SCIENTIFIC RESULTS OF THE HASSLER EXPEDITION. BRYOZOA. BARBADOS NO. 1. JUDITH E. WINSTON1 AND ROBERT M. WOOLLACOTT2 CONTENTS Abstract 240 Steginoporella connexa Harmer, 1900 ..... 254 Introduction 240 Superfamily Cellarioidea 256 Study Area and Materials 241 Family Cellariidae 256 Results 242 Genus Cellaria 256 Systematics 242 Cellaria louisorum new species 256 Class Stenolaemata 242 Infraorder Ascophora 257 : Order Cyclostomata 242 "Grade" Acanthostega 257 Family Crisiidae 242 Superfamily Cribrilinoidea 257 Genus Crista 242 Family Cribrilinidae 257 Crisia sp. 242 Genus Puellina 257 Family Oncousoeciidae 243 Puellina smitti Winston, 2005 257 Genus Stomatopora 243 "Grade" Umbonulomorpha 259 Stomatopora sp. _ 243 Superfamily Lepralielloidea 259 Genus Proboscina 245 Family Romancheinidae 259 Proboscina robusta ... 245 Genus Exochella 259 Family Terviidae 245 Exochella tropica new species 259 Genus Tervia 245 "Grade" Lepraliomorpha 260 Tervia sp. 245 Superfamily Smittinoidea 260 Family Lichenoporidae 247 Family Smittinidae 260 Genus Patinella 247 Genus Smittoidea 260 Patinella sp. 247 Smittoidea reginae new species 260 .. Class Gymnolaemata 248 .; Genus Parasmittina 262 Order Cheilostomata 248 Parasmittina barbadensis new species 262 Suborder Neocheilostomina 248 Family Bitectiporidae 262 Infraorder Flustrina 248 Genus Parkerrnavella 262 Superfamily Calloporoidea 248 Parkermavella salebrosa new species 262 Family Antroporidae 248 Genus Hippoporina 264 Genus Antropora 248 Antropora typica....(Canu & Bassler, 1928) 248 Hippoporina rutelliformis new species 264 Genus Metroperiella 266 Superfamily Buguloidea 252 Family Candidae 252 Metroperiella agassizi new species _____ 266 Superfamily Schizoporelloidea 268 Genus Caberea 252 Caberea hassleri new species 252 Family Schizoporellidae 268 Genus Stylopoma 268 Superfamily Microporoidea 252 Stylopoma smitti Winston, 2005 268 Family Steginoporellidae 252 Stylopoma haytvardi new species 271 Genus Steginoporella 252 .... Family Gigantoporidae 273 Steginoporella magnilabris (Busk, Genus Barbadiopsis new genus 273 1854) 252 Barbadiopsis trepida new species 273 Family Teuchoporidae 274 Virginia Museum of Natural History, 21 Starling Genus Lagenicella 274 Avenue, Martinsville, Virginia 24112. Lagenicella verrucosa (Canu & 2Museum ofComparative Zoology, Harvard Univer- Bassler, 1928) .. ..... 274 sity, 26 Oxford Street, Cambridge, Massachusetts Family Microporellidae 274 02138. Genus Microporella 274 Bull. Mus. Comp. Zool., 159(5): 239-300, November, 2009 239 240 Bulletin ofthe Museum ofComparative Zoology, Vol. 159, No. 5 Microporella protea Winston, 2005 274 INTRODUCTION Family Escharinidae 275 Genus Bryopesanser 275 This report concerns a small collection of Bryopesanser pesanseris 275 bryozoans from the island of Barbados Superfamily Mamilloporoidea 277 located in the Lesser Antilles chain border- Family Cleidochasmatidae 277 ing the Caribbean Sea. The specimens were Genus Gemelliporina 277 Gemelliporina hastata new sp_ecies .:. 277 obtained as part of an expedition conducted Superfamily Celleporoidea 277 in 1871-1872 and initiated by Benjamin Family Celleporidae 277 Peirce, then superintendent of the United .,.. Genus Buffonellaria 277 States Coast Survey (Anonymous, 1871). Buffonellaria ensifera new species 277 Peirce commissioned construction on the Genus Buskea 281 Buskea minutiporosa (Canu & Atlantic Coast of the iron-hulled 165-foot Bassler, 1928) 281 steamer Hassler for use in hydrographic .. Genus Cigclisula 281 surveys on the Pacific Coast of North Cigclisula gemmea new species 281 America. The ship, therefore, needed to Genus Trematooecia 285 Trematooecia turrita (Smitt, 187_3) __ 285 be moved to San Francisco, the site of its Family Phidoloporidae 285 first positioning. Jean Louis Rodolphe Genus Bhynchozoon 285 Agassiz (founder of the Museum of Com- ... Bhynchozoonsexaspinatumnewspecies 285 parative Zoology, Harvard University, Cam- Genus Stephanollona 287 bridge, Massachusetts, USA) accepted an Stephanollona propinqua new species 287 Genus Beteporellina ... 289 invitation from Peirce to lead a scientific Reteporellina directa new species 289 expedition of Agassiz's design during the Discussion 290 ship's circumnavigation of South America Acknowledgments 297 from Boston to San Francisco. The ship was Literature Cited :...... 297 staffed with Navy Department personnel headed by Commander P. C. Johnson, but Abstract. An unidentified collection of bryozoans Agassiz was permitted to select nonmilitary made by L. F. Pourtales and L. Agassiz during the scientists and assistants for the expedition Hassler Expedition (1871-1872) was recently discov- ered in the teaching collection of the Invertebrate (Peirce, 1871). The scientific contingent Paleontology Department at the Museum of Compar- included Agassiz; Mr. James Blake, assistant ative Zoology, Harvard University. Bryozoan samples and artist; Dr. Thomas Hill, a physicist and found included stations from Barbados to Brazil and former president of Harvard College; Louis around the coast of South America to La Jolla, California. Francois de Pourtales, Esq., scientist in The first and most successful deepwatercollectingof charge of dredging and collecting; Dr. bryozoans was done early in the expedition at two Franz Steindachner, an ichthyologist; and stations (146 m and 183 m) offBarbados, on December Mr. William White, assistant. Agassiz's 29 and 30, 1871. Thirty-one taxa of bryozoans were J. collected at the Barbados stations: five cyclostomes and wife, Elizabeth Cary Agassiz, as well as the 26 cheilostomes, includingone newgenus. None ofthe wife of Commander P. C. Johnson, also cyclostomes was reproductive, so taxa could not be accompanied the expedition (Agassiz, E.C., identified to species level. Of the 26 cheilostomes, 16 1872; Peirce, 1871). represent new species: Caberea hassleri, Cellaria louisorum, Exochella tropica, Smittoidea reginae, Louis Agassiz had grand visions for the Parasmittina barbadensis, Parkermavella salebrosa, significance of this expedition. In letters Hippoporina rutelliformis, Metroperiella agassizi, Sty- soliciting funds for costs of the scientific lopoma haywardi, Barbadiopsis trepida, Gemelliporina component ofthe expedition, Agassiz wrote: hastata, Buffonellaria ensifera, Cigclisula gemmea, Bhynchozoon sexaspinatum, Stephanollona propinqua, ... the results of this voyage will be as and Beteporellina directa. All species found are important for the increase ofour knowledge described and illustrated with scanning electron of the characteristics of the sea, as the microscope photographs. Bryozoans of the Hassler voyages of Capt. Cook were, a century ago, Expedition are nowincorporated into the collections of the Department of Marine Invertebrates of the for the improvement of navigation and Museum ofComparative Zoology. geography" (Agassiz, L., 1871). Time would Scientific Results of the Hassler Expedition • Winston and Woollacott 241 prove, however, that Agassiz's dreams were mens of particular sponges, crinoids, echi- not realized for the most part. noids, and an extremely rare living specimen The voyage began in Boston on 4 of the gastropod genus Pleurotomaria (Mal- December, 1871, and concluded in San acology Catalogue number MCZ 119057). Francisco on 30 August 1872. During the These hauls also yielded 31 species of course of the cruise numerous stops were bryozoans, which are described for the first made, some planned and some unplanned. time in this report. Sixteen ofthe 31 are new That portion ofthe expedition relevant to species, and there is one new genus. The this study concerns circumstances sur- Hassler departed its unscheduled detour to rounding collections made off Barbados on Barbados about 5:00 p.m. on 30 December 29 and 30 December, 1871. Unless noted and sailed toward Rio de Janeiro, the next otherwise, the account given here was taken destination on the expedition itinerary. in large measure from E. C. Agassiz's narrative of the voyage (Agassiz, E. C, STUDY AREA AND METHODS 1872) and L. F. de Pourtales' official The small (34 km long by 18 km wide) summary of the expedition written for the coral-limestone island of Barbados is the U.S. Coast Survey (Pourtales, 1875). easternmost island of the Lesser Antilles The Hassler departed St. Thomas on 22 volcanic arc, an area extending from the December and headed toward the islands of Virgin Islands to Tobago. Santa Cruz, Montserrat, and Guadaloupe to The Barbados lots studied here were conduct benthic sampling at the deepest among the specimens ofbryozoans collected depths possible. Heavy seas, however, made such dredging impossible. On 26 December, during the Hassler Expedition (1871-1872). Bryozoan material from the Hassler was one ofthe airchambers ofthe auxiliary steam pump failed and could not be repaired at sea. discovered in the paleontology teaching The captain electedtohead foran unplanned collection in the Museum of Comparative destination, Barbados, where in the city of Zoology (MCZ) in the early 2000s and transferred to the Department of Marine Bridgetown the engine could be refurbished. They sighted Barbados around 5:00 p.m. that Invertebrates. All of the material was uncatalogued and specimens were uniden- afternoon and anchoredlaterthat eveningoff Bridgetown. Repairs began the following tified with the exception of a single lot. morning and were completed on the after- Remaining bryozoan lots obtained from noon of 29 December. During this interval, other locales visited by this expedition will Agassiz's group went ashore for touring and be treated in forthcoming publications. collecting. The Barbados lots were received as On the afternoon of 29 December, the several boxes of dry specimens with labels Hassler steamed 5 or 6 miles up the coast indicating the depth at which they had been from Bridgetown to Sandy Bay. At this time, collected (80 or 100 fm), but otherwise measurements of the amounts of light unsorted except as "Bryozoa". Specimens present with varying depth were made by from each depth were rinsed in freshwater Dr. Hill and a series of dredge hauls were and dried. They were then examined under undertaken. The ship returned to Bridge- a dissecting microscope and sorted, or town that evening, but traveled again the detached from larger substrata when nec- following day back to Sandy Bay for an all- essary. The processed specimens were given day dredging trip (Blake, 1871-1872). The preliminary taxonomic identifications and dredge hauls were wonderfully successful in placed in plastic boxes approved for collec- the eyes of Agassiz. Blake reports "Prof A. tions storage. From each sorted lot one or said he would be contented if we got more colonies or colony fragments were nothing else" (Blake, 1871-1872). Especial- selected for study and digital imaging using ly important finds to Agassiz were speci- scanning electron microscopy (SEM). Stan- 242 Bulletin ofthe Museum ofComparative Zoology, Vol. 159, No. 5 dard measurements were made on at least Parasmittina barbadensis one colony of each taxon found. Measure- Reteporellina directa ments were made using a Wild stereomi- Smittoidea reginae = croscope with 20X oculars (magnification Steginoporella magnilabris 100X). The measurements included dimen- Stephanollona propinqua sions of the following characters if present: Stomatopora sp. zooid length (Lz), zooid width (Wz), opesia SYSTEMATICS length (Lop), opesia width (Wop), primary orifice length (Lo), primary orifice width (Wo), orifice diameter (Diam.o) for round Class Stenolaemata orifices, secondary orifice length (Lo2), Order Cyclostomata secondary orifice width (Wo2), ovicell Family Crisiidae Johnston, 1838 length (Lov), ovicell width (Wov), avicular- Genus Crisia Lamouroux, 1812 ian length (Lav), avicularian width (Wav). Crisia sp. RESULTS Figure 1 Description. Colony erect, white, jointed, Species Found slightly curving biserial branches (Fig. 1A) Barbados 80 fm (146 m) and long internodes (12-15 zooids per Antropora typica internode). Zooids long and tubular, their Buffonellaria ensifera calcified walls sparkling with relatively large, Caberea hassleri evenly spaced pseudopores. Distal portion of Cellaria louisorum zooid tubes curving slightly up and outward Crisia sp. from the branch, orifices round to oval Exochella tropica (Fig. IB). Bases rami, the initiation points Hippoporina rutelliformsis of side branches, occur above the third to Metroperiella agassizi seventh zooid from the base ofan internode. Microporella protea Chitinous joints between internodes black. Patinella sp. Tubular attachment kenozooids about 0.9- mm Proboscina robusta 0.10 in diameter growfrom the abfrontal Puellina smitti side of some of the branches. The colony Rhynchozoon sexaspinatum fragments found did not have gonozooids. Steginoporella connexa Measurements Stomatopora sp. Stylopoma haywardi. Range Mean IV Stylopoma smitti Lz 0.491-0.819 0.646 6 Wz 0.109-0.127 0.121 6 Tervia sp. Lo 0.055-0.073 0.067 6 Trematooecia turrita Wo 0.073-0.082 0.074 6 Wbranch 0.264-0.309 0.281 6 Barbados, 100 fm (183 m) Barbadiopsis trepida Notes. The morphology and dimensions Bryopesanser pesanseris of zooids and branch internodes are quite Buffonellaria ensifera similar to those described for Crisia denti- Buskea minutiporosa culata (Lamarck, 1816). However, as Hay- Cigclisula gemmea ward and Ryland (1985, p. 54) point out, Gemelliporina hastata "Despite the distinctive appearance of this Lagenicella verrucosa species, with the jet black joints and Parkermavella salebrosa wedged-in basis rami being especially clear Metroperiella agassizi characters, the name denticulata has been Microporella protea widely applied to almost any coarsely Scientific Results of the Hassler Expedition • Winston and Woollacott 243 Figure 1. Crisia sp. Hassler Box 4. A. Branches of colony. Scale bar = 500 |xm. B. Zooids, showing size of tubes and orifice shape. Scale bar = 100 urn. MCZ 100108. straggling, long internode species, causing pores that show a sieve platelike ultrastructure confusion on a par with that for C. when highly magnified (5,000X, Fig. 2F). eburnea." The Barbados material is also Orifices small, round, raised slightiy in short similar in colony and zooid size, internode peristomes from the recumbent portions of length, and dark joint coloration to Crisia zooids. No gonozooids found. ficulnea (Buge, 1979) from Brazil. However, Measurements since the material examined lacked gono- zooids, we identify it here only as Crisia sp. Range Mean N Lz 0.892-1.183 0.995 3 Distribution. Barbados. MCZ Wz 0.182-0.200 0.194 3 Specimens Examined. 100108. Hass- Diam.o 0.109-0.164 0.133 3 ler Box 4, Barbados, 80 fm. December 1871. Notes. The Stomatopora trahens de- Family Oncousoeciidae Canu, 1918 scribed by Lagaaij (1963, p. 208) is very similar to the Barbados material in morphol- Genus Stomatopora Bronn, 1825 ogy. Only one branching point occurs in the Stomatopora sp. Barbados colony, and it is broken on one Figures 2, 4D side, but the angle ofbranching is similar, as are the proportions of zooids and orifices. Description. Colony white, encrusting, However, it has since been shown (Hayward uniserial, sparsely dichotomously branching, and Ryland, 1985, p. 60 and Fig. 39B) that S. spreading over and among colonies of bryo- trahens of Couch (1841, p.71) is merely an zoans and other calcareous substrata (Figs. 2 earlygrowth stage ofEntatophoroecia deflexa A-D). Zooids narrow, elongate, concentrically (Couch) 1842. True Stomatopora species, wrinkled tubes, impossible to measure more mostly from deepwater habitats, have a than approximately because oftheir curvature gonozooid in the form ofa small sac budded or their distortion by underlying substratum from the end ofthe peristome, with a narrow topography (Fig. 2E). Calcification with trans- oeciostome projecting from it (Harmelin, verse striations and numerous small pseudo- 1974, Hayward and Ryland, 1985). The 244 Bulletin ofthe Museum ofComparative Zoology, Vol. 159, No. 5 Figure2. Stomatoporasp. (A-E HasslerBox2. MCZ 100109)A. Stomatoporacolonycreeping over Proboscina robustacolony. Scalebar = 200|im. B. Overlapofthetwospecies. Scalebar = 100|iim. MCZ 100109. C. Anothercolony, scalebar = 200|im. D. Partly overgrown colony. Scale bar = 200 nm. E. Close-up of single zooid of Stomatopora being overgrown by Hippoporina rutelliformis. Scale bar =100 (am. F. (Hassler Box 6. MCZ 100112) Pincushion pore plates. Scale bar = 2 jam. Scientific Results of the Hassler Expedition • Winston and Woollacott 245 specimen from Barbados, and Lagaaij's ofadjacent zooid tubes ofthe two species at specimen also, have proportionally longer 700X for comparison. tubes and smallerdiameter orifices than seen Distribution. Caribbean, Brazil. MCZ in the encrusting phases of other erect Specimens Examined. 100111. tubuliporids. They may represent a new Hassler Box 2, Barbados, 80 fm. December MCZ species of Stomatopora, but lacking gono- 1871. 100112. Hassler Box 6, Barba- zooids they cannot be described further. dos, 100 fm. December 1871. Distribution. Barbados. Specimens Examined. MCZ 100109. Family Terviidae Canu & Bassler, 1920 Hassler Box 2, Barbados, 80 fm. December Genus Tervia Jullien, 1882 MCZ 1871. 100110. Hassler Box 6, Barba- Tervia sp. dos, 100 fm. December 1871. Figure 5 Genus Proboscina Audouin, 1826 Description. Colony white, erect, and Proboscina robusta Canu & Bassler, 1928 dichotomously branching (Fig. 5A). Zooid Figures 3, 4 tubes opening on frontal and frontal-lateral surface, and with an abfrontal surface Proboscina robusta Canu & Bassler, 1928a: 157, pi. 30, marked by crescentic transverse striations. fig. 7, 1928b: 100, pi. 9, fig. 4. Zooids tubular, their boundaries obscured Proboscina robusta, Osburn, 1947: 3. Buge, 1979: 219, by their curving and transversely striated pi. 2, fig. 2. calcification, but with distal ends raised and Description. Colony white, encrusting, curved away from the branch axis as oval to composed of a series of large tubular zooids subangular peristomes. Pseudopore ultra- in Diserial to multiserial arrangement structure consisting of simple angular to (Figs. 3A-C). Zooids adherent to substratum rounded openings (Fig. 5D). The colony formostoftheirlength,butwithdistalvertically fragment present in the Barbados material projecting peristomes, ending in round to oval is not complete but shows on its abfrontal openings. Calcification heavy relative to some surface a swelling (broken off basally) that other cyclostomes (e.g., the Crisia sp. above), may be a gonozooid, with a possible with wrinkled appearance due to numerous oeciostome at its distal end (Figs. 5C-E). transverse striations, and a surface speckled by Measurements pseudo-pores, which at a high magnification (5,000X) show a spokelike arrangement of Range Mean N denticles (Figs. 4B, C). Gonozooids formingin Lz 0.746-1.056 0.904 6 Wz 0.328-0.382 0.349 6 lobatemultiserialexpansionsofthecolony,with Lo 0.191-0.246 0.224 6 same pseudopore ultrastructure as autozooid Wo 0.164-0.182 0.179 6 tubes (Figs. 3D-F). Those collectedherewere found on a very abraded colony, and the SEM Notes. images appear flatter than oeciostome could not be determined with the viewofthe same specimen through a light certainty, but is likely the narrow proximally microscope. In light microscope view the facing tube showing in Figures 3E and F. curving horizontal striations are much more Measurements noticeable as is the ridged very three- dimensional shape ofthe branches. No Tervia Range Mean N Lz 0.437-0.801 0.576 6 species have been described from the West- Wz 0.309-0.382 0.352 6 ern Atlantic-Caribbean region. According to Diam.o 0.109-0.146 0.129 6 Harmelin (1976) and Hayward and Ryland (1985), the eastern Adantic species, Tervia Notes. Both zooid dimensions and pseu- irregularis (Meneghini) 1844, prefers deep dopore ultrastructure contrast with those of water (100 m+) and detritic deposits of the Stomatopora sp. Figure 4D shows portions continental shelves, which agrees well with 246 Bulletin ofthe Museum ofComparative Zoology, Vol. 159, No. 5 Figure3. Proboscinarobusta-1. HasslerBox6. MCZ 100112. A. Portionofencrustingcolony. Scalebar = 200|jm. B. Expansion ofzooid rows into lobes. Scale bar = 500|am. C. Viewoftwo lobes ofcolony. Scale bar = 500|im. D. Close-up of left lobe. Scale bar = 200 ^im. E. Lobe with possible gonozooid. Scale bar = 200 |am. F. Close-up of gonozooid. Scale bar = 50 urn. Scientific Results of the Hassler Expedition • Winston and Woollacott 247 Figure4. Proboscinarobusta-2. A. Close-upofporesonzooidperistomesandcolonysurface. Scalebar = 100|im. B. Magnified viewof pores showing radial denticles. Scale bar =10 (xm. C. Highly magnified pore. Scale bar = 1 urn. D. Direct comparison of pore structure of Stomatopora (on left) and Proboscina (on right) zooids. Scale bar = 10 ^im. the depth and habitat in which the Barbados knobby keels, their terminal openings with specimen was found. scalloped projections (Fig. 6B). Colony Distribution. Barbados, 100 fm. skeleton, including that of the basal lamina, MCZ Specimens Examined. 100113. Hass- with a pustulose texture due to tiny rounded ler Box 7, Barbados, 100 fm. December 1871. bumps of calcification (Fig. 6C). No gono- zooids present on Barbados specimens. Family Lichenoporidae Smitt, 1867 Genus Patinella Gray, 1848 Measurements Patinella sp. Range Mean N Figure 6 Lz 0.291-0.364 0.328 6 Wz 0.109-0.146 0.123 6 Lo 0.073-0.100 0.088 6 Description. Colony rounded in outline, Wo 0.055-0.091 0.070 6 the raised central area of spiny zooids interspersed with alveolar spaces is sur- Notes. Figure 6A shows what appear to be rounded by a flat peripheral lamina two subcolonies regenerated from a dam- (Figs. 6A, B). Zooid tubes radiating outward aged or senescent older colony. As there are from colony center, upper surfaces with no gonozooids on this composite colony, it is 248 Bulletin ofthe Museum ofComparative Zoology, Vol. 159, No. 5 Figure 5. Terviasp. Hassler Box7. MCZ 100113. A. Frontal viewof colony branch. Scale bar = 500 |im. B. Abfrontal surfaceof same branch. Scale bar = 500 ^m. C. Possible gonozooid. Scale bar = 100 urn. D. Opening to possible gonozooid. Scale bar = 20 ^m. E. View of broken proximal end of branch. Scale bar = 100 \im. not possible to identify it further. Its Genus Antropora Norman, 1903 morphology is very similar to the Patinella Antropora typica (Canu & Bassler, 1928) MCZ sp. From Smitt's Florida material, as Figures 7, 8 shown in Figures 348-352 of Winston (2005). But, as with that juvenile colony, Membrendoecium strictorostris Canu & Bassler 1928a: there are no gonozooids on which to base a 23, pi. 2, fig. 7. species identification and description. Dacryonella tijpica Canu & Bassler 1928a: 57, pi. 5. MCZ figs. 4-8, pi. 32, figs. 11-12, text-fig. 8a. 1928b: 65, Specimens Examined. 100114. Hass- pi. 1, fig. 10. ler Box 1-2, Barbados, 80 fm. December 1871. Antropora typica, Lagaaij, 1963:171, pl.l, fig. 3. Win- ston 1986: 5, figs. 1-2. Tilbrook, 1998: 37, fig. 3A. Class Gymnolaemata Description. Colony encrusting in one or Order Cheilostomata more layers (Fig. 7A). Zooids with a large Suborder Neocheilostomina d'Hondt, 1985 irregularly polygonal gymnocyst surround- (part) ing a raised subtriangular to pear-shaped Infraorder Flustrina Smitt, 1868 (part) cryptocyst with wide striated rows of Superfamily Calloporoidea Norman, 1903 granularcalcification,wideproximally,becom- Family Antroporidae Vigneaux, 1949 ing very narrow at the distal end around the