the more abundant species found. To judge Flores, 0-40 m. 72C, Siboga Station 303, by the number ofspecimens listed in Harmer Haingsisi, Samau Island, W. Timor, 0-36 m. (1957), it was also one ofthe most abundant 81B, Siboga Station 105, Sulu Archipelago, species in Siboga collections. 275 m. 150E, 518A, Siboga Station 204, be- DISTRIBUTION: Indonesia, W. Africa, Aus- tween Wowoni and Buton, S.E. Celebes, 75- tralia, Philippines, India, Red Sea. 94 m. 295G, Siboga Station 273, Jedan, Aru Island, 13 m. 376K, Siboga Station 213, Sal- eyer, S. of Celebes, 0-36 m. 412C, Siboga Parasmittina hastingsae Station 310, Sumbawa, E., 73 m. 377E, Si- Soule and Soule, 1973 boga Station 240, Banda, Banda Sea, 0-45 Figures 59-63 m. Harmer list, no. 1394, Malay Peninsula, Parasmittina hastingsae Soule and Soule, 1973, p. Skeat, reg. Nov. 30, 1899, lb. Harmer list, 417. 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 27

DESCRIPTION: The colony is small and en- orifice, and a rounded suboral umbo. Paired crusting on coral. Zooids are diamond shaped. avicularia locatedjust proximal to the umbo, The surface ofthe zooid is smooth in younger their mandibles directed crosswise. Ovicell portions ofthe colony. There is a row ofmar- with curved median band of pores. ginal pores which are small on zooids near ETYMOLOGY: The species name is derived the growing edge and become larger in older from the Greek karsios = crosswise, and the zooids by incorporation in larger pits. Two Latin seta = bristle, in an attempt at paral- spines are located at the distal edge of the leling the other descriptive names used in this orifice, which is almost square. The lyrula is genus. broad and slopes down into the orifice open- HOLOTYPE: AMNH 612. ing. Older zooids develop a broad secondary DESCRIPTION: The colony is encrusting. sinus. The peristome is raised laterally, form- Zooids have 10 marginal spines and a frontal ing two lappets which flare out on the sides. wall that is imperforate except for marginal There are three kinds of avicularia: two are areolae. The frontal wall surface is faintly located on the zooid, and the third is inter- granular and slightly convex with a pro- zooecial. One type ofavicularium is oval and nounced suboral umbo. The orifice is trifo- directed distally. It is located on the side or liate, rounded distally, and arcuate proxi- slightly below the orifice. The second type is mally, its distal edge raised so that it is hooded triangular, directed outward and distally, lo- in side view. A pair of small triangular avic- cated on the side of the orifice, and raised ularia face each other across and proximal to slightly. The interzooecial avicularium has a the suboral umbo, their mandibles directed triangular and serrated rostrum with a round- proximomedially (crosswise). The ovicell is ed proximal end. The two parts are divided globular and imperforate except for a curved by a cross-bar. The ovicell is globose, its sur- median row of pores (fig. 65). face covered with many pores. Measurements Measurements Range Mean N Range Mean N Lz 0.396-0.540 0.436 5 LZ 0.288-0.540 0.356 10 Wz 0.270-0.324 0.299 5 wz 0.252-0.396 0.306 10 Lo 0.090-0.108 0.104 5 Lo 0.090-0.144 0.115 10 wo 0.090 0.090 5 wo 0.108-0.144 0.115 10 Lov 0.162 0.162 Lov wov 0.180 0.180 0.144-0.180 0.162 7 Lav 0.036-0.054 0.040 5 wov 0.216-0.234 0.221 7 WaV 0.018 0.018 5 Small oval avicularia Lav 0.090-0.144 0.111 7 DISCUSSION: Brown (1954) revised the WaV 0.054 0.054 7 species of the genus Crepidacantha. Of the Medium triangular avicularia species he discussed, C. carsioseta most Lav 0.144-0.216 0.162 7 closely resembles C. poissonii and C. solea, WaV 0.054-0.090 0.072 7 two species which have avicularian mandi- Interzooecial avicularia bles directed inward and an ovicell with a Lavz 0.216-0.450 0.354 9 band-shaped porous area. Harmer (1957) Wavz 0.162-0.270 Lav 0.144-0.396 0.274 9 found two species of Crepidacantha, C. pois- Wav 0.072-0.162 0.118 9 sonii and C. crinispina, in the Siboga collec- OCCURRENCE: Two colonic.s were found at tions. Brown did not have access to Harmer's Komodo. Siboga specimens (Harmer's ascophoran DISTRIBUTION: Hawaii, In(donesia. manuscript had not yet been published), so did not discuss them. However, examination FAMILY CREPIDACANTHIDAE LEVINSEN, 1909 of Harmer's specimen ofpoissonii showed it Genus Crepidacantha Leivinsen, 1909 to be that species and not carsioseta. In C. poissonii, the avicularia are located much Crepidacantha carsioseta, new species closer to the orifice and to the lateral edge of Figures 64-66 the zooid, and there is no pronounced suboral DIAGNOSIS: Crepidacanthcz with 10 mar- umbo. In avicularian position and in the ginal spines, a trifoliate proiximally arcuate presence of the suboral umbo, C. carsioseta 28 AMERICAN MUSEUM NOVITATES NO. 2847 is very similar to C. solea, a species described Measurements by Canu and Bassler (1929) from Tizard Reef, Range Mean N South China Sea. However, in that species Lz 0.450-0.540 0.508 15 the proximal lip ofthe orifice is squared rath- Wz 0.396-0.522 0.441 15 Lo 0.072-0.126 0.099 15 er than rounded. Wo 0.144-0.216 0.167 15 ADDITIONAL SPECIMENS ExAiNED: (BMNH) Lov 0.234-0.270 0.257 10 Wov 0.270-0.324 0.306 10 Crepidacantha poissonii, 249Q, Siboga Sta- Lasc 0.036 0.036 15 tion 310, Sumbawa, E., 73 m. Wasc 0.036-0.054 0.049 15 OCCURRENCE: Sanur. Only one colony found. It was heavily fouled by diatoms and DISCUSSION: A number of species and sub- encrusting algae. species of Fenestrulina have been described, particularly from the Southern Hemisphere. We feel our specimens differ enough from FAMILY MICROPORELLIDAE HINCKS, 1879 these to warrant specific designation. Zooids Genus Fenestrulina Jullien, 1888 of Fenestulina malusii have the central area of the frontal wall proximal to the ascopore Fenestrulina harmeri, new species imperforate, and the orifice has two or three Figures 67-69 spines. Two specimens illustrated in the lit- Fenestrulina malusii Harmer, 1957, p. 966, in part. erature are similar to Fenestrulina harmeri in having a completely porous frontal wall: DIAGNOSIS: Fenestrulina with an evenly the F. malusii of Wass and Yoo (1983, figs. porous frontal wall, one or two rows ofpores 67-69) and F. catastichos (described by Gor- distal to the orifice, a single oral spine, and don, 1984, pl. 40D, E). The specimens from three or four pore chambers in distal halves south Australia described by Wass and Yoo of zooids. Ascopore crescentic, denticulate, have two to three oral spines and the ovicell with a proximally raised rim. Ovicell with has a bumpy rather than a ridged texture. In radiating ridges and a marginal row ofpores. addition, though the frontal wall is porous, ETYMOLOGY: The species is named in hon- there appear to be only narrowed areolae dis- or of Sir Sidney F. Harmer, who made the tal to the orifice. According to the descrip- study ofbryozoans from the Indonesian area tion, F. catastichos has a row of pores distal a major part of his life's work. to the orifice (though they are not visible in HOLOTYPE: AMNH 613. specimens illustrated), but differs from F. PARATYPES: AMNH 614, 615. harmeri in having no orificial spines, having DESCRIPTION: The colonies are encrusting a raised buttonlike disc rimming the ascopore on pieces of shell. Zooids are hexagonal, the all around, and, at least in the specimen il- frontal wall slightly convex and granular in lustrated, having the outer row of pores of texture, its surface evenly perforated by large the ovicell occluded by calcification. Our pores. In just budded zooids these pores are specimen most closely resembles the speci- round, but in older zooids they are stellate, men illustrated by Harmer (1957, pl. LXII, their centers filled by tissue (fig. 69). Three fig. 32) from the Siboga collection (no. 489C). or four pore chambers can be seen in distal Of the other Siboga material ofF. malusii in halves ofzooids (fig. 68). One or two rows of the collection of the British Museum, speci- pores occur distal to the orifice, and there is men no. 55 1A-3 is also F. harmeri, but spec- no imperforate proximal area. The orifice is imen 371G is not. Although Harmer stated semicircular, with a smooth rim from which that the specimen illustrated in figure 32 is one distal oral spine (fig. 68) emanates. The like "the Torres Straits specimen and oth- median ascopore (fig. 69) is crescentic and ers," examination of the Torres Strait spec- denticulate, with a proximal raised rim. There imen showed it to belong to a species with a are no avicularia. Ovicells are prominent, re- large imperforate frontal area. cumbent, and circular in outline, closed by ADDITIONAL SPECIMENS EXAMINED: the zooecial operculum. Ridges of calcifica- (BMNH) Fenestrulina malusii, 489C, Siboga tion radiate from the center ofthe ovicell past Station 136, Ternate, 23 m. 551A-3, Siboga the row of pores just inside its outer rim. Station 310, Sumbawa E., 73 m. 371, Siboga 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 29

FIGS.64-69. 64. Crepidacantha carsioseta, Bali (holotype, AMNH 612). Scale = 200 ,m. 65. C. carsioseta, including ovicelled zooid. Scale = 200Am. 66. C. carsioseta, close-up ofzooid showing orifice shape and marginal spines. Scale = 200 Am. 67. Fenestrulina harmeri, Komodo (AMNH 613, holotype), zooids and ovicells. Scale = 200Am. 68. F. harmeri, close-up of growing edge showing pore chambers. Scale = 100 ,um. 69. F. harmeri, close-up of ascopore and two pores. Scale = 20 ,um.

Station 273, Jedan, Aru Island, 13 m. Harm- lection, reg. Feb. 24, 1898, 54. Harmer list, er list, no. 1669, Torres Strait, Haddon Col- no. 1666, Japan, Uraga Channel, off Tokyo, 30 AMERICAN MUSEUM NOVITATES NO. 2847

,':,U71 .1. I IC:a 4, 'Sd

M , it; -v.1-0 '01 ,AZXI,-^, FIGS. 70-72. 70. Margaretta triplex, Komodo (AMNH 616), colony branch, showing chitinousjoint. Scale = 1 mm. 71. M. triplex, close-up ofzooids. Scale = 200 ,tm. 72. M. triplex, cross-section ofbranch. Scale = 200,um.

20-30 fm, Owston Collection, reg. June 23, OCCURRENCE: One colony was collected at 1902. Harmer list, no. 1667, Japan, deep Komodo. water, Prof. Mitsukuri, reg. Sept. 23, 1896. DISTRIBUTION: Torres Straits, Zanzibar, OCCURRENCE: Komodo. Australia, Indonesia. DISTRIBUTION: Indonesia.

FAMILY MARGARETTIDAE HARMER, 1957 SUPERFAMILY CELLEPOROIDEA LAMOUROUX, 1821 Genus Margaretta Gray, 1843 FAMILY CELLEPORARIIDAE Margaretta triplex Harnier, 1957 HARMER, 1957 Figures 70-72 Genus Celleporaria Lamouroux, 1821 Tubucellaria cereoides Kirkpatrick, 1 890b, pp. 604, 61 1, in part. Celleporaria sibogae, new species Tubucellaria cereoides var. chuakensis Waters, Figures 73-78 1907, p. 130. Livingstone, 1926, p. 87. Celleporariafusca Harmer, 1957, p. 680, in part. Marearetta trinlex Harmer. 1957. t. 841. DIAGNOSIS: Celleporaria with primary ori- DESCRIPTION: The colony is very young, fice orbicular and without a sinus. Frontal small, and erect, with chitinouIsjoints. Zooids, wall granular in texture and imperforate ex- almost rectangular in shape, are arranged in cept for marginal and a few distal pores. Peri- four longitudinal rows. The firontal surface is stome blunt, sometimes bifid, with ovoid covered by rows of pores ex(cept for a small suboral avicularium on its side. Scattered area proximal to the secondary orifice. The small triangular avicularia and large inter- peristome is ribbed longitudinally. The sec- zooecial avicularia with a smooth, spatulate ondary orifice is circular and raised. Spine rostrum of variable length. bases can be seen on the edge of the orifice. HOLOTYPE: AMNH 617. No avicularia occur. No feirtile peristomes PARATYPES: AMNH 618, 619. were present on the colony collected. Ac- ETYMOLOGY: The species is named in hon- cording to Harmer (1957) these are truncated or ofthe Siboga Expedition which from 1899 cones, transversely directed with swollen to 1900 made collections of marine organ- bases. isms at over 300 stations throughout the In- Measurementss donesian region. Range Mean N DESCRIPTION: Colonies are encrusting in LZ 0.090-1.026 0.965 5 multilaminar patches on coral and are a dull wz 0.360-0.486 5 LO 0.108-0.127 0.410 greenish or grayish color when dried. Zooids wo 0.144 0.144 5 are rectangular, decumbent, and regularly ar- 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 31

wF *

k... 0 9 * . 4-

FIGS. 73-78. 73. Celleporaria sibogae, Komodo (AMNH 617, holotype). Scale = 1 mm. 74. C. sibogae, zooids at growing edge of colony. Scale = 400 ,m. 75. C. sibogae, close-up showing shape of primary orifice and triangular adventitious avicularia. Scale = 100 ,um. 76. C. sibogae, Bali (AMNH 618), a more heavily calcified orifice. Scale = 100 ,um. 77. C. sibogae, showing raised interzooecial avicularium. Scale = 200 ,m. 78. C. sibogae, showing suboral avicularium, and large interzooecial avicularium. Scale = 100,m. ranged at the growing edge (fig. 74), but par- primary orifice is orbicular, without a sinus tially or wholly erect in inner regions. The (fig. 75), although the peristome as it develops 32 AMERICAN MUSEUM NOVITATES NO. 2847 includes a sinus. The frontalI surface is im- indicates that several species may be in- perforate except for marginalI areolae (occa- volved, but most of the Siboga specimens sionally a few additional pores are at the dis- belong to the species described above. Torres tal edge of the zooid) and has a regularly Strait specimens he cites as C. fusca appear granular pebbled-glass textur(e that continues to belong to another species. Specimens from on the peristome. The perisstome is raised the Great Barrier Reefand Queensland show into a blunt or bifid point on its side. Facing some similarities with C. sibogae, but be- into the peristomial sinus is a vertically po- cause those we examined lacked interzooecial sitioned ovoid adventitious anvicularium with avicularia, positive identification was im- a serrated rostrum. Other ad,ventitious avic- possible. Specimens in our collection from ularia with blunt triangular mandibles and Moreton Bay, Queensland, have small zooids, raised rostra develop from some marginal but have interzooecial avicularia more sim- pores (fig. 75). Large interzooecial avicularia ilar to Bass Strait C. fusca. Thus, three (or are scattered over the colon) . These have a more) species may be involved, all pigment- spatulate mandible and a rraised, smooth, ed, all with small triangular avicularia in curved rostrum with a thicl( cross-bar (fig. marginal positions, and all with nonsinuate 78). No ovicells were found in our specimens. primary orifices. ADDmONAL SPECIMENS ExMAmNED: (BMNH) 1963.9.8.13 pt. 71C-1, 71C-9, 71C-2, Siboga Measurements Station 99, N. Ubian, Sulu Archipelago, 16- Range Mean N 23 m. 1963.9.8.12, 71C-3, same station, fig- Lz 0.378-0.594 0.488 15 ured specimen, Harmer (1957), pl. XLIII, fig. Wz 0.360-0.576 Lo 0.108-0.162 0.140 15 2. 1963.9.8.10, 472M, same station. 1963. Wo 0.126-0.180 0.168 15 9.8.16, 472K-2, same station. 1963.9.8.17, Lov 0.126-0.162 0.144 6 466A, Siboga Station 91, Muaras Reef, E. Wov 0.180-0.270 6 0-54 m. Small avicularium 0*228 Borneo, 1963.9.8.15, 376G, Siboga Lav 0.054-0.108 0.094 15 Station 213, Saleyer, S. of Celebes, 0-36 m. Wav 0.036-0.072 0.054 15 1963.9.8.28, Torres Strait, 15-20 fm, Kirk- Interzooecial avicularium patrick, reg. May 26, 1900. 1963.9.8.23, Port Lavz 0.252-0.360 0.318 3 Denison, Queensland, Busk, October 23, Wavz 0.198-0.270 0.234 3 1899. 32.4.20.84 pt., Great Barrier ReefExp. Lav 0.180-0.324 Wav 0.108 0.208 3 Station XXII, Hastings. 99.5.1.1316, Bass Strait, Hincks Collection. OCCURRENCE: Komodo, Sanur, and Lom- DISCUSSION: Harmer (1957) placed a large bok. group ofpigmented Celleporaria from the Si- DiSTRIBUTION: Indonesia. boga and other collections in Busk's (1854) species, Celleporaria fusca, which was described from Bass Strait. There appears to be Celleporaria aperta (Hincks), 1882 no type material for this species. However, Figures 79-84 the illustration (fig. 9.17d) of Bass Strait Cel- Schizoporella aperta Hincks, 1882, p. 126. leporariafusca in Bock (1982) shows a species Mucronella serratimargo Ortmann, 1890, p. 48. with interzooecial avicularia raised on a con- Schismopora cucullata Maplestone, 1905, p. 389. ical rostrum and having a rostrum with a Holoporella aperta Waters, 1909, p. 161. Living- striking serrated edge. A British Museum stone, 1926, p. 97. specimen from Bass Strait (99.5.1.1316, ?Holoporella discoidea Canu and Bassler, 1929, p. Hincks Collection) appears to be the same. 426. In addition to the very different form of the Celleporaria aperta Harmer, 1957, p. 673, in part. interzooecial avicularia, it also had consid- Cook, 1968, p. 175. erably larger zooids. In view ofthis evidence DESCRIPTION: Colonies are encrusting on we believe that Harmer's specimens cannot coral. Zooids are ovoid, becoming erect in belong to Busk's species. Examination of frontally budded regions. The surface is cov- specimens available at the British Museum ered with irregularly granular calcification and 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 33

I1 f -h;b.10 i -) >'' 'i 'i

183

FIGS. 79-84. 79. Celleporaria aperta, Komodo (AMNH 620). Scale = 200 gm. 80. C. aperta. Scale = 400 MAm. 81. C. aperta, close-up of primary orifice. Scale = 100 ,im. 82. C. aperta, close-up of rostral avicularium. Scale = 20 im. 83. C. aperta, showing triangular interzooecial avicularium. Scale = 200 ,um. 84. C. aperta, another triangular avicularium. Scale = 100 Am. perforated by marginal and occasional frontal orifice is semicircular with a small, rounded pores. There is no high peristome, and the proximal sinus. There are two to four long- shape of the orifice may be easily seen. The jointed oral spines, though these are often 34 AMERICAN MUSEUM NOVITATES NO. 2847 broken, and even their bas es may be ob- Jedan, Aru Island, 13 m, figured specimen, scured by secondary calcificaltion. Small ad- Harmer (1957) pl. XLII, fig. 11. 1936.12. ventitious avicularia are locaited on the side 30.125A, India, Thornely. 1963.3.10.19, of the small conical rostrum, just below the Singapore, Hannitsh Collection. 1963.2. orificial sinus. This aviculariuim is ovoid and 12.84, Panama, off Colon, Dundee Collec- its rostrum has a scalloped (edge. Large in- tion. 1963.3.10.18, Skeat, Malaya, reg. Nov. terzooecial avicularia are comimon. They have 30, 1899. Celleporaria tridenticulata: 1963. a subtriangular mandible andl a serrated ros- 3.10.17, Torres Strait, figured specimen, tral margin. Ovicells are capllike and imper- Harmer (1957), pl. XLII, fig. 12. 1963.3. forate, with a curved opening above the zooid 10.24, 336B, Siboga Station 37, Sailus Ketjil, orifice. Paternoster Island, 0-27 m. 1963.3.10.21, 1963.3.10.22, 385C-2, Borneo Bank, Strait Measurements ofMakassar, 59 m. Celleporaria sp. (3): 1963. Range Mean N 3.10.16, 295F, Siboga Station 273, Jedan, Lz 0.270-0.540 0.413 15 Aru Island, 13 m. 1962.2.20.9, China Sea, Wz 0.216-0.468 0.317 15 1 19035'E, 23032'N, Professor Ma. Other Cel- Lo 0.090-0.126 Wo 0.108-0.144 0.134 15 leporaria species: 26.9.6.151,26.9.6.154, Suez Lov 0.180 0.180 1 Canal, Hastings. 1963.3.10.20, Japan, Uraga Wov 0.090 0.090 1 Channel, off Tokyo, 150 fm, reg. June 23, Oral avicularium 1902. Lav 0.036-0.090 0.065 15 OCCURRENCE: Komodo. Wav 0.036-0.054 0.038 15 DISTRIBUTION: Singapore, Philippines, Red Interzooecial avicularium Lavz 0.270-0.540 0.414 15 Sea, Indonesia, W. Africa, Cape Verde Is- Wavz 0.144-0.360 0.258 15 lands, Caribbean. Lav 0.180-0.458 0.311 15 Wav 0.090-0.162 0.140 15 FAMILY CLEIDOCHASMATIDAE DISCUSSION: Our specimens agree best with CHEETHAM AND SANDBERG, 1964 that illustrated in Harmer's (1957) plate XLII, Genus Cleidochasma Harmer, 1957 figure 11. Harmer (1957) noted that the orifice in his specimens sometimes has three Cleidochasma porcellanum (Busk), 1860 denticles. However, upon reexamination of Figures 85-87 some of the material he cited, it became ap- Lepralia porcellanum Busk, 1860, p. 283. parent that he had combined several species Lepralia cleidostoma Smitt, 1873, p. 62. under the name C. aperta, including colonies Hippoporina cleidostoma Canu and Bassler, 1928a, of Celleporaria tridenticulata. The specimen p. 80; 1928b, p. 104. from Torres Strait belongs to C. tridenticu- Hippoporina porcellana Hastings, 1930, p. 721. lata, as do the Siboga specimens from Pa- Marcus, 1937, p. 96. Osburn, 1940, p. 422; 1952, ternoster Island and Borneo Bank. Some Si- p. 344. Shier, 1964, p. 633. boga specimens do belong to C. aperta (see Cleidochasma porcellanum Cheetham and Sand- below); another group appears to belong to a berg, 1964, p. 1032. Cook, 1964b, p. 11; 1968, third species which has a tridenticulate orifice p. 197. Long and Rucker, 1970, p. 19. Powell, 1971, p. 771. Winston, 1982, p. 148; 1984, p. and low pointed avicularia. Other specimens 28. Harmer lists under C. aperta, e.g., those from Japan and the Suez Canal, appear to belong DESCRIPrION: Colony is encrusting on a to entirely different species of Celleporaria. piece ofcoral. Zooids are rectangular in shape. ADDmIoNAL SPECIMENS EXAMiNED: (BM7NH) There are three to five marginal pores on a Celleporaria aperta: 97.5.1.720, Singapore or zooid; the otherwise imperforate frontal sur- Philippines, type specimen. 1963.3.10.25, face is somewhat granular. A thin line ofcal- 360I, Siboga Station 240, Banda, Banda Sea, cification marks the boundary of each zooid. 0-45 m. 1963.3.10.26, 322J, Siboga Station An umbo is present at the base ofthe orifice. 50, Badjo Bay, W. Flores, 0-40 m. The orifice is keyhole shaped, with proxi- 1963.3.10.15 pt. 298D-2, Siboga Station 273, mally directed condyles. One avicularium is 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 35

IW 'L.4 a .' w i

i. a, a A _w

- 87' a

.s ^i

I.,

V.7-. ~~MRI_ q-qTWA S te.. oUft.I FIGS. 85-90. 85. Cleidochasma porcellanum, Komodo (AMNH 621), ovicelled zooids. Scale = 100 ,um. 86. C. porcellanum. Scale = 200 ,um. 87. C. porcellanum, close-up of ovicell plate. Scale = 40 ,um. 88. C. peristomarium, Komodo (AMNH 622). Scale = 200 ,um. 89. C. peristomarium. Scale = 200 ,um. 90. C. peristomarium, close-up of orifice. Scale = 100 ,m. present on each zooid, usually oriented dis- are helmet-shaped, deeply immersed, and tolaterally. The avicularium is located at the imperforate. At the opening of the ovicell is base of or slightly below the orifice. Ovicells a calcified plate, with slitlike openings into 36 AMERICAN MUSEUM NOVITATES NO. 2847

. 93 -11.

'IN.-,-K I, w s I _ sv~~~~~~ I Is. a FIGS. 91-94. 91. Rhynchozoon verruculatum, Komodo (AMNH 623), zooids near growing edge. Scale = 200 gm. 92. R. verruculatum, zooids from inner region of colony. Scale = 200 ,m. 93. R. verruculatum, close-up of orifice. Scale = 20 ,um. 94. R. verruculatum, showing oral avicularium. Scale = 40 ,um.

the ovicell at the top and bottom and at- Cleidochasma peristomarium tached to the ovicell at each side. (Canu and Bassler), 1929 Figures 88-90 Measurements Range Mean N Gemelliporaperistomaria Canu and Bassler, 1929, Lz 0.234-0.450 0.344 10 p. 310. Wz 0.252-0.360 0.302 10 Cleidochasma peristomarium Harmer, 1957, p. Lo 0.108-0.144 0.130 10 1047. Wo 0.090-0.126 0.106 10 Lov 0.162-0.198 0.180 6 DESCRIPTION: The colonies were found en- Wov 0.216-0.234 0.228 6 LaV 0.072-0.108 0.079 10 crusting on a piece of shell. The zooids are WaV 0.036-0.073 0.047 10 rhomboidal, their surfaces granular and imperforate except for a row of marginal pores. OCCURRENCE: One colony was found at A thin line of calcification marks each zooid Komodo. boundary. The orifice is keyhole shaped. A DISTRIBUTION: Florida, Gulf of Mexico, raised peristome flares out on each side of Caribbean, Bermuda, Madeira, Brazil, Co- the orifice. The avicularia are located below lombia, Panama, Galapagos Islands, Ceylon, the orifice and directed transversely inward. Philippines, West Africa, Indonesia, Red Sea. They are almost oval in shape with a cross- 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 37

wtAb 97 4'.

I. 4W.

4"L .1

FIGS. 95-98. 95. Rhynchozoon rostratum, Bali (AMNH 624), zooids at growing edge of colony. Scale = 100 Am. 96. R. rostratum. Scale = 200um. 97. R. rostratum, close-up of orifice. Scale = 20,m. 98. R. rostratum, close-up of oral aviculanum and developing ovicell. Scale = 40 ,um. bar in the middle. Five to six pores perforate Genus Rhynchozoon Hincks, 1895 the elongated proximal portion of the avicularium. The ovicells are globose and per- Rhynchozoon verruculatum (Smitt), 1873 forated, their surfaces beaded and heavily Figures 91-94 calcified. Cellepora verruculata Smitt, 1873, p. 50. Osburn, 1914, p. 214. Measurementss Escharoidea verruculata Busk, 1884, p. 150. Range Mean N Rhynchozoon verruculatum Canu and Bassler, LZ 0.378-0.504 0.439 10 1928a, p. 88. Hastings, 1930, p. 728. Marcus, Wz 0.324-0.450 0.376 10 1939, p. 153. Osburn, 1940, p. 444. Winston, Lo 0.108-0.144 0.128 10 1984, p. 31. Wo 0.108-0.162 0.119A .1 in 10,,\ , . Lov 0.180-0.216 0.196 9 DESCRIPTION: Colonies encrusting and Wov 0.144-0.270 0.236 9 Lav 0.126-0.162 0.135 10 multilaminar. Zooids at the growing edge are Wav 0.090 0.090 10 subrectangular in shape; in the older regions ofthe colony they become irregular. There is OCCURRENCE: Two colonies were found at a series of marginal pores on each zooid and Komodo. additional pores in older zooids. The frontal DiSTRIBUTION: Indonesia, Philippines. surface of the zooid is smooth. The orifice is 38 AMERICAN MUSEUM NOVITATES NO. 2847 semicircular distally, with a beaded margin, rected. The front of the ovicell is covered by and shallowly U-shaped proximally, with a a subtriangular to oval plate and is buried in plain margin. There is a peristome around secondary calcification. However, the bumps the orifice with a proximolateral avicularium caused by the ovicells are visible on the fron- bordering an asymmetrically placed second- tal surface; thus the ovicells are more prom- ary sinus. In older sections of the colony the inent than those of many Rhynchozoon calcification increases greatly. The tubercles species. around the orifice are more pronounced and the peristome becomes immersed in the fron- Measurements tal shield. An elongate diamond-shaped avic- Range Mean N ularium is located on almost every zooid in Lz 0.342-0.558 0.472 15 Wz 0.270-0.396 0.336 15 a median-lateral position. The avicularia Lo 0.108-0.144 0.130 15 point in either a distolateral or lateral direc- Wo 0.108-0.180 0.138 15 tion. Ovicells are semicircular and are Lov 0.108-0.162 0.134 9 embedded in the secondary calcification, so Wov 0.171-0.234 0.202 9 that the Suboral avicularium only ovicell plate can be seen within Lav 0.054-0.144 0.110 15 the peristome (fig. 91). Wav 0.036-0.072 0.047 15 Measurements Adventitious avicularium Range Mean N Lav 0.072-0.126 0.106 10 WaV 0.036-0.072 0.054 10 Lz 0.306-0.450 0.365 15 Wz 0.216-0.342 0.276 15 Lo 0.090-0.126 0.106 15 DiSCUSSION: Osburn (1952) synonymized Wo 0.090-0.144 0.118 15 Lov 0.090-0.144 Smitt's Cellepora verruculata (1873) and Wov 0.126-0.216 0.162 5 Busk's Lepralia rostrata (1855). Study ofCa- Suboral avicularium ribbean and Indonesian specimens has shown Lav 0.072-0.126 0.095 11 that, in fact, these are two distinct species, Wav 0.036-0.054 0.044 11 which may co-occur. They are superficially Adventitious avicularium quite similar, and, like most species ofRhyn- Lav 0.090-0.216 0.154 13 chozoon, exhibit marked astogenetic and on- Wav 0.036-0.072 0.051 13 togenetic differences in zooid morphology. OCCURRENCE: Komodo. Zooids of R. verruculatum are somewhat DISTRIBUTION: Indonesia, Indian Ocean, larger than those ofR. rostratum. In addition Caribbean, Florida to Brazil. to size, the most useful characters for distin- guishing the two are the shape ofthe primary Rhynchozoon rostratum ((Busk), 1855 orifice (the sinus is deeper and narrower in R. rostratum), the shape of the frontal avic- Lepralia rostrata Busk, 1855, p. 4; 1856, p. 178. ularia (less elongate in R. rostratum), and the Rhynchozoon rostratum Hastings, 1930, p. 728. nature ofthe ovicell, which is completely im- Soule, 1961, p. 43. Soule an(d Soule, 1964, p. mersed in the frontal surface in R. verrucu- 33. Long and Rucker, 1970, I . 178. latum, but slightly protruding in R. rostra- DESCRIPTION: Colonies are encrusting. tum. Zooids are hexagonal. The zooid surface is ADDmONAL SPECIMENS ExAmiNED: (BMNH) heavily calcified and divided by a line ofcal- 92.9.6.10,92.9.6.9,99.7.1.2237, 99.7.1.1844, cification. Zooids in older regions ofcolonies 99.7.1.1845, 99.7.1.1846, 99.7.1.1847, all develop prominent peristomial knobs and Mazatlan, Busk Collection, paratype mate- processes. The primary orifice has a beaded rial. 29.4.26.169 pt., 29.4.26.229 pt. (two margin distally and a deep, narrow, rounded slides), Gorgona 3, St. George Collection, sinus proximally. A distolaterally suborally Hastings. 1961.11.2.6, Campeche Bank, Gulf directed avicularium is located in the peri- of Mexico, off Alacran Reef, Deep Water stome. This avicularium, conspicuous in ear- #194, Coll. L. Komicker, det. A. H. Chee- ly ontogeny, has a curved triangular rostrum tham. and a moderately developed uncinate pro- OCCURRENCE: Sanur. cess. Adventitious avicularia are diamond DISTRIBUTION: Indonesia, E. Pacific, Ca- shaped, often paired, and distolaterally di- ribbean, Gulf of Mexico. 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 39

im *.__

FIGS. 99-102. 99. Rhynchozoon detectum, Komodo (AMNH 625), zooids at growing edge ofcolony. Scale = 400 ,um. 100. R. detectum, zooids at angle to show length of the rostrum. Scale = 400,Am. 101. R. detectum, orificial avicularia. Scale = 100 Mum. 102. R. detectum, close-up of orifice. Scale = 40 Am.

Rhynchozoon detectum Harmer, 1957 ed base. One is located suborally at the base Figures 99-102 of the rostrum. It has a sharp, pointed uncinate process. The other avicularium is lon- Rhynchozoon crenulatum Philipps, 1899, pp. 440, ger and narrower. It is located medially on 448 (not Rhynchopora crenulata Waters, 1887, the frontal wall of the zooid and directed P. 195). proximally. No ovicells were present on our Rhynchozoon detectum Harmer, 1957, p. 1067. specimens. According to Harmer, ovicells are DESCRIPTION: The colony is encrusting on large, with a finely tuberculate frontal plate. coral. The zooids' rectangular shape can be Measurements seen clearly at the growing edge, while it is Range Mean N obscured in the older portions of the colony. Lz 0.324-0.486 0.407 15 The zooid surface is convex and has a row Wz 0.198-0.342 0.258 15 ofmarginal pores. The orifice is semicircular Lo 0.090-0.144 0.101 15 and opens into a very broad sinus. The inner Wo 0.090-0.198 0.115 15 distal margin of the orifice is beaded. A long Suboral avicularia Lav 0.072-0.126 0.097 15 blunt or bifid-ended rostrum is present below Wav 0.054-0.072 0.061 15 the orifice opening. There are two types of Longer avicularia avicularia which differ in size and location. Lav 0.162-0.252 0.209 15 Both have a triangular mandible and a round- Wav 0.054-0.072 0.063 15 40 AMERICAN MUSEUM NOVITATES NO. 2847

_4 Aprb ..It 1. 11 ,r

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FIGS. 103-106. 103. Drepanophora verrucosa, Lombok (AMNH 626, holotype), primary orifice and peristomial avicularium in zooids at growing edge. Scale = 100 ,um. 104. D. verrucosa, showing uncinate process in peristome, and developing ovicells. Scale = 100 ,um. 105. D. verrucosa, ovicelled zooids. Scale = 200 Am. 106. D. verrucosa, close-up of orifice and avicularium. Scale = 40 ,m.

ADDmONAL SPECIMENS ExAmINED: (BMNH) peristome with a subcircular opening and a 483C, 483C-1, 483C-2, 483C-3, Siboga Sta- hooded, immersed ovicell lacking the typical tion 117, Kwandang Bay, N. Celebes, 80 m. lateral pores. Harmer's list no. 1871, Torres Strait, Had- HOLOTYPE: AMNH 626. don Coll., reg. Feb. 24, 1898, 189. Rhyncho- PARATYPE: AMNH 627. zoon crenulatum, Lifu, Loyalty Islands, Phil- ETYMOLOGY: The species name is derived ipps (Willey Collection), reg. Mar. 1, 1898. from the Latin verrucosus, meaning "full of OCCURRENCE: Komodo. warts," because of the rough, tuberculate DISTRIBUTION: Lifu, Loyalty Is., Indonesia, zooid surface. Torres Strait. DESCRIPTION: Colonies are encrusting on shell. Zooids are small and ovoid in shape. The frontal wall is convex; it and the tubular Genus Drepanophora Harmer, 1957 peristome become thickened, roughened, and Drepanophora verrucosa, new species covered by small tubercles. Five or six large Figures 103-106 areolae are found at zooid margins. The secondary orifice is subcircular. A sharp-beaked DIAGNOSIS: Drepanophora with rough peb- triangular avicularium is located inside the bled texture to frontal surface, short tubular peristome on a lateroproximal wall. Hood- 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 41 shaped ovicells are embcPdded in the DISCUSSION calcification of the distal zooiid. Ovicells are covered with the same thick tubercular cal- Thirty-three species of cheilostome bryo- cification as zooids; they have a curved rim zoans were found in these collections: 13 which shows both calcified layers. In our species occurred at Sanur, Bali, 6 species were specimens, which had heavy secondary cal- found at Lombok, and 23 were found at Ko- cification, the two lateral pores characteristic modo (table 1). Although Wallace's Line of Drepanophora were not apiparent. Ovicell passes between Bali and Lombok, there is no pores seemed to be identical with the prox- evidence of any influence on the bryozoan imal areolae ofthe distal zooidI andwere often distributions. This probably reflects the fact completely occluded. that Lombok Strait, between Bali and the Lesser Sunda chain, is not particularly deep. Measurements In view ofthe various changes in sea level in Range Mean N the region during the last 3 million years Lz 0.306-0.450 0.387 10 (Potts, 1983), Lombok Strait does not rep- Wz 0.252-0.306 0.279 10 resent a barrier to most marine Lo 0.090-0.144 1 organisms. Wo 0.108-0.144 0.1 10 The differences between the three sites can Lov 0.090-0.126 0.108 10 be better explained ecologically. The back- Wov 0.144-0.216 0.180 10 reef area at Lombok was the shallowest site, Lav 0.036-0.072 0.054 10 Wav 0.036-0.054 0.040 10 ranging from 1 to 1.5 m, and may have re- ceived considerable subaerial exposure at DISCUSSION: Examination 4of British Mu- times. Only six species were collected there. seum specimens of the thre.e Indo-Pacific Four of them were also found in at least one species described by Harmer (1957) showed ofthe other two sites; the other two are known that the specimen found in our collection did from elsewhere in the region. Although low, not correspond to any ofthem.. It is somewhat this degree ofdiversity is probably higher than similar to Drepanophora ttiberculatum, a for equivalent depths on Caribbean reefs. For species described by Osbum ( 1914), from the example, at Carrie Bow Cay in Belize, only Tortugas. However, zooids ofFDrepanophora one species, Trematooecia aviculifera, oc- tuberculatum are slightly largevr, with numer- curred in water less than 1 m in depth, while ous marginal areolae. In additiion, the peristo- only three species, Trematooecia aviculifera, mial avicularium is smaller;and is oriented Hippopodina feegeensis, and Steginoporella frontally, rather than transve: rsely, while the sp. were found in 1-2 m depths (Winston, ovicells are initially globose, Mvith distinct lat- 1984). Hippopodinafeegeensis was also found eral pores, although in older regions of col- at Lombok (and at Bali and Komodo); it ap- onies they may become increasingly embed- pears to be one of the most successful reef- ded in secondary calcificatior dwelling species worldwide (Winston, 1986). ADDMONAL SPECIMENS EXAM 4INED: (BMNH) The most characteristic Caribbean shallow Drepanophora incisor. 06.1 i2.3.13, 1969.1. water species, Trematooecia aviculifera, does 10.1, Ceylon, Thornely, 19)05, paratypes. not occur in the Indo-Pacific. Its ecological 1936.12.30.53, Ceylon, Thornely, 1935, 53, replacement may be Stylopoma viride, a specimen illustrated in Harn mer (1957), pl. species which, like Trematooecia, has mas- LXX, figs. 26-28. Drepanop)hora corrugata: sive well-calcified frontally budding colonies 1969.1.10.2, 348M, Siboga S tation 213, Sa- which appear to be less cryptic than those of layer, S. ofCelebes, 0-36 m, sypecimen figured most reef cheilostomes. in Harmer (1957), text fig. 1 14. 06.12.3.12, At Sanur, in depths between 2 and 3 m, 13 Ceylon, Thomely, paratype. 1 936.12.30.52D, species were collected. At Komodo, in 1.5 to Ceylon, Thomely. 1938.2.22.2, Maldive Is- 2 m depths, 23 species were collected. This lands (note-this does not aLppear to be D. diversity is also higher than for equivalent corrugata). Drepanophora lotngiuscula: 99.7. depths in the Caribbean. 1.2012, Bay of Suez, 7 fm, B'usk, holotype. Five of the 33 species reported are new. OCCURRENCE: Two colonie,s were collected Four species, Watersipora edmondsoni, Par- at Lombok. asmittina hastingsae, Rhynchozoon verrucu- 42 AMERICAN MUSEUM NOVITATES NO. 2847

TABLE 1 Distributions of Cheilostome Bryozoans Found at the Three Sites Studied Lom- Komo- Indo-W. Red E. Carib.- E. Bali bok do Pac. Sea Atl. W. Atl. Pac. Austra. 1. Cranosina coronata x x x x x 2. Parellisina curvirostris x x x x x 3. Caulibugula mortenseni x x x 4. Scrupocellaria spatulata x x x 5. Thalamoporella stapifera x x x x 6. Thalamoporella komodoensis* x 7. Steginoporella buskii x x x x x x 8. Smittipora harmeriana x x 9. Cribrilaria flabellifera x x x x 10. Exechonella brasiliensis x x x 11. Coleopora verrucosa x x 12. Hippopodina feegeensis x x x x x x x x x 13. Petraliella chuakensis x x 14. Watersipora "subovoidea" x x 15. Watersipora edmondsoni x x x 16. Calyptotheca wasinensis x x x 17. Schizomavella inclusa x x x 18. Stylopoma duboisii x x x x x x 19. Stylopoma viride x x x x x 20. Parasmittina signata x x x x x x x 21. Parasmittina parsevalii x x x x x x 22. Parasmittina hastingsae x x x 23. Crepidacantha carsioseta* x 24. Fenestrulina harmeri* x x 25. Margaretta triplex x x x 26. Celleporaria sibogae* x x x x 27. Celleporaria aperta x x x x x 28. Cleidochasma porcellanum x x x x x x 29. Cleidochasma peristomarium x x 30. Rhynchozoon verruculatum x x x 31. Rhynchozoon rostratum x x x x

32. Rhynchozoon detectum x x x 33. Drepanophora verrucosa* _ x 13 5 25 30 10 8 1 1 9 10 * = new species. latum, and Rhynchozoon rostratum are re- and 33 percent from the Caribbean. These ported for the first time from Indonesia; the results are in agreement with Jackson et al.'s first two were previously known only from (1985) finding that about one-third of Carib- Hawaii, the latter two from the Caribbean. bean reef-dwelling cheilostome species are As table 1 shows, many of the other species also known from the Indo-West-Pacific, and have broad distributions: 91 percent occur provide further indication that ecological tol- elsewhere in the Indo-West-Pacific region, 30 erances of reef-dwelling cheilostome species percent reach the Red Sea, and 30 percent are generally much greater than those oftheir occur in Australia, primarily on the Great coral substrata. This finding does not mean Barrier Reef. Almost a quarter of the species that reef-dwelling cheilostomes can never be listed have also been reported from the east- useful in biogeographic studies. It does mean ern Atlantic (Mediterranean, West Africa), that we need to be able to determine which and 28 percent are reported from the eastern species occupy a narrow ecological niche and Pacific (Hawaii, Galapagos, Baja California), which species a broad one, something that 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 43 can only be accomplished through careful lection of the British Museum, pt. 2. ecological and taxonomic studies of addi- London, British Mus., pp. 55-120. tional bryozoan faunas. 1855. Class Bryozoa. In P. P. Carpenter, Cat- alogue of the Reigen Collection of Ma- zatlan Mollusca in the British Museum. ACKNOWLEDGMENTS Warrington, pp. 1-6. We are most grateful to Miss Patricia L. 1856. Zoophytology. Quart. Jour. Micros. Soc., Cook of the British Museum (Natural His- vol. 4, pp. 176-180. tory) for making accessible to us the Siboga 1860. Zoophytology: catalogue ofthe Polyzoa and other specimens used by Harmer that are collected by J. Y. Johnson, Esq., at Ma- presently in the collections ofthe British Mu- deira, in the years 1859 & 1860, with seum. We would also like to thank Miss Cook descriptions ofnew species. Quart. Jour. Micros. Soc., vol. 8, pp. 280-285. and Dr. Alan H. Cheetham ofthe Smithson- 1884. Report on the Polyzoa collected by ian Institution for carefully reviewing and H.M.S. Challenger during the years criticizing the manuscript; errors that remain 1873-1876. Report on the scientific re- are our own. sults of the voyage of H.M.S. Chal- lenger during the years 1873-1876. LITERATURE CITED London, Zoology, vol. 10, pt. 30, pp. 1- Audouin, J. V. 216. 1826. Explication sommaire des planches de Canu, F., and R. S. Bassler polypes de l'Egypte et de la Syrie, pu- 1917. A synopsis of American Early Tertiary bliees par Jules-Cesar Savigny. Pp. 225- cheilostome Bryozoa. Bull. U.S. Natl. 244 in "Description de l'Egypte, ou re- Mus., Bull. 96, pp. 1-87. cueil des observations et des recherches 1924. Contribution et I'etude des Bryozoaires qui ont ete feutes en Egypte pendant d'Autriche et de Hongrie. Bull. Soc. l'Expedition de l'Armee francaise ... Geol. Fr., ser. 4, vol. 24, pp. 672-690. Histoire Naturelle," tome 1, pt. 4. Paris, 1927. Classification of the cheilostomatous Imprimerie Imperiale, 339 pp. Bryozoa. Proc. U.S. Natl. Mus., vol. 64, 1828. Ibid., Ed. 2, vol. 23. pp. 1-42. Banta, W. C., and R. J. M. Carson 1 928a. Bryozoaires du Bresil. Bulletin de la So- 1977. Bryozoa from Costa Rica. Pacific Sci., ciete des Sciences de Seine-e-Oise, Tome vol. 31, no. 4, pp. 381-424. 9, Fasc. 5, pp. 58-100, 9 plates. Bassler, R. S. 1928b. Fossil and Recent Bryozoa of the Gulf 1953. Bryozoa. In R. C. Moore (ed.), Treatise of Mexico region. Proc. U.S. Natl. 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1968. Bryozoa (Polyzoa) from the coasts of 1932. The Polyzoa, with a note on an asso- tropical west Africa. Atlantide Rept. no. ciated hydroid. Great Barrier Reef Ex- 10,pp. 115-262. pedition, 1928-1929, Scientific Re- Gordon, D. P. ports, vol. 4, pp. 399-458. 1984. The marine fauna of New Zealand: Haswell, W. A. Bryozoa: Gymnolaemata from the Ker- 1880. On some Polyzoa from the Queensland madec Trench. New Zealand Oceanogr. coast. Proc. Linn. Soc. New South Wales, Inst. Mem. 91, pp. 1-198. vol. 5, pp. 33-44. Gray, J. E. Hayward, P. J., and J. S. Ryland 1843. Additional radiated animals and An- 1979. British ascophoran bryozoans. London, nelides. In E. Dieffenbach, Travels in Academic Press, 312 New Zealand. London, John Murray, pp. vol. 2, Appendix, pp. 292-295. Hincks, H. 1848. List ofthe specimens ofBritish animals 1862. Catalogue of the zoophytes of South in the collection ofthe British Museum. Devon and Cornwall. Ann. Mag. Nat. Part 1. Centroniae or radiated animals. Hist., ser. 3, vol. 9, pp. 200-207. London, Trustees of the British Mu- 1879. On the classification ofthe British Poly- seum (Natural History), 173 pp. [Poly- zoa. Ann. Mag. Nat. Hist., ser. 5, vol. zoa, pp. 91-151]. 3, pp. 153-164. Harmelin, J. G. 1880. A history ofthe British marine Polyzoa. 1970. Les Cribrilaria (Bryozoaires Chilo- London, Van Voorst, 2 vols. stomes) des Mediterranee; systematique 1881. Contributions towards a general history et ecologique. Cah. Biol. Marine, vol. of the marine Poloyzoa. IV. Foreign 11, pp. 77-98. Membraniporina. Ann. Mag. Nat. Hist., 1978. Sur quelques Cribrimorphes (Bryozoa, ser. 5, vol. 7, pp. 147-161. Cheilostomata) de l'Atlantique oriental. 1882. Contributions towards a general history Tethys, vol. 8, no. 2, 1976 (1978), pp. of marine Polyzoa. IX. Foreign Chei- 173-192. lostomata (miscellaneous). Ann. Mag. Harmer, S. F. Nat. Hist., ser. 5, vol. 9, pp. 116-127. 1900. A revision ofthe genus Steganoporella. 1884. Contributions towards a general history Quart. Jour. Micros. Sci., vol. 43, pp. of the marine Polyzoa. XII. Polyzoa 225-297. from India (coast of Burmah). Ann. 1915. The Polyzoa of the Siboga Expedition. Mag. Nat. Hist., ser. 5, vol. 13, pp. 356- Part I. Entoprocta, Ctenostomata and 362. Cyclostomata, vol. 28a, pp. 1-180. 1887a. Critical notes on the Polyzoa. Ann. Mag. 1923. On cellularine and other Polyzoa. J. Nat. Hist., ser. 5., vol. 19, pp. 150-164. Linn. Soc., Zool., vol. 35, pp. 293-361. 1887b. On the Polyzoa and Hydroidea ofMer- 1926. The Polyzoa of the Siboga Expedition. gui Archipelago. J. Linn. Soc. (Zool.), Part II. Cheilostomata, Anasca. Siboga- vol. 22, pp. 121-135. Exped., vol. 28b, pp. 181-501. 1895. Contributions towards a general history 1957. The Polyzoa of the Siboga Expedition. ofthe marine Polyzoa. Index. Published Part 4. Cheilostomata, Ascophora II. privately (? for Hincks) by Ann. Mag. Siboga-Exped., vol. 28b, pp.641-1147. Nat. Hist., London, pp. i-vi. Hassall, A. H. Jackson, J. B. C. J., J. E. Winston, and A. G. Coates 1842. Remarks on the genus Lepralia of Dr. 1985. Niche breadth, geographic range, and Johnston with descriptions of six un- extinction of Caribbean reef-associated described species. Ann. Mag. Nat. Hist., cheilostome Bryozoa and Scleractinia. vol. 9, pp. 409-414. In Proc. Fifth Internat. Coral Reef Hastings, A. B. Symposium, Tahiti, vol.4, pp. 151-158. 1927. Zoological results ofthe Cambridge Ex- Jullien J. pedition to the Suez Canal, 1924, XX, 1882. Note sur une nouvelle division des Bry- Report on the Polyzoa. Trans. Zool. Soc. ozoaires Cheilostomiens. Bull. Soc. Zool. London, vol. 22, pp. 321-354. Fr., vol. 6, pp. 271-285. 1930. Cheilostomatous Polyzoa from the vi- 1883. Dragages du Travailleur. Bryozoaires; cinity ofthe Panama Canal collected by especes draguees dans l'ocean Atlan- Dr. C. Crossland on the cruise of the S. tique en 1881. Bull. Soc. Zool. Fr., vol. Y. "St. George." Zool. Soc. London 7, pp. 497-529. Proc., 1929, pp. 670-740. 1888. In Bryozoaires, Mission scientifique du 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 45

Cap Horn, 1882-1883. Paris, vol. 6, Victoria. Trans. Roy. Soc. Victoria, vol. Zoologie, pt. 3, pp. 1-92. 4, pp. 1-166. Kirkpatrick, R. Maplestone, C. M. 1888. Polyzoa of Mauritius. Ann. Mag. Nat. 1905. Lord Howe Island Polyzoa. Proc. Roy. Hist., ser. 6, vol. 1, pp. 72-85. Soc. Victoria, n.s., vol. 17, pp. 386-390. 1890a. Report upon the Hydrozoa and Polyzoa Marcus, E. collected by P. W. Bassett-Smith, Esq., 1921a. Results ofDr. E. Mjobergs Swedish Sci- Surgeon R.N., during the survey of the entific Expeditions to Australia 1910- Tizard and Macclesfield Banks in the 13. XXIV. Bryozoen. K. Svenska Ve- China Sea, by H.M.S. "Rambler," tensk.-Akad. Handl., vol. 61, pp. 1-34. Commander W. U. Moore. Ann. Mag. 1921b. Indo-Pacifische Bryozoen Aus Dem Nat. Hist., ser. 6, vol. 5, pp. 11-24. Riksmuseum in Stockholm. Ark. Zool., 1890b. Reports on the zoological collection vol. 14, pp. 1-23. made in Torres Straits by Professor A. 1922. Bryozoen von den Aru-Inseln. Abh. C. Haddon, 1888-1889. Hydroidea and Senckenbergisch. Naturforsch. Ge- Polyzoa. Roy. Dublin Soc. Sci. Proc., sellsch., vol. 35, pp. 421-446. n.s., vol. 6, pp. 603-626. 1925a. Bryozoa. In P. Schulze (ed.), Biol. Tiere Lagaaij, R. Deutschlands, pt. 47 (Lief. 14), pp. 1- 1963. New additions to the bryozoan fauna of 46. Berlin. the Gulf of Mexico. Publ. Inst. Marine 1925b. Uber Stirpariella mortenseni und das Sci., Univ. Texas, vol. 9, pp. 162-236. Genus Stirpariella-papers from Dr. Th. Lamouroux, J. V. F. Mortensen's Pacific Expedition, 1914- 1821. Exposition methodique des genres de 1916, XXII. Vidensk. Medd. Dansk Nat. l'orde des polypiers, avec leur descrip- For., vol. 81, pp. 37-55. tion et celle des principales especes, fi- 1937. Bryozoarios marinhos brasileiros, 1. Bol. gurees dans 84 planches; les 63 pre- da Faculdade de Filosofia, Ciencias e mieres appartenant la l'Histoire Letras, Univ. Sao Paulo, vol. 1, Zool- naturalle des Zoophytes d'Ellis et So- ogia, no. 1, pp. 1-224. lander. Paris, Mme. Veuve Agasse, Im- 1938. Bryozoariosmarinhosbrasileiros, 2.Bol. primeur-Libraire, 115 pp. da Faculdade de Filosofia, Ciencias e Levinsen, G. M. R. Letras, Univ. Sao Paulo, vol. 4, Zool- 1909. Morphological and systematic studies ogia, no. 2, pp. 1-137. on cheilostomatous Bryozoa. Copen- 1939. Bryozoariosmarinhosbrasileiros, 1. Bol. hagen, Natl. Forfatteres Forlag, 431 pp. da Faculdade Filosofia, Ciencias e Livingstone, A. A. Letras, Univ. Sao Paulo, vol. 3, Zool- 1926. Studies on Australian Bryozoa. No. 3. ogia, no. 3, pp. 111-299. Records ofthe Australian Museum, vol. 1949. Some Bryozoa from the Brazilian coast. 15, no. 1, pp. 79-99. Comun. Zool. Mus. Hist. Nat. Monte- Long, E. R., and J. B. Rucker video, vol. 3, pp. 1-33. 1970. Offshore marine cheilostome Bryozoa 1955. Notas sobre briozoos marinhos brasi- from Fort Lauderdale, Florida. Marine leiros. Arch. Mus. Nat. Rio de Janeiro, Biol., vol. 6, pp. 18-25. vol. 42, pp. 273-324. MacGillivray, P. H. Neviani, A. 1860. Notes on the cheilostomatous Polyzoa 1895. Nota preliminare sui Briozoi fossili del ofVictoria and other parts ofAustralia. postpliocene antic della Farnesina e Trans. Phil. Inst. Victoria, vol. 4, pp. Monte Mario. Boll. Soc. Romana stud. 159-168. Zool., vol. 4, pp. 65-74. 1869. Descriptions of some new genera and Norman, A. M. species of Australian Polyzoa. Trans. 1903. Notes on the natural history ofEast Fin- Proc. Roy. Soc. Victoria, vol.9, pp. 126- mark Polyzoa. Ann. Mag. Nat. Hist., 148. ser. 7, vol. 11, pp. 567-598. 1879. Polyzoa. In F. McCoy, Prodromus of D'Orbigny, A. the zoology of Victoria. Melbourne, 1851. Paleontologie Fran9aise, terrains Cre- Decade IV, 1879, pp. 21-35. taces. Paris, vol. 5, "Bryozoaires," pp. 1891. Descriptions of new or little known 1-188. Polyzoa. Proc. Roy. Soc. Victoria, n.s., Ortmann, A. vol. 3, pp. 77-83. 1890. Die japanische Bryozoen-Fauna. Arch. 1895. Monograph of the Tertiary Polyzoa of Naturgesch., vol. 1, pp. 1-74. 46 AMERICAN MUSEUM NOVITATES NO. 2847

Osburn, R. C. Smitt, F. A. 1914. Bryozoa of the Tortugas Islands, Flor- 1872. Floridan Bryozoa collected by Count L. ida. Carnegie Inst. Washington Publ., F. de Pourtales, pt. I. K. Svenska Ve- no. 182, pp. 181-222. tensk.-Akad. Handl., vol. 10, pp. 1-20. 1927. The Bryozoa of Cura9ao. Bijdr. Dier- 1873. Floridan Bryozoa collected by Count L. kunde, vol. 25, pp. 123-132. F. de Pourtales, pt. II. K. Svenska Ve- 1940. Bryozoa ofPorto Rico with a resume of tensk.-Akad. Handl., vol. 1 1, pp. 1-83. the West Indian Bryozoan fauna. N.Y. Soule, J. D. Acad. Sci., Sci. Survey Porto Rico and 1959. Results of the Puritan-American Mu- Virgin Islands, vol. 16, pp. 321-486. seum of Natural History Expedition to 1952. Bryozoa of the Pacific coast of North Western Mexico. 6. Anascan Cheilo- America. Part 2. Cheilostomata Asco- stomata (Bryozoa) of the Gulf of Cali- phora. Allan Hancock Pacific Exped., fornia. Amer. Mus. Novitates, no. 1969, vol. 14, pp. 271-611. pp. 1-54. Philipps, E. G. 1961. Results of the Puritan-American Mu- 1899. Report on the Polyzoa collected by Dr. seum of Natural History Expedition to Willey from the Loyalty Islands, New Western Mexico. 13. Ascophoran Chei- Guinea and New Britain. Willey's Zool. lostomata (Bryozoa) ofthe Gulfof Cal- Res., pp. 439-450. ifornia. Amer. Mus. Novitates, no. 2053, Poluzzi, A. pp. 1-66. 1977. Contributi per una revisione del genere Soule, D. F., and J. D. Soule Teuchopora Neviani 1895 (Bryozoa, 1964. The Ectoprocta (Bryozoa) of Scam- Cheilostomata). Bull. Soc. Paleontol. mon's Lagoon, Baja California, Mexico. Ital., vol. 16, pp. 69-77. Amer. Mus. Novitates, no. 2199, pp. 1- Potts, D. C. 56. 1983. Evolutionary disequilibrium among 1968. Bryozoan fouling organisms from Oahu, Indo-Pacific corals. Bull. Mar. Sci., vol. Hawaii with a new species of Watersi- 33, pp. 6 19-632. pora. Bull. So. Calif. Acad. Sci., vol. 67, Pouyet, S., and L. David pp.203-218. 1979. Revision systematique du genre Stegi- 1970. New species of Thalamoporella (Ecto- noporella Smitt, 1873 (Bryozoa Chei- procta) from Hawaii, examined by scan- lostomata). Geobios, vol. 6, pp. 763- ning electron microscopy. Amer. Mus. 817. Novitates, no. 2417, pp. 1-18. Powell, N. A. 1973. Morphology and speciation of Hawai- 1967a. Polyzoa (Bryozoa). Ascophora from ian and Eastern Pacific Smittinidae North New Zealand Discovery Rpt., vol. (Bryozoa, Ectoprocta). Bull. Amer. Mus. 34, pp. 199-393. Nat. Hist., vol. 152, pp. 365-440. 1967b. Bryozoa (Polyzoa) from the South Red 1975. Species groups in Watersiporidae. Do- Sea. Cah. Biol. Marine, vol. 8, pp. 161- cum. Lab. Geol. Fac. Sci. Lyon h.s. 3 183. (fasc. 2), pp. 299-309. 1971. The Marine Bryozoa near the Panama Stach, L. W. Canal. Bull. Mar. Sci., vol. 21, pp. 766- 1936. Studies on Recent Petraliidae (Bryo- 778. zoa). Rec. Australian Mus., vol. 19, pp. Ryland, J. S., and P. J. Hayward 355-379. 1977. British anascan bryozoans. London, Thornely, L. R. Academic Press, pp. 611-619. 1905. Report on the Polyzoa collected by Pro- Savigny, J.-C. fessor Herdman, at Ceylon in 1902. 1809-1829. Description de l'Egypte, ou recueil Ceylon Pearl Oyster Fisheries Suppl. des observations et des recherches qui Rpt. 4, no. 26, pp. 107-130. ont 6te faites en Egypte pendant l'Ex- 1906. Additions and correction. Report. Pearl pedition de l'Armee francaise ... His- Oyster Fisheries, Gulf of Manaar, IV, toire Naturelle. Imprimerie Imperiale, Suppl. Rpt. XXVI, pp. 449-450. Part. 2 vols. & Atlas (Br-yozoa, vol. 2, 1907. Report on the marine Polyzoa in the pls. 6-13). collection of the Indian Museum. Rec. Shier, D. E. Indian Mus., vol. 1, pp.- 179-196. 1964. Marine Bryozoa from northwest Flori- 1912. Marine Polyzoa of the Indian Ocean. da. Bull. Mar. Sci. Gulf Caribbean, vol. Trans. Linn. Soc. London, ser. 2 (Zool.), 14, pp. 603-662. vol. 15, pp. 137-157, pl. 8. 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 47

Uttley, G. H., and J. S. Bullivant smaller than the supporting autozooid 1972. Biological results of the Chatham Is- (feeding zooid). lands 1954 Expedition. Part 7. Bryozoa areolae marginal pores in the frontal wall Cheilostomata. New Zealand Oceanogr. of a zooid. Inst. Mem., no. 57, pp. 1-61. Verrill, A. E. ascopore a median frontal pore, marking the 1900. Additions to the Tunicata and Mollus- entrance to the ascus, the sac which permits coidea of the Bermudas. Trans. Conn. the protrusion of the polypide in some as- Acad. Arts Sci., vol. 10, pp. 592-594. cophorans. Vigneaux, M. avicularium a modified cheilostome zooid 1949. Revision des Bryozoaires neogenes du in which the internal organs are rudimen- Bassin d'Aquitaine et essai de classifi- tary and the operculum is enlarged into a cation. Mem. Soc. Geol. Fr., n.s., vol. blade-, spoon-, or bristle-shaped mandible, 28, pp. 1-153. opened and closed by hypertrophied mus- Wass, R. E., and J. J. Yoo cles. 1983. Cheilostome Bryozoa from the southern Australian continental shelf. Australian caliper a calcareous spicule, shaped like the Jour. Marine Freshwater Res., vol. 34, measuring device called an outside caliper, pp. 303-354. found beneath the frontal membrane of Waters, A. W. members of the genus Thalamoporella. 1887. Bryozoa from New South Wales, North compass a calcareous spicule in the form of Australia, etc., pt. II. Ann. Mag. Nat. two straight pointed rodsjoined at one end, Hist., ser. 5, vol. 20, pp. 181-203. resembling a drafting compass. These oc- 1889. Bryozoa from New South Wales. Ann. cur beneath the frontal membrane ofmem- Mag. Nat. Hist. ser. 6, vol. 4, pp. 1-24. bers of the genus of Thalamoporella. The 1907. Tubucellaria: its species and ovicells. morphology of compasses and calipers is Jour. Linn. Soc. (Zool.), vol. 30, no. 196, important in species-level taxonomy. pp. 126-133. 1909. Reports on the marine biology of the condyles paired small skeletal projections Sudanese Red Sea ... 12. The Bryozoa. on which the operculum of autozooids or Part 1. Cheilostomata. Jour. Linn. Soc. the mandible of avicularia pivot. (Zool.), vol. 31, pp. 123-18 1. costae the modified spines which curve and 1913. The marine fauna ofBritish East Africa meet over the frontal membrane in cribi- and Zanzibar, from collections made by morph cheilostomes, forming a frontal Cyril Crossland, M.A., B. Sc., F.Z.S. in shield. the years 1901-1902. Bryozoa-Chei- cross-bar a bar ofcalcification on which the lostomata. Proc. Zool. Soc. London, vol. mandible is hinged in many avicularia. 2, pp. 458-537. frontal skeletal reinforcement 1918. Some collections of the littoral marine cryptocyst fauna of the Cape Verde Islands, made formed by calcification of an interior wall by Cyril Crossland, M.A., B. Sc., F.Z.S., which grows out parallel to, but beneath, in the summer of 1904. Bryozoa. Jour. the frontal membrane of a zooid. In living Linn. Soc. (Zool.), vol. 34, pp. 1-45. zooids there is a coelomic fluid-filled space Winston, J. E. between the cuticle-covered frontal mem- 1982. Marine bryozoans (Ectoprocta) of the brane and the underlying cryptocyst. The Indian River Area (Florida). Bull. Amer. cryptocyst contains uncalcified spots for Mus. Nat. Hist., vol. 173, pp. 99-176. passage of muscles and connections to the 1984. Shallow-water bryozoans ofCarrie Bow main body cavity of the zooid. Cay, Belize. Amer. Mus. Novitates, no. decumbent lying flat (in reference to the po- 2799, pp. 1-38. 1986. A checklist ofcoral associated bryozoan sition of zooids at the growing edge of a species. Amer. Mus. Novitates. colony). distal the direction toward the growing edge ofthe colony and away from the ancestrula GLOSSARY or metamorphosed larva, the origin ofcol- adventitious avicularium an avicularium ony growth. which develops on the lateral, frontal, or erect (1) colonies with bushy or treelike basal wall of an autozooid. It is usually forms or (2) zooids with deep body cavities 48 AMERICAN MUSEUM NOVITATES NO. 2847

and relatively short frontal walls, often with are protruded through the larger distal por- the orifice displaced medially. They are tion. characteristic of the secondary layers in ovicell brood chamber, usually mostly cal- many species with frontal budding. cified, located at the distal end of the ma- frontal in the direction of the orifice-bear- ternal zooid. ing surface of a zooid or colony. palate the space beneath the avicularian frontal membrane uncalcified frontal body mandible equivalent to the orifice or opesia wall, consisting ofan outer noncellular cu- of an autozooid. In life it is covered by ticle, produced by underlying epidermis. frontal membrane and has a central open- frontal wall calcareous frontal covering ing to the polypide rudiment. which may be formed in several different peduncle the stalk of a bird's head avicu- ways and modified during the life of the larium, formed by the proximal portion of zooid by secondary calcification. The mor- that modified zooid. phology and texture of the frontal wall is peristome a raised rim or collar of calcifi- an important component of species de- cation surrounding the primary orifice. Its scriptions. opening is called the secondary orifice. gymnocyst frontal wall formed by deposi- polypide tube a square-sided or semicir- tion of calcification between the outer cu- cular section ofcryptocyst which covers the ticle and the epidermis ofthe frontal mem- retracted tentacle sheath in some families brane. It may be completely calcified or of anascans, e.g., Steginoporellidae, Thal- have gaps (pores) covered only by cuticle. amoporellidae. intercalary cuticle the outermost layers of pores gaps in calcification of frontal walls lateral walls ofadjoining zooids. The fron- of zooids, ovicells, and (rarely) avicularia. tal trace ofthis cuticle and the rims of cal- Though in the skeletal preparations used cification on either side of it often form a in taxonomy these are open holes, in life very noticeable line of demarcation be- they are covered by cuticle and may be tween zooids. plugged by tissue as well. interzooecial avicularium one which is primary orifice the opening in the body wall placed in the budding series between zooids, through which the feeding organs are ex- communicates with at least two zooids, but truded, located directly beneath the oper- is smaller than a feeding zooid in size. culum. The shape of the primary orifice is lappets lateral extensions of the calcifica- of great importance in cheilostome tax- tion of the peristome. onomy. However, in species with long peri- lyrula a median toothlike projection ofcal- stomes, the primary orifice may be impos- cification at the proximal end ofthe orifice. sible to see without breaking away the mandible the enlarged and modified oper- peristome. culum ofan avicularium, capable ofopen- proximal in the direction of the ancestrula, ing and closing by muscular action. the origin of colony growth. mural rim raised inner edge of the gym- rostrum (1) a sharp or blunt pointed suboral nocyst, often carrying marginal spines. spike of solid calcification, usually sup- opesia a noncalcified area under the frontal porting an avicularium (e.g., in Cellepor- membrane, bordered by cryptocyst. aria) or (2) the distal calcified beaklike end opesiule groove or hole in cryptocyst of an avicularium on which the closed through which parietal muscles pass. mandible rests. oral in the vicinity of the orifice. scutum a modified (usually enlarged and orifice opening through which the lopho- flattened) marginal spine which overarches phore protrudes for feeding. In almost all the frontal membrane in members of the cheilostomes this is closed by a hinged or Scrupocellariidae. pivoting operculum. In many ascophorans secondary calcification additional deposi- the orifice is sinuate, the proximally locat- tion ofskeletal material occurring as a zooid ed sinus marking the opening to the ascus, ages. In some species this can completely through which water enters as the tentacles transform a zooid's appearance; thus in 1986 WINSTON AND HEIMBERG: BRYOZOANS FROM BALI 49

taxonomic work it is often necessary to ex- umbo a suborificial moundlike projection amine recently budded zooids as well as of solid calcification on the frontal wall of those in older regions ofthe colony in order a zooid. See also rostrum (1). to determine identity. vibraculum an avicularium with an elon- setae the long bristlelike mandible of a vi- gate bristlelike mandible. braculum. vicarious avicularium an avicularium which sinus (1) an indentation or slit in the prox- replaces an autozooid in the budding series imal end of the peristome or (2) the U- or and is in the same size range as an auto- V-shaped proximal portion ofthe primary zooid. orifice in some ascophorans. zooid the individual of a bryozoan colony. spatulate shaped like a spoon or a broad Autozooids are the regular feeding zooids blunt-ended blade (of avicularian mandi- ofthe colony. Additional specialized poly- bles). morphic zooids: avicularia, gonozooids, spine a tubular or flattened projection of rhizooids, and kenozooids are found in calcified body wall and coelomic cavity, most cheilostome species. sometimes jointed.

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