SY +. * seathan ee NV freee ‘ vn an asa re Ran eert scat ongeimntete Wa wears uh Raced Catt Ria ore a aa ee Sein ige lanes Rei are Me Sein anak mR ara EME a St selacsmt ner erate ent ee See) Prete a hatord © sea 4 our ‘utes apa a ae a mate oe ‘ th ; cher PEP EP TOR aaah aio AE haha eas iaidbabeta eed tees pita ' | ’ St z th = if ny Ah ey i > bat ¢ F A : sh x + i %, a n t lay 1 fe mae ” i ' : i eos { & saa * : f why By ' Ves ¢ t 9 f =) tp: 1 8 Tr ' = } ; 4 ; } ; 5 f i te A ’ ips » : j ‘ Issued by NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C. U.S.A. APRIL 1996 ATOLL RESEARCH BULLETIN NOS. 435-442 RESEARCH BULLETIN ATOLL RESEARCH BULLETIN NOS. 435-442 NO. 435. NO. NO. NO. NO. NO. NO. NO. 436. 437. 438. 439. 440. 441. 442. MORPHOLOGY AND MARINE HABITATS OF TWO SOUTHWESTERN CARIBBEAN ATOLLS: ALBUQUERQUE AND COURTOWN BY JUAN M. DIAZ, JUAN A. SANCHEZ, SVEN ZEA, AND JAIME GARZON-FERREIRA CORAL FAUNA OF TAIPING ISLAND (ITU ABA ISLAND) IN THE SPRATLYS OF THE SOUTH CHINA SEA BY CHANG-FENG DAI AND TUNG-YUNG FAN FIRST OBSERVATIONS ON THE FISH COMMUNITIES OF FRINGING REEFS IN THE REGION OF MAUMERE (FLORES- INDONESIA) BY MICHEL KULBICKI GROUPER DENSITY AND DIVERSITY AT TWO SITES IN THE REPUBLIC OF MALDIVES BY ROBERT D. SLUKA AND NORMAN REICHENBACH EFFECT OF TYPHOONS ON THE LIZARD COMMUNITY OF A SHELF ATOLL BY MICHAEL JAMES MCCOID FLOWERING AND FRUITING IN THE FLORA OF HERON ISLAND, GREAT BARRIER REEF, AUSTRALIA BY R.W. ROGERS NAMU ATOLL REVISITED: A FOLLOW-UP STUDY OF 25 YEARS OF RESOURCE USE BY NANCY J. POLLOCK CRUSTACEA DECAPODA OF FRENCH POLYNESIA (ASTACIDEA, PALINURIDEA, ANOMURA, BRACHYURA) BY JOSEPH POUPIN ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. APRIL 1996 ACKNOWLEDGMENT The Atoll Research Bulletin is issued by the Smithsonian Institution to provide an outlet for information on the biota of tropical islands and reefs and on the environment that supports the biota. The Bulletin is supported by the National Museum of Natural History and is produced by the Smithsonian Press. This issue is partly financed and distributed with funds from Atoll Research Bulletin readers and authors. The Bulletin was founded in 1951 and the first 117 numbers were issued by the Pacific Science Board, National Academy of Sciences, with financial support from the Office of Naval Research. Its pages were devoted largely to reports resulting from the Pacific Science Board's Coral Atoll Program. All statements made in papers published in the Atoll Research Bulletin are the sole responsibility of the authors and do not necessarily represent the views of the Smithsonian nor of the editors of the Bulletin. Articles submitted for publication in the Atoll Research Bulletin should be original papers in a format similar to that found in recent issues of the Bulletin. First drafts of manuscripts should be typewritten double spaced and can be sent to any of the editors. After the manuscript has been reviewed and accepted, the author will be provided with a page format with which to prepare a single-spaced camera-ready copy of the manuscript. COORDINATING EDITOR Ian G. Macintyre National Museum of Natural History MRC-125 ASSISTANTS Smithsonian Institution Kasandra D. Brockington Washington, D.C. 20560 William T. Boykins, Jr. Theodore E. Gram EDITORIAL BOARD Stephen D. Cairns (MRC-163) National Museum of Natural History Brian F. Kensley (MRC-163) (Insert appropriate MRC code) Mark M. Littler (MRC-166) Smithsonian Institution Wayne N. Mathis (MRC-169) Washington, D.C. 20560 Victor G. Springer (MRC-159) Joshua I. Tracey, Jr. (MRC-137) Warren L. Wagner (MRC-166) Roger B. Clapp National Museum of Natural History National Biological Survey, MRC-111 Smithsonian Institution Washington, D.C. 20560 David R. Stoddart Department of Geography 501 Earth Sciences Building University of California Berkeley, CA 94720 Bernard M. Salvat Ecole Pratique des Hautes Etudes Labo. Biologie Marine et Malacologie Université de Perpignan 66025 Perpignan Cedex, France PUBLICATIONS MANAGER A. Alan Burchell Smithsonian Institution Press ATOLL RESEARCH BULLETIN NO. 435 MORPHOLOGY AND MARINE HABITATS OF TWO SOUTHWESTERN CARIBBEAN ATOLLS: ALBUQUERQUE AND COURTOWN BY JUAN M. DIAZ, JUAN A. SANCHEZ, SVEN ZEA, AND JAIME GARZON-FERREIRA ‘ Pr EM MISUNIATN = ¥ \ Ql ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. APRIL 1996 RMON MMOQ AA NM STAND NANT DAT RDNA BIA DTT Ny S| . \ S \ N S N NN N N \ h N S| ‘| N N N hy RK N S N 's N N ny S ‘| N ‘ Ss N S 'S N S \ N N N Ss N N NY N NN Ss N N N N S| N N N yy N N N \ N S| N N Ss NN N N N N N N 's N N N N NY N \ N \ NN N N AMD DE MMMM MLL QUITASUENO BANK PROVIDENCIA Is. SONI Honduras ALBUQUERQUE ATOLL Nicaragua COLOMBIA ZL SSLSS EA. LA GLIDOV ISAS IETS SE Z LFS. Z EIS. FHV GALS LS PLIST. LPS. LIDS. Gil He VA LED Vids f the Colombian h the location o 2. wit bbean Sea ri ines are iSO 1. Map of the Southwestern Ca igure F In m baths ( lands, atolls and banks. Contour li 1S MORPHOLOGY AND MARINE HABITATS OF TWO SOUTHWESTERN CARIBBEAN ATOLLS: ALBUQUERQUE AND COURTOWN BY JUAN M. DIAZ}, JUAN A. SANCHEZ , SVEN ZEA!” and JAIME GARZON-FERREIRA' ABSTRACT Albuquerque and Courtown are two small, uninhabited oceanic atolls, located in the southwestern Caribbean Sea, belonging to the San Andrés and Providencia archipelago, Colombia. These atolls have a volcanic basement and are surrounded by deep water. Based on photo-interpretation of geomorphological and ecological features as well as on data collected during field work, the gross morphology, marine bottom habitats and reef structures of both atolls are described down to a depth of 50 m. Distributions of morphological and bottom habitat units are presented in thematic maps showing the overall zonational patterns in the two atolls. Morphological and ecological zonations in both atolls are primarily controlled by both wave exposure in a windward-leeward gradient and depth. The presence of an ample windward fore-reef terrace, a well developed windward barrier reef with spur-and-groove system, an extensive lagoonal terrace with sudden transition to the lagoon basin, and profuse development of ribbon and anastomosing patch reefs in the lagoon are characteristics common to both atolls. As in other Caribbean atolls, the outer slope in Albuquerque and Courtown is outlined by a sandy step or bench at 35 to 45 m depth. Significant differences between the two atolls exist in the degree of development and structure of leeward peripheral reefs, as well as in the amplitude of the leeward fore-reef terrace. At Albuquerque, peripheral reefs grow on a shallow flat and enclose the lagoon along a wide semicircle, whereas at Courtown such reefs have in part developed algal ridge-like structures and are unevenly distributed, leading to an open lagoon to the east. The broad leeward terrace in Albuquerque contrasts markedly with the rapidly dipping leeward slope towards the outer shelf margin in Courtown. Accumulations of sand and rubble have led to the formation of cays and small islands on the lagoonal terrace in both atolls, but also on leeward peripheral reefs in Instituto de Investigaciones Marinas y Costeras, INVEMAR, Apartado 1016, Santa Marta, Colombia * Universidad Nacional de Colombia (Departamento de Biologia) Manuscript received 31 March 1995; revised 21 Novermber 1995 Courtown, some of which have experienced remarkable changes in their size and shape in the last 25 years. Biological composition and structure of reefs in both atolls show a great resemblance to one another and to the better-known reef complexes around the nearby islands of San Andrés and Providencia. Although no urban development exists in these atolls, recent decline of living coral and over-exploitation of marine resources were evident. INTRODUCTION There are about 425 atolls worldwide and only 15 of them are located in the Atlantic, of which four are part of the San Andrés and Providencia archipelago in the Southwestern Caribbean Sea (Milliman, 1973; Geister, 1983). This archipelago comprises a series of islands, atolls and coral shoals running in SSE-NNE direction, parallel to the Nicaraguan Rise for more than 500 km. It is separated from the Central American continental shelf by the San Andrés Trough (Fig. 1). The southernmost reefs of the archipelago, Albuquerque and Courtown (the latter are also called Bolivar Cays) are two small atolls lying about 200 km east of the Nicaraguan coast. Although geographically closer to Central America than to the South American continent, the archipelago has belonged to the Republic of Colombia since 1822. The accurate date of human discovery of these atolls is uncertain but their locations were well known to the Spanish sailors of the 16th century and were probably occasionally visited by Miskito Indians from the Central American coast, who came for fishing and turtling (Parsons, 1956). None of the tiny sand cays on the atolls has sufficient land to warrant permanent settlement, but one of them on each atoll serves presently as a military post for the Colombian navy, and they are visited regularly by fishermen and tourists in chartered yachts from nearby San Andrés. Briefly mentioned by Darwin (1842) in his interpretation of Caribbean reef structures and their origin, the reefs of Albuquerque and Courtown have since received little scientific attention in comparison with those around the nearby islands of San Andrés and Old Providence (Geister, 1969, 1973,1975,1992; Kocurko, 1977; Marquez, 1987; Diaz et al., 1995) and other West Atlantic and Caribbean atolls such as Hogsty Reef (Milliman, 1967), Alacran Reef (Kornicker and Boyd, 1962; Bonet, 1967), Chinchorro (Jordan and Martin, 1987) and those off Belize (Stoddart, 1962; James and Ginsburg, 1979: Riitzler and Mcintyre, 1982; Gischler, 1994). Albuquerque and Courtown were briefly visited by the Fifth George Vanderbilt Expedition in 1941. Published observations include reports on the birds (Bond and DeSchauensee, 1944), fishes (Fowler, 1944) and crustaceans (Coventry, 1944). The R/V GERDA, of the University of Miami, stopped in May 1966 for few days at Albuquerque and Courtown and conducted observations on the ecology, morphology and oceanography of the atolls. From this visit, Milliman and Supko (1968) made preliminary conclusions on the geological origin, and Milliman (1969) described the general characteristics of the reefs and commented on hydrography. Further oceanographic findings from the waters surrounding the atolls have been recorded during research cruises by the Colombian navy (Gonzalez, 1988; Téllez et al., 1988). Aspects of the terrestrial environment and fauna were more recently discussed by Chirivi (1988). However, very little is known about the distribution of marine bottom habitats and the zonation of the reefs constituting these atolls. Therefore, the purpose of this paper is to give the first detailed systematic description of the gross morphology and the marine habitats of Albuquerque and Courtown atolls, with emphasis on the reef structures. REGIONAL SETTING Albuquerque and Courtown are the southernmost reef complexes of the San Andrés and Providencia archipelago. Albuquerque (12° 10' N and 81° 51' W) is located 37 km south of San Andrés Island and about 190 km east of the Nicaraguan coast. It is nearly circular in shape, about 5.5 km E-W and 4.5 km N-S. Two small islands, North Cay and South Cay, rise up to 2 m above mean sea level behind the seaward barrier reef and are separated from each other by a 250 m shallow channel. Courtown (12° 24' N and 81° 28' W) lies 30 km southeastward from San Andrés and 47 km northeast of Albuquerque. It is kidney shaped, about 3.5 km E-W and 6.5 km SSE-NNW (Fig. 3). Although this atoll presently bears two cays (East Cay and West or Bolivar Cay) and a tiny sand spit, their size, shape and number seem to be quite variable in the course of time, as can be easily inferred from an earlier description and map of the atoll by Milliman (1969). Toward the north and eastern sides of both atolls, an almost continuous barrier reef is well developed, whereas the leeward peripheral reefs are absent or ill defined and are separated by wide gaps and channels. Both atoll foundations rise from the surrounding sea floor more than 1000 m deep, and apparently have a volcanic basement. Unequivocal evidence for the volcanic origin of these atolls and nearby islands comes from the magnetic anomalies detected at San Andrés Island and Courtown, one volcanic pebble dredged from Albuquerque basement (Milliman and Supko, 1969), as well as the volcanic rocks of Providencia (Geister, 1992) and the Corn Islands (McBirney and Williams, 1965). Further aspects of the geological origin of the archipelago are discussed by Geister (1992: p. 56-58) Available meteorological data recorded from nearby San Andrés between 1959 and 1986 (Diaz et al., 1995) are used here, as there are no recorded observations from either atoll. The mean annual air temperature is 27.4°C, with a 1°C range in monthly values. The annual rainfall measured at San Andrés is about 1900 mm, of which over 80% falls between June and November. Winds are trades, from the ENE, with a mean annual intensity of 6.1 m/s and mean monthly variations between 4.5 m/s (May, September- October) and 6.6 m/s (December-January, July). Sporadic storms occur mostly in the second half of the year, with westerlies or northwesterlies attaining speeds over 20 m/s. Albuquerque and Courtown lie in the Caribbean hurricane belt. Hurricanes were recorded in 1818, 1876, 1877, 1906, 1940, 1961, 1971 and 1988 (cf. Barriga et al., 1969: 23; Geister, 1992: 7; Diaz et al., 1995: 112). The latter, 'Joan', on October 20-22 1988, passed westwards 90 km south of San Andrés (about 50 km south of Albuquerque); its eye attained a diameter of about 35 km and the wind reached speeds over 210 km/h (Geister, 1992). The Caribbean Current reaches Albuquerque and Courtown from the NE with speeds of 0.5-1 m/s and passes over the atoll shelf in a SW to W direction, being highly affected by the irregular bottom topography of the shallow-water zones. Waves are generated by the trade winds and approach the atolls from the NE to E, the effective fetch extending for nearly 2,000 km over almost the entire width of the Caribbean Sea. Hence, the considerable amplitude and height of waves breaking on the barrier reef along the windward side of the atolls. The sea surface temperature averages 27.5°C, with mean monthly values ranging between 26.8 (February-March) and 30.2°C (September-October). Surface salinity fluctuates between 34.0 and 36.39/00 (Gonzalez, 1988). Tides on the atolls are mixed with a strong diurnal component. Tidal ranges between 0.3 and 0.6 m are recorded from nearby San Andrés (Geister, 1975). METHODS A preliminary photo interpretation of geomorphological and ecological features of both atolls was done on panchromatic total coverage air photography taken in 1971 and 1984 by the Colombian Geographical Institute (Instituto Geografico 'Agustin Codazzi') approximately 1:22,500, 1:23,000 and 1:30,000, which was then used as basis for field sampling. Preliminary morphological and habitat distribution maps at 1:20,000 scale were drawn combining reef and lagoon photo-patterns defined on the basis of tone, texture and location, as well as bottom topography inferred from bathymetric charts 1:20,000 COL- 203 (Albuquerque) and COL-204 (Courtown). Further detail of the spur-and-grove system of the barrier reefs and lagoonal patch reefs was obtained from oblique aerial colour slides taken on September 29, 1994 from a chartered aircraft at altitudes of 200 to 500 m. During a cruise to the atolls in May-June 1994 aboard the R/V ANCON of the Instituto de Investigaciones Marinas y Costeras (Santa Marta, Colombia), 8 days (May 20-27) were spent at Courtown and 12 (May 28 - June 8) at Albuquerque. A total of 23 (Courtown) and 25 (Albuquerque) observation and sample sites were visited (Figs. 2-3). Location of sample sites included several examples of each of the photo-pattern units, and their exact geographical placement was carried out with an accuracy of 20 m with the aid of a portable Geographic Positioning System (GPS) instrument. SCUBA was used for depths over about 6 m SCUBA was used, otherwise observations were made while skin diving or walking for shallower areas. Observations of bottom types, depth, direction of currents, dimensions and distribution patterns of the reef structures, as well as species composition of dominant biota were recorded on acrylic data sheets. Complementary depth profiles were recorded with the ship's echosounder (28 khz). Final thematic maps at 1:20,000 (bathymetry, geomorphology, bottom habitats, wave exposure) were entered via a digitizing table into a geographic information system (GIS-ILWIS) for storage, processing and further analysis. Morphology and _ habitat classification and terminology vary considerably between authors, and the terms used here to define morphological units and reef zones follow those of Geister (1975, 1977, 1983). Marine habitats are named, where possible, after the substrate dominating macrobiota or substrate features, as was done by Duyl (1985) for the reef environments of the Netherland Antilles. RESULTS Both atolls have the same basic morphological features (Figs. 2 and 3) and, with minor differences, the same marine environments (Figs. 4 to 7). To save space, a general description of each of the morphological units is given below with comments on the bottom habitats found there (map units on Figs. 6 and 7) and, where necessary, on the pecularities of each atoll. Table 1 includes a brief description of the habitats (map units) and allows cross referencing to morphological units. Figures 4 and 5 are representative profiles of the atolls and show the morphological features and bottom habitat distribution along a windward-leeward (right to left) gradient. FORE-REEF TERRACE AND OUTER SLOPE The windward margin of the atolls is characterized by the presence of a gently dipping terrace, descending at low angle (from 6 to 9 degrees) to -24 to -30 m (somewhat deeper in Albuquerque than in Courtown), where a topographical break gives way to a subvertical slope below -30 m. The break marks the transition to the outer slope of the atoll shelf. From a depth of 4-8 m seaward of the barrier reef, to about -15 m, this flat, calcareous platform is, with the exception of scattered gorgonians (Pseudopterogorgia sp.) and large sheets of excavating sponges (Cliona aprica and C. caribbea), mostly devoid of sessile organisms and sediments (‘bare calcareous hard bottom' unit). Low relief calcareous ridge-like structures, with a parallel layout similar to the spur and groove system of the barrier reef (see below), are found along the entire width of the terrace and are more conspicuous on the northeastern section of the atolls. Shallow furrows between these low ridges are filled with coarse sediments and rubble below 18 m. Toward the outer margin of the fore reef terrace, faunal richness and diversity increase gradually, at first especially with brown algae (Sargassum sp., Stypopodium sp.), green algae (Halimeda spp.), massive scleractinians (Diploria spp., Porites astreoides, Siderastrea siderea) and many branching octocorals (Pseudopterogorgia spp., Pterogorgia citrina, Eunicea spp., Plexaurella spp.) (‘Gorgonaceans on hard bottom’ unit). Below 18 m more and more hemispherical scleractinians (Montastraea spp., Colpophyllia natans, and others) and sponges come into sight, as well as coarse sediments that accumulate in shallow hollows. Although coral heads often attain considerable size, they are mostly solitarily, tens of meters apart. In contrast to this, a narrow belt along the transition zone to the outer slope (24 to 30 m) exhibits a well developed coral community, and the calcareous platform appears therefore almost totally covered by corals (Vontastraea spp., Colpophyllia natans, Agaricia agaricites, Dichocoenia stokesii, Stephanocoenia intersepta, among others) , algae (Lobophora sp., Halimeda spp.), sponges and octocorals (‘mixed corals’ unit). The windward outer slope was visited only in Courtown, but its morphology seems to be similar in both atolls, as could be inferred from the recorded bathymetric profiles. The outer slope dips gently (ca. 40-50°) to a sand step beginning at -30 to -35 m. Since this step can be easily recognized on the aerial photographs as a narrow, light grey band along the windward margin of the atolls, thus it seems to be covered by high- reflectance sediments. Below this step, the outer slope decreases subvertically to -400 m and then at lower angle to depths beyond 1,000 m. WINDWARD BARRIER REEF The barrier reef does not completely encircle the atolls, but extends only along the inner shelf from the NNW, N, NE, E, and SE almost continuously for about 5.6 km at Albuquerque and 7.5 km at Courtown. The continuous reef segments are 50-250 m across, being formed by more or less coherent ridges rising from the upper margin of the fore-reef terrace at 5 to 6 m to a reef flat near low tide level. The barrier reef is normally deeply penetrated by surge channels oriented perpendicular to the reef front, forming a typical spur-and-groove system which is easily recognizable on the aerial photographs. Also scattered coral pinnacles rise in some places from about -4 m, just windward of the surf zone, often breaking the surface. The spurs rise 0.5 to 2 m above the adjacent grooves, the latter being 1 to 5 m or more wide and often exhibiting anastomosing bifurcations (Plate 1). At Courtown, the barrier reef is indented at two places, giving the atoll its distinctive kidney shape. Here, the reef crest becomes discontinuous, and a well developed buttress-groove system appears instead (Plate 2), creating a transition zone 300 to 500 m wide between the fore-reef terrace and the lagoonal terrace in its lee. The 2-3 m depth surge channels in this area allow small boats to pass the barrier during calm days. At Albuquerque, the barrier reef is virtually continuous, but on its NE margin a few unusually wide grooves interrupt the reef flat for 10 to 20 m, permitting some waves to pass undisturbed into the lagoonal terrace. At this place, a second, discontinuous barrier reef, located 100 to 200 m behind the former and nearly parallel to it, generates a displaced surf zone clearly observable from the air. The main framework builder in the windward reef flat is the hydrocoral Millepora complanata, which is commonly associated with incrustations of coralline algae. Millepora and the zoanthid Palythoa sp. overgrow the shallowest zone of the barrier reef flat and the upper surfaces of the spurs (‘Millepora-Palythoa' unit), the high surf splashing and washing permanently the emergent colonies. In the buttress-groove area in Courtown, as well as in the second barrier at the NE margin of Albuquerque, Palythoa is generally replaced by Porites porites (growing usually within the Millepora colonies) and crustose forms of Porites astreoides and Diploria clivosa, which overgrow with Millepora the upper parts of the buttresses and the reef flat (Willepora-P. porites' unit). The upright sides of the spurs and buttresses are encrusted with Diploria spp., Porites astreoides and Agaricia agaricites, often assuming a flat form. The hydrocoral Stylaster roseus, the green alga Halimeda, as well as coralline red algae (Porolithon sp.) are also common elements in this zone. Large (up to 2-3 m in diameter) sheet-like excavating sponges (Cliona aprica, C. caribbea) may be fairly common at the sides and bottom of grooves. Leeward of the reef crest, following the ‘Millepora-Palythoa’ unit, cushion-like colonies of Porites porites as well as massive P. astreoides and Diploria strigosa occur at some places among small ridges of Millepora and calcareous boulders (‘Millepora-P. porites’ unit). The displaced rear barrier reef on the NE side of Albuquerque consists likewise of extensive ridges with Millepora complanata and Porites porites rising from - 1.5 to -2.5 m. In some places, like in the NE barrier of Albuquerque and the SE section of Courtown, the coral growth on the rear reef zone extends for about 250 m. There, the end of the barrier reef is marked lagoowards by patchy thickets of Acropora palmata, accompanied by small colonies of Diploria strigosa, Montastraea spp. and occasionally also by cushion shaped colonies of Porites porites (‘Diploria-A. palmata’ unit). In Courtown, the southernmost portion of the barrier reef becomes discontinuous after it bends westward. Numerous pinnacles, constituted mostly by a framework of Millepora at their upper parts, rise in this area from -4 to -5 m reaching up usually to a few centimeters below the surface (Plate 3). At the base of the pinnacles are massive colonies of Diploria spp. commonly more than 2 m in diameter, small thickets of Acropora cervicornis and branching octocorals. The pinnacles are generally arranged in groups, separated by anastomosing sandy channels, with a characteristic wave-induced pattern of ripple marks. Coral rubble (mostly of Acropora cervicornis) accumulates at the sides of the channels. LAGOONAL TERRACE The leeward margin of the reef flat leads down to the lagoonal terrace usually with an abrupt, 0.6 to 1.5 m high, steep slope. The lagoonal terrace is a flat platform attaining a width of 200 to 900 m and increasing in depth from 1 to 3 m towards its inner margin. The lagoonal terrace is one of the most discernible features from the air due to its pale hue. Close to the rear reef, the terrace is covered by rubble (‘hard bottom and rubble' unit), which is gradually replaced lagoonwards by gravel and coarse sand (‘sand and rubble’ unit). The rubble zones are usually arranged in elongated layers perpendicular to the barrier reef, apparently related to the grooves and depressions of the reef crest. The innermost rubble areas on the terrace are overgrown by green (Halimeda, Padina), brown (Dictyota, Turbinaria) and red algae (Amphiroa, Neogoniolithon), as well as scattered encrusting scleractinians (P. astreoides, Siderastrea)('‘rubble with algae’ unit). Some portions of the sandy bottom, particularly in Albuquerque, are sparsely colonized by green algae (Penicillus, Rhipocephalus, Udotea), where juvenile individuals of the gastropod Strombus gigas are fairly common. The lagoonal terrace normally terminates on its lee with a steep 'sand cliff, leading down into the lagoon basin with slopes up to 40°. It represents an accretionary fore-set of fine-grained sediments transported from the reef area to the leeward margin of the terrace The two cays existing in Albuquerque (North Cay and South Cay, Plate 4), as well as East Cay in Courtown (Plates 5 and 6), are sand and rubble accumulations on the lagoonal terrace. Coconut palms, Ficus trees, Scaevola bushes and Tournefortia shrubs are the dominant vegetation. North Cay, at Albuquerque, serves today as military post for the Colombian navy. Several bands of beachrock, paralleling the windward shoreline of these cays, extend eastward on the lagoonal terrace for about 15 (both cays in Albuquerque, Plate 7) to 70 m (East Cay in Courtown), suggesting the location of previous shorelines and thus a lagoonward migration of the cays. The two cays at Albuquerque are presently very close to the leeward margin of the lagoonal terrace. Where submerged beachrock is not covered by rubble and sand, it is mostly overgrown by encrusting coralline and green algae (Halimeda, Rhipocephalus) that contain dense populations of boring sea urchins (Echinometra lucunter). The only sea grasses on the atolls occur on the sheltered leeward side of North Cay in Albuquerque and East Cay in Courtown, where they cover the shallow sandy bottom of the terrace (‘sea grass' unit) The dominant grasses in Courtown are Syringodium and Halodule, whereas Thalassia is more abundant in Albuquerque. The edible urchin, 7ripneustes ventricosus, is abundant in these grass meadows. LAGOON WITH PATCH REEFS The depth of the lagoonal basin is as much as 18 m (in Albuquerque, see below) but generally it varies between 8 and 10 m. Where corals and coral reefs are lacking, the lagoon floor is covered by white calcareous sediments, the coarser fractions of which consist mostly of fragments of coral, molluscs, foraminifera, coralline algae and Halimeda, and rubble. Numerous burrows, mouds and faecal pellets throughout the deeper parts of the lagoon evidence an active bioturbation of the bottom (‘bioturbated sediments’ unit). Green algae (Rhipocephalus, Udotea, Halimeda) grow sparsely around coralline areas forming small patches, where one or more individuals of the Queen Conch, Strombus gigas, as well as patchy aggregations of garden eels (7aeniconger sp.) are occasionally found. A significant portion of the lagoon is occupied by coral reefs, which are highly variable in shape and size, as well as in the dominant scleractinian species, depending mainly on the depth and wave exposure. Reefs occur as solitary mounds and miniatolls, or as ribbon and anastomosing patch reefs. In order to simplify the nomenclature, we divided the patch reefs found inside the lagoon into three main types (map units, see Table 1), according to the dominant scleractinian species: a) emergent to very shallow 'Diploria-A. palmata’ reefs dominated at their summit by Diploria strigosa and Acropora palmata, b) 2-5 m deep 'A.cervicornis' reefs dominated by thickets of Acropora cervicornis, and c) 4- 16 m deep '‘Montastraea spp'.-reefs dominated by one or more species of the Montastraea annularis species complex (see Weil and Knowlton, 1994). At Courtown, lagoon depths vary between 7 and 15 m. Patch reefs cover about 30% of the lagoon floor. In the northern half of the lagoon, where the average depth is about 10 m, a dense net of anastomosing reefs (Montastraea spp.) covers nearly 50% of the bottom. Most of them are low-lying, rising no more than 4 m above the bottom (Plate 8), but some are nearly emergent and form a wave-breaking zone of thickets of A. palmata. The relative coverage of living scleractinians composing these reefs ranges between 10 to 50% from one patch to another. In many places, heads of Montastraea annularis are extent overgrown by filamentous and brown algae (mainly Lobophora variegata), and scattered thickets of Acropora cervicornis are up to 90% devoid of living tissue. Although the bottom in the central and southern portions of the lagoon is predominantly covered by sand, solitary mounds and scattered coral heads are common. In some places of the central area, large aggregations of single coral heads and small thickets of A. cervicornis occur (at present largely dead), forming diffuse, non-cohesive reef communities. The lagoon is rather open to the E and NE, lacking a well defined sill. Nearly 25% of the lagoon floor at Albuquerque is covered by patch reefs. The lagoon exhibits two distinctly depth levels, which are easily recognized from the air because of their different blue hues (Plate 9). A first level, with an average almost constant depth of 9 m, takes up the N and E parts of the lagoon and about 65% of its whole area. The second depth level averages about 15 m and takes up the leeward half of the lagoon to the W and S. Both levels are separated by a meandering ribbon reef of '‘Montastraea spp.', which wanders for nearly 6 km, attains 10 to 30 m in width and rises up to -4 m. On the upper lagoon level there are also several nearly circular shaped miniature atolls which break the surface. These reefs are of type Montastraea spp. at their base but show a typical zonation to the 'Diploria-A.palmata' type towards the summit. Anastomosing patch reefs (Montastraea spp.), with the same basic structure as those at Courtown, are found in the northeastern and southeastern parts of this lagoon level. The deeper level is more sparsely covered by reefs. These are mostly low-lying, isolated patch reefs of the ‘Vontastraea spp' type. The depth of the lagoon diminishes leewards to about -5 m or less and the bioturbated sediments of the bottom give way to a gravel-rubble zone, representing the lagoon sill and the transition zone to the western terrace. According to our observations, lagoonal currents are completely wind-driven and perceptible over the entire water column. Although some differences in direction and 10 intensity were noticed from one location to another, average current velocities of about 2.5 m/min were estimated on the surface at an almost constant wind intensity of 3 m/s in Courtown, and of about 3.5 m/min (wind velocity: 6.5 m/s) in Albuquerque. Considering the rather small size as well as the shallow and open nature of the lagoons, the residence time of lagoonal water masses are thus apparently short, probably not exceeding 24-36 hours. LEEWARD PERIPHERAL REEFS Leeward peripheral reefs are poorly developed in both atolls. In Courtown, the absence of such reefs for more than 2 km results in a widely open lagoon to the west. The northernmost portion of the barrier reef becomes interrupted after it curves southwestward semi-enclosing the northern part of the lagoonal terrace. Southwards, detached reef flats rise from 5 to 7 m depth and break the surface in irregular intervals of 50 to 400 m for about 1.4 km, building the northern peripheral reefs. Wave refraction around the north end of the atoll results in colliding surf from both the NE and the NW. Similarly, beginning at the southwestern tip of the atoll, a series of detached reefs and Shoals semi-enclose the southern third of the lagoon. Some of these reefs are partly emergent at low tide and most of them are almost completely coated by calcareous red algae (Porolithon sp., 'coralline algae! unit), resembling the algal ridges characteristic of Pacific atolls. The algal crust usually exhibits numerous bores caused by chitons (Choneplax lata) similar to the systems described elsewhere in the Caribbean (Littler ez al., 1995). Scattered colonies of Diploria strigosa and Millepora encrust the reef flat, whereas on the subvertical to overhanging walls Dendrogyra cylindrus, Agaricia agaricites, branching octocorals (Plexaura sp., Pseudoplexaura sp.) and bunches of Halimeda are common. Wave turbulence, swift currents and the presence of an intricate system of caves in the northern and southernmost peripheral reefs in Courtown create a bizarre and attractive environment. Sand and rubble accumulations over the larger leeward peripheral shoals at Courtown led to the formation of one island (formerly two, see discussion) and a small sand spit. The island serves today as military post for the Colombian navy (Cayo Bolivar). At Albuquerque, leeward peripheral reefs grow on a shallow, wide sand flat, which represents the lagoon sill. A series of small, low lying reefs enclose the lagoon basin along a wide semicircle between the northwestern tip of the barrier reef and the southern margin of the lagoon. Two navigable channels on the NW and SW breach the flat into the lagoon basin. The peripheral reefs are constituted mainly by large thickets of Acropora palmata, as well as isolated heads of Diploria strigosa and Porites astreoides. Crustose coralline algae (Porolithon sp.), coating large areas of the coral framework, are also major constituents of these reefs. Octocorals and dense beds of brown algae (Dictyota) extensively cover the reef flat bottom. In some places, the scleractinians are dead and overgrown by Dictyota or encrusted by coralline algae. The patch reefs in the southwestern edge, at both sides of the navigable channel, are particularly affected. Here, large thickets of A. palmata were found broken and even overthrown. Large amounts of coral debris were dispersed around the reef flat, including fragments of A. cervicornis, at present an uncommon species in Albuquerque's reefs. This perturbation may have been 11 caused by hurricane ‘Joan’, whose eye passed westward in October 1988 only a few kilometers south of Albuquerque, with winds of more than 200 km/h, which produced very abrasive swells from the south. LEEWARD TERRACE AND OUTER SLOPE In the leeward margin of Courtown Atoll there is not a well defined fore-reef terrace. A slope descends in a distance of no more than 200 to 300 m from the shallow reef flat or the lagoon sill to 17 to 20 m, giving rise suddenly to a subvertical sand slope or to a vertical cliff with locally overhanging ledges. The sand-covered slope of the terrace acts as sedimentary ramp, across which reef detritus falls to greater depths. In the northern and central sections, extensive but somewhat diffuse coral carpets cover as much of the bottom, forming elongated low buttresses in an E-W direction and alternating with rather broad sandy channels. Much of the coral (ca. 75% of the bottom) is at present dead and overgrown by fleshy brown algae (Lobophora, Dictyota), whereas living scleractinians cover no more than 10% of the bottom. Although in the southern half of the terrace coral carpets are scantier and have a patchy distribution, they are better developed and form a distinct hardground on the sandy slope, showing a coverage of nearly 70% of living tissue (‘scattered corals’ unit). At the outer edge of the terrace, the angle of the sandy slope increases to nearly 45°, whereas the reef slope drastically changes to a near vertical wall at about -15 m. Species richness and abundance of scleractinians are very high on the outer margin of the terrace, where massive Montastraea annularis, M. franksi, M. cavernosa and Colpophyllia natans form especially in the southern part, large dome-like structures rising up to 3 m above the bottom (‘mixed corals’ unit). Between these structures usually run ‘sand rivers', which continue as sand falls on overhanging locations along the drop-off. Apart from scattered, small plate-like agariciids such as Agaricia undata, the vertical cliff is mostly devoid of corals and the only organisms attached to the rather smooth substratum are large tube-like and ramose sponges (Agelas conifera, Aplysina spp., Totrochota birotulata), antipatharians and clumps of Halimeda. At the southern locality visited, the cliff remains vertical to about -45 m, where a slanting sand-covered step, about 40 m wide, lines the outer slope of the atoll shelf. In this area, the sand-covered bench deepens at an angle of nearly 30° to about -55 m, where a steep slope continues to greater depths. The loose sand on the slope is composed of Halimeda with accessory shell and coral grains. Large plate-like corals (probably Agaricia and Montastraea) and antipatharians could be observed from above growing along the outer margin of the sandy slope. At another locality, situated in the central section, the drop-off is subvertical to -28 m and is mostly covered by plate-like scleractinians (Agaricia, Montastraea), the sand step is much wider and dips at a lower angle. It seems probable that such a sandy step does occur along the entire leeward margin of the atoll, although the indicative lighter photo-pattern is not always visible on the aerial photographs, possibly due to its variable slope angle and width. In contrasting to Courtown, the leeward fore-reef terrace at Albuquerque is broader, extending for 1 to 1.6 km, and reaching depths greater than 30 m. It is an 12 extensive, gently dipping platform, descending at a low angle (4 to 7 degrees) to about 15 m and then gradually steeper to nearly 40 m, where the subvertical drop-off of the outer slope begins. The bottom in the upper portions of the terrace is mostly covered by ripple marked sand and rubble, although the calcareous hardground appears at certain locations as elongated buttresses, about 1.5 m high, being thus sparcely overgrown by brown algae (Stypopodium, Dictyota) and branching octocorals. Scleractinians are very scarce to depths of about 12 to 15 m (living coral coverage: 5-20%), but their abundance and species richness increase gradually with a simultaneous increase of the slope angle. At the two localities visited, the outer margin of the terrace is marked respectively at -18 and -27 m by a subvertical escarpment, densely covered by plate-like and pagoda-like scleractinians (Agaricia spp., Montastraea franksi), sponges and antipatharians, which descend to nearly -35 m and give way to the accustomed sand step. Such a sand-covered bench or step at 40-45 m depth was recorded on bathymetric profiles at other places of the leeward outer margin of Albuquerque (Fig. 8), and can be distinguished on aerial photographs as a lighter narrow band, outlining almost the entire outer slope around the atoll shelf. DISCUSSION Rather than by its origin (e.g. Darwin, 1842), an atoll is defined by its geomorphic features (Milliman, 1967, 1973; Geister, 1983). Hence, Albuquerque and Courtown may be called atolls. When Milliman (1969) first described the gross morphology and environmental features of the southwestern Caribbean atolls, he was impressed by their close climatologic, oceanographic and geologic resemblance to many Pacific atolls: surrounded by deep water, little seasonal change, appreciable windward fetch, and a Millepora-Palythoa zone that emerges at low tide, resembling somehow the leeward portions of the algal ridge found in Pacific reefs. Besides this, the atolls belonging to the archipelago of San Andrés and Providencia are supposedly the only ones in the Caribbean atolls with a volcanic basement (cf. Milliman and Supko, 1968; Geister, 1992). The atolls of Albuquerque and Courtown share with nearby San Andrés Island and other reef areas of the archipelago, the geological foundations upon which they rest and a similar set of environmental conditions. Other Atlantic atolls, such as the ones found off Belize and the Yucatan Peninsula (Lighthouse Reef, Glover's Reef, Turneffe Islands, Chinchorro Bank, see Stoddart, 1962; James and Ginsburg, 1979; Jordan and Martin, 1987), in the Gulf of Mexico (Alacran Reef, Kornicker and Boyd., 1962) and the Bahamas (Hogsty Reef, Milliman, 1967) show indeed some analogies with Albuquerque and Courtown in their basic morphology, but they have different geological histories. The presence of an extensive windward fore-reef terrace in Albuquerque and Courtown is a characteristic common to most Caribbean atolls. As in the Belizean atolls, the outer margin of the fore-reef terrace is defined by a sudden change of the slope angle at about -20 to -25 m, where the nearly vertical cliff of the outer slope begins. The fore- reef terrace or seaward bank is likely one of the essential morphological differences between Caribbean and Pacific atolls. In the latter, the exposed reef margin margin is the site of most active coral growth, leading to the development of characteristic shelf-edge reefs.(cf. Wiens, 1962). The existence of a sandy step or bench at -35 to -45 m, that outlines the outer slope of the atoll shelf, is also a common feature of the Belizean atolls (cf. James and Ginsburg, 1979). This step, called by some authors the '-40 m Terrace’, is a widespread characteristic of Caribbean reefs. It occurs also in the Bahamas (Zankl and Schroeder, 1972), Jamaica (Goreau and Land, 1974), Curagao (Focke, 1978), San Andrés (Geister, 1975), Providencia (Geister, 1992) and other Caribbean islands. The present morphology of the outer margin in Caribbean reefs has been interpreted in relation to the fluctuations of sea level in the last 80,000 years. As did James and Ginsburg (1979) for the Belizean reefs, and Geister (1975, 1992) for the fore-reef terraces of San Andrés and Providencia, respectively, we may assume that the outer margin of Albuquerque and Courtown, indicated by the '-20 m Terrace’, corresponds to a truncation of the former marginal reef area that occurred before the last interglacial (Sangamon, about 125,000-80,000 years b.p.). In the period between Sangamon and 10,000 years b.p. sea level was not constantly low (about -120 m under present sea level). At least three high stands of sea level took place during that time, reaching to nearly -25 to -40 m below present sea level (Bowen, 1988). The coincidence of the sandy bench in present morphology at -35 to -40 m around both atolls, as well as the occurrence of a deep intertidal notch at this level on the vertical cliff (at least at the visited locality in Courtown), led us to explain this topography as a truncation of the emerging shelf margin during a Pleistocene sea-level stand at about -40 m that may be regarded primarily as an erosional feature. Unlike other Caribbean reefs, such as those off Belize (James and Ginsburg, 1979) and Jamaica (Goreau and Land, 1974), where this feature is now subdued by overgrowing modern facies, no significant accretion to the reef margin seems to have occurred during the Holocene rise of the sea level either in Albuquerque or in Courtown, or in the reefs surrounding San Andrés (cf. Geister, 1975) and Providencia (cf. Geister, 1992), where a truncation of the outer margin at -35 to -40 m and an intertidal notch are very distinctive. The uppermost part of the reef front in both atolls shows a well developed spur- and-groove system, similar to other reef complexes in the western Caribbean, such as those off Belize, Yucatan, San Andrés and Providencia (cf. Stoddart, 1962; James and Ginsburg, 1979; Jordan and Martin, 1987; Geister, 1975, 1992). In some localities, such as the northeastern barrier of Courtown, where the relief between the spurs and grooves often attains more than 3 m, and the grooves penetrate deeply into the reef flat, they are apparently cut into Pleistocene rock, indicating an essentially erosional origin of this system. It acts as an effective baffle for the immense energy expended by incoming surf (Roberts, 1974; Geister, 1982). In other parts of the Caribbean, where the effective windward fetch and the energy of the incoming surf are not as great, the spur-and-groove system may owe much of its relief to differential rates of scleractinian growth (cf. Goreau, 1959). The presence of an extensive lagoonal terrace between the reef crest and the lagoon basin on the windward side, as well as its abrupt transition into the lagoon in the form of a 'sand-cliff, are also characteristics common to most oceanic reefs with a considerable windward fetch, due to active movement of debris associated with the 14 prevailing northeasterly winds and waves. The presence of seagrasses on the lagoonal terrace is conditioned by shelter created on the leeward side of the cays and islands. The depth of the lagoon floor in Albuquerque and Courtown is not very different from most Caribbean atolls, whose average lagoon depth ranges between 10 and 15 m (Milliman, 1973). A singular feature is the existence of two well defined lagoon depth- levels in Albuquerque. It is likely a consequence of the barrier effect of the 'Montastraea spp.'-ribbon reef, which restricts leeward transport of bottom sediments to fill the lagoon basin evenly. The occurrence of anastomosing and ribbon patch reefs covering unusually large portions of the lagoon floor seems to be a rather common feature of oceanic reef complexes in the Caribbean, such as Serrana Bank (Milliman 1969) and Alacran Reef (Kornicker and Boyd, 1962). The NE portion of the lagoon bottom in Providencia Island exhibits also several coalescing patch reefs (J.M. Diaz, J.A. Sanchez and S. Zea, pers. obs., Sept. 1994). It seems likely that the greatest development of anastomosing patch reefs is attained always on the windward side of the lagoon. Contrasting with Pacific atolls, the absence or poor development of leeward peripheral reefs is a characteristic common to most Caribbean atolls (Milliman, 1973). However, Chinchorro Bank and some of the Belizean atolls exhibit a discontinuous leeward reef crest which almost completely encloses the lagoon. Residence time of lagoonal water may hence undergo a notable prolongation in these atolls. Coincidentally, the abundance and development of lagoonal patch reefs in these atolls is apparently much reduced in comparison to Albuquerque and Courtown (cf. Stoddart, 1962; Jordan and Martin, 1987). It seems probable that the residence time of lagoonal water plays an important part in the luxuriance and relative bottom coverage of patch reefs in Caribbean atolls along with other physical factors, such as substrate availability and depth. As stated by Milliman (1969), it seems probable that leeward peripheral reefs in Albuquerque have originated from coalescing patch reefs. On the aerial photographs, most peripheral reefs and the rubble zones surrounding them are arranged in a meander- like fashion. Former ribbon and cellular reefs on the leeward lagoon margin have apparently been damaged again and again by storms and hurricanes, leaving only the most resistant frameworks of Acropora palmata and coralline algae, which built such peripheral reefs. In Courtown, leeward peripheral reefs have developed only in the NW and SW parts of the atoll, where the windward barrier reef bends southwestward at its northern end and northwestward at its southern end. They are heavily exposed to colliding surf from both the NW and the NE (or SW and NW) and are formed mainly by a framework of coralline algae (Porolithon sp., Titanoderma spp., Lithophyllum sp.) comparable to that of algal ridges. Although algal ridges had been thought characteristic of the Indo- Pacific region until recently (Frost and Weiss, 1975), the southernmost leeward peripheral reef in Courtown, with its emergent crest, represents in fact a true algal ridge, such as those described recently elsewhere in the Caribbean (Glynn, 1973; Adey, 1975; Adey and Burke, 1976). This feature was apparently overlooked by Milliman (1969), who refers to it as a ‘small rocky spit, composed of massive coral debris’. Although not so well developed, similar structures also have been recognized adjoining the NW end of the barrier reef in nearby San Andrés by Geister (1975). Interesting discussions concerning the existence and development of Caribbean algal ridges are found in Adey and Burke (1976), Stoddart (1977) and Littler et al (1995). At present, two cays exist in Albuquerque, both lying on the lagoonal terrace. Their position, size, and shape have not changed significantly in the last 25 years, except that North Cay has currently a more rounded shape than in the map of Milliman (1969) and on the aerial photograph taken 1971. On the 1984-photograph it exhibits approximately the current shape and size. The western and southern shores of this cay have been dammed with piles of Strombus shells by the marines of the Colombian navy. On the other hand, islands and cays in Courtown experienced remarkable changes in number, size and shape since that time, and it seems likely that further changes are even now taking place. Milliman (1969) mentioned four small cays, a sand spit and a rocky spit. Sand Cay and East Cay lay close together on the lagoonal terrace and have currently coalesced in an arrow-shaped island (about 800 m long), which seems to grow further to the NW by accretion of sand and rubble (Plates 5 and 6). Of the formerly two cays sitting on leeward peripheral reefs, Middle Cay was the only one visited by Milliman, who noticed the presence of Yournefortia and Scaevola bushes and even some native fishermen living on it. This cay might have disappeared between 1966 and 1971, since no trace of it can be seen in the aerial photographs taken in August 1971. On the contrary, West Cay (currently called Cayo Bolivar and serving as military post) and the sand spit have experienced little change. The shallow bottom (1-2 m depth), where Middle Cay lies, is currently covered with rubble and coral debris. It is not known if the.disappearance of the cay was a slow erosional process that took place within five years or a rapid loss produced by a forceful weather event. The latter seems less probable, since the only hurricane recorded between 1966 and 1971 affecting this area, 'Irene' in 1971, had only trivial consequences in nearby San Andres (IGAC, 1986). Although a detailed checklist of scleractinians from Albuquerque and Courtown has not yet been published, our survey indicates no noteworthy differences in species composition and structure between the reefs of both atolls. It can be stated however that the reefs in both atolls show a highly diverse fauna of about 40 species, not significantly diverging from those known from neighbouring San Andrés and Providencia, where 44 and 43 species have been respectively recorded (Geister, 1975; 1992). The distribution pattern of reef framework associations in both atolls, at least in shallow-water to about 15 m, is highly controlled by wave-energy and corresponds well to the ‘wave zones’ model postulated by Geister (1977). With the exception of a ‘Porites zone’, each of the most important reef framework associations recognized in the Caribbean Sea were found in Albuquerque and Courtown. Only the names employed by Geister (1977) for his 'Melobesiae-zone' has been modified to designate the 'Coralline algae! unit (including the algal ridges) in our maps. In spite of a generally similar distribution pattern of reef framework associations, there are some qualitative differences between Albuquerque and Courtown. Neither 'A.cervicornis' reefs nor a '‘coralline algae’ (or algal ridges) unit occur in Albuquerque. Due to the interrupted windward reef crest in Courtown (ie.,discontinuity of the ‘Millepora-Palythoa” unit), medium-energy waves can penetrate in some places into respective rear reef and lagoonal areas, leading to a better development of 'Diploria- A.palmata" and 'A.cervicornis' reefs in this atoll. Contrary to Albuquerque, Courtown lacks a gently dipping and extensive leeward terrace, which represents a highly abrasive environment during storms and hurricans coming usually from the SW. This is seemingly the main reason for a much reduced ‘scattered corals' unit and the lack of a ‘bare calcareous hard bottom’ unit there. Although detailed information about the current conditions of reef health in these atolls will be presented and discussed elsewhere, some preliminary statements can be made here. In the description of habitats presented above, we mentioned several signs that are indicative of some degradation of the coral reef environment in both atolls. Proliferation of algae overgrowing scleractinian colonies, low proportions of living coral cover at several sites, abundance of heaps of skeletons of recently dead scleractinians (i.e. Acropora spp.), as well as a noticeable depletion of commercial organisms, such as queen conchs (Strombus gigas), lobsters (Panilurus spp.), snappers (Lutjanidae), groupers (Serranidae) and turtles, are the most evident signs of degradation. Although no human development exists in the atolls, they have been visited for many years by San Andrean and Providencian natives for fish and turtles. In contrast to the condition in 1944, when Fowler reported abundant fish and lobsters, by 1966 the populations of these resources seemed to be low at Courtown, possibly a result of the increasingly fishing pressure caused by overexploitation of Strombus, lobster and fish stocks at San Andrés (Wells, 1988). The health condition of reefs in these atolls is even at several sites not significantly different from those around the densely populated San Andrés island (cf. Diaz ez. al., 1995), indicating that, besides local human factors (i.e. sand mining, siltation, pollution) and local natural agents (i.e. hurricanes), recent coral mortality is highly associated rather to a generalized phenomenon of coral decline occurring in the Caribbean from beginning of the 1980's (Hallock ez al., 1993; Ginsburg, 1994). Overfishing has also been recently recognized as an indirect agent of coral mortality (Hughes, 1994). ACKNOWLEDGMENTS The authors express their gratitude to Luz S. Mejia, Guillermo Diaz (INVEMAR, Santa Marta) and the crew of the R/V 'Ancon' for assistance in the field surveys. We extend our appretiation to Dr. Jorn Geister (University of Bern, Switzerland) for his helpful discussions and encouragement to make possible the flight over the atolls, as well as friendly loan of the photos included in Plates 2, 4, 5 and 9. We thank Martha Prada for her friendly hospitality at San Andrés. For their help in map digitizing and improvement of computer drawings we are indebted to the students P. Sierra, J.A. Pulido and N. Ardila. This study has been funded by the Instituto Colombiano de Ciencia y Tecnologia (COLCIENCIAS, Grant No. 2105-09-023-93), the Instituto de Investigaciones Marinas y Costeras (INVEMAR, Santa Marta) and the Universidad Nacional de Colombia. REFERENCES ADEY, W.H., 1975. 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Coral reefs of the world. Vol 1: Atlantic and Eastern Pacific. UNEP/IUCN, Gland, Switzerland, 373 pp. WEIL, E. and N. KNOWLTON, 1994. A multi-character analysis of the Caribbean coral Montastraea annularis (Ellis & Solander, 1786) and its two sibling species, M. faveolata (Ellis & Solander, 1786) and M. franksi (Gregory, 1895). Bull. Mar. Sei, SS) rls IES» WIENS, H.J., 1962. Atoll environment and ecology. Yale Univ. Press, New haven and London, 532 pp. ZANKL, H. and J.H. SCHROEDER, 1972. Interaction of genetic processes in Holocene reefs off North Eleuthera Island, Bahamas. Geol. Rundschau, 61(2): 520-541. 21 Table 1. Marine habitats (Map units) of Albuquerque and Courtown atolls with their corresponding geomorphological units and absolute and relative area. Habitat unit ‘pioturbated sand’ ‘rubble with algae’ ‘sand and rubble’ ‘A.cervicornis’ ‘scattered corals’ ‘rubble on hard bottom’ ‘Gorgonaceans on hard bottom’ ‘mixed corals’ ‘Diploria-A. palmata’ Geomorphol. units (depth) Lagoon (6-18m) Lagoonal terrace (1-2m) All zones Lagoon (3-5m) Leeward terrace (15-30m) Laggonal terrace (1-2m) Fore-reef terrace (15-30m) Fore-reef and Leeward terraces (25-37m) Lagoon and Lagoonal terrace (0.5-3 m) Brief description Calcareous sand (Halimeda, coral, shells) with many burrows and mounds (Arenicola, Callianasa). Coral debris with rodoliths formed by coralline algae mostly overgrown by brown algae. Bare coarse to medium sand with scattered coral rubble and algal rodoliths. Patch reefs dominated by thickets of Acropora cervicornis, scattered coral heads (Siderastrea, Montastraea) and plexaurid octocorals. Scattered massive and hemispheric scleractinians (Siderastrea, Colpophyliia, Diplona, Montastraea), gorgonaceans and fleshy brown algae. Rather barren bottom with coral debris sometimes encrusted with coralline algae. Rather flat bottom with luxuriant growth of gorgonaceans (Pseudopterogorgia spp., Pterogorgia, Plexaurella, Eunicea, Munceopsis, etc), scattered massive scleractinians, many fleshy algae and large sponges. Diverse scleractinians (Montastraea franksi, Diploria, Colpophyllia, Pontes, Mycetophyliia, etc.) gorgonaceans, sponges and Halimeda. Moderate wave exposed reefs dominated in the shallow zones by thickets of Acropora palmata, massive Diplona stngosa and encrusting Porites astreoides. Area (Ha) Courtown Albuquerque 631.8 841.2 (12.8%) (11.7%) 127 122.4 (2.5) (1.7) 1270 237 (25.5) (33.2) 28.2 - (0.6) 226.6 357.8 (4.6) (5) ' 361.3 260.2 (7.3) (3.6) 910 327.4 (10.3) (4.5) 134.1 180.2 (2.7) (2.5) ] 29.6 107.6 (0.6) (1.5) 22 Table 1. continued. ae iad eos Brief description Area (Ha) uni units (depth) Courtown Albuquerque ‘ : Patchy seagrass meadows with Thalassia, sealgiiass goonal Halodule and/or Syringodium growing on 35.5 3.8 aay sandy bottom. (0.7) (<0.1) -om ‘ , Emerging sand and rubble accumulations, land (cays) Fee mostly vegetated with shrubs (Scaevola, 9.2 78 eared Tournefortia), coconut palms or Ficus trees. (0.2) (0.1) eewarl peripheral reefs ‘Millepora- Barrier reef Highly Wave-exposed reefs dominated by 195 195.4 ; 0-3 m) Millepora complanata and Palythoa sp., (4) (2.7 Palythoa ( mostly accompained by crustose coralline 1) algae. ‘Millepora- Barrier reef ear surf zone of the barrier reef. Millepora 107 59.2 pila 0-3 m) complanata, Portes ponites, P.astreoides (2.2) (0.8) P.porites ( and Diploria strigosa. ; : ‘Montastraea Lagoon Ribbon and anastomosing patch reefs 325 1 304.1 ‘ 5-15 m) dominated by massive Montastraea (6.5) (4.2) Spp. ( annulans and M. faveolata, brown algae : : (Lobophora-Dictyota) and some octocorals. bare Fore-reet's War otioctigiemanidalytitcl) noggiiS.6 trafi224 calcareous hard terrace algae, scattered sea fans, brown algae. (8.4) (17.2) bottom (3-15m) Heavily excavated by sheet-like sponges (Cliona spp.). ‘outer slope’ Fore-reef Vertical to subvertical drop-off of the atoll 522.4 806.2 shelf. Sedimentary ramp or subvertical 10.4 11.2 and calcareous wall (covered or not with plate-like (10.4) (11.2) Leeward scleractinians, sponges and antipatharians). terraces (>35m) ‘ : ’ Wave-exposed reefs, almost completely z SEU I US cele Leeward covered by encrusting algae (Porolithon) - - Peete building algal-ridge-like emerging crests. (0.5) reets (0-5m) eee 4953.3 7172.5 a 23 BB RNK EEA ACS @) es CE HOS A ASKS —S YA Y SP YS LE PAWN ae Lagoon with patch reefs Windward barrier reef ca Cays oa Fore-reef terrace and outer slope KA Leeward terrace and outer slope ¢| Lagoonal terrace Leeward peripherical reefs Sample sites Figure 2, Geomorphological units and visited stations at Albuquerque Atoll. Straight lines mark the location of the schematic profiles of Fig. 4. 24 81° 28'W 12° 28’N Lagoon with patch reefs | Ninaward barrier reef ow Cays [+ Fore reef terrace and outer slope RY Leeward terrace and outer slope 0 500 1000 1500 m Figure 3. Geomorphological units and visited stations at Courtown Atoll. Straight lines mark the location of schematic profiles of Fig. 5. A. OUTER LEEWARD TERRACE PR LAGOON LAGOONAL TERRACE] BR | fendez Ores 0 1 2 3 T4 15 16 7 km PEN Fk Ce eRe od pe ta eer a pr ere erence REE level depth = MO7TLLG aed SYP? seattered corals — mixed OTa 60 m L— B. OUTER LEEWARD TERRACE lP.r| LAGOON Sty’ | LAGOONAL TERRACE B.R| F.R.TERRACE OUTER (0) 1 2 3 4 5 6 7 8 9 10km se siti on ear ee ee depth bb with a ip Montastraea spp. gorgonian on hard bottom re < QO mixed corals 60 mL_— Figure 4. West-East schematic profiles (straight lines in Fig. 2), showing the different geomorphological and habitat units of Albuquerque Atoll. P.R- peripheral reefs, B.R- barrier reef, F.R.- fore-reef. 25 26 § 3 > © [ste | ee | LAGOON LAGOONAL Terrace] 8.«| Seb 3) i 2 4km seq depth Millepora—Palythoa Le coralline algae rubble with algae i gorgonians on.hard bottom ool of Montastraea spp. eee scattered corals mixed corals 50 m B. Lev [eri ¢ said GOON MMA Lee 0 Fr 19 aie 4 5 km On == eye Oe SO ee ie Nim oe depth Millepora—Palythoa = cervicornis = algae or grass meadows = coralline algae gorgonians on hard bottom il Montastraea spp. = scattered corals mixed corals 50) tm == Figure 5. West-East schematic profiles (straight lines in Fig. 3), showing the different geomorphological and habitat units of Courtown Atoll. P.R- peripheral reefs, B.R- barrier reef. 77) = Outer slope Z ward bottom with rubble Fra Scattered corals Sea grasses E24 Rubble with algae —] Gorgonaceans on hard bottom Ej Diploria—A.palmata 12°07’N = lap) w 2 2] Bare calcareous hard bottom Montastraea spp. 4 Millepora—Palythoa | Bioturbated sediments ai Millepora—P.porites EC] Sand and rubble Figure 6. Distribution of bottom habitats and reef types at Albuquerque Atoll (for brief description of map units see Table 1). 28 Diploria—A.palmata Bioturbated sediments = Millepora—Palythoa A.cervicornis fj Gorgonaceans on hard bottom Eel Millepora—P.porites (2) Bare calcareous hard bottom Scattered corals [_] Sand and rubble E2 Montastraea spp. @4 Coralline algae [] Sea grasses —] Outer slope 22 Mixed corals Rubble with algae Hard bottom with rubble 500 1000 1500 m 0 Figure 7. Distribution of bottom habitats and reef-types at Courtown Atoll (for brief description of map units see Table 1). Figure 8. Echosounder bathymetric profile of the leeward terrace and outer slope at Albuquerque atoll. Note the presence of a truncation (sandy bench or step) at about -40 m depth on the outer slope. Plate 1. The spur-and-groove system of the windward barrier reef. The spurs are overgrown on the top by Millepora complanata and by Porites spp. and crustose coralline algae on the sides, whereas the narrow groove is filled with sand (Courtown, 22 May, 1994). we 30 Plate 2. Oblique aerial view to the N, showing the buttress-groove system on the central portion of the windward barrier reef at Courtown atoll (Sept. 29, 1994). Plate 3. Rounded pinnacle (left) and narrow pillar formed by Millepora spp. at the SW section of Courtown atoll, where the barrier reef becomes discontinuous (27 May, 1994). 31 Plate 4. Oblique aerial view to the W of Albuquerque atoll showing the two cays lying close to the leeward margin of the lagoonal terrace (Sept. 29, 1994), Plate 5. Oblique aerial view to the NW of Courtown atoll. The arrow-shaped island in the center right is East Cay, which currently is connected with Sand Cay by a sand bar. Sand Cay grows seemingly further to the NW by recent gradual addition of sand (Sept. 29, 1994). 32 Plate 6. East Cay, Courtown atoll, looking SE along the sand bar which at present connects this cay with Sand Cay (21 May, 1994). Plate 7. East shore of South Cay, Albuquerque atoll. Note the conspicuous band of beachrock parallelling the shore line ( June 6, 1994) 33 Plate 8. Lagoonal patch reef in the upper depth level at Albuquerque atoll, made up mostly by Montastraea annularis and M. faveolata (June 7, 1994). Plate 9. Oblique aerial view to the SE of Albuquerque atoll. Note the two different hues of the lagoon basin denoting the two depth-levels of the lagoon floor (Sept. 29, 1994). ) now tga ‘A ots i ‘en iil ) aimbogeal, MM bas ‘svn voruncnnahs yd avid sal! OA jlote ie) #8 ie iShabt ut eu inet ATOLL RESEARCH BULLETIN NO. 436 CORAL FAUNA OF TAIPING ISLAND (ITU ABA ISLAND) IN THE SPRATLYS OF THE SOUTH CHINA SEA BY CHANG-FENG DAI AND TUNG-YUNG FAN ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. APRIL 1996 115°E Taiwan ¢ Pratas I. Hainan Paraceles I. a o rh) %2¢ (a ‘ 1S) 15°N Vietnam SOUTH CHINA SEA ~ cA ~ 7 2 : ~ , os XN Taiping I. \ y Palawan ping fe] : 114°21E Taiping Island E Fig. 1. Locations of the survey sites (A-G) at Taiping Island in the Spratlys of the South China Sea. CORAL FAUNA OF TAIPING ISLAND (ITU ABA ISLAND) IN THE SPRATLYS OF THE SOUTH CHINA SEA BY CHANG-FENG DAI AND TUNG-YUNG FAN ABSTRACT The coral fauna of the Taiping Island (Itu Aba Island) in the Spratlys of the South China Sea was surveyed on April 19-23, 1994. A total of 163 species of scleractinians in 15 families and 56 genera; 15 species of alcyonaceans in three families and five genera; and six species of gorgonaceans in four families and five genera were recorded. The coral communities of the Taiping Island were dominated by scleractinian corals with high species diversity and coral cover found on the lower reef flat at depths between 1 and 3 m. Alcyonaceans and gorgonaceans are mainly distributed on the reef slopes at depths below 15 m. Wide reef flats and reef terraces exist on the east and west sides of the island indicating that the reef development is better in these areas. Species diversity of coral communities was the highest on the east side and the lowest on the west side of the island. The depauperate coral fauna on the west side is possibly related to the strong SW monsoon during summer and autumn. In comparison with other tropical coral reefs, species diversity and abundance of coral communities of Taiping Island are relatively low. Dead coral skeletons and debris were widely spread on the reefs below 3 m deep and only small colonies were found. These facts indicate that coral communities of Taiping Island may have been heavily damaged by natural catastrophes or artificial destruction during the last decade. The possible destruction forces are typhoon disturbances and sea warming events. INTRODUCTION The South China Sea, situated between the Indian and Pacific Oceans, has an historical importance in politics, economics, military affairs and transportation (Gomez, 1994). As the South China Sea is surrounded by continental Asia and many islands, it is generally recognized as the major marginal sea in Asia. Major islands in the South China Sea such as Tungsha Island (Pratas Island), Xisha Islands (Paracel Shoals) and Nansha Islands (Spratly Islands) are reef islands. Most reef islands are atolls or emergent islands, which are mainly composed of coral debris and sand. The emergent islands constitute only a small portion of the reefs; the major parts are underwater reefs, shoals and banks. Institute of Oceanography, National Taiwan University, P.O. Box 23-13, Taipei, Taiwan, R.0.C. Manuscript received 8 September 1995; revised 8 March 1996 The Spratly Islands, consisting of some 600 coral reefs and associated structures scattered across an area north of Sabah and southern Palawan stretching for more than 500 km, are a group of atolls, islets, and sea mounts in the South China Sea. The structures which protrude above the sea surface at high tide include at least 26 islands and seven exposed rocks (McManus, 1992). Taiping Island, or Itu Aba Island, is one of the major islands in the Spratly Islands. The Indo-Pacific region, which includes the Spratlys, is characterized by a high diversity of marine organisms. Among reef building corals, for example, the region in which the Spratlys reside includes at least 70 genera (Veron, 1986, 1993). Inthe coral reef ecosystem alone, more than 400 species of corals (Veron and Hodgson, 1989), 1500 species of reef fishes and 200 species of algae are found in this area (McManus, 1994). The exact number of all marine species in the South China Sea is difficult to estimate given the inadequate state of taxonomy, but the total number of species to be found at all depths in the Spratlys certainly ranges to the tens of thousands (McManus, 1992). The marine ecosystem of the South China Sea can be assumed to be dependent on the Spratlys, at varying levels, for sources of larvae of renewable resources. Due to prevailing monsoonal currents, the Spratly reefs may serve as sources of larvae that could recruit to the disturbed coral reefs in the South China Sea (McManus, 1994). The semi- enclosed nature of the South China Sea and hydrodynamic patterns prevailing in the area could explain this linkage of coastal ecosystems in terms of nutrient level and fauna. It is very likely that the Spratly Islands and similar groups of uninhabited reefs serve as a mechanism for stabilizing the supply of young fish and invertebrates to these areas. This becomes increasingly important wherein coastal populations of adult fish decline, as appears to be the case in many coastal reefs of the Philippines and elsewhere. The dispersal of larvae from the Spratlys possibly contribute to the coral reef fishery in the region. The contribution of coral reef fishery to the national fish production of countries bordering the South China Sea varies between 5-60% (McManus, 1994). Thus, the Spratlys could be considered as a “saving bank” where commercially important fish and invertebrates are saved from overharvest and supply a constant flow of larvae to areas of depletion. Coral reefs are widely distributed in shallow water areas in the South China Sea. The high spatial heterogeneity and productivity of coral reefs provide not only various habitats for marine organisms but also feeding and nursery grounds for fishery resources such as fish, shells, crustaceans and cephalopods. Flourishing coral reefs also constitute beautiful underwater scenery that are valuable resources for the development of touristic industry. As corals play a key role in marine ecosystems of the South China Sea, a better understanding of the coral fauna in this area is necessary for conservation and management of the marine resources in the future. Several scientific expeditions in the South China Sea over the last 50 years have provided oceanographic information and taxonomic listing of marine organisms, mainly fishes. Although corals are widely distributed in the South China Sea, the coral fauna of 3 this area is poorly documented because of its remoteness and difficulty of access. Bassett-Smith (1890) first described corals from Tizard Bank. Ma (1937) studied the growth rates of scleractinian corals from Tungsha Island (Pratas Island). In recent years, a few expeditions have been conducted to investigate the fauna and flora of the South China Sea (Yang et al., 1975; Zou, 1978a, b; Fang et al., 1990). These studies have provided valuable information for a preliminary understanding of the coral fauna of this area. However, in comparison with the vast area of the South China Sea, these studies have only covered a very restricted area. Studies on the coral fauna in other areas are thus necessary. We sought to provide baseline information for resource conservation and exploitation of Taiping Island (or Itu Aba Island). The objectives of this work were to survey and to describe the distribution of coral reefs and reef topography, to provide an inventory of coral species and their estimated relative abundance, and to identify special coral biotopes. STUDY SITE AND METHOD Seven sites around Taiping Island (Fig. 1) were surveyed on April 19-23, 1994. Taiping Island (10°23'N, 114°22'E), located on the northwest side of Tizard Bank, is one of the major islands on the west side of the Spratly Islands (Nan-sha Islands). The island, with an area of 0.49 km’, is about 1300 m long and 350 m wide (Fig. 3). The climate is tropical oceanic. The average water temperature is about 28-29°C. The island is influenced by seasonal monsoons. The northeast monsoon blows from October to March, the southwest monsoon from May to October. The current flows southeast during the former and east or north during the latter (UNEP/ITUCN, 1988). Coral reefs were surveyed by snorkeling and scuba diving. Reef topography, coral Species, community types and estimated coral cover were recorded. The relative abundance of each coral species was estimated according to the number of colonies encountered during each survey as common with more than 50 colonies, occasional with about 10-50 colonies, or rare with less than 10 colonies. Underwater camera and video were used to record photographs of coral colonies and reef topography. Coral species were identified in the field. Whenever confronted with an uncertain species identification, a piece of coral skeleton was detached and brought to the laboratory for further identification. The identification of species was based on Veron and Pichon (1980, 1982), Veron and Wallace (1984), Veron et al. (1977), Veron (1986), Dai (1989), Hoeksema and Dai (1991), and Dai and Lin (1992). RESULTS AND DISCUSSION Description of Reef Topography and Coral Community Site A is located on the south side of the island. The substrate of the upper reef flat at 1-2 m depth is covered with sand and seagrasses. On the lower reef flat at 2-4 m depth, there are abundant massive and stoutly branching colonies of Porites, Acropora and Pocillopora spp. Below the reef flat at depths between 4 and 15 m, there is a steep slope; only a few foliaceous Montipora and branching Acropora colonies were found on the surface of the slope. At depths between 15 and 21 m, it is a reef terrace. The substrate is flat and composed of coral debris with some ridges and grooves (Fig. 2a). The coral cover is less than 5%; only a few small colonies are scattered on the substrate. The species diversity was quite high, more than 67 species were recorded. The most abundant species at this site is the octocorallian, /sis sp. (bamboo coral). They can form large colonies of 1 m long and in dense assemblages at some locations. Scleractinians found here are mainly species of Montipora, Favia, Favites, Goniastrea and Cyphastrea. They typically exist as small colonies with a diameter less than 10 cm. The widespread coral debris covering the substrate was mainly Acropora and Pocillopora skeletons indicating that there were flourishing branching coral communities in the past. The scarcity of coral species and scattered small colonies indicate that the community might have been destroyed recently and that recovery is slow. Site B is located on the southeast of the island. Reef topography is similar to Site A. There is a reef flat about 50 m wide at depths between 0 and 4 m. Living coral cover on the reef flat exceeds 50%, but a trend of decrease toward the west is evident. Scleractinian corals of about 120 species were found. Species commonly occurring on the reef flat were stoutly branched colonies of Pocillopora damicornis, P. verrucosa, P. eydouxi, Acropora monticulosa and A. gemmifera (Fig. 4). Colonies of A. digitifera, A, palmera, Favia speciosa, Leptoria phrygia, Platygyra lamellina and the hydrocoral, Millepora platyphylla were also commonly found on the reef flat. These species generally form large colonies with diameters greater than 1 m. Corals existing on the flat are mainly massive, encrusting and stoutly branched forms. The colony morphology of corals of this area indicates that the reef flat is exposed to strong wave action. Below the reef flat on the seaward side between 5 and 18 mis a steep slope on which coral cover was less than 5%; only a few coral colonies were found to grow on the surface of the slope. A few solitary corals of Fungia spp. and several large colonies of the blue coral, Heliopora coerulea, were found on the sandy grooves. Below 18 m the bottom is sandy and no coral was found. Site C is situated on the west of the island. It is characterized by a wide reef flat that extends westward to over 500 m from shore with depths about 3-8 m (Fig. 2b). On the surface of the flat, there are low reef ridges alternating with shallow grooves running in the NE-SW direction. Currents of this area are generally strong especially during flood and ebb tides. This area is also exposed to strong waves during the summer monsoon. The 5 substrate on the upper reef flat was characterized by a dense seagrass bed. The lower reef flat was covered with dead coral skeletons; some of them were clearly identifiable based on skeletal features. Few small colonies were found and the coral cover was less than 2%. These phenomena indicate that the coral communities might have been destroyed during the past decade. Some small soft coral colonies such as Sarcophyton spp. and Lobophytum spp. were scattered on the substrate (Fig. 5); few attained a diameter of 50 cm. Site D is located on the northwest side of the island. The reef flat has a width about 100 m and stretches from 1 to 6 m deep (Fig. 2c). Coral communities on the reef flat can be divided into two zones. In the upper zone between | and 3 m deep, coral cover is higher than 50%. Species common in this zone are Favia, Favites, Goniastrea, Coeloseris mayeri and Pavona spp. Some large colonies with diameters greater than 1 m were found. In the lower zone between 3 and 6 m deep, coral diversity is low and coral cover is less than 10%. The reef surface is covered with dead coral skeletons and algae. Below 6 m, there is a steep drop-off, descending at a nearly perpendicular angle to a depth about 60-80 m. On the wall of this drop-off, there are colonies of Dendronephthya spp., Junceella fragilis and Isis sp. Scleractinians were rare; only few small colonies of foliaceous corals were found to grow on the slope. The coral cover is less than 5%. However, sponges, bryozoans and other sessile invertebrates are abundant. Site E is situated on the northeast side of the island. Reef topography and coral fauna of this site are similar to those of Site D. On the upper zone of the reef flat, the coral cover was higher than 50% and approximately 100 scleractinian species were found. Among the most abundant species are Pocillopora verrucosa, P. eydouxi, Acropora digitifera, Heliopora coerulea, and Millepora platyphylla (Fig. 6). Species of Montipora, Porites, Favia, Favites and Goniastrea are also common in this zone; most of them are massive, encrusting or stoutly branched forms, with colony sizes often less than 30 cm in diameter. At the lower zone between 3 and 6 m deep, the substrate 1s covered mainly by dead coral skeletons and green algae, Caulerpa spp. The coral cover is less than 5% in this zone. There is a steep drop-off below 6 m; many large gorgonian and antipatharian colonies were found overhanging on the slope. Sponges, bryozoans, crinoids and other groups of marine invertebrates are abundant, which comprise a rich benthic fauna and colorful scenery (Fig. 7). Below 35 m the bottom 1s sandy and no coral was found. Site F is located on the east side of the island. The reef flat is wider in the north where it extends seaward to approximately 500 m from shore but becomes narrower to the south (Fig. 2d). Dense coral cover (>50%) and high species diversity were found on the upper part of the reef flat at depths between 1 and 3 m. More than 100 scleractinian Species were recorded, most of them were small colonies. Species commonly present in this area are Pocillopora damicornis, P. verrucosa, Acropora digitifera, Cyphastrea chalcidicum and Favites abdita. Coral cover and species diversity are low on the lower part of the reef flat. Less than 5% of the substrate was covered by corals and only few small colonies were found. Below 6 m there is a steep drop-off that extends to about 30 m and reaches the sandy bottom. The most peculiar organisms on the surface of the slope are 6 many colorful soft corals, Dendronephthya spp. hanging on the wall. Other corals are rare and scattered. Below 35 m the bottom is sandy and no corals were found. Site G is located on a reef ridge on the southeast of the island. The reef ridge is separated from the island by a trough approximately 20 m deep (Fig. 2e). The surface of the ridge is smooth and about 7 m deep. More than 70 species of scleractinian corals were found on the top of the ridge, mainly species of Acropora, Favia, Favites, Goniastrea, and Fungia. The coral cover is about 30-40%. Many colonies of solitary corals such as Fungia cyclolites, F. costulata, F. tenuis, F. fungites, F. scutaria and Herpolitha limax were found on the sandy grooves. The edge of the reef ridge is about 8 m deep. Below 8 m there is a steep slope down to approximately 37 m. There are several 7ubastraea micranthus colonies growing on the upper part of the slope. The lower part of the slope between 20 and 37 m deep is covered by thick patches of Dendronephthya colonies (Fig. 8). These colorful soft corals, when fully extended, form a gorgeous underwater "flower wall”. The slope reaches the sandy bottom at 37 m. Coral Fauna A total of 163 species in 15 families and 56 genera of scleractinians; 15 species in three families and five genera of alcyonaceans; and six species in four families and five genera of gorgonaceans were recorded during this survey (Table 1). The results showed that coral communities of the Taiping Island are dominated by scleractinian corals with high species diversity and abundant coral cover found on the reef flat between 1 and 3 m deep. Alcyonaceans and gorgonaceans are relatively rare and their distributions are limited to reef slopes at depths below 15 m. Although the coral fauna varied slightly among the surveyed sites, species compositions of the coral communities are similar and can be regarded as typical of tropical reef communities. The abundance of small coral colonies indicates that coral communities are in their early stages of succession (Grigg, 1983). As early succession communities generally have high species diversity (Connell, 1978), this conditions may also relate to the high diversity of coral communities at Taiping Island. In comparison with the known coral fauna of other reefs in the South China Sea, the number of scleractinian species recorded during this study exceeds those of Tungsha Island (Pratas Island, 101 species; Dai et al., 1995) and Xisha Islands (Paracel Shoals, 127 species; Zou and Chen, 1983). In general, the species composition of the coral fauna among these islands is similar. Biogeographically, these coral fauna belong to the Indo- Pacific province. Because Taiping Island is situated at a lower latitude and closer to the area of highest coral diversity, it is natural that its coral fauna is more diverse than those of other reefs in the South China Sea. According to the biogeographical location of Taiping Island, this island is expected to have more than 70 genera and 400 species of scleractinians (Veron, 1993). However, during our brief survey to the island, only 51 genera and 163 species were recorded (Table 1). Further intensive surveys of adjacent islands may reveal more species. 7 The coral reef of Taiping Island is a typical oceanic reef. It has a wide, shallow reef flat and a steep drop-off on the edge of the flat. The reef flat is a site of intensive coral calcification that forms the reef framework. The substructure of this region is invariably composed of large, massive, interlocking colonies of hermatypic corals cemented by calcareous algae. The drop-off borders the reef framework and generally descends to depths below 30 or 60 m. At the base of the drop-off there are abundant coral debris and accumulation of sediment. These facts indicate that physical and biological destruction of the reefs is relatively high and debris produced through these processes are transported to a deeper zone at which accumulation occurs. The development of reefs on the southwest and northeast sides of Taiping Island is better than that of other areas. On both sides there are wide reef flats extending beyond 500 m from shore which basically conform to the shape of the island. Such a pattern of reef development is likely related to the water flow of the reef as both sides are located in the path of tidal current entering and leaving Tizard Bank. Reef growth is usually better where there is strong water flow (Stoddart, 1969; Goreau and Goreau, 1973) because this flow brings food and raw materials at the same time that it removes sediments and waste products. In terms of species diversity, coral communities on the east, southeast and northeast sides of the island are higher than in other areas. The depauperate coral fauna on the west and southwest sides are possibly related to the strong SW monsoon during summer and fall. Zou et al. (1978) reported that coral communities of Xisha Islands (Paracel Shoals) were well developed on the northeast side and poorly developed on the southwest side of the islands and that such distribution patterns are likely related to local flow patterns. Due to the influence of the prevailing SW monsoon during summer, such distribution patterns of coral communities are likely common in the South China Sea. The tropical reef environment of Taiping Island implies that its coral fauna is rich and the reef is highly developed. However, in comparison with other tropical Indo-Pacific coral reefs, the species diversity and abundance of coral communities at Taiping Island are relatively low. Dead coral skeletons were widely spread on the reef surface below 3 m and only small coral colonies were found. These facts indicate that the coral communities of Taiping Island have suffered severe damage during the last decade. The cause of such extensive coral death is uncertain. Many natural and anthropogenic stresses on coral reefs have been reported (see reviews by Brown and Howard, 1985; Grigg and Dollar, 1990). According to the current status of the reef environment, the possible disturbances are likely include artificial destructions, pollution, storms, predation of Acanthaster planci, and El Nifio events. Artificial destructions including blast fishing and underwater bombardment may have caused heavy destruction in certain areas. The presence of idle troops at Taiping Island is also of concern because they may engage in environmental damaging activities such as shooting and fishing with explosives. Substantial damage may also come from occasional parties of blast fishers and coral-smashing muroami fishers from the Philippines and Vietnam (McManus, 1992). The possibility of oil pollution is also of concern because the Spratlys lie near to major shipping lines for oil and nuclear waste. Oil and nuclear waste could be released in the event of a tanker accident in these reef-studded waters (McManus, 1992). However, we found no substantial record or evidence of these pollutants. The tropical position of Taiping Island places it within the area of frequent typhoon disturbances. The typhoon-generated waves and storm surges may erode reef crest corals and sediments down to about 20 m depth (Stoddart, 1985; Scoffin, 1993). The recognition of past storm disturbances may rely on several features such as the deposits of coral debris, the assemblages of corals and other reef biota, the reef framework structure, and the existence of reef flat storm deposits (Stoddart, 1971; Scoffin, 1993). During this survey, widespread coral debris were found to accumulate as talus at the foot of the fore-reef slope, on submarine terraces and in grooves on the reef front. In addition, on the shallow reef flat there are mainly massive, encrusting or stout branching corals that are basically wave-resistant forms. These facts indicate that typhoon disturbances are possibly the major destructive forces that have caused severe damage to the coral communities of Taiping Island. The population outbreak of the crown-of-thorn starfish, Acanthaster planci, has been recognized as the most potent biotic disturbance affecting coral communities on many Indo-Pacific reefs (Endean and Cameron, 1990). However, on reefs where marked destruction of hard-coral cover was not apparent, A. planci was either not observed or found at very low populations densities. Since we did not find any individual of A. planci during this survey, it was unlikely that the crown-of-thorn starfish was the major destructive force to the coral communities of Taiping Island. Global sea warming associated with El Nifio events has caused widespread coral bleaching in the Caribbean and the Pacific (Glynn, 1984, 1988; Williams and Bunkly- Williams, 1990; Gleason, 1993). The ecological consequences of bleaching events include widespread mortality with resultant decreases in coral cover, changes in species composition, reduced growth rates and reproductive output of corals (Szmant and Gassman, 1990; Gleason, 1993). Mortality rates in bleaching events have ranged from zero (Hoeksema, 1991) to very severe (50-98%) as on the eastern Pacific during the 1982-83 El Nifio event (Glynn, 1988). This severe event also had other associated secondary disturbances following coral mortality such as a subsequent increase in number of grazers and bioerosion rates (Glynn, 1988). Whether the widespread mortality of corals at Taiping Island is related to the El Niio-Southern Oscillation (ENSO) events need to be studied. Analysis of the environmental record in coral skeletons and marine environmental data are thus needed to answer this question. In conclusion, the coral fauna of Taiping Island is dominated by scleractinian corals, distributed mainly on the shallow reef flat at depths of 1-3 m on the east, south and north sides of the island at which flourishing coral communities were found. Few 9 gorgonaceans and alcyonacean species were found mainly on deeper reef slopes. Coral cover and species diversity of Taiping Island are relatively low in comparison with other tropical Pacific coral reefs indicating that the coral communities of Taiping Island may have been destroyed by artificial or natural disturbances. Since flourishing of coral communities and reef-building activities are the basis of sustained development of this island, we propose that reef conservation and protection are urgent and should be enforced immediately by reducing artificial destruction and pollution to the reefs. In addition, the changes of reef environment and biotic communities should be monitored. On a broader scale, the Spratly Reefs, including Taiping Island, are ecologically important, with abundant and relatively unexploited resources and where endangered species still abound. The Spratlys may also serve as a pool of larvae for fishes and other marine organisms that recruit to depleted fringing reefs and coastal habitats of the South China Sea. For these reasons, it is worthwhile to conserve the ecosystem and genetic diversity of the Spratlys by establishing a marine park in the Spratlys as proposed by McManus (1992). ACKNOWLEDGEMENTS We are grateful to Dr. L.-S. Fang, National Museum/Aquarium of Marine Biology for his support and to Mr. D.-S. Chen for his assistance with field work. Special thanks are due to the captain and crew of the Fishing Training Ship No. 2, Deep Sea Fishing Training Center, Council of Agriculture. This study was supported by a grant from the Council of Agriculture, Executive Yuan, R. O. C. (83-S.T.-2.15-F.-13). REFERENCES Bassett-Smith, P.W. 1890. Report on the corals from Tizard and Macclesfield Banks. Ann. Mag. Nat. Hist. 6(6):353-374, 443-458. Brown, B.E. and L.S. Howard. 1985. Assessing the effects of “stress” on reef corals. Adv. Mar. Biol. 22:1-63. Connell, J.H. 1978. Diversity in tropical rain forests and coral reefs. Science 199:1302- 1B 10; Dai, C.-F. 1989. Scleractinia of Tarwan. I. Families Astrocoentidae and Pocilloporidae. Acta Oceanographica Tatwanica 22:83-101. Dai, C.-F., T.-Y. Fan and C.-S. Wu. 1995. Coral fauna of Tungsha Tao (Pratas Islands). Acta Oceanographica Taiwanica 34:1-16. Dai, C.-F. and C.-H. Lin. 1992. Scleractinia of Taiwan. III. Family Agariciidae. Acta Oceanographica Taiwanica 28:80-101. Endean, R. and A.M. Cameron. 1990. Acanthaster planci population outbreaks. In: Dubinsky, Z. 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Preliminary study on the geographical distribution of shallow-water scleractinian corals from China. Nanhai Studia Marina Sinica 4:89- 96. (in Chinese) 12 Table 1. Distribution and relative abundance of shallow water corals at seven study sites (A-F) of Taiping Island. Relative abundance, +: rare, ++: occasionally, +++: common. species / Site A B (C D E F G SUBCLASS ZOANTHARIA ORDER SCLERACTINIA Family ASTROCOENIIDAE Stylocoeniella armata + 3 + + + S. guentheri + 3 + + Family THAMNASTERIIDAE Psammocora profundacella ar ase + + ++ ++ + P. digitata 3° Te + + P. contigua 2 35 25 + + Family SIDERASTREIDAE Pseudosiderastrea tayami = + Coscinarea columna + + ~ C. exesa + Famliy POCILLOPORIDAE Pocillopora damicornis =r + ar a7 +++ ++ P. eydouxi 3 sete ++ + + P. meandrina + 3 ae + 4 + P. verrucosa oF states + ++ 40 cm. The size frequencies for the most abundant commercial species are given on Figure 3. Most Serranidae were juveniles or small species. The Lethrinidae, Caesionidae and Scaridae were small in size (sizes at least 30% less than average reproductive size). This could be due to fishing pressure, but the high densities observed indicate that other factors could possibly be involved. DISCUSSION The data set presented here are minimal and one should be cautious in generalizing these results to a large area. In the absence of other comparable data from the Flores Islands or even Indonesia, it is difficult to assess how representative are these results. In particular, it is noteworthy that the stations were sampled in a leeward zone and that on the windward side of the island the morphology of the reefs is very different, and it is likely that the reef fish communities there would be different also. However, data from New Caledonia (Kulbicki et al. , 1994a) indicate that even in a wide zone, reef fish communities from the same type of reef habitat share much in common in species richness, density, biomass and structure. The substrate found on the stations is typical of many fringing reefs in the region. Indeed, in many cases terrestrial runoffs bring very fine sediment, and wave action induces the formation of rubble and coarse sediment. The very low algae and coral cover is not unusual either, especially in turbid areas. It is difficult to compare the total number of species with other areas, because the sampling effort was low. However, this number (255) is higher than observations made on fringing reefs in Hawaii, 81 - 187 species (Hayes et al. , 1982) or French Polynesia, 80 species (Galzin, 1985), which have been sampled much more thoroughly. These numbers are comparable to the highest diversities found in New Caledonia, 168 - 252 species, but with a much larger sampling effort (Kulbicki, 1992). The number of species /station is a better indicator, if the stations are sampled in a similar manner. The only data (Table 7) that have been collected according to the same methods are from Kulbicki et al. (1989, 1994a). The species richness observed in Flores is higher than in any of the New Caledonian areas. It is estimated that there are 1140 reef and lagoon fish species in the Maumere area (Kuiter and Allen, unpublished), whereas there are 940 species in the SW lagoon of New Caledonia (Rivaton et al. 1989), with 550 species in the Chesterfield Islands (Kulbicki et al. , 1994b) and 630 in Ouvéa (Kulbicki et al, 1994a). The families that are best represented in Flores exhibit considerable species diversity in most parts of the tropical Pacific, but some families that contain many species elsewhere (Apogonidae, Holocentridae, Scaridae, Acanthuridae) (Thresher, 1991) did not exhibit similar diversity in our observations. The densities observed in Flores are very high, especially for fringing reefs. Such densities have not been recorded in this type of environment in the tropical Pacific (Kulbicki, 1991). However, most of this density is due to only one species, Pomacentrus coelestis, a 6 planktivore. Large densities of planktivores are common on reefs (Kulbicki et al. , 1994a), and these species are usually short lived and experience large temporal variations. The other components of the density in Flores are usually found on fringing reefs in the Pacific, in particular, the Acanthuridae, Pomacentridae and small Labridae. This is confirmed by the few published studies on fringing reefs in the Pacific that give a detailed account of the contribution of the various species to density. In Hawaii (Hayes et al., 1982), the dominant species were two Acanthuridae (A.nigrofuscus, Ctenochaetus striatus), followed by small Labridae (Thalassoma duperrey, Gomphosus varius), the Pomacentridae being the third major component of the Hawaiian reef communities. In French Polynesia, Galzin (1985) also found a majority of Ctenochaetus striatus on the fringing reefs, the second most abundant species being another herbivore, the Pomacentridae Stegastes nigricans. In New Caledonia, the composition of the density varied from one zone to another. In Ouvéa (Kulbicki et al., 1994a) the most abundant fish were Acanthurus nigrofuscus and Stegastes nigricans, followed by three planktivorous Pomacentridae (Pomacentrus coelestis, Chromis chrysura, Chrysiptera cyanea). In the Chesterfield islands (Kulbicki et al., 1989) the most abundant species were Mulloides flavolineatus, juvenile Scaridae, Acanthurus nigrofuscus, Ctenochaetus striatus, three species of Caesio and three Pomacentridae, all herbivores (Pomacentrus molluccensis, Stegastes nigricans, Pomacentrus vaiuli). On the main island of New Caledonia (Kulbicki, unpubl.data), the major contributor to density were planktivorous Caesionidae (Pterocaesio diagramma, P.tile), several Pomacentridae (the two major ones being Chromis viridis and Dascyllus aruanus, which are mainly planktivores), Acanthurus nigrofuscus, small Labridae (Thalassoma lunare, T.lutescens) and juvenile Scaridae. The biomass (187 g/m?) found in the Flores is high for fringing reefs. In Hawaii Brock et al. (1979) found 106 g/m?, on the GBR (inshore reefs) Williams and Hatcher found 92 g/m?; the results for New Caledonia are given in table 7. The distribution of the biomass can be compared only to the studies from New Caledonia. There, the major contributors varied greatly from one zone to another. In Ouvéa (Kulbicki et al., 1994a) the top three species in terms of biomass were herbivores (Hipposcarus longiceps, Acanthurus blochii, Acanthurus xanthopterus); in the Chesterfield Islands (Kulbicki et al., 1989) the top species were two herbivores (Kyphosus vaigiensis, Naso unicornis) and a carnivore (Mulloides flavolineatus); and on the mainland the main species were planktivores (Pterocaesio tile, P.diagramma) and herbivores (Acanthurus nigrofuscus, Scaridae spp.). The similarity between Flores and New Caledonia is the presence of Acanthuridae and Scaridae as major contributors to the biomass. The differences are in the species involved, with larger species in New Caledonia than in the Flores Islands. The comparison of some length frequencies (Figure 3) between Flores and New Caledonia show that there is usually no difference in the size range. However, no small Siganus doliatus were observed in Flores, which could be due to the season, small Siganus doliatus (less than 15 cm) being found mainly during the dry season in New Caledonia. Monotaxis grandocculis did not exceed 22 cm in Flores, whereas this species was found to reach 38 cm in New Caledonia, with the largest sizes found on the barrier reef. It is often assumed that the number of species contributing in an important manner (major species; more than 2% in the present case) to the density or biomass decreases as diversity increases (Richards, 1952 and Whittaker, 1964 in McIntosh, 1967; Spight, 1977; Wahington, 1984). The relationship is not clearcut, because it is often not specified which diversity is taken into account: the observed diversity (number of species in the sample) or the potential diversity (number of species in the region). The correlation between density and biomass for major species exists both for the observed diversity and the potential diversity, but is not as good for the latter gi (Table 8 and Figure 4). This result suggests that highly diverse communities have lower numbers of dominant species. In other words, one would expect the resources to be better shared and utilised in these communities that in less diverse ones. Analysis of the trophic structure and of distribution of the life-history strategies will in part answer this question. It is difficult to compare the trophic structure found in Flores with most of the findings in the literature, because the methods were very different from one study to another (Kulbicki, 1991). The data from New Caledonia were collected and analysed with the same methods used in the present study and are, therefore, comparable (Figure 5). The distribution of species among trophic categories (Figure 5a) is very similar in all 4 studies. However, Flores had more zooplankton feeding species than the fringing reefs of New Caledonia. In density (Figure 5b) and biomass (Figure 5c) the results from Flores and mainland New Caledonia are almost identical. The latter two islands differ from Chesterfield and Ouvea, both of which are offshore islands, in having larger numbers of zooplanktivores, lower abundances of microherbivores and carnivores, and larger biomasses of zooplanktivores. This larger importance of zooplanktivores in the Flores and mainland New Caledonia could be linked with high terrestrial runoffs (these islands have similar land masses -10 000 and 20 000 km? - and average rainfall - 1500 to 2000 mm/ year). There are also trends common to all four studies. In particular, coral feeders form 2-7% of the species but account for very little in density or biomass. Detritus feeders and "other planktivores" are never an important component of the trophic structure, whereas they form between 10 and 15% of the abundance or weight for the coastal (mangroves and estuaries) areas in New Caledonia (Thollot, 1992). Fringing reefs and coastal areas are often adjacent in New Caledonia, thus indicating that the trophic structure is greatly influenced by the substrate. Very few studies on reef fishes have treated life-history strategies (Kulbicki, 1991; Kulbicki et al., 1992, 1994a) or assimilated structures (ecological categories x size classes) (Harmelin-Vivien, 1989). Kulbicki (1992), based on original data, compared life-history strategies from several types of reefs across the Pacific using the same classification. The data of the present study can be compared with data processed in the same way for fringing reefs in New Caledonia (Figure 6). The distribution of species among life-history strategies is almost identical for all reefs (Figure 6a). This result could be expected from the findings of Kulbicki (1992), who demonstrated that within the Western Pacific there were little differences in this structure at the species level. Flores and mainland New Caledonia also have very similar structures in terms of density and biomass (Figures 6b, c). In particular, they differ from the fringing reefs of the islands of Ouvea and Chesterfield by having more class-1 species, which have the fastest turnover. Conversely, Flores and mainland New Caledonia have a low proportion of biomass represented by long living fishes (classes 5 and 6) which are important on the Ouvea and Chesterfield islands. This suggests that in Flores the fish communities of the fringing reefs should be more sensitive to short term variations than they would be on isolated islands such as Ouvea or the Chesterfield. This is logical since most of these class 1 and 2 fish feed mainly on zooplankton and microalgae, which are variable food sources, depending on primary production and mineral inputs. Our findings indicate, therefore, that the functioning of the fringing-reef fish community of Flores is very similar to what is observed on mainland New Caledonia where ecological conditions are similar. Conversely, fringing reef fish communities from isolated islands of New Caledonia, despite their similar species composition, have different structures. Diversity alone does not account for the major differences in the structure of these fish communities. ACKNOWLEDGEMENTS The author wishes to thank the following persons and organisations: Prof. Dr. Kasijan Romimohtarto and the organizing committee of the Pre Indo-Pacific Fish Conference workshop held in Maumere (November 20-25, 1993), R.Kuiter, Dr.G.Allen, G.Moutham, P.Dalzell and the two anymous reviewers. 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Harmelin-Vivien M. 1989 Reef fish community structure: an Indo-pacific comparison. in Ecological studies - Vertebrates in complex tropical systems (Harmelin-Vivien M., Bourliére F. eds) Springer Verlag N.Y. 69: 21-60 Hayes T., Hourigan T., Jazwinski S., Johnson S., Parrish J., Walsh D. 1982 The coastal resources, fisheries and fishery ecology of Puako, West Hawaii - Hawaii Cooperative Fishery Research Unit Technical Report 82-1: 159 + Annexes Kulbicki M. 1991 Present knowledge of the structure of coral reef fish assemblages in the Pacific - in Coastal resources and systems of the pacific basin: investigation and steps toward a protective management - UNEP Regional Seas Report and Studies : 147: 31-53 Kulbicki M. 1992 Distribution of the major life-history strategies of coral reef fishes across the Pacific. Proc. 7th Intern. Coral Reef Symp. - Guam 1992 : 918-929 Kulbicki M., Doherty P., Randall J.E., Bargibant G., Menou J-L., Mou-Tham G., Tirard P. 1989 - La campagne Corail 1 du N.O. Coriolis aux iles Chesterfield (du 5 aoit - 4 sept. 1988) : données préliminaire sur les peuplements ichtyologiques ORSTOM Nouméa. Rapp. Sci. Tech. Sci. Mer Biol. Mar. 57 : 88 p. Kulbicki M., Thollot P., Wantiez L. 1992 Life history strategies of fish assemblages from reef, soft bottom and mangroves from New Caledonia. Seventh Intern Coral Reef Congress - Guam June 1992 abstract Kulbicki M., Dupont S., Dupouy C., Bargibant G., Hamel P., Menou J.L., Mou Tham G., Tirard P. 1993 Caractéristiques physiques du lagon d'Ouvéa - in Evaluation des ressources en poissons du lagon d'Ouvéa: 2éme partie: l'environnement physique: sédimentologie, substrat et courants - Convention Sciences de la Mer ORSTOM Nouméa 10: 47-150 Kulbicki M., G. Bargibant, Menou J.L., Mou Tham G., P.Thollot, L. Wantiez, Williams J.T. 1994a Evaluations des ressources en poissons du lagon d'Ouvéa. in Evaluation des ressources en poissons du lagon d'Ouvéa: 3éme partie: les poissons; Convention Sciences de la Mer ORSTOM Nouméa 11: 448 p. Kulbicki M., Randall J.E., Rivaton J. 1994b Checklist of the fish from the Chesterfield islands. Micronesica - 27 (1/2): 1-43 Kuiter R., Allen G. submitted Fishes of Maumere Bay, Flores Indonesia - Tropical Diversity Indonesian Journal McIntosh R.P. 1967 An index of diversity and the relation of certain concepts to diversity - Ecology 48 (3) : 392 - 404 Rivaton J., Fourmanoir P., Bourret P., Kulbicki M. 1989 - Catalogue des poissons de Nouvelle- Calédonie. Catalogues Sciences de la Mer, ORSTOM Nouméa 2: 170 p. Spight T.M. 1977 Diversity of shallow water gastropod communities on temperate and tropical beaches - The American Naturalist 111 (982): 1077-1097 Thollot P. 1992 - Les poissons de mangrove du lagon sud-ouest de Nouvelle-Calédonie - écologie des peuplements, relations avec les communautés ichtyologiques cotiéres. Ph.D. Thesis University of Aix-Marseille II (France), 406 p. Thresher R.E. 1991 Geographic variability in the ecology of coral reef fishes : evidence, evolution, and possible implications - in The ecology of fishes on coral reefs (P.Sale ed.) Academic Press Inc. New York 754 p. Washington H.G. 1984 Diversity, biotic and similarity indices. A review with special relevance to aquatic ecosystems - Water Research 18 (6): 653-694 Williams D.McB., Hatcher A. 1983 Structure of fish communities on outer slopes of inshore, mid-shelf and outer shelf reefs of the Great Barrier Reef - Marine Ecology Progress Series 10: 239-250 10 Piscivores Macrocarnivores Life-history strategy classes Life-history strategy classes Microcarnivores Zooplanktivores 70 + Oce2p eo Bic2B 50 + 40 + x 30 20 + 10 + (0) 1 2 3 4 5 6 Life-history strategy classes Life-history strategy classes Microherbivores Life-history strategy classes Figure 2: distribution of trophic categories according to life-history strategies. D: density; B: Biomass; Pi: piscivores; C1: macroinvertebrate feeders; C2: microinvertebrate feeders; Zoo.: zooplanktivores; Mi.: microalgae feeders Pterocaesio tile number 16 Size (cm) Scolopsis bilineatus o Ynumber = a = Ss Sette Sea zz Size (cm) Ctenochaetus striatus 11 Monotaxis grandocculis © Flores N=320 BI NC N=3825 Scarus fasclatus O Flores N= 29 O Flores N= 23 BANC N= 1421 FNC N=295 Size (cm) Figure 3: size distribution of the most abundant commercial species (NC: Caledonia) Y=10.13 -0.023X r1r?=0.60 12 i= 2 10 o E =, 8 ore 06 ea ow iS o a o Nn BR DD 100 Number of species in sample 200 300 Size (cm) data for New InY=6.95 -0.75InX 17=0.69 2.50 = In%B 2.00 5 |In%D 1.50 1.00 0.50 In % major species 0.00 6.00 400 7.50 In Number of potential species 6.50 7.00 8.00 Figure 4: correlation between number of species ("major species") contributing to more than 2% of density (%D) or biomass (%B) and number of species in sample, or number of reef species known in region. Data from Table 8. Note that for second figure a log scale is used. WA _] Flores Sp. ES Ouvea Sp. [J Chest. Sp. NC Sp. % species number a) b) ” 7) Oo £ 2 fe} 2 : N . =N % EN =f i oo 5 3) S o <= io} Ss N 5 s O fo) C) Figure 5: comparison of trophic structure (a: species, b: density, c: biomass) of fringing reefs Flores with New Caledonia: Ouvéa (Kulbicki et al., 1994a), Chesterfield islands (Kulbicki et al., 1989), main island (NC) (Kulbicki (1991). Pi: piscivores; Cl: macrocarnivores; C2: microcarnivores; Zoo: zooplankton feeders; Other P.: other plankton feeders; MaH.: macroalgae feeders; MiH.: microalgae feeders; Cor.: coral feeders; De.: detritus feeders 13 Flores S. ES Ouvea S. Chest. S. NC S. Life-history strategy classes a) L] Flores D. ES OuveaD. [J Chest.D. N NCD. 70 7 60 | > 50 @ 40 ® So 307 32 20 10 (0) ce ss CoE a 1 2 3 4 5 6 Life-history strategy classes b) Flores B. 3 OuveaB. [:] Chest. B. NC B. Life-history strategy classes c) Figure 6: comparison of life-history strategy classes in Flores and New Caledonia. Key same as Figure 5. 14 Table 1: definition of the 6 life-history strategy classes used for defining structure. Life length can be considered as life expectancy (LSO after recruitment) Class Size Reproduction Behavior Growth Mortality Life length 1 Small to Very early in life Most species Very fast High 0.5 to 3 medium __—~ Very high gonado-somatic school years < 30cm index or reproductive Simple sexual effort behavior 2 Small to 1-3 years old at first Often schools, Rapid initially Medium 3 to7 years medium reproduction may be < 30cm High gonado-somatic territorial index Sexual behavior may be complex 3 Medium to 2-3 years old at first Often schools, Rapid initially Medium 3 to7 years large reproduction seldom or through > 30 cm High gonado-somatic territorial life index Simple sexual behavior 4 Small to Late in life Seldom Slow after Low 7 to 12 medium - Usually > 50 % maximum schools first years < 30cm size at first reproduction Often reproduction Medium gonado-somatic territorial _— initial growth index often fast 5 Medium to Late in life Seldom Slow after Low 7-12 years large Usually > 60% maximum schools first > 30cm size at first reproduction Often reproduction usually Low gonado-somatic index __ territorial Often rapid >50cm initial growth 6 Large to Very late in life Almost never Veryslow Verylow > 12 years very large Usually > 60% maximum schools especially > 50cm size at first reproduction except for after usually > 1m Often ovoviviparous reproduction reproduction Low gonado-somatic index 15) Table 2: composition of substrate. Depths in m. All other numbers are percentages. STATION NUMBER 1 2 3 4 5 6 Total SUBSTRATE Sand - muddy 12 6 8} Sand - fine Sand - coarse WZ, 5) 5 17 11 8 10 Gravel and Debris 3 7 10 24 7 36 16 Small boulder 3 3 2 10 SY 16 14 Large boulder 23 3 4 7a) 18 34 22 Rock 47 41 71 28 3 5 33 Beachrock 8 8 3 TOTAL 100 100 100 100 100 100 100 ORGANISMS Algae 5) 1 Coral 13 <1 <1 D, Alcyonarians 15 DEPTH RANGE 3/9 2/9 2/4 2/10 1/12 7/9 1/12 Table 3: major fish families and their contribution to total diversity and comparison with New Caledonia (NC) Family Number of %total Species in | Family Number of %total Species in species species common with NC Serranidae : Labridae : Caesionidae 7 Da, 5 Scaridae 15 5.9 13 Mullidae 8 3a 7 Acanthuridae 16 6.3 £5 Chaetodontidae 15) D9) 13 Siganidae Pomacanthidae 7 De 5 Balistidae Pomacentridae 49 19.2 42 Total 197 77 170 16 Table 4: density (fish/m?) and biomass (g/m?) of the major families and species. FAMILIES DENSITY BIOMASS SERRANIDAE 0.099 6.36 Pseudanthias squamipinnis 0.047 0.17 Cephalopholis urodeta 0.013 0.99 Epinephelus fasciatus 0.010 1.10 LUTJANIDAE 0.021 3.95 Lutjanus decussatus 0.015 2S LETHRINIDAE 0.025 Sle Lethrinus harak 0.006 E92 Monotaxis grandocculis 0.012 1.85 NEMIPTERIDAE 0.040 4.28 Scolopsis bilineatus 0.021 1.67 MULLIDAE 0.042 9.48 Parupeneus indicus 0.003 4.53 Parupeneus trifasciatus 0.021 1.10 CHAETODONTIDAE 0.049 1.67 POMACANTHIDAE 0.044 29, POMACENTRIDAE 4.954 18.4 Chromis amboinensis 0.163 0.64 Chromis xanthura 0.226 0.23 Neopomacentrus azysron 0.139 0.48 Pomacentrus amboinensis 0.074 0.31 Pomacentrus brachialis 0.103 0.63 Pomacentrus coelestis 3.468 10.4 LABRIDAE 0.374 7.86 Cirrhilabrus cyanopleura 0.027 0.11 Cirrilabrus sp. 0.027 0.06 Halichoeres melanurus 0.056 0.29 Novaculichthys taeniourus 0.004 1.09 Thalassoma amblycephalum 0.048 0.23 SCARIDAE 0.106 33:1 Scarus spp. juvenile 0.052 13.0 Scarus fasciatus 0.016 5.56 Scarus quoyi 0.014 6.94 ACANTHURIDAE 0.132 18.4 Acanthurus leucocheilus 0.033 2.50 Ctenochaetus striatus 0.059 5.60 Naso hexacanthus 0.008 2.14 SIGANIDAE 0.023 4.85 BALISTIDAE 0.065 4.84 TOTAL fas 187 Table 5 : trophic structure. All numbers are percentages. CATEGORY DIVERSITY DENSITY BIOMASS Piscivores 11.9 Mp 8.4 Macrocarnivores DD 43 19.3 Microcarnivores 14.2 6.5 3.8 Zooplanktivores Dey Se) 29.9 Other planktivores 0.1 0.1 0.1 Macroherbivores 2 0.1 0.8 Microherbivores 20.5 ee 34.9 Coral feeders Soe! 0.5 0.9 Detritus feeders 2.0 9.2 D0) Table 6: distribution of the life-history strategies. All numbers are percentages. Classes refer to the classification given in table 2. LIFE-HISTORY STRATEGY DIVERSITY DENSITY BIOMASS CLASS 1 10.0 61.6 8.2 D, 39.8 eS Sled! 3 16.1 5.8 36.4 4 Zk 3.8 3} 5) 10.0 1S 10.0 6 2.8 0.1 Med Table 7: species richness (species /transect), density (fish/m?), biomass (g/m?) from fringing reefs in New Caledonia (SW lagoon, Chesterfield and Ouvéa)(Kulbicki, 1991; Kulbicki et al., 1989, 1994a). REGION SPECIES RICHNESS DENSITY BIOMASS Chesterfield 64 QBS) 90/200 Ouvéa 85 2.4 340 SW Lagoon 55 2.2/5.8 61/155 18 Table 8: number of species (N) contributing to more than 2% of density or biomass for Flores and other fringing reefs in the Pacific. Sampled species: number of species sampled. Potential species: number of reef species known in the area; %N: percentage of N in the number of species recorded during the survey. 1: Kulbicki unpublished; 2: Kulbicki et al., 1994a; 3: Kulbicki et al. 1989; 4: Galzin, 1985; Hayes etaliy1982 Region N density %Ndensity Nbiomass %Nbiomass Sampled Potential Land are species species (km?) Flores 6 3) 10 3.9 255 1140 ~10 000 New Caledonia (1) 10 29 11 3.2 348 940 20 000 Ouvéa (2) 14 ies 8 4.3 152 630 130 Chesterfield (3) 14 10.8 10 7.8 130 550 10 Moorea (4) 6 I>) 80 630 130 Hawaii (5) 9 4.8 187 460 =500 19 Appendix 1: list of species observed. St: number of stations where species was observed; N: total number of individuals seen; Sch.: average size of schools; Size: average size in cm NAME Taeniura lymma Plotosus lineatus Saurida gracilis Synodus variegatus Synodus dermatogennis Synodus spp. Sargocentron caudimaculatum Aulostomus chinensis Pterois antennata Pterois volitans Pseudanthias squamipinnis Pseudanthias tuka Anyperodon leucogrammicus Cephalopholis argus Cephalopholis cyanostigma Cephalopholis leopardus Cephalopholis microprion Cephalopholis miniata Cephalopholis sexmaculatus Cephalopholis spiloparea Cephalopholis urodeta Epinephelus cyanopodus Epinephelus fasciatus Epinephelus hexagonatus Epinephelus merra Variola louti Variola albomarginata Pseudochromis exquisitus Pseudochromis paccagnellae Apogon fraenatus Apogon nigrofasciatus Cheilodipterus lineatus Malacanthus latovittatus Carangidae spp. Caranx para Caranx tille Caranx spp. Gnathanodon speciosus Lutjanus decussatus Lutjanus fulvus Lutjanus rivulatus Lutjanus vittus Macolor niger Caesio cuning Caesio lunaris Pterocaesio chrysozona Caesio xanthonota 2 NPN NY HH KH KN DH WK KP KN KN KH RK KN WK K DN WK KK WWNnN DN WH OK NH RK We Ke eS Nn i=) KS NOK DK KK WH eS 100 — WD We WD HAwW NY NM VY — lon Se UMW pe HP OW HN NY OY 18 232) 17.3 15.2 W267) NAME Pterocaesio diagramma Pterocaesio teres Pterocaesio tile Plectorhinchus picus Lethrinus olivaceus Lethrinus harak Lethrinus rubrioperculatus Monotaxis grandoculis Pentapodus caninus Scolopsis affinis Scolopsis bilineatus Scolopsis lineatus Scolopsis margaretifer Mulloides flavolineatus Parupeneus barberinus Parupeneus bifasciatus Parupeneus cyclostomus Parupeneus indicus Parupeneus macronema Parupeneus trifasciatus Upeneus tragula Platax orbicularis Chaetodon adiergastos Chaetodon baronessa Chaetodon citrinellus Chaetodon kleinii Chaetodon lineolatus Chaetodon lunula Chaetodon melannotus Chaetodon ornatissimus Chaetodon pelewensis Chaetodon rafflesi Chaetodon trifascialis Chaetodon trifasciatus Chaetodon vagabundus Chaetodon xanthurus Heniochus varius Centropyge bicolor Centropyge tibicen Centropyge vrolicki Genicanthus lamarcki Pomacanthus imperator Pomacanthus xanthomethopon Pygoplites diacanthus Abudefduf saxatilis Acanthochromis polyacanthus Amblyglyphidodon aureus St KBWNWNNK DHE NPE UWHWHE NEP HE NEP KP PEP SPE NHK DPEPRWRKE WHE DRE UEP NHK EP wD eG = S| ne i) i) — ins DNDN WOOWWN KH WD NO nA © 20 NAME Amblyglyphidodon curacao Amblyglyphidodon leucogaster Amphiprion clarkii Amphiprion melanopus Amphiprion perideraion Chromis amboinensis Chromis atripectoralis Chromis atripes Chromis viridis Chromis chrysura Chromis flavicauda Chromis flavomaculata Chromis margaritifer Chromis retrofasciata Chromis vanderbilti Chromis spp. Chromis xanthura Chromis weberi Chrysiptera rex Chrysiptera rollandi Chrysiptera talboti Dascyllus aruanus Dascyllus melanurus Dascyllus reticulatus Dascyllus trimaculatus Discistodus melanotus Neopomacentrus azysron Neopomacentrus nemurus Neopomacentrus violascens Paraglyphidodo nigroris Neoglyphidodon crossi Plectroglyphidodon dicki Plectroglyphidon lacrymatus Pomacentrus alexanderae Pomacentrus amboinensis Pomacentrus bankanensis Pomacentrus brachialis Pomacentrus coelestis Pomacentrus lepidogenys Pomacentrus philippinus Pomacentrus reidi Pomacentrus simsiang Pomacentrus sp. Pomacentrus taeniometopon Pomacentrus vaiuli Cirrhitichtys falco Paracirrhites forsteri Sphyraena barracuda Sphyraena japonica Anampses caeruleopuncta Bodianus mesothorax 2) road BWOrrP Ke HB PWWNYK WN DWW WH PP KP WWE NH WNKH KH UNDUADWK KEP NNN KH WHEN KEN HK HK NH WwW NAME Cheilinus celebicus Cheilinus chlorourus Cheilinus diagrammus Cheilinus fasciatus Cheilinus trilobatus Choerodon anchorago Cirrhilabrus exquisitus Cirrhilabrus cyanopleura Cirrhilabrus sp. Coris gaimard Coris schroederi Diproctacanthus xanthurus Epibulus insidiator Gomphosus varius Halichoeres argus Halichoeres chrysus Halichoeres hortulanus Halichoeres melanurus Halichoeres miniatus Halichoeres prosopeion Halichoeres podostigma Halichoeres nebulosus Halichoeres scapularis Hemigymnus fasciatus Hemigymnus melapterus Hologymnosus annulatus Hologymnosus doliatus Labrichthys unilineatus Labroides bicolor Labroides dimidiatus Macropharyngod meleagris Macropharygodo ornatus Novaculichthys taeniourus Pseudocheilinu evanidus Pseudocheilinu hexataenia Pseudocheilinu octotaenia Pseudodax mollucanus Stethojulis bandanensis Stetholulis interrupta Stethojulis strigiventer Stethojulis trilineata Thalassoma amblycephalum Thalassoma hardwicke Thalassoma janseni Thalassoma lunare Scarus spp. Cetoscarus bicolor Scarus bleekeri Scarus altipinnis Scarus dimidiatus Scarus flavipectoralis ~ oo NK KE DK AADANNWNNRKP KF KF WK NWN DPRK KP PEN RP RP UN WNnNNK PK NWWN WD KN WHY RWW We PNY W oo Wo —_ (Gey ey ee ee Nn os) 10.4 — (oe) —_— ~~) — 00 — ee eo ee dO oe, oe, a eet et — ee Won fp ~TIN 1.3 NAME Scarus fasciatus Scarus forsteni Scarus microrhinos Scarus niger Scarus oviceps Scarus psittacus Scarus quoyt Scarus prosognathos Scarus sordidus Parapercis clathrata Parapercis cylindrica Parapercis multiplicata Parapercis tetracantha Ecsenius bandanus Ecsenius bicolor Ecsenius midas Plagiotremus rhinorhynchos Amblygobius rainfordi Istigobius decoratus Ptereleotris evides Ptereleotris heteroptera Valenciennea strigatus Acanthurus mata Acanthurus fowleri Acanthurus dussumieri Acanthurus nigricans Acanthurus blochii Acanthurus lineatus Acanthurus nigrofuscus Acanthurus leucocheilus Acanthurus olivaceus Acanthurus pyroferus Ctenochaetus binotatus Ctenochaetus striatus Naso hexacanthus Naso lituratus Paracanthurus hepatus Zebrasoma scopas Siganus argenteus Siganus canaliculatus Siganus corallinus Siganus doliatus Siganus puellus Siganus vulpinus Zanclus cornutus Rastrelliger kanagurta Amanses scopas Aluterus scriptus Balistapus undulatus Balistoides viridescens Melichthys vidua n oo BEA HBB BPWHAN WR DHY HE UANN HK HNN KKH KH eK Ke PN HEP Hanne aernweyr Oe WHE WDA | 23 NAME St Odonus niger Pervagor melanocephalus Rhinecanthus verrucosus Sufflamen bursa Sufflamen chrysopterus Arothron meleagris Arothron nigropunctatus WN NY Fe W Canthigaster solandri N 65 Sch. 16.2 aN Size 13.7 $4 nies on vga) ninnabayg: oma rvyiny ava te a wise Te MAA t Be ag 54 senor aR, ys vivaess thea sgh ahh Pee janis ran as rete ea) preg re ee wears Bs lanes ay, Soyk ce apilinnd SORA 9 A a r Ney od t ey " Mi, 1 ae Re nih a ue, eee | ane ah Aled in ads ass Ruf. fl S: ! Sey f 4 aly, Aiea atgeanee M j ert | if ili Bad rabbi ot.” ays ny an GY Cpe s eee ae a a oh 4 é a % % i ‘ th ‘a ey i 1 ; i wi & , : | j i } A ; ’ New “ ty a om, {2 t | 4 Beh or) y By a ub f deeb ! WA A uta rierver ath Woe fae white — | 0 ig tal knee i = ss iP sr > es — = ae —_—_— — = - & — = = a oe oe - “ Jy a5 ti aber a J vin mab apse tuntniycanid yee se : al Fo j nleyag tat . ATOLL RESEARCH BULLETIN NO. 438 GROUPER DENSITY AND DIVERSITY AT TWO SITES IN THE REPUBLIC OF MALDIVES BY ROBERT D. SLUKA AND NORMAN REICHENBACH ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. APRIL 1996 Tp) a) > — (am) =] << = io GaN noe otal? GROUPER DENSITY AND DIVERSITY AT TWO SITES IN THE REPUBLIC OF MALDIVES BY ROBERT D. SLUKA! AND NORM REICHENBACH? ABSTRACT The density and diversity of shallow-water groupers at Gaagandu, North Male Atoll and Olhugiri, Thaa Atoll, Republic of Maldives was enumerated using visual transects. Four different habitat types were surveyed: reef lagoon, reef crest, reef slope, and a well- developed lagoonal reef. Twenty-two species in seven genera were recorded. Median densities ranged from 7 to 23 grouper 240 m*. At Gaagandu Island, the reef slope was repeatedly sampled using 20-m belt transects to estimate the efficiency and accuracy of the sampling methodology. Fifteen transects were necessary to estimate the median density of all species within 10% of the reference value and to develop a species list containing 80% of the total number of species observed. The species observed varied in their degree of site attachment. Those species which were most closely tied to their habitat exhibited clumped spatial distributions while those species which ’roamed’ over large areas had random spatial distributions. The number of transects necessary to adequately characterize the median density of a species was related to the degree of clumping in its spatial distribution. INTRODUCTION Groupers are an important fishery resource throughout the world and are important predators in coral reef ecosystems. Approximately 30 grouper species occur in the Republic of Maldives. Maldivians prefer to eat tuna and have not developed extensive reef fish fisheries (Anderson et al. 1992). Total reef fish catch is approximately 3000 tons per year (Anderson et al. 1992). At present, we are aware of only two operations exploiting groupers, one of which has had little effect on the grouper population (Sluka unpublished data). A market has developed exporting groupers to other southeast Asian countries and to supply many of the resorts located around North Male Atoll. It is therefore likely that reef fish, especially groupers, will come under increasing exploitation in the near future in the Republic of Maldives. Differences in catch 1 University of Miami, Department of Biology, P.O. Box 249118, Coral Gables, Florida 33124 USA 2 The Oceanographic Society of Maldives, Male, Republic of Maldives Manuscript received 19 July 1994; revised 19 November 1994 2 composition during exploratory fishing were found between a southern atoll (Laamu) and more northern atolls (Alifu and Shaviyani) (Anderson et al. 1992). Shepherd et al. (1992) reported that the abundance and biomass of all species combined was lower on reef flats that were mined than on unmined reef flats. However, the abundance and biomass of fish on slopes adjacent to mined flats was greater than on slopes adjacent to unmined flats. ‘Four grouper species were among the 20 fish which showed the most dissimilarity between these slopes. Cephalopholis miniata and Variola louti had higher biomass on slopes adjacent to mined flats, while Plectropomus pessuliferus and Gracila albomarginata had higher biomass on slopes adjacent to unmined reef flats. The difficulties in using visual survey methods such as transects has been reviewed by other authors (De Martini and Roberts 1982; Bortone et al. 1986; Sanderson and Solonsky 1986; Greene and Alevizon 1989). Various techniques for solving problems such as transect width (Sale and Sharp 1983), transect length (Fowler 1987), duration of the survey (St. John et al. 1990), and sample size (Sale and Douglas 1981) have been developed. However, these studies usually involved sampling the whole community and in many cases were specifically directed towards sampling patch reefs. Methodologies for surveying serranids were examined by the Great Barrier Reef Marine Park Authority (1979) and Craik (1981) for the Great Barrier Reef region. Groupers are relatively sedentary and site attached. Survey methods must take into account their cryptic behavior and the likelihood of having a patchy or clumped dispersion pattern. This clumped dispersion could lead to misleading results if only a few samples are collected. The number of samples necessary to accurately assess population density will depend on the degree of clumping in their dispersion pattern. The density and diversity of groupers was studied at two sites in the Republic of Maldives and related to habitat preferences of the different species. The sample size necessary to accurately estimate the density and diversity of groupers in a specific area was examined using visual belt transects. METHODS Habitat characterization: The atolls were divided into three habitat zones: 1) lagoon, 2) reef crest, and 3) reef slope. The habitat was characterized by recording the coverage class of dominant substrate (sand, sand-mud, rubble, and hard reef) and lifeforms (seagrass, algae, sponges, octocorals, and hard coral). Substrata and lifeform information were collected by visually estimating the coverage in a belt of 1 m* quadrats. Coverage was scored in the following categories: 1) < 10%, 2) 10 - 30%, 3) 30 - 70%, and 4) > 70%. In order to convert to cm’ the midpoints of each coverage class were summed for each quadrat and averaged. Visual surveys: Prior to observation, the observer was trained to accurately estimate length using models of fish with a known size-frequency distribution (Bell et al. 1985). Visual surveys were conducted similarly to GBRMPA (1979). A 20-m transect was placed in a haphazard fashion along a particular depth gradient (parallel to shore). An 3 area 6 m out from one side of the transect was intensively searched for all grouper species and then the diver searched the other side in a similar fashion. The number and size of all groupers observed were recorded. Groupers were placed in one of five size categories: <5 cm, 5-15 cm, 15-25 cm, 25-35 cm, and >35 cm. The depth and time of each survey were recorded. All of the habitat zones had similar sampling effort except the reef slope at Gaagandu, which was more intensively surveyed. A distance of approximately 300 m along the reef slope from 6 to 20 m depth was repeatedly sampled in order to assess the number of transects necessary for reliable estimates of density and diversity. Species identifications were made using Heemstra and Randall (1984), Randall (1992), Randall and Heemstra (1991), and Allen and Steene (1987). When information on species identification differed between sources, Randall and Heemstra (1991) was used. Species presence/absense data was collected at Chicken Island, near Gaagandu, for comparison. Statistical analysis: Descriptive statistics, histograms, correlations and other calculations were performed using Microsoft Excel® software. The frequency distributions of numbers of groupers observed per transect (240 m”) exhibited various degrees of skewing to the right (Figure la, b). Because of the skewed distributions, medians were considered to characterize the densities better than means. Performance curves based on cumulative medians and species-sample curves were used to determine the number of transect replicates needed to obtain adequate density and diversity estimates for groupers observed in the 48 transects from the slope area (Brower et al. 1990). Medians were compared statistically using a Chi-square procedure (Zar 1984). For species with median density estimates greater than zero, performance curves were calculated. The performance curves calculated were considered to stabilize when all subsequent cumulative medians fell between the 40th and 60th percentiles calculated from the entire set of 48 transects. The least number of transects required to stabilize the performance curve was considered the number of replicates required for a reliable density estimate. This process was repeated 20 times, with the order of the 48 transects entering the cumulative median calculation being randomized each time. Medians were then calculated from the 20 estimates of replicates required to obtain a reliable density estimate. The median estimates for required replicates were then correlated with the species dispersion pattern using Morisita’s Index of Dispersion (I,) (Brower et al. 1990). For the density and diversity of all species combined on the reef slope, performance curves and species-sample curves were calculated. The number of replicates required for a stable density estimate was determined in a fashion similar to that noted for individual species except for the criteria used to determine performance curve stability. Instead of using one level for determining stability, i.e. the 40th and 60th percentiles, several levels were evaluated. These levels included 20% of the median (30 and 70 percentiles), 15% (35 and 65 percentiles), 10% (40 and 60 percentiles), and 5% (45 and 55 percentiles). If the median estimated from all 48 transects is considered to be the reference median density, then these different levels for assessing performance curve stability would indicate the accuracy of the median estimated from a given number of transects. The number of replicates based upon the species-sample curves were also 4 48). is urodeta (n 48) and (b) Figure 1: Frequency distribution of number of grouper observed per transect for (a) = Cephalopholis Aethaloperca rogaa (n a) Asuenbel4 11S SMS IS 10 A. rogaa (#/transect) b) Prrrrrs OOO II RrEDLCEetirLeceeeenereLereeLieeLereeLee eles PETRELELEMERILEL EERE LULLEL ELE LE LeELeerery OOK OOK HO HHH III A MHI ANIPIPIID IDI L ILI SL LL LIN IID LLLP LLDPE DO SESASNAASSA XMAS Asuenbel4 C. urodeta (#/transect) 5 assessed at various levels of percent of species observed. The levels included >70%, >80%, =90%, and 100%. This process was repeated 20 times, randomizing the order of the transects each time. Medians were then calculated from the 20 estimates of replicates required for each level of percentage of species observed. RESULTS Habitat characterization: Gaagandu Island is located inside the main atoll ring of North Male Atoll. The northern and western sides of the island are surrounded by a lagoon approximately 50 m wide and approximately 2 m deep at high tide. The lagoon was primarily rubble with very small areas of sand (Figure 2a). The rubble areas of the lagoon were covered by turfing algae, had no soft coral or sponges, and very little hard coral (Figure 2b). The reef crest consisted of large, eroded coral heads covered by algal turf. The crest had only slightly higher hard coral cover than the lagoon and had very low coverage of sponges and soft coral. From the crest, the reef sloped down steeply to a sand flat at 30 m depth. The reef slope appeared to be divided into areas of high vertical relief separated by ’landslides’ of rubble with sand. The reef slope had the highest percentage cover of hard coral (approximately 30%) and low numbers of sponges and soft coral. The southwestern portion of the island had a well-developed reef consisting of a huge bed of Acropora sp. interspersed by massive coral colonies. This reef is designated as reef 1 for further analyses. The depth ranged from 1-10 m at reef 1 and no substrate/lifeform data was taken at this site. Olhugiri island is located on the northern edge of the outer ring of Thaa Atoll, approximately 2.35 N latitude, 73.05 E longitude. The lagoon of the atoll stretches approximately 50 m in each direction around the island. The northern side of the island is open to the sea and has a reef crest which slopes steeply down to 50 m where the slope becomes much gentler. The western portion of the island is lagoonal connecting to another island without any deep passages. The inner side of the island has a reef crest which slopes gently to about 10 m into a sand flat. The eastern portion of the island has a channel about 10 m in depth which allows passage of water into the atoll. The outer and inner reef crests were sampled for grouper density and diversity No quantitative habitat data was collected at Olhugiri. Density _and_ diversity of groupers: There was no correlation between any species abundance, nor total abundance, with depth (minimum, maximum, or mean) or time of day (p> 0.05) along the reef slope. There was a significant difference in the median number of grouper observed per transect between sites (X? = 44.84, df = 4, p < 0.001, Table 1). The slope at Gaagandu had the highest median density with 23 grouper observed per transect. Excluding the slope data, the other sites had no significant differences in the median number of grouper observed per transect (X? = 4.74, df = 3, p > 0.05). The lagoon at Gaagandu had a median density of 5 and the lagoon at Olhugiri 16. These two sites were not included in the density comparisons due to the low sample size (2 and 4 transects, respectively). 6 Figure 2: Substrata (a) and Lifeform (b) coverage of the site at Gaagandu Island, North Male Atoll. Open bars represent the slope area (n=100 1 m? quadrats), solid bars represent the reef crest (n=100), and striped bars represent the reef lagoon (n=40). (a) S = sand, RB = rubble, and HR = hard reef. (b) AT = algae, SP = sponge, SC = octocoral, and HC = hard coral. a) Percent Cover Substrata b) Percent Cover \ \ N N \ \ \ \ \ NN: NY Lifeforms qj Table 1: Median, maximum, and minimum number of grouper observed per 240 m? transect within each zone at the two island sites. GAAGANDU OLHUGIRI INNER OUTER CREST SLOPE REEF 1 CREST CREST MEDIAN 7 7a) 2 10 JUGS) MAXIMUM 13 50 18 24 15 MINIMUM 3 11 4 3 7 The lagoon at Gaagandu was characterized by low diversity (4 species). There were 7 species observed on the reef crest, dominated by Cephalopholis argus and C. urodeta (Table 2). Reef 1 was dominated by C. argus and Epinephelus merra. The slope had the highest diversity with 17 species (also the largest sample size). Cephalopholis miniata, C. leopardus, C. urodeta, E. spilotoceps, and C. argus dominated numerically in decreasing order of importance. Along the slope the densities of ’roving’ species, such as G. albomarginata, Variola louti, and Plectropomus spp., were probably underestimated; these species were frequently observed swimming along the reef slope, but outside transect boundaries. Overall, the species of Cephalopholis tended to dominate numerically with many Epinephelus spp. being rarely observed. The Epinephelus groupers commonly observed (E. spilotoceps, E. merra, and E. macrospilos) were similarly colored, a white to cream background with brown spots or hexagonal markings. The inner reef crest of Olhugiri had 16 species present and the outer reef crest 15. The dominant species on both reefs was C. argus, with a median number per transect of 7 inside and 6 outside (Table 2). C. leopardus and E. spilotoceps were the second most abundant species on the inner crest, whereas C. urodeta was second most abundant on the outer slope. Length-frequency distribution: The majority of grouper observed in the lagoons at Gaagandu and Olhugiri were small (5-15 cm Total Length (TL)). No groupers were observed over 25 cm TL. The reef crest and slope had similar size - distributions (X? = 7.07, df = 3, p > 0.05). The < 5 cm and 5-15 cm categories were combined due to an expected value < 1 (Everitt 1992). The majority of grouper observed were 5-25 cm TL. On the slope the smaller grouper (5-15 cm) were dominated numerically by Cephalopholis leopardus and C. urodeta. The largest fish observed on the slope (> 35 cm) were Anyperodon luecogrammicus, Aetheloperca rogaa, C. argus, Variola louti, E. polyphekadian, and C. miniata. Fish observed were mostly less than 50 cm TL. Fish greater than 50 cm were mostly V. louti and P. laevis. The larger grouper observed on the reef crest (25-35 cm) were C. argus. Reef 1 had similar numbers of fish in the 5-15 cm, 15-25 cm, and 25-35 cm categories when compared to the other sites at Gaagandu (X? = 0.43, df = 2, p > 0.05). Reef 1 had a larger percentage contribution of the > 8 Table 2: Median and maximum number of groupers observed per transect (median, maximum) for Gaagandu slope (GS), Gaagandu crest (GC), Gaagandu lagoon (GL), Gaagandu reef 1 (GR), Olhugiri inside crest (OI), Olhugiri outside crest (OO). The minimum number observed per transect was zero except * = 3, + = 1, and # = 2. % = species observed outside boundaries of transects SPECIES GS GC” Gl. (GEKea Or OO Number of transects 48 11 2 12 13 6 Aethaloperca rogaa 156 --- --- = 0,1 ee Anyperodon luecogrammicus 1,4 0,2 --- 12 0,2 --- Cephalopholis argus 3,11 Bhi vatgaas 5: naeOs te (OF C. leopardus 4.5,16 --- --- 0,1 1,6 Ua C. miniata 5: aes) ele - 15 meters deep. Variola louti was not observed in caves or hiding in the Society Islands, but swam off the bottom (Randall and Brock 1960). Gracila albomarginata was observed frequently in shallow water 5-10 m, however, Randall and Heemstra (1991) reported that this species was more abundant in depths greater than 15 m. This species tended to swim along the slope and did not appear to hide when frightened, but swam away, as is consistant with Randall and Heemstra’s (1991) observations. Smith-Vaniz et al. (1988) also indicated that this species was an active swimmer, not resting on the reef substratum. Plectropomus areolata appeared more substrate attached; the younger ones were observed swimming among the Acropora thickets on reef 1. The species of Plectropomus feed mainly on fishes and tend to be less sedentary than most groupers (Randall and Hoese 1986). Aethaloperca rogaa tended to be intermediate between these free-roaming species and the more substrate attached species. Individuals tended to swim about freely, but would often hide under coral heads and ledges when approached. They did not traverse long distances as did the previously mentioned species, but would remain near a large coral structure in the water column. The reef slope contained areas with high coral relief, in between which occurred *landslides’ of coral rubble and sand. Stoddart (1966) documented these same features of Maldivian reefs. These rubble patches were frequently inhabited by small Cephalopholis urodeta and, especially, Epinephelus spilotoceps. The latter species was usually observed on the edge of these rubble patches near high coral relief rather than out in the open. Epinephelus merra was abundant in the lagoons of the islands and at reef 1. This species is similar to E. spilotoceps, being a demersal carnivore living under ledges near the bottom of coral mounds and rubble (Hiatt and Strasburg 1960). E. merra is typically found in shallow water on patch reefs in lagoons and bays (Heemstra and Randall 1993). Many C. urodeta observed had a coloration with the posterior 1/3 to 1/2 of the fish black. Species descriptions of this fish indicate that the Indian Ocean variety has only a dark caudal fin, but that in "dark habitats" in the Comoros Islands it was uniformly brown (Randall and Heemstra 1991). Small specimens (< 10 cm) of C. urodeta were observed in shallow water that appeared uniformly black or with a red head region and black body posteriorly. Most of the individuals of this species conformed to the species description in Randall and Heemstra (1991), however many followed this pattern of more extensive black coloration on the posterior 1/3 to 1/2 of the body and the soft dorsal and anal fins. Cephalopholis urodeta is strongly demersal and rarely ventures away from shelter (Hiatt and Strasburg 1960). The most site attached of the slope species was C. leopardus. It was always seen within patches of coral with closely set *finger’ arrangements. When approached it would dart into the coral head. Anyperodon luecogrammicus was often seen in pairs. Cephalopholis sexmaculata was observed only in Caves as iS consistent with the observations of Randall and Ben-Tuvia (1983). 13 C. argus tended to have a higher density at shallower depths and dominated the diversity on the reef crest. This species is one of the most common food fishes (Randall et al 1985), and is generally one of the most abundant piscivores at most locations thoughout the Indo-Pacific (Randall and Ben-Tuvia 1983). It is more common on exposed rather than protected reefs (Randall and Brock 1960) and prefers depths of 1-10 m (Heemstra and Randall 1993). Shpigel and Fishelson (1989) found this species on the shallow reef table and reef wall in the Gulf of Elat. Harmelin-Vivien (1977) observed C. argus at depths of 6-18 m on spur and groove reefs and 18-25 m on the lower sloping platform at Tulear. Cephalopholis miniata is abundant in deep lagoons and dominates coral knolls that are isolated at depths of 17-33 m (Randall and Brock 1960). At one knoll off the slope at Gaagandu at 30 m depth, this species was the most numerous of the groupers observed. The grouper species observed on the reef crest and lagoon were in close association with structural features such as overhangs and crevices (with the exception of C. argus, which roamed about freely while darting into cover when approached). The species observed in the lagoon were all similarly colored (brown spots or hexagons on a light background) and tended to blend into the background of algal covered rubble. Hiatt and Strasburg (1960) found E. macrospilos under large coral heads and rock ledges, seldom far from cover. Our observations on this species in the lagoon at Gaagandu support their findings. Epinephelus fasciatus was observed in the lagoon closely associated with shelter. Fishelson (1977) observed this species near rocks in the lagoon of the Gulf of Eilat (Aqaba) as well as in the fore reef. The number of transects required to adequately characterize grouper density and diversity is dependent upon the dispersion patterns and the desired levels of precision, accuracy, and percent of the species observed in the community. A single visit to a reef is not likely to record all species present, especially cryptic ones (Sale and Douglas 1981). An analysis similar to that conducted here could be done on a preliminary set of transects in order to determine the number of transects required. The number of transects should be determined not only by the dispersion patterns of the species of interest, but also by logistical constraints on effort. Collecting a large sample might increase accuracy minimally and use time that could be applied to other sites (Bros and Cowell 1987). In addition, if only species densities are required, the level of effort devoted to a particular species could be tailored to the degree to which a species is clumped. Only a few transects would be required to characterize the density of a randomly dispersed species, while a species which is clumped would require more transects. The groupers observed in this study appeared to have specific habitat requirements or preferences. The dispersion of the groupers throughout the site is probably related to the dispersion of their preferred habitat. Cephalopholis leopardus is strongly substrate attached and its distribution was significantly clumped (Table 4). The clumped distribution of the species is likely due to a clumped distribution of its preferred habitat. Thirteen transects would be needed to adequately characterize the density of this species whereas a species such as Aethaloperca rogaa which had a random distribution (Table 4), would need only 3 transects. A. rogaa is a species which is not strongly substrate attached. However, our data on Anyperodon luecogrammicus does not follow this pattern as it was randomly dispered, but appears to be strongly substrate attached. A 14 more detailed investigation of its habitat might reveal that it is a generalist in its association with the substrate. ACKNOWLEDGEMENTS We gratefully acknowledge the help of Mohamed Haleem, Omar Maniku, Ahmed Shakeel, and Steve Holloway. Without their contributions this research could not have been accomplished. We also thank the men of Gaagandu and Olhugiri Islands for helping with the research and providing a great living environment. The manuscript was significantly improved by two anonymous reviewers. This project was sponsored by the Oceanographic Society of Maldives. LITERATURE CITED Allen, G.R. and R.C. Steene. 1987. Reef fishes of the Indian Ocean. T.F.H. Publications, New Jersey. 240 pp. Anderson, R.C., Z. Waheed, M. Rasheed, and A. Arif. 1992. Reef fish resources survey in the Maldives - Phase II. Bay of Bengal Program BOBP/WP/80, Madras, India. Bell, J.D., G.J.S. Craik, D.A. Pollard, and B.C. Russel. 1985. Estimating length- frequency distributions of large reef fish underwater. Coral Reefs 4:41-44. Bortone, S.A., R.W. Hastings, and J.L. Oglesby. 1986. Quantification of reef fish assemblages: a comparison of several in situ methods. Northeast Gulf Science 8:1-22. Bros, W.E. and Cowell, B.C. 1987. A technique for optimizing sample size (replication). J. Exp. Mar. Biol. Ecol. 114:63-71. Brower, J., J. Zar, and C. von Ende. 1990. Field and Laboratory Methods for General Ecology. Wm. C. Brown Publishers, Dubuque, IA, 237pp. Craik, G.J.S. 1981. Underwater survey of coral trout Plectropomus leopardus (Serranidae) populations in the Capricorn section of the Great Barrier Reef Marine Park. Proc. 4th Int. Coral Reef Symp. 1:53-58. De Martini, E.E. and D. Roberts. 1982. An empirical test of biases in the rapid visual technique for species-time censuses of reef fish assemblages. Mar. Biol. 70:129-134. Everitt, B.S. 1992. The Analysis of Contingency Tables, Second Edition. Chapman & Hall, New York. 164pp. Fishelson, L. 1977. Sociobiology of feeding behavior of coral fish along the coral reef of the Gulf of Elat (= Gulf of Aqaba), Red Sea. Isr. J. Zool. 26:114-134. 15 Fowler, A.J. 1987. The development of sampling strategies for population studies of coral reef fishes: a case study. Coral Reefs 6: 49-58. Great Barrier Reef Marine Park Authority (GBRMPA). 1979. Great Barrier Reef Marine Park Authority workshop on reef fish assessment and monitoring. Workshop Series No. 2 GBRMPA, Townsville, Australia. 64pp. Greene, L.E. and W.S. Alevizon. 1989. Comparative accuracies of visual assessment methods for coral reef fishes. Bull. Mar. Sci. 44:899-912. Harmelin-Vivien, M.L. 1977. Ecological distribution of fishes on the outer slope of Tulear reef (Madagascar). Proc. Int. Coral Reef Symp. 3rd 1:289-295. Heemstra, P. and J.E. Randall. 1984. Serranidae. In: Fischer, W. (Ed.), FAO Species Identification Sheets for Fishery Purposes, Western Central Atlantic (fishing area 31). Vol. 4,5. FAO, Rome, Italy. Heemstra, P.C. and J.E. Randall. 1993. FAO Species Catalogue. Vol. 16. Groupers of the world (Family Serranidae, subfamily Epinephelinae). An annotated and illustrated catalogue of the grouper, rockcod, hind, coral grouper and lyretail species known to date. FAO Fisheries Synopsis No. 125, Vol.16. Rome, FAO. 382pp. Hiatt, R.W. and D.W. Strasburg. 1960. Ecological relationships of the fish fauna on coral reefs of the Marshall Islands. Ecol. Monogr. 30:65-127. Randall, J.E. 1992. Diver’s guide to fishes of Maldives. Immel Publishing, London. 193 pp. Randall, J.E., M.L. Bauchot, and A. Ben-Tuvia. 1985. Cephalopholis argus Schneider, 1801 and Cephalopholis sexmaculata (Ruppell, 1830) (Ostiechthyes, Serranidae: Proposed conservation by suppression of Bodianus guttatus Bloch, 1790, Anthius argus Bloch, 1792 and Serranus zanana Valenciennes, 1828 Z.N.(S.)2470). Bull. Zool. Nom. Vol 42 pt.4:374-378. Randall, J.E. and A. Ben-Tuvia. 1983. A review of the groupers (Pisces: Serranidae: Epinephilinae) of the Red Sea, with description of a new species of Cephalopholis. Bull. Mar. Sci. 33:373-426. Randall, J.E. and V.E. Brock. 1960. Observations on the ecology of epinepheline and lutjanid fishes of the Society islands, with emphasis on food habits. Trans. Am. Fish. Soc. 89:9-16. Randall, J.E. and P. Heemstra. 1991. Revision of Indo-Pacific groupers (Perciformes: Serranidae: Epinephelinae), with descriptions of five new species. Indo-Pacific Fishes 20: 1-332. 16 Randall, J.E. and D.F. Hoese. 1986. Revision of the groupers of the Indo-Pacific Genus Plectropomous (Perciformes: Serranidae). Indo-Pacific Fishes 13:1-31. Sale, P.F. and W.A. Douglas. 1981. Precision and accuracy of visual census technique for fish assemblages on coral patch reefs. Env. Biol. Fishes 6:333-339. Sale, P.F. and B.J. Sharp. 1983. Correction for bias in visual transect censuses of coral reef fishes. Coral Reefs 2:37-42. Sanderson, S.L. and A.C. Solonsky. 1986. Comparison of a rapid visual and a strip transect technique for censusing reef fish assemblages. Bull. Mar. Sci. 39:119-129. Shepherd, A.R.D., R.M. Warwick, K.R. Clark, and B.E. Brown. 1992. An analysis of fish community responses to coral mining in the Maldives. Env. Biol. Fishes 33:367-380. Shpigel, M. and L. Fishelson. 1989. Habitat partioning between species of the genus Cephalopholis (Pisces, Serranidae) accross the fringing reef of the Gulf of Aquaba (Red Sea). Mar. Ecol. Prog. Ser. 58:17-22. Smith-Vaniz, W.F., G.D. Johnson, and J.E. Randall. 1988. Redescription of Gracila albomarginata (Fowler and Bean) and Cephalopholis polleni (Bleeker) with comments on the generic limits of selected Indo-Pacific groupers (Pisces: Serranidae: Epinephelinae). Proc. Acad. Nat. Sci. Philad. 140(2):1-23. St. John, J., G-.R. Russ, and W. Gladstone. 1990. Accuracy and bias of visual estimates of numbers, size structure and biomass of a coral reef fish. Mar. Ecol. Prog. Ser. 64:253-262. Stoddart, D.R. 1966. Reef studies at Addu Atoll, Maldive Islands. Atoll Res. Bull. 116. 122pp. Zar, J.H. 1984. Biostatistical Analysis, 2nd Edition. Prentice-Hall, Inc., Englewood Cliffs, N.J. 718 pp. ATOLL RESEARCH BULLETIN NO. 439 EFFECT OF TYPHOONS ON THE LIZARD COMMUNITY OF A SHELF ATOLL BY MICHAEL JAMES MCCOID ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. APRIL 1996 Cocos I. ~ 4 Km is es ae | Figure |. Map of the Mariana archipelago with the location of the study site. EFFECT OF TYPHOONS ON THE LIZARD COMMUNITY OF A SHELF ATOLL BY MICHAEL JAMES MCCOID!.2 ABSTRACT Two major typhoons hit the southern Mariana Islands within an 11 month span and provided a- unique, unplanned opportunity to investigate storm influences on the herpetofauna of an atoll. Habitat specialists (Emoia atrocostata and Cryptoblepharus poecilopleurus) endured the largest population declines because of habitat destruction. All other species, particularly scincids, suffered less drastic population declines. The highest population declines for all species occurred on the developed (resort) end of the island, suggesting that removal and restructuring of typhoon-adapted vegetation allowed complete overwash and local extirpations. Cumulative effects of typhoons suggest a resilience to storm influences by atoll-dwelling reptiles. The Mariana Islands comprise an archipelago of volcanic origin oriented north-south roughly equidistant between New Guinea and Japan. There are 15 major islands, with the northernmost (Farallon de Pajaros = Uracas) located at approximately 20°N, 145°E and the southernmost (Guam) at 139N, 145°E (Fig. 1). Two km south of Guam, situated on the southern portion of a coral lagoon, is Cocos (Dano) Island. This atoll has a maximum elevation of 2 m and is approximately 100 m by 2 km. As of September 1992, forest vegetation on Cocos Island was dominated by Cocos nucifera (Coconut Palm), Hernandia sonora (no common name) and Casuarina equisetifolia (Australian Pine). Understory vegetation in the forest was dominated by Carica papaya (Papaya) with ground cover dominated by unidentified grasses and Ipomoea pes-caprae (Railroad Vine). Bordering the surf / tidal splash zone on the windward side of the atoll were dense thickets of Pemphis acidula (no common name). Vegetation on the developed (resort) northeastern 1/3 end of the atoll was dominated by C. equisetifolia, C. nucifera, and ornamental trees and shrubs. An historical record detailing vegetation was provided by Neubauer and Neubauer (1981). The atoll has undergone substantial changes during the past half-century including the development of a coconut plantation prior to WWII, construction of a U. S. military installation (formerly occupying approximately 1/4 of the island), two resorts (occupying a total of 1/2 the island; the present resort occupies only 1/3 the atoll), and at least three typhoons since 1949 that overwashed the island (Neubauer and Neubauer, 1981; per. obs.). Only about 1/3 of the island remains as atoll forest, albeit regenerated. The climate of the southern Marianas is tropical with annual diurnal temperatures ranging between 22° and 31°C (Anon., 1990). Rainfall is seasonal (Anon., 1990) with 1Division of Aquatic and Wildlife Resources, P. O. Box 2950, Agana, Guam 96910, USA. ; 2Present Address: Caesar Kleberg Wildlife Research Institute, Texas A&M University, Kingsville, Texas 78363, USA. Manuscript received 19 July 1994; revised 25 April 1995 Z most occurring between June and December. Typhoons in the western Pacific are common and have been recorded on Guam in most months of the year (Myers, 1991). The typhoon season on Guam is between June and December. Information on the effects of typhoons on the fauna of atolls is minimal; Jackson (1967) reported that insects and vertebrates persist despite catastrophic impacts and that lizards "somehow have found sufficient protection". Damage to and recovery of vegetation is better documented, with estimates of as long as ten years for a marked recovery (Wiens, 1962). In this unplanned study, I document changes in the herpetofauna of Cocos Island after the cumulative effects of two major typhoons. While typhoons are a yearly event in the Mariana Islands, two storms of severe magnitude recently hit Guam within an 11 month span. Typhoon Russ hit Guam in December 1990 and Typhoon Yuri in November 1991. Minimum sustained wind speeds to attain classification as typhoons are the same as hurricanes (>74 MPH = 119 KPH) but these storms had sustained wind speeds recorded at 175 MPH (281 KPH). Along southeastern exposures (including Cocos Island), the direction that typhoons usually approach Guam, maximum estimated wave heights were 9 m. Damage caused by high winds and waves, in both typhoons, were substantial on Guam and catastrophic on Cocos Island. Typhoon Russ totally overwashed the atoll, defoliated all broadleaf vegetation, and downed an unknown, but large number of trees, particularly C. equisetifolia along the windward side of the island. Typhoon Yuri inflicted similar damage including loss of a substantial portion of the remaining C. equisetifolia on the windward side of the atoll. An estimated 40-60% of C. equisetifolia on Cocos Island were cumulatively lost during the typhoons. Another cumulative overt vegetation change observed was the virtual elimination of the P. acidula thickets bordering the high energy zone on the windward side of the atoll. An estimated 95% of the thickets were destroyed by Typhoon Yuri. Between typhoons, dominant forest vegetation releafed, seeded, and a dense understory of C. papaya and C. nucifera developed. Also during this period, the remaining P. acidula thickets releafed. Due to Typhoon Yuri, the papaya and coconut palm understory was destroyed and tremendous amounts of debris from the resort were strewn throughout the forest. The dominant understory vegetation that emerged after the second storm was /. pes-caprae. The herpetofauna of the Mariana Islands has been characterized as depauperate (Rodda, et al. 1991) consisting of a pre-western contact terrestrial reptile fauna of 13 species (McCoid, 1993). Ten of these species occur on Cocos Island (Gehyra mutilata, G. oceanica, Lepidodactylus lugubris, Perochirus ateles, Cryptoblepharus poecilopleurus, Emoia cyanura, E. caeruleocauda, E. atrocostata, E. slevini, and Varanus indicus) and an additional two species (Hemidactylus frenatus and Carlia cf. fusca), both introduced to the Marianas (McCoid, 1993), are established on Cocos Island. At present, Cocos Island possesses the most diverse reptile fauna (12 species) of any island in the Mariana archipelago. Declines in the herpetofauna of the Mariana Islands were discussed by Rodda, et al. (1991) but most species formerly found on Guam still occur on Cocos Island. Although there are no native amphibians on the Mariana Islands, Bufo marinus is established on Guam and Cocos Island. The pre-typhoon reptile fauna on Cocos Island was not uniformly distributed in all habitats. The gekkonids G. oceanica, H. frenatus, and P. ateles, were found in both developed and forested areas (McCoid and Hensley, 1994), but differences in densities between these habitats were not investigated. Gehyra mutilata and L. lugubris, however, were far more common in the relatively undisturbed forested areas; | encountered only two L. lugubris in the resort area during nocturnal surveys and no G. mutilata (G. Rodda, pers. com., recorded these species in the forest). Scincids were also not evenly distributed in all habitats. Carlia cf. fusca, perhaps introduced as recently as the late 1980's to Cocos Island (T. Fritts, pers. com.) was found only at a boat landing and public park on the western end 8 of Cocos and at the resort on the eastern end on the island in early 1989. By mid-1990, the species was observed in intervening habitats on Cocos Island. By early 1991 (see below), the species was abundant in all areas. Cryptoblepharus poecilopleurus was most conspicuous on the windward (east) side of the island where it commonly occurred on tree trunks in C. equisetifolia groves (Hensley and McCoid, 1994). Generally, any tree with a trunk diameter > 2.5 cm had at least one resident C. poecilopleurus. Emoia cyanura was found in both resort and forest areas but was associated with sunlit, open habitat. Expansive areas of dense undergrowth harbored few individuals. Emoia caeruleocauda favored heavily shaded areas and was common in the forest and resort, but was occasionally found in open areas. Emoia atrocostata was restricted to the high energy P. acidula zone (total habitat 4 ha) on the windward side of Cocos Island. Emoia slevini only occurred in forest (total habitat 9 ha) (McCoid, et al. 1995). Qualitative surveys of the herpetofauna of Cocos Island were initiated in April 1989 and initially consisted of nocturnal surveys for gekkonids, diurnal surveys for arboreal scincids (both time-constrained surveys), and diurnal surveys for terrestrial scincids using rubberbands. Time-constrained surveys (N = 5, between April 1989 and December 1991) for C. poecilopleurus were limited from 15 to 30 min during which all lizards seen while walking through C. equisetifolia groves were recorded. Time-constrained surveys for gekkonids were conducted on the resort and lasted between 1.5 and 2 h during which all lizards encountered along a predetermined route were either collected or recorded. In September 1990, sticky traps (see Rodda, et al. 1993), which provide a mechanism to estimate relative abundance, were first employed to sample terrestrial reptile faunas in forested, resort, and beach areas of Cocos Island. Traps (10-80) were placed at five m intervals and checked every 15 min at which time any lizards captured were removed. Generally, sticky trapping spanned the time between 0700 and 1200 h. Rubber-banding was only rarely employed after September 1990. After the December 1990 typhoon, nocturnal surveys were discontinued (see below) and only arboreal diurnal and sticky trapping survey techniques were used. Pre-typhoon Russ herpetological surveys of gekkonids in the resort yielded a qualitative estimated community structure (expressed as percentage of total number of lizards) of P. ateles (4.5 %), G. oceanica ( 6.2 %), L. lugubris (0.6 %), and H. frenatus (88.8%) (N = 315 lizards in 30 person-hours survey effort). Unfortunately, the survey route for gekkonids was completely destroyed by the cumulative effects of both typhoons. This was exacerbated by the clean-up efforts of the resort corporation in which remaining debris was removed. Thus, no comparable post-typhoon data could be generated. Surveys immediately after Typhoon Yuri yielded no lizards of any species on the approximately 1/3 of the island occupied by the resort. This portion of the island was subjected to the most intense vegetation / structural loss from typhoons. Although gekkonids were common in the resort prior to the typhoons, population densities of gekkonids in the relatively unsurveyed forest sections of Cocos Island are unknown; I can only assume that a sizable fraction of the gekkonids on Cocos Island were lost because of typhoons. Post-Typhoon Yuri diurnal surveys in forest areas targeting gekkonids revealed the persistence of all previously recorded species on Cocos Island. Pre-typhoon sticky trapping surveys for E. atrocostata yielded a Catch-Per-Unit-Effort (CPUE) of 0.304 lizards/trap hr (N = 51 lizards, trap hrs = 168). Trap-hours are defined as one trap set for one hr = one trap hr. CPUE's are the number of lizards captured/trap hr. Post-typhoon surveys yielded a CPUE of 0.022 (N = 2 lizards, trap hrs = 90). This is a decline of an order of magnitude in catch rates and suggests that the population on Cocos Island declined by over 90% due to cumulative typhoon effects. The remaining Emoia species (cyanura, caeruleocauda, and slevini) and C. cf. fusca can be discussed as a group as no changes in ranking of species collected (see below) in the forest area were noted after or between typhoons. These four species were initially 4 sampled in forest using rubber-banding in early 1989 through late 1990 and sticky trapping in September 1990. Initial levels of efforts were low (total trap hrs = 22) or not quantifiable (rubber-banding). Numbers of lizards collected, ranked in terms of most to least abundant, indicated that C. cf. fusca was the most common followed by E. caeruleocauda, E. cyanura, and E. slevini. All sticky trapping surveys in the forest after December 1990 (N = 5) were conducted along the same transects and yielded the same ranking in abundance as above. Trapping (N = 1400 trap hrs) was conducted in January, June, October, and December 1991, and September 1992. Two surveys (January 1991 and December 1991) were conducted within two weeks after typhoons. Percentage composition for each of the species (grand total = 365 skinks) in the five forest surveys ranged between 57.6 and 68.9 for C. cf. fusca, 20.7 and 30.3 for E. caeruleocauda, 2.6 and 12.9 for E. cyanura, and 0.0 and 2.6 for E. slevini. Changes in percentage compositions between surveys were tested using a R X C test of independence with a William's correction and were not significantly different (X7cale,12,.05 = 9.197). This suggests that responses of individual species to typhoon effects were not statistically different. Similarly, CPUE's for all surveys were within the same order of magnitude (range 0.171 - 0.475) indicating that the cumulative effects of the typhoons did not dramatically decrease catch-rates of forest-dwelling scincids. Since at least 1/3 of the island was devoid of any lizards after Typhoon Yuri (see above), it is safe to assume that total population declines were greater for E. cyanura, E. caeruleocauda, and C. cf. fusca than for E. slevini, which occurred only in forest. Numbers of C. poecilopleurus were gauged by sightings per min (range 0.33 - 1.1). These sighting data, including both pre- and post-typhoon observations, are within the same order of magnitude suggesting that typhoon effects were minimal on survivorship of C. poecilopleurus. importantly though, post-typhoon observations were made on existing trees and since sighting rates after typhoons did not increase on these trees, perhaps indicating emigration of surviving lizards from felled trees to existing trees, it is assumed that if a tree was lost during a typhoon, the resident lizards were also lost. The ability of a herpetofauna to persist on an atoll after substantial environmental perturbations are also highlighted by observations on two species not directly surveyed in this report. Varanus indicus, although found on Guam, was probably introduced to Cocos Island in the late 1980's (pers. obs.) and managed to persist through two major typhoons. By December 1991, in addition to a number (3 - 5) of 200 to 450 mm snout-vent length (SVL) lizards, a small (ca. 100 mm SVL) individual had been observed on Cocos Island. These observations suggest that successful reproduction had occurred and monitor lizards had survived the typhoons. Bufo marinus was probably introduced to Cocos Island in 1989 and successful reproduction (large numbers of tadpoles in rain pools) was observed in September 1989. In September 1992, after both typhoons, two adult (ca. 830 mm SVL) B. marinus were observed in a freshwater pool. Observations of the herpetofauna on Cocos Island after typhoons suggest a resilience to environmental perturbations. Terrestrial forest-dwelling scincid populations appeared to persist relatively unscathed despite substantial typhoon impacts. Habitat specialists (E. atrocostata and C. poecilopleurus) were more susceptible to population declines due to habitat destruction. All gekkonid species also persisted after the substantial effects of the typhoons. Besides C. poecilopleurus and E. atrocostata, the largest localized population declines of other species are associated with the developed (resort) section of the atoll. This may be related horticultural / architectural practices that restructure typhoon adapted vegetation allowing complete overwash and loss of most structures, soil, and sand during severe storms. Considering the absence of all lizards on the resort 1/3 of the atoll, a substantial fraction of the lizard population was lost because of the cumulative effects of typhoons. Habitat 5 specialists E. atrocostata and C. poecilopleurus probably suffered much greater population declines, which is related to susceptibility of these habitats to typhoon damage. Despite that, the data suggest that relatively undisturbed atolls will tend to retain herpetofaunal components despite substantial typhoon influences. ACKNOWLEDGMENTS Assistance in the field was provided by Rebecca Hensley, Robert Cruz, and Earl Campbell III. Gordon Rodda and Thomas Fritts generously provided unpublished data. Rebecca Hensley, Gordon Rodda, Thomas Fritts, and Kevin de Queiroz commented on a version of the manuscript. Portions of this study were funded by the Endangered Species Conservation Program, Project E-4 (to Guam) and by the U. S. Department of the Interior, National Biological Survey. LITERATURE CITED ANONYMOUS. 1990. Local climatological data. Annual summary with comparative data. Guam, Pacific. NOAA Natl. Clim. Data Center, Asheville, NC. 8 p. HENSLEY, R. A. and M. J. MCCOID. 1994. Cryptoblepharus poecilopleurus (Snake- eyed Skink). Activity. Herpetol. Rev. 25:121. JACKSON, W. B. 1967. Productivity in high and low islands, with special emphasis to rodent populations. Micronesica 3:5-15. MCCOID, M. J. 1993. The 'new' herpetofauna of Guam, Mariana Islands. Herpetol. Rev. 24: 16-17. .and R. A. HENSLEY. 1994. Distribution and abundance of Perochirus ateles (Gekkonidae) in the Mariana Islands. Herpetol. Rev. 25: 97-98. ,G. H. RODDA, and T. H. FRITTS. 1995. Distribution and abundance of Emoia slevini (Scincidae) in the Mariana Islands. Herpetol. Rev. 26: in press. MYERS, R. F. 1991. Micronesian Reef Fishes. 2nd ed. Coral Graphics, Barrigada, Guam. 298 p. NEUBAUER, C. P. AND D. R. NEUBAUER. 1981. The vegetation of Cocos Island (Mariana Islands). In L. Raulerson (ed.). Plant biogeography of Guam. Univ. Guam Mar. Lab. Tech. Rep. 69. pp. 23-39. RODDA, G. H, T. H. FRITTS, AND J. D. REICHEL. 1991. The distributional patterns of reptiles and amphibians on the Mariana Islands. Micronesica 24: 195- 210. , M. J. MCCOID, AND T. H. FRITTS. 1993. Adhesive trapping II. Herpetol. Rev. 24:99-100. WEINS, H. J. 1962. Atoll Environment and Ecology. Yale Univ. 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BO cio can ‘ Re. eae “i. a UMA © a TAG ey daha mn ee Mad dad Cohdeanueniaten i wd civithaneloconehinal jive baa p ti wut a eee NM A eget cc Wes} sree) aise i my Me We a MAR oe Une Piet «yas wren’ Ie Tyee Hinata tt ub al P| pW tea ia Vahudieh. ay) Viper Med ji tae 25 bated veo mena pom i Selene ted anak ste wr nfo lasiagtiom twirl aot mon ss , med ec Miami mie Fk tle Lorast boaathe od Lat | ier are atvaniahead whee abagalenpmadl Gieaet seek ak ee ail Central skubieg tevep gems ty vr mala eal RRR Bt b SUVA Meal ATA AWD nh Magia PVA SC a aD: Deh a i renin Maines nf MAL reasty ce beled » aut Ts tie re (acetate eh Shik iA) | nut | “atuinet wl Me ee ee ATOLL RESEARCH BULLETIN NO. 440 FLOWERING AND FRUITING IN THE FLORA OF HERON ISLAND, GREAT BARRIER REEF, AUSTRALIA BY R.W. ROGERS ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. APRIL 1996 yi \ Genie? See WITAIIUG HORAGEAS JI01 ay co a Abi KONTO AMOI TIT M1 OATTIUNT GA OVIAWORE iaiasebern 0 AMLAAR TARA SHADOW Ya (ae (OTA JARUTAM FO MUU AAMONT AM VOITUTTP SYR FART Ee en al, OT OMIRA AT | ove) JADA FLOWERING AND FRUITING IN THE FLORA OF HERON ISLAND, GREAT BARRIER REEF, AUSTRALIA. BY R.W.ROGERS' ABSTRACT The plant species in flower and fruit on Heron Island, a sandy cay on the Great Barrier Reef, Australia, were observed at intervals of three months for three and a half years. At no time were less than 20 nor more than 36 of the 49 species monitored found to be in flower, nor were less than 20 nor more than 41 of the 50 species monitored found to bear fruit. Despite a strongly seasonal climate there was not a strong seasonal pattern evident in the number of species in flower or fruit, although some species were themselves strongly seasonal. A principal components analysis of all flowering records, however, demonstrated a seasonal polarity with March and September representing the two extremes. Fleshy fruited species, important for frugiverous birds such as silvereyes, bore fruit throughout the year. INTRODUCTION Temporal patterns in flowering and fruiting are significant attributes of vegetation, for these are attributes subject to selection as are any others. Variation in seasonal flowering patterns has proved to be significant in understanding of heathlands in Australia (Specht et a/. 1981) and Europe (Woolhouse & Kwolek 1981), both in terms of ecophysiology, and in terms of the evolutionary derivation of the floras. The availability of flowers and fruit is manifestly important to those animals which depend on fruit, seed and nectar as food resources, and an interaction between plant phenology and the birds responsible for seed dispersal has been postulated (Herrera 1986). There has been little previous study of the temporal variations in flowering and fruiting of the plants on the cays of the Great Barrier Reef, although Heatwole (1981) noted flowering times for a few species on One Tree Island. There has apparently been little if any study of seasonality on similar islands elsewhere. Opportunity to visit Heron Island on a regular basis was therefore used to collect information on the flowering and fruiting patterns of the flora of that island. Heron Island is a small coral cay about 800 m x 300 m situated 70 km offshore from the Australian continent (Lat. 23°26' Long. 151°5S'E). Much of the island is clad in a dense forest of Pisonia grandis trees, with some fringing grasslands (Fosberg 1961). The island has been the subject of extensive weed invasions during its recent history of intense human activity, and this has been documented in particular for a spoil dump established on the island in 1987 (Rogers 1993) and in more general terms by Chaloupka & Domm (1986). ‘Botany Department, The University of Queensland, Queensland 4072, Australia. Manuscript received 4 May 1994; revised 21 April 1995 Heron Island has a strongly seasonal climate, with mean rainfall varying from as little as 20 mm in September to as much as 145 mm in February. The four months July to November receive in total less than 15% of the annual rainfall (Fig. 1). A consequence of this strong seasonality is that Pisonia grandis and Ficus opposita often lose many of their leaves by December, and the native grass cover of the island dies. 160 140 120 100 80 60 Mean rainfall (mm) 40 20 Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Month Figure 1 Distribution of mean monthly rainfall through the year for Heron Island (from Walker 1991a). Methods Heron Island was visited at intervals of about three months between September 1990 and December 1993, and the flora surveyed to determine which species were represented by individuals in flower or fruit at the time. These data were recorded and analysed in terms of total species in flower and fruit, native plants in flower and fruit, weedy plants in flower and fruit, native dicots in flower and native and weedy grasses in flower, and numbers of species bearing fleshy fruit. Information was collected in September, December, March and July, for convenience 3 referred to as spring, summer, autumn and winter. Records were made by walking a series of transects on the island which traversed all the vegetation types and the habitats of all the plant species known for the island. To detect pattern in a multivariate system such as that studied here where it is proposed to detect seasonality of flowering in a system comprising 49 species requires a multivariate analysis. One such analysis which is suitable for the purpose is Principal Components Analysis, and in order to reveal underlying patterns in the data a principal components analysis was executed on the flowering data (Wilkinson 1990). Table 1 Seasonality of flowering of plants on Heron Island, Great Barrier Reef, Australia, between September 1990 and December 1993. N = total number of plants in the category. Number of species in flower year 90 90 91 91 91 91 92 92 92 92 93 93 93 93 Month Si) De Mads) Die Med Sab) YM.) Si iD N All species natives 2X5 ALO) 4s IS ass ah aL fs} La US) LG L114) aly 9) weeds AS} 6 iO) WS 7 US) a7 12 WS) Qi 16 14 1 2B 22 Total 49 28 24 32 35 30 34 20 30 36 32 26 33 40 41 Dicots Native 7x 0) 13} Ws 1G 12 16 7 YF Ws les ail 12 WH il6 Weeds TS 7) 9S Be pelea 7S a nO MO, 2 aS) aS Total 2) 22 20) 23 2y 2A Bil 14 22 2 2 ZO AX BO Sil Grasses Native Vat dL, wR TUITE 2), Geen ORES My goal aOR (2) dialer er Lee On eA ee Weeds Sh See Sy ime GwG. GS.) Cho ei wig eae 7M iBict 7 Total ib 6 4 SF Bo 6 FW. 6 tS) th By) 8) ao). ako) Table 2 Seasonality of fruiting of plants on Heron Island, Great Barrier Reef, Australia, between September 1990 and December 1993. N = total number of plants in the category. Number of species in fruit year SOR90 NST eS 1 Oi 91929269259 21493593.9393 Month Sap Oto. DOM ed So. DOM wd. 6S. 6D N All species natives PAI) Wyo De Th ALS) PAO) AUG) AUS) TLD oll Ah ENT ood E'S) TL fg ILS} hI IAS Dae weeds 28) Gh MoD may iG Oe ISteGw20Na Tei A TS 220 total 50,29..26536936136 .40,,20,.30)\37,36. 29..37:1:40 41 Fleshy fruits Att AAT EREUGARAMA AMAR YAN 41S edie Bit & 45 40 Species in flower S90 $91 $92 $93 Season Figure 2.The total number of plant species in flower (4), number of native species in flower (CQ), and number of introduced species in flower(™) Heron Island, Great Barrier Reef, Australia, at three month intervals from September 1990 until December 1993. RESULTS Flowering was recorded for 46 of the 49 species examined for flowers (appendix 1): Ipomoea pes-caprae and Salsola kali were not seen in flower, and flowering of Ficus opposita was not determined because of the syconium within which flowers are produced. No distinct patterns of number of species in flower around the year was apparent (Fig. 2, Table 1), whether the flora was taken as a whole or divided into components. The patterns of variation shown by all groups of species are essentially the same. A depression in the number of species in flower is apparent in July (southern winter) of 1992 and 1993, but July 1991 shows a high number of species flowering. Patterns can be detected in individual species (appendix 1). Apiwm leptophyllum, Bromus catharticus and Wollastonia biflora flower only in spring-summer, whereas Cordia subcordata may be found flowering in any season except summer. Pisonia grandis flowers in summer-autumn. The small herb Gnaphalium luteoalbum appears to flower in any season but autumn. Of the remaining species 17 were recorded flowering in 12-14 of the times surveyed, and 4 in three or less occasions. ar iN eet aE iin a 5 The principal components analysis showed that over 50% of the variance in the flowering matrix was explained by the first two components (36.3% and 15.4% respectively), and that the replicate seasonal collections for March and September represented poles of an ordination, with the summer and winter collections (December and June) falling into an intermediate position (fig.2) Component 2 6 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 Component 1 Figure 3. Plot of the first two components from a Principal Components Analysis of three- monthly flowering records from Heron Island. Each record is identified by month (M = March, J = July, S = September, D = December) and year. Fruiting was observed in 47 of the 49 species (appendix 2), Ipomoea pes-caprae and Salsola kali not being observed in fruit during the study period: expanded syconia of Ficus opposita were treated as fruits. No distinct seasonal patterns were evident in numbers of species fruiting around the year, and the total number of species in fruit closely paralleled the number flowering (fig.3). At least four fleshy fruited species were in fruit at any time, and three of these, Tournefortia argentea, Ficus opposita and Pipturus argenteus bore fruit every time they were observed. DISCUSSION Observations made four times per year do not permit a study of seasonality of flowering in any great detail. However, it is sufficiently frequent to detect the more striking patterns 6 which may be present. It is apparent that the very strong seasonality evident in the rainfall pattern for the Island is not reflected in the number of species in flower or fruit. The polarity demonstrated by the principal components analysis between March and September, however, is quite clear. Thus, although the seasonal differences are diffuse, involving numbers of species showing quite different patterns, the overall view is of well developed seasonality in flowering pattern. At all times studied there was a higher proportion of weed species in flower than of native species, and at all times except December 1993 a higher proportion of weed species in fruit than native species. This is similar to the finding of Odgers and Rogers (1993) that weedy grass species growing together with native Australian grass species showed longer flowering seasons than did the native species. It is not surprising that weedy species, characteristic of frequently disturbed sites, are more likely to have individuals in a reproductive condition than are those species which characteristically occupy relatively undisturbed natural systems. Some of the native species, however, flower and fruit continuously. The continual fruiting of Ficus opposita, Pipturus argenteus and Tournefortia argentea, all of which have fleshy structures associated with their fruits, is probably of great importance to the resident population of silvereyes (Zosterops lateralis) and bar-shouldered doves (Geopelia humeralis) on the island, indeed, Walker (1991b) ascribed the extinction of bar-shouldered doves on Lady Elliot Island to the loss of Ficus opposita from the flora of that island. It is also likely to be important for the plants that a population of seed dispersers is maintained. A continuous flowering is probably also important to the plants in that it permits the maintenance of populations of pollinators in an otherwise isolated habitat. The nectar associated with flowering may also be important to the silvereyes (Barker & Vestjens 1989) and raises the possibility of bird pollination in Tournefortia argentea which has strongly nectiferous flowers, and perhaps in Pipturus argenteus too. The strongly seasonal native trees Cordia subcordata and Pisonia grandis appear not to depend on permanently resident birds for seed dispersal. P. grandis seeds are commonly associated with the migratory sea bird populations which nest amongst them during the fruiting season, especially the white-capped noddy (Anous minutus), and are presumably dispersed adhering to birds (Walker 1991a), although the noddies may suffer heavy fatalities from high P. grandis fruit loads in those years in which P. grandis fruiting is heavy. Cordia subcordata produces a large and rather corky fruit which is often found in drift along beaches and is apparently dispersed by flotation. Boerhavia tetrandra is in the same family (Nyctaginaceae) as Pisonia grandis, but is a prostrate herb which bears sticky fruit similar to that of P. grandis, but displayed only a few centimetres above ground level. Seedlings of Boerhavia tetrandra are commonly observed in disturbed areas of Heron Island, whereas those of Pisonia grandis are seen very rarely, but then in very large numbers within the forest (e.g. March 1972 when several hundred could be found in a square meter). The sticky fruit of Boerhavia tetrandra, however, are more likely to be dispersed by ground feeding birds such as the doves and rails, in contrast to the dispersal of Pisonia grandis seeds by marine bird species. The possibility of coevolution of the frugivorous birds and the flora cannot be dismissed, especially in terms of selection for plants which have an extended fruiting season, for Herrera (1986) has observed that changes in phenology are amongst those most likely to occur in response to frugivory and contingent dispersal. It is recognised that the silvereyes of the Capricorn Group are a distinct variety, differing from their mainland relatives. However, it is not possible to argue for any close co-evolution between the birds of the cays and the flora in the way in which Reid (1991) argued for coevolution of mistletoes and mistletoe birds, although in arid zones mistletoe birds may be dependent on very few species of mistletoes for 7 survival, just as frugivores are on a coral cay. Comparative studies of the eating habits of the mainland and cay varieties of silvereyes and of the seasonality of flowering in Tournefortia argentea may be profitable in this context. Acknowledgements I am grateful to the Queensland National Parks and Wildlife Service for permission to work on Heron Island, and to the staff of the Heron Island Research Station of the University of Queensland for assistance. REFERENCES Barker, R.J. & Vestjens, W.J.M. 1989. 'The Food of Australian Birds. 2. Passerines'. CSIRO, Canberra. Chaloupka,MY., and Domm, S.B. 1986. Role of anthropochory in the invasion of coral cays by alien flora Ecology 67, 1536-1547. Fosberg, F.R. 1961. Description of Heron Island. Attol Research Bulletin 82, 1-4. Herrera, C.M. 1986. Vertebrate dispersed plants: why they don't behave the way they should. In 'Frugivores and Seed Dispersal' (eds A Estrada & T.H.Fleming) pp 5-18. Junk, Dordrecht. Odgers, B.M. and Rogers, R.W. 1993. Contrasting diaspore and vegetation attributes from natural and disturbed habitats in an urban eucalypt forest reserve. Australian Journal of Botany 41, 637-648. Reid, N. 1991. Coevolution of mistletoes and frugiverous birds. Journal of Ecology 16, 457- 69. Rogers, R.W. 1993. Plant colonization of a rubble bank on Heron Island, Great Barrier Reef, Australia. Atoll Research Bulletin 384, 1-8. Specht, R.L., Rogers, R.W. & Hopkins, A.J.M. 1981. Seasonal growth and flowering rhythms: Australian Heathlands. In 'Ecosystems of the World 9B: Heathlands and Related Shrublands' (ed R.L. Specht) pp 5-13. Elsevier, Amsterdam. Walker, T.A. 1991a. Pisonia Islands of the Great Barrier Reef. Part 1. The distribution, abundance and dispersal by seabirds of Pisonia grandis. Atoll Research Bulletin 350, 31-39. Walker, T.A. 1991b. Pisonia Islands of the Great Barrier Reef. Part 3. Changes in the vascular flora of Lady Musgrave Island. Afoll Research Bulletin 350, 31-39. Woolhouse, H.W. & Kwolek, A.V.A. 1981. Seasonal growth and flowering rhythms in European heathlands. In ‘Ecosystems of the World 9B: Heathlands and Related Shrublands' (ed R.L.Specht) pp 29-38. Elsevier, Amsterdam. Wilkinson, L. 1990. SYSTAT: "The System for Statistics'. Evanston, Systat Inc. 8 Appendix 1: Flowering calendar for Heron Island. 1 = plant seen in flower; 0 = no plant seen in flower. Year 90) 9091 191 91194" 92192 92%92)\93).98. 93493 Month S cD. pM AS Dow. oT Si) Di iS ED Species Abutilon indicum Achyranthes aspera Amaranthus viridis Apium leptophyllum Bidens pilosa Boerhavia tetrandra Brachiaria subquadripara Bromus catharticus Cakile edentula Calyptocarpus vialis Capsella bursa-pastoralis Cassytha filiformis Casuarina isetifolia Celtis paniculata Cenchrus echinatus Commicarpus insularum Conyza bonariensis Cordia subcordata Coronopus didymus Cynodon dactylon Digitaria ciliaris Eleusine indica Euphorbia atoto Euphorbia cyathophora Euphorbia prostrata Euphorbia tannensis Ficus opposita Gnaphalium luteo-album Ipomoea pes-caprae Lepidium virginicum Malvastrum coromandelianum Pandanus tectorius Pipturus argenteus Pisonia grandis Poa annua Portulaca oleracea Salsola kali Scaevola taccada Sisymbrium orientale Solanum americanum Sonchus oleraceus Sporobolus virginicus Stenotaphrum micranthum Suriana maritima Thuarea involuta Tournefortia argentea Tribulus cistoides Wollastonia biflora (je) SOPPORHP OPRPERPOOROPROO POP BROOPRHPORHPOOHOOO OF OFPHO BHRHPHPO PRERPRPRP OCOFPOPOOORPOFP HOO BPROORPHFPOFPACDD0D00GO CF COCO HORHOP SCPRPRPOO BPRORFPOOROBRKBPP OPO BEBRBBPBPEPBPEPBPEPEBPHOO CO BHROP HRHROHOPE PRERPRPRP OF PORPORPPRPPOO COP BPRORPOHPOHHPORHPHOO BH BHOHP BPHOPRPE FPOPORPFP OFPPRPRORRPRPPOO POP BPRORPOPOHHPOPRHHOO OF COHFO BPRERPHOP PRRORPP OPPRPRPORRPBPRPOP POP BPRPRPORBPHBHBBPHPOHOO HO C000 BHREHOP CPRRPROO FPODOOFOFPOOFODO COO BERPRPORRPEHOPRHPOHOGO CO C000 COOrPOP SCPRPRPBPEP OCORPPRPRPRPRORPOP POP BRORPOPRBPEPHOHOODOO OF OFOH BHHEOPROE PREPEPRPRP OPPRPRPORPRPOROP POP BHROPROBRBHBPHOBHPOOO OF ORPBRH BRBHHEE PRPPEPRP OFPFPORPOFPOOPORP POP BERPRORBROBRBHEHOO CO HOOP BHRHBHOP SCPRPRPOO OCOPFPORPOFRPOOBHEHP COO BERPRPOBRBEHBHHOHOO CO CGO00 BHHROKOEH COPPRPERRP OFPFPRPOCORPORPORP POP BEBBBPBEBBPEBHOHROHOO OF BHOPR BROPREPE PRRPRPRPRP OFPPRPRORBPBHBOP BOP BRBPBHBBPEEPHPEODOOHOP OF BHBHBHB PREP HEBE PREPRPRPRP PRBPRPRPORPORPRPORP POR BRBBBBBPBBBHBHBEHBHO CO BHBBHP BPRPRBPHPOP ee ee a. = Appendix 2: Fruiting calendar for Heron Island. 1 = plant seen bearing fruit; 0 = no plant seen bearing fruit. Year 90 90 91 91 91 91 92 92 92 92 93 93 93 93 Month Se'DO MiG Ss) DM U.S DM UJ S D Species Abutilon indicum i Achyranthes aspera Amaranthus viridis Apium leptophyllum Bidens pilosa Boerhavia tetrandra Brachiaria subquadripara Bromus catharticus Cakile edentula Calyptocarpus vialis Capsella bursa-pastoralis Cassytha filiformis Casuarina equisetifolia Celtis paniculata Cenchrus echinatus Commicarpus insularum Conyza bonariensis Cordia subcordata Coronopus didymus Cynodon dactylon Digitaria ciliaris Eleusine indica Euphorbia atoto Euphorbia cyathophora Euphorbia prostrata Euphorbia tannensis Ficus opposita Gnaphalium luteo-album Ipomoea pes-caprae Lepidium virginicum Malvastrum coromandelianum Pandanus tectorius Pipturus argenteus Pisonia grandis Poa annua Portulaca oleracea Salsola kali Scaevola taccada Sisymbrium orientale Solanum americanum Sonchus oleraceus Sporobolus virginicus Stenotaphrum micranthum Suriana maritima Thuarea involuta Tournefortia argentea Tribulus cistoides Wollastonia biflora kh OPFPOOO OCOPREFPRPODOORPOPRRO POP BPREROORPORHOOHHOH OF OFPBPH BPRELHO PRROPF OCOFPPOOOOORPBRHE POP BEOORPPORPOCOHODOH OH OHOO HOOrOH SCOPPRPOO BPRORPFPOORPOREBH COO BRBBPBPBBPBPBPEPEPEPEPEPHEH OF BRPOP BHROoPPLE OPPRPRE OFOORPOOPORPRHP POP BRORPORPBPEBPEPEPPORBH BH BPHOP BPROPRPE PRROPP OPRPEPRPORPRPORBRHEP COP BPROPRPOPOPBPBPBPBBEHP OF OFFPO BREEHPOPE PRRPOPP OPRPREPRPORPRPRPRPEBEP POP BRRPRPOPBPEBHPORPOPRHBPHE BPH OPRHPO BPREBPHPOPE SCPFPFPODOD BPODDDODDCOOPRHBHP COO BEBPRPORRPEOOFPOPEHEHR CO C000 HBOOROLH SCOPPRORP DCORPFPRPOOCOOREBHP POO BRORPORPBPEBERPHOFPOHP OF OFOP BRORHE PREPRPRP OPRPFOOOPORPRHP POP BPROPOPRPBPEBEBPEBEBPEBPHOH OF OFPRBHP BPREBPHPOHP PRERPRPRP ORFPORPORPORBPBH POP BEBPRPORPRPRPOOPRHBPHPOHP OO HOOHR BHREPPHEE SOPFPFOO ORPFPORPOFRPOREBPHEPHPR COO BPRBPRPORRPHPORHPOBRBHHE CO C000 BPROPRBPE OPPRPRPRP OPROFPOOORPORPBHR POP BEBBBBPBPEPEPEPPORHBHP OF BROP BHROPRPE PRERPRPRP OPRPFOOORPORPBH POP BPRBBBBPBPBPEPEPHPOOPRBH OF BRBPBHB BPHBRPPPL, PRRPRPRPRP PREPRPOOORBPEBPHRP POP BPREBBPBPEBPRPRPRHPORPHEHBH OO BHRBBP BRPRPHOP ‘eee elaalline pats lors? oS sting (Okie iT piteed nese tial mt ce ee £2 6 beeen seit ™ rere neon “93 99 Serie = te = Ege! ey fo fa ty Pe eS he es SS eet Pee rep cs Pe ont oe 2 Pe oe Sh Shh et + es a 228 Sie es Bee oe eye ha & @ 0 Wrage ae La) ETS Po, hei ier eal Sen fa ius So cyet wey Lowe ’ Word fot 2 Sona int aBraes \nivedse i St or is ke oma t op Mei uesta | no r.” bee res kas grey [Me Me Labctosaioes alta OF Be Leer LTS ON ‘ Soe pi eee aiode “Ad pet 0’ f Lars = 8 Ae St 5 rune s HG $ ie oe 1) A Teodess’ fru iie Liners O* re Ae 0: akte % in fs 2} ogi * s hochpctss! wi pio aie yore aN? eicyor po arragan | “ie Hered ye HL Dateae os Laxelt ft 0 ge See, opie kL sackegene i 4 is) ee eee “ike . a hae sag & ray | po fio cies be mcd ES rs Tae a hota 0 Lnpig sala ssi SITIO | C hewiteottoed acca : at Fo pe et Wi ES ¢ Ce tamesp 6. hock | t ho) Varnes ae ME Nw ag om) Ne Un 9 1 WS Aes elon \) . wk ayy By Livy { otoattet sisal L MUISUBT pe meen LO f i wie SAIS: ) BITES EOR or RU yr ty tel MmrigQn sores? Averery> tim ATOLL RESEARCH BULLETIN NO. 441 NAMU ATOLL REVISITED: A FOLLOW-UP STUDY OF 25 YEARS OF RESOURCE USE BY NANCY J. POLLOCK ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. APRIL 1996 NAA lh, AAA RA IPRA SARE RRA DRA A BAA EAE NAR A AA RBA te Bokak AASARAAR SARS AAARBANARRAPABARAIR SNA DADADED ASI BI nnnnpnedapannnncansssnscnssapeces ceeeers WwW AAS RSS SAY SADA AAA ARR AA RRA at te PARRA REDD A RR Y RODD $ + eens anc ranacernres ccreccnntees soon annnannnnesnamnnnatan | $ $ 3 3 3 A ASAAASAS SSA RAA SAAR SAAT AR SAA AARAAAA SARA SAE AA 3 weer $ A, 3 pence ssstnnancanannannnennnnnancersnannanane 3 AAA An 20 18 16 ABRADED RAR A AA RARE ERA R REARS Rt RRR : : AAS RAR RRS tt tn name a a neta a Bs ARR ARR RRA A Ante Mn nn enn nnn nnn nnn nie Bikar 3 PARANA AA EE EN ENN LIL NOE EEE EES $ i $ $ z ; $ F i ‘Rongelap ~~” RO, prrreener heen renner ne BW Likiep a ist i=] ae CANNON Gd Annas, QO Wotho eeeencee werrenereresy ve RE Kwajalein : . Wotje cs linglaplap Aili AAA DARA D AOE wee 4 Ame T ene enenneeannee 2 asdenne RAAAAE ADR RAASA KERR A RS Ree Re eee me ae erreeeeceererec ts A AAA ARR ORR n eee AAMAS P PAE Peeerono oOo cnet eet eee Peeonereeete try PDEA ABDEADDASLD ERS AADS BID AED DAD EIR RS ARDS IRD SOO ES 68 rw. 166 164 160 : nautical miles The Marshall Islands NAMU ATOLL REVISITED: A FOLLOW-UP STUDY OF 25 YEARS OF RESOURCE USE BY NANCY J. POLLOCK ABSTRACT Reliance on local resources for food remains even though the population of Namu has doubled in size. Breadfruit, pandanus and fish are still the main subsistence foods on an atoll in the northern Marshalls. A restudy of household food uses 25 years after the original study revealed few changes in the supply, but dependency on those foods had doubled, thereby increasing the risks from both natural and market failures. Marshall Islands government support will be necessary to maintain such a population on their home atoll. INTRODUCTION Atolls are ever-changing environments. Nature changes them with all her complex interactions. Human activity also brings many additional changes. This paper documents the impact of human lifestyle over 25 years on one islet of an atoll in the Northern Marshal Islands. It aims to add to the work of a number of scientists who have studied the Northern Marshal Islands over a forty year period. The paper is a vote of thanks to Dr. Ray Fosberg whose work contributed greatly to my attempts to understand the interaction between people and their island environment, particularly their food resources. Data on food resources were gathered over a period of fifteen months spent on Namu Atoll in 1968 and 1969 with the aim of understanding the interrelationships between food use and social organization (Pollock 1970; 1992). Similar data have been the subject of a year long study during 1992-3 in the Marshal Islands, that enabled me to pay two return trips to Namu for a week each, one in November 1992 and another in August 1993. Most of my work has concentrated on one islet, Majkin on the eastern side of the atoll. Namu lies in Ralik or the western chain of the Marshal Islands, some 25 miles southwest of the large atoll of Kwajalein. The lights of Kwajalein can be seen from the northern shores of Namu Namu islet, the northernmost piece of land in Namu Atoll. There are some 56 islets in Namu Atoll, only three of which are regularly inhabited. Namu Namu was the first settled, according to local legend then Majkin to the eastern side, and finally Mae-Leuen in the south. The latter is two islets joined together by a sand bar that formed after a Japanese ship wrecked on the reef during World War II. The whole of Namu Atoll stretches over some 25 miles from north to south, and is 15 miles at its widest point. Dept. of Anthropology, Victoria University, Wellington, New Zealand Manuscript received 25 February 1994; revised 13 December 1994 Rainfall amounts to some 80" - 120" p.a., so that Namu lies near the northern limit of adequate moisture for breadfruit trees to flourish. Today they dominate the upper canopy, as seen from the air, with a few old Lukwej (Calophyllum inophyllum) trees reaching similar heights. The coconut trees are tall too, but are becoming spindly, as many are now 50 and 60 years old. The atoll has been hit by several cyclones in recent years, the latest in December 1992 caused considerable damage to the vegetation; FEMA aid was provided to the people of Namu. Each islet is divided by the Marshallese into three use zones, lik or oceanside, eolap or middle of the island, and iar, or lagoonside. These three zones refer to particular kinds of vegetational zones as determined by human activity. These activities have resulted in differential soil fertility. The usage of these zones is determined in part by the orientation of the piece of land to wind and waves. In addition each islet is divided into two parts, jittoen and jittoken, terms that originally referred to the ways a canoe could head. The lik side of Majkin is exposed to the prevailing northeast wind and thus is battered by salt spray, even though there is a wide protecting reef. Hence only salt tolerant strand vegetation such as Scaevola or Morinda citrifolia (nin) grow here. This side of the islet is mainly used by adults for defecation on the reef; that usage places taboos on visiting the area for other reasons. Eolap, or the middle of the island is quite a small area at either extremity of Majkin, but in the centre of the atoll this area can cover a quarter of a mile. Here the greatest proportion of the coconut trees have been planted, and pits excavated to grow the form of taro best suited to atolls, namely iarej, Cyrtosperma chamissonis. Pandanus trees in a number of varieties, have also been planted on the lagoon side of this area mainly for use of the leaves in making handicrafts. Lar, or the lagoon side of the island, is the most heavily used part of the atoll. Residences, the road and useful trees are concentrated here. The main reason is that this area gives best access to the fresh water lens accessed by means of wells. This fresh water supply is also tapped by the breadfruit trees which have been planted around the residence areas both to give shade and to provide fruit and leaves. Residential areas are readily distinguishable by the white coral which is gathered from the lagoon shore to spread around the house sites to keep them clean and improve drainage. The lagoon shore is also the landing site for any visiting ships, whether sailing canoes, one man dug-out canoes, faster launches with outboard engines, or occasional deep water field trip ships. Landholdings reflect the tripartite use of the land. Each landholding or weto, extends from oceanside to lagoonside in more or less straight lines. This means that the landholders can make best use of the various parts of the land. Each weto is named and is controlled by a particular matrilineage. Members of that matrilineage have the rights to live there and to make copra on that land and to plant new vegetation. There are fourteen named weto on Majkin, each running across the island; they vary in width, some being very wide. The two weto at either end of the islet are not as productive as those in the middle of the islet because of limited access to fresh water. The core social group that shares a residence weto on Namu is two or more sisters and their descendants, together with their nuclear families. That group forms one household. They may have several sleeping houses, but they share a common cookhouse where two or three women 3 of the group cook for everyone. Households on Majkin vary in size from 12 to 75 people, even getting as large as 110 when special events, such as a church conference, are held on this islet. Chief decision maker for a weto , known throughout the Marshal Islands as the alab, is usually the oldest brother of the lineage. He may not be resident in the particular household he controls but is usually resident on the islet at another weto. He manages all affairs connected with the plot of land, including planting new crops and digging wells and erecting new structures. He thus is an important influence on the productivity of a weto. In 1968 when the whole islet was short of food and one weto group had none at all, a comment was made that their alab in times past had not planted well for their lineage, and thus he was to blame for their plight. The main starch food sources on Namu are breadfruit and pandanus, with some arrowroot. These are eaten with some fish or shellfish or coconut as an accompaniment (jalele). Rice is heavily used, being purchased from field-trip ships with money earned from the sale of copra. Some flour is also purchased to make a type of loaf, or dumplings. The pattern of food use has not changed drastically over the twenty five year period as discussed below, even though the population of the atoll has doubled. It has just become more dependent on rice to balance the supply when there is little or no breadfruit or pandanus. Breadfruit Breadfruit trees dominate the house sites. These are mainly the seedless variety (Artocarpus altilis) of the Bitaakdak, Bukdol/Bukarel varieties, though seeded varieties (Artocarpus mariennensis) such as Mejwaan are also planted. A house by house assessment of breadfruit trees on Majkin in July 1993 revealed an average of 6 trees per house, with one house having 27 trees and another only 4. Most of the breadfruit trees were 60 to 70 feet tall and had a base trunk circumference of between 8 and 12 feet. They grow around the residence site with at least one being fairly central to provide shade. A few smaller trees have been planted more recently by taking a cutting from the root of an old tree. These young plants are carefully nurtured until they can stand alone. People on Majkin cannot remember when particular trees were planted, but judging by the similarity in size of the largest trees, there must have been a concerted effort to propagate new trees in the 1930s to 1950s, but after that few new trees were planted until the late 1980s. As result food resources will be severely curtailed when those older trees pass the peak of their bearing lifespan. In 1993 most of them were bearing heavily. Breadfruit trees are so valuable as a food source for the whole residence group on one weto that one is cut down only out of extreme necessity. Indeed the Marshallese term for breadfruit, ma, is also the generic Marshallese name for a plant, indicating that breadfruit has prime status in their categorization of plants. In former times a tree might be cut down if a canoe was sought by the paramount chief for a particular reason. The green, globular fruit of the seedless variety is round like a boy's head, as Dampier (1616) described it, and weighs on average 3 to 5 Ibs. On Namu, a breadfruit tree will produce fruit three times a year in a good season, that is, one without drought or cyclones. The main season is May through August when the greatest number and largest fruit are produced. Two secondary 4 seasons of fewer and smaller fruit do occur in October/November and January/February. The ten or so different varieties mature at slightly different times, thereby lengthening the breadfruit season. The fruit of the seeded variety, Mejwaan, is very different from the seedless variety. It is of irregular shape and weighs about 2 or 3 pounds. One crop matures in late April and May, and may have a small second season. Both the pulp and the seeds are eaten. The flesh has a slightly more tangy flavour than that of the seedless breadfruit (NJP Namu fieldnotes 1968) (For a full discussion of breadfruit varieties throughout the Pacific, see Ragone 1987). Namu people cook the seedless variety in a number of ways. The most common way is to roast the whole fruit in the coals and then scrape off all the charcoal before serving (kwonjen). Breadfruit may also be boiled and coconut cream added at the end to make a dish called bwilitudek, or baked in the earth oven as a whole fruit to which coconut cream has been added in the centre(beljij). Another five ways of cooking breadfruit were recorded during the 1993 season. At the end of the season, particularly the main season, the ripe fruit are picked just before they are ready to fall in order to make them into fermented breadfruit paste (bwiro). The process involves all the families associated with a household, even those living elsewhere but strongly attached such as a brother or sister; even 8 or 9 year olds and those in their 70s help in the peeling stage. The fruit is first peeled, then quartered, and placed in sacks to be soaked in the lagoon for several hours. Then it is left for two nights resting in a tree to drain and begin the fermentation process. Once fermentation has commenced and the fruit has become mushy, four or five sacksful are tipped into one pit. These pits are made by excavating a hollow in the sand and lining it with old breadfruit leaves; each pit is then covered with old breadfruit leaves and weighted down with several coral slabs. These storage pits are usually placed near the cookhouse. The paste stays there fermenting for a month or more before it is ready to be processed for eating. Each household on Majkin was very active making bwiro in early August 1993. Some families had filled five or six pits already and expected to add still more. The people were rejoicing because the season had been so good. By October and November they will begin to use the fermented paste for their daily food supply, and also send some of the loaves baked from the paste away to their relatives in Kwajalein. On other atolls in the Marshal Islands, however, breadfruit is being allowed to fall and rot and is not being made into bwiro. Laura, at the western end of Majuro Atoll, is one such islet that has a multitude of breadfruit trees and yet the fruit are being wasted. There are two possible reasons; firstly the people prefer store food because it is quicker to prepare, and secondly money is more readily available as many households have one or two persons working for wages at the other end of the atoll. The Ministry of Resources and Development has therefore accepted external aid money to build a factory in Laura to make breadfruit chips. These will be made from the fruit at the height of the season, those people owning several trees selling the fruit to the factory. The chips will be marketed like potato chips in small packets as a snack food. Such projects have been successful in Western Samoa and other parts of the Pacific. The project will 5 thus utilize a resource that is being wasted at the moment, and also produce something locally as a Substitute for an imported product. The managers of the project aim to extend utilization of the plant for the same chipping process with other crops such as iarej and bananas; these snack foods will be sold locally and exported. Besides its fruit, the breadfruit tree has several other uses. Its leaves, both green and brown are used extensively for wrapping food to be placed in the earth oven, and to cover the earth oven before earth is piled on top. The leaves can be used as an instant plate, and to cover food left in a container. The sap of the breadfruit tree is a well known form of glue used in calking canoes and in handicrafts. The trunk was formerly the most favoured wood from which to make a dugout paddling canoe (korkor) or to make planks for the larger sailing canoe (tibnil). Few trees have been cut down recently, however, fishermen preferring to cannibalize old canoes to patch up one that is broken. Dead wood from breadfruit and other trees contributes to the fuel supply. The detritus including leaves forms valuable mulch on land that has very little humus. Pandanus The pandanus tree (bob) is much smaller than the breadfruit, standing only some ten or fifteen feet high at the most. Its distinctive feature is its prop roots which in an old tree may be three or four feet long. Two distinct varietal groups have been propagated over time to meet local needs on an atoll such as Namu, one for the production of fruit, and one that produces the best leaves for making handicrafts. The two uses mean that new varieties are selected on the basis of their appropriate qualities. Many varieties are named by local people, and their attributes clearly distinguished. In 1968 I recorded 24 different varieties growing on Majkin; in 1993 | was told that double that number now exist, though | did not record the names. Pandanus fruit are large and globular. They weigh some 20 to 35 lbs. Each fruit consists of some 50 or 60 drupes that are attached to a central stem. Each drupe has a hard generally green exterior and a fibrous ‘brush’ interior surrounded by an orange pulp when ripe. The fruit bearing qualities of the pandanus have been carefully selected for by atoll populations across the Pacific; for other Pacific societies the pandanus is considered rubbish food. The plant is grown by vegetative propagation using a slip from one of the prop roots. The fruits seldom contain seeds, and if they do, the resulting plant will not be fruit-bearing. The pandanus season on Namu begins in August or September and lasts through to December. The fruits are eaten when ripe by breaking off a drupe from the central stem and rubbing the fibres between the teeth. Other varieties may be cooked to soften the fibres. The process of eating pandanus resembles sucking on a shaving brush. Since the edible part is so fibrous, the eater ends up with many strings between the teeth. The paste is high in vitamin A and thus is a valuable addition to the diet. Pandanus is not considered a main source of food by the Marshallese, though it is eaten extensively in season. Rather it is used as a snack by both adults and children alike. Formerly the pulp was extracted from the fibres by pressing them against a v-shaped object made of shell, wood or (today) metal. This juice was then boiled until it thickened, and the paste set out on mats to dry in the sun for three or four days. This dried product was rolled and tied to form a 6 product known as mokwan; it was carried by sailors as it did not deteriorate in its leathery form. Such preservation is seldom carried out today as it is time consuming and there are more convenient imported foods. An alternative recipe was to add arrowroot starch to the boiled pandanus paste, plus some coconut cream to yield a food known as peru. This was considered a delicacy and so was only made on special occasions, or as a gift for the paramount chief. It required a lot of time to prepare. It did not keep, so was not a form of preservation. Today it is made only very occasionally. A second major use of the pandanus tree is in the manufacture of handicrafts (amimono). The green leaves are cut, dried and processed into strips which can be woven into sleeping mats, or smaller objects, or are boiled to produce a very fine white fibre for special basketry. Alternatively the dried leaves, once the spine is removed, may be processed into coarser mats, or into thatch. Handicrafts have become such a mainstay of the economy that considerable effort has gone into finding the right plant for the particular product desired. This trade has led to a diversification in the species of pandanus grown. Fishing Fish are the third major resource which is heavily utilized on Namu. Fish are considered a highly desirable complement to the starch portion of the daily meal, but on Majkin they are are a luxury. They are not easily caught, and with a large population to be fed there are never enough to satisfy everyone. In part this shortage is due to difficult access across the reef on the oceanside of the islet, and in part to the shortage of fish on the lagoon side. Whether the latter is due to over-fishing, or some ecological anomaly is not clear. Every household aims to have some fish for the Saturday evening meal. So those men who have access to some form of boat, whether one-man dug-out or a launch with an outboard motor, spend the day fishing in the hopes of providing a decent supply for everyone in the household to have a small portion on Saturday night, and hopefully some left over for Sunday. But they are not always successful, so that fish is available for only about half the weekends of the year (Pollock fieldnotes 1968.). The greatest amounts are caught by those who use the launch to go farther out into the lagoon, or even out the pass on the other side. But that requires gas, and gas is a scarce commodity. It can only be purchased from fieldtrip ships, and then only five or six drums at a time. So frequent trips across the lagoon are not possible and men tend to fish close to the lagoon shore. Even the good fishermen will tell you they can sit there for hours and catch only four or five fish. But when the launch goes out they will catch some 30 or 40 Ibs of fish. The people of Majkin would like to have a more regular supply of fish. A new fisheries facility (built by MIMRA) has just been completed in the centre of Majkin islet right on the lagoon shore. It consists of storage facilities, a fresh water tank, two large launches, and a tractor for launching them. This is one of three such facilities already built in the Marshalls with aid money to assist the people to become more self sufficient in fish. The aim i is to provide the facilities for catching fish for their own needs, and also to catch fish to sell to Ebeye, the urban concentration on Kwajalein. In August 1993 the project was awaiting the appointment of a director to begin operations. It will be interesting to see how successful this operation is, given the ongoing difficulties of catching fish on Majkin. In contrast, Namu Namu islet to the north of the atoll has a plentiful supply of fish as they can use both the ocean side as well as the lagoon side of their islet. On the ocean-side they catch flying fish (jojo) in season. Thus it is surprising that MIMRA (Marshall Islands Marine Resources Association) did not see fit to build the new fisheries project plant at Namu Namu. Time will tell. Continuities and Changes These basic resources have remained unchanged. What has changed is the demands upon these resources. The population of Majkin islet has doubled to about 440 people from 200 in 1968. People move constantly between the three islets of the atoll, and also out beyond the atoll, so it is difficult to give an exact figure at any one time. The household survey we conducted in late October 1992 yielded a total of 517 people, but a similar survey in August 1993 yielded a total of only 347. The reason for the large number in 1992 was that many people had travelled from Namu Namu and Mae-Leuen, and some from Ebeye to take part in a church conference on Majkin. In addition, the numbers in August 1993 were down by about 50 people who were away at another church conference in Majuro during July 1993. So the demands of the population on the resources do fluctuate quite considerably. Tree crops and fish are admirably suited to such irregular demands. Copra is still the main source of cash as it was in 1968. The price of copra as paid to the producer has fluctuated considerably over the years; it was so abysmally low in 1968 at 2 cents per pound that Namu people (along with other Marshallese) were seriously questioning whether it was worth the effort. But without copra money they could not buy rice, flour, tea and sugar, so it was better than nothing. In 1990 the Marshallese government agreed to support the price of copra at 15 cents per pound in an effort to draw people away from the urban areas where there was little likelihood of their finding jobs, and back to their home atolls where they could participate in a more subsistence oriented economy. Namu people living in Majuro in July 1993 agreed that life on their home atoll was better than living in Majuro "because you don't have to buy food there". For younger people that is not so much a concern as it is for older people with families. Another concern is that many of the copra trees on Majkin are old and approaching the end of their productivity. Unless they are replaced soon there will be a severe shortage of cash for families to buy the necessary foodstuffs to balance out the times when little or no local foods are available. Even the Copra Support scheme will be of little use in keeping people on their home atolls unless the coconut trees are producing enough nuts to be sold to bring in sufficient cash to feed the increasing population. Breadfruit trees produce enough fresh fruit to feed the population of Majkin through three or four months of the year, if eaten at only one meal a day, and supplemented by rice. The 8 fermented paste, bwiro, extends the subsistence base by approximately one month to six weeks, if eaten only once a day, and if only moderate amounts are given away or sold on Ebeye. As the number of mouths to be fed increases so these time frames are correspondingly reduced. If breadfruit is severely hit by cyclone, drought or disease, then this subsistence base is hard pressed. During 1993 the trees appeared to provide a strong subsistence base. But in another year the picture may be less rosy. Coconuts are thus crucial as the intermediary between subsistence and cash food sources. They must be renewed if the population is to continue to maintain at least a measure of subsistence. New foods have been added to the inventory since 1968. Pumpkins now are grown successfully by almost every household and used as an additive to rice, thereby enhancing its nutritive value. Bananas too have been planted on the borderline between the eolap and iar areas of each weto. They are eaten as a nutritious snack, in their ripe form. The green banana, widely used elsewhere in the Pacific is not a familiar source of starch to Marshallese and thus may not be acceptable, whereas the yellow bananas are. They bring in much desired cash when sold by the stem on Ebeye. The planting of taro (iarej) has been encouraged by the Ministry of Resources and Development in Majuro as an additional starch source. But even though it was growing well on several weto on Namu, it was not included in the daily diet during 1993. This was partly because the taste was not wholly acceptable, and partly because it was used formerly as a feast food only, and not for everyday use. Another drawback was that it takes time, and considerable fuel to cook. So plants regenerate but are seldom used. However its use may increase with time. Fuelwood is an increasing problem. At times throughout 1968 fuel for the cooking fires was in short supply. The people rely mainly on coconut husks for cooking the large pots of rice, or for roasting the kwonjen form of breadfruit. Such fires are lit at least once a day. As the coconut trees get older and the number of nuts produced diminishes, so too will the amount of fuel for cooking fires diminish. Renewal of the coconut trees, plus some attempts to identify suitable fuelwood trees are two urgent aspects of the subsistence support plan. The alternative cooking fuel is a kerosene stove. Eleven of the fourteen households had these in 1993, whereas only four households had them in 1968. This indicates a greater reliance on kerosene, and thus on cash, for small cooking jobs such as boiling water in a kettle or frying pancakes. Otherwise the open fire, or the earth oven are used. The kerosene stove is a measure of a modern lifestyle, as relatives living in Majuro or Ebeye tend to cook on these. Communication systems have also proliferated. Today every household has one or more transistor radio, whereas in 1968 only one or two were operating at any one time on the whole islet, due to shortage of batteries, and/or the radio being broken. As the radio was the only source of information about field trip ships, it had a major impact on the economy. People relied on field trip ship itineraries to know when one was coming to Namu so they could make copra. If they made it too long before the ship arrived, the copra dried out too much and so they lost money. If they did not have enough time to make copra before the ship arrived they also lost money. In addition to the transistor radio, four c.b. aerials are conspicuous additions to the household 9 sites. These are used mainly to talk to friends and relatives in Ebeye and to arrange visits of people and goods. One of them belongs to Air Marshalls and is used for receiving and sending information about air traffic. A weekly air service linking Majkin, Namu to Majuro and Kwajalein, the two urban centres of the Marshalls, is another major innovation in the communication system. Land on the ocean side of three weto in the northern part of the islet has been cleared to create a coral runway as a landing strip. The Air Marshalls 18 seater Dornier lands twice in the same day, once on its way from Majuro to Namu and then to Ebeye, and once on its return from Ebeye to Majuro via Namu. Passengers and freight travel regularly, the most heavy traffic being that between Namu and Ebeye. An additional airstrip exists on Mae-Leuen at the southernmost tip of Namu Atoll. Majkin people are using the air service as a means of supplying the market for island produce in the urban centre of Ebeye on Kwajalein Atoll. Boxes of kwonjen, whole pandanus fruits, and boxes of bwiro, and fish if available, were sent on the plane to Ebeye, either to relatives, or to be sold. This use of the plane as a means of marketing subsistence produce provides small but welcome returns to families who have few other alternative sources of cash. Conclusions The plant and fish resources of Majkin islet, Namu atoll have continued to be heavily used over the past twenty five years. The diversity of species, particularly of pandanus, has been increased to meet specific needs of the handicraft industry, and attempts are being made to increase the amount of local fish available. Local resources thus directly provide about 40 per cent of the total local needs today. Copra, and the cash earned from it indirectly provide another 40 per cent, the cash being spent to buy the same four basic items as in 1968, rice, flour, tea and sugar. Thus local resources are under greater pressure today as demands increase. And there are high risks of failure. The balance of the needs are met by other means, such as support by relatives working in urban centres, wage labour jobs (though these are still extremely limited) and community support. The proportion of foodstuffs used locally has remained about the same as that in 1968. But the possibilities for selling local produce are entirely new, due to new communication systems, such as c.b. and the airstrip. The pressure on locally produced goods is thus much greater. Two major differences influence this pattern. Firstly the population of the islet has doubled. It still maintains its movement patterns from islet to islet and beyond the atoll, so numbers are not constant; however there are at least twice as many mouths to be fed as in 1968. Secondly the increased demand for cash has impacted on species other than the coconut for copra. Today breadfruit, both uncooked and cooked, and its cooked fermented paste as well as pandanus fruits are highly marketable items among the urban populations. Namu people tend to sell more to Ebeye because Kwajalein atoll is much closer and communications are better. Some of the goods are sent by air freight, while other goods are sent in a launch with outboard motor. The cash realized enables them to buy more rice. Copra remains as the main source of cash income, but as the trees decline in productivity, and alternative sources of cash become available, its overall contribution to the economy is 10 declining. There is an urgent need to replace the vast number of older coconut trees, if local subsistence levels are to be maintained. The population of Majkin "manages" to get by with their current lifestyle, but would like to have more. Unless rapid and severe measures of population control are introduced, that lifestyle will not be maintained as there are more mouths to be fed and educated. More young breadfruit trees also need to be planted as they are slow growing and will be needed in the future. The plant resources of Majkin are currently at the interface between subsistence and a cash economy. More varieties of the same tree crops could be planted, and the coconut trees should be renewed. These actions would enable the current pattern of about 35 per cent of the food needs to be met directly from local resources. The other 65 per cent will continue to come from cash, earned partly from copra, and partly from handicrafts and food sales. By extending plantings of pandanus and breadfruit further into the middle zone of each weto yields could be increased. By renewing coconut trees more cash would be available. But so would the risks of both natural and market disasters. So whereas local produce in the 1960s was used only for local needs, today that produce is committed both to local needs and to sales outside the atoll. And with such a rapid increase in the number of mouths to feed, the atoll is running out of options based on local resources. The people of Majkin are becoming more and more dependent on outside sources of food, but with a diminishing supply of cash as copra returns decrease. Highly polished rice is not as nutritious as breadfruit and pandanus, and it costs money, though it is very popular with the Majkin people. The people of Majkin face a bleak future. The Marshall Islands government wants to encourage people to stay on their home islands, and thus reduce the urban pressure. But the resources on an outer island such as Namu cannot support any further population increase. The limit has been reached between supply and demand for food. Urgent attention is needed by the central government to replace coconut trees, increase the increase the number of breadfruit trees, and to introduce an acceptable means of restricting population size. Otherwise outer island living will no longer be the idyllic option. 11 BIBLIOGRAPHY DAMPIER, William, 1697.— A new voyage around the world. Reprint. London 1937. POLLOCK, Nancy J., 1970. — Breadfruit and Breadwinning on Namu, a Marshallese atoll. Ph. D thesis, Anthropology. Univ. of Hawaii. = 1992. — These Roots Remain. Food Habits in islands of the Central and Eastern Pacific since Western contact. Hawaii: The Institute for Polynesian Studies. RAGONE, Diane, 1988. — Breadfruit varieties in the Pacific atolls. UNDP Project Series. N.Y.: United Nations Development Program. yt _techiaing ‘Theme jammy tine en boptine dna: vert nuchber al oldar cxcidnat tredis, if ah 7 * alate ir hen * af ‘hy A sey Fe ww hag ey ee wee Aw a! ig i 4 peraulat Fi mS ai Ki alee 'y ATM E tpgat ky, wither ; tin Ti Picker nt ry rope amd sewer® names od ompily 7. | pen iti wet} ns: ther maaintnined us theme ales Krunye cnet oy |, ah Y Ce i ee ae AMIS es bred rue faeetesiys Hine: ricigel ay tus i shan bexd an they dau steew gra DA. VORT. soot eth te. ae Oe ee we thie nna cit eet coer ) Meroe By ‘| ese Pipi panna ot Gish slay Peso id. son reset We sali sin PN OPN RAREST tesa drotoqocahadieroodt Ga% sioter «en td | ws chet AaweRy Thon ooal.esourpes, The cther 5 ge waet aid! contin te va ~ weit Oo Wem Le? Th scineea neni AMM aie Beet) Ha SOPs, Bye flea nascar Witt inept pisenpiacignntevwerciennete merentho | vchteg Hii STU S gew! ; MPU: URINE Was © ie ae vh vinta erNRRGeTNSER SS PL cl Sale tt * nhdln ation inthe vanssiant surttbierrs cals ees saris ih ‘Woe sh ei i VERT DAE eee ml sai e whol 1s TanMay one of aphid based On local ram tenth ia rae and. more deperilie| any guitsice ax MICOS off ADAP ES DECI die renaee. bing! iy polished roe is eka aed fier. atte ard CoRtemomey, Thanet ie vey popdian wink thn momple at Majkic Geo wi ipa sear Thre Macshad) Istoade governmned We tO Cay bt Te SRR, aa Sea recheawie. shat urteac. pte PRE Lodo Se aS eons A nappa “ey feorther Penyidhananh am Rittiad Der een Shy eal eich lon ANE, ingont attention im DN ae Tie ita eget, § % eA) ge ree bir Ht tag) PTs sree the: martther ot tr Picton iin gomeptable cca OE cam h in gop tying ele. iconiebaes ene fi ; ae Ta en) ae Cy CRY eal \ Fy s Fj I i 1 vue angel oe * a ‘ i-gupet ip ‘tal ui ey n j oe” eae ‘iv - ts i) 74 r aero i : vio i" UL eal a ~ et * Ler ' . q i os | Yo | ae, | ou F ee ij ws) ‘7 ae om ay » Joe a ATOLL RESEARCH BULLETIN NO. 442 CRUSTACEA DECAPODA OF FRENCH POLYNESIA (ASTACIDEA, PALINURIDEA, ANOMURA, BRACHYURA) BY JOSEPH POUPIN ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. APRIL 1996 STIVERS (pi a a ELT Re een eee LINGER © 10) UW) STN @ Niece oe ees cacao tees eae e sea ee ee cree eR ee a cae So Ce aan cc sass Sat MeN svsuaceeaetaswosecoiuousies ELS AN@ RR CATE ie a oie eae Oe attri ace denne sea ese cial sce cen eater tO Bits aside be RM eo le Ja des scadabe GONWVE NGTONS eee ee ee ec ED EMT oo abd sctcedesssenceess LETS IK OP ANEIE}S PE CIES eee ettiaaretees neers oeteterereccereccctera ei ccoscnces soueeeee treo eases peels cs spiuecctsgoustvecuesssecvesss INERAG ORDER PASTA GIDEA tres csee carer eoccuscetal strstr scien vasursaaeaa tesrvesecansusasnesesauhs seveeevsiieseestavenss tauscéesvesses IFA MIEYPENOPEOMETOPIDAE jazcactevecectass ccc ctlessccoesvaas sabes suvencesncecss steoses cnan cesses calvusutisacesaaessisteetedsecd INERAZ ORDERS PD ATINURID EA cote gar tessa ce tase ase nc cence viak sestucsebeca sabe sse anes saatatescectesvassesadee cues svaessscsesadevedoxeevsszcs IBAMIIEY PATINURIDAE ¢rccs-teccct tes Sessa vases Se sesscssessatesesd oscdedcehasgescogadanssnededuecsess ssusedatucossesscisssees JOVI AUTEN 2 SSN INIAOSTID YN Bhat ie RE ence One eoca COUR eee eica occa eco Eeeecaceerrere 10 IRAMIEYGES GYIDIEARIIDA BE pevccpateevatesnaersccseectescsesetsdssecrcccdas sae scavedsoccneeestaisatanesusestieeesustsessdessavedevevis 10 INFRACORDERSANOMU RiAvcssccctecsecrosetectacecacestecse Sis scivendavotuceee iuscsewet secs sueesocnaces ao naan stoaniva wanes vondeene ctee ee 11 RAMEY C OENOBIMIDVAR rere eee teeta eens saeco vc sea tecr esa os ovat dua cesedcsseeaue et cannes canta alacusscadenznansessetek 11 IPA NITEY@e DIOGENTD AE tarerreretaetccenscce cen coc cnacesarcorc sce cocecer soc ee ns aactncues ey ocuraue sesaserecenuneieweibesss asesue 13 RAMTEYS AGURED AE Reet ee eee seen eee eee erage eave rstasesseceeven lussvousian faneseeis 18 RAIMTTAYS GATEATIEET A Eee ease rere eae eee ee ee eee ee raccedecerecsdetsasesnsssedtt 19 AMIS Ye PORCEDIUANID AB itera sueecd acetals sanea sie Sis shaun ots sence tnhs RLULUEAA SAAS BCs 30 Ee ND ces 20 AIMIIEYAPATEB UNEIDAE bercrtrteesnedasccortestecee ls rat cecatpucesstueeasrasaeesubtereesierpvetcwseccusecdusuuvee sour areculesieee 22 RAIMI VeRIIPPHIDAE etc staecosstass cree ato gacsouet ins casducuoues busewvetesbus ones Saseaeh suvsbhs oot sen sus da vcvaseveteuusdetcesdevessce 23 INERAG ORDE RUB RACEIYURA vrcrsttercee tere tace ra cerse cee sue aecsers acs sce saceavsensuteseceses sunuesues duoeesusascoeesetaseinsscetuveest 23 LUN TOES? IDI KON TID DYN Bcc cenecer tre ser ror rec Sc LaCe cere cor DEE EEC EER Soe eeESCEE SCC PeCE eeREeee eee eee ec 23 ANY VINO MENIDVAE yestccreascteccccenssstrncsscoteusssuk cocsoecsnsssaecc seat ace santos ror tuctnate nevaconecvoussatessios suvsevs 24 ES ANMITIB Yas VA NINIDDAE tecarerscrscessco tacirsccecocecoas'asescaooesada Sovse tates couee See te secre nantes suey sa cetaedivadeus eetiaeeate 24 ANALG CAAA PPI) AE omen tie ee acne Menace este sauvce savteuues fewstiv ween atalssel anes avsesteessivecevecsusstontiee 25 JOLIE ILTSLU COS) TIDY Eh scp eacsoceoeae Recon sheccecirchocaasar machaec ec crea He CGC CREPE Ree nce GREE Ree eer eRe reer ee 26 AIMEE DIVE AE ee eee seecenee tant ae acai ROOM BU RESO oy sbacedaaacavecvandsavses sovesswceuones 26 RAMEY) PA RTHE NOPIDVAB recs cpeeen secs eceeency seecosss ceca lent culcunc tense teenteecads scessectucessesiceterseessueviceseeessese 28 VAIMIIYAE, UMEDONIDDAE wresese ets cess ccscersscscr oesee les ves cbseatetoe ses Se sarees peo watotate 380k onsen cubcencvadenddaceveswssosue 29 AMIE YAR ORTUNIDDVAI prmeresseecae na cuck seve sncseccoccstacss coast usceveseeoe cc oeeaeee Scene pasate eS t a dueea aes eones boos tmuraeee 29 SUBEAMIIBYS CATIOPMRINAE serccsssccesecseascotcess canciones stars sca cre sosesaseceee canes este natal exe dant dc weeeestucsevesescoteososs 29 IS WBEAIMMIEYA | GABETVIRINAE Spccscscossia- sess cctae eoeetaceecaa anc teeicve scenes seat sbgecsnastnrGesnsasesuat sav cstewacat Wastetows 29 SUWBRA MIE VG PORMUNINAB seteresrcrsticse ters iuces Secesssztvesssesoacaeswesseeea hoarse age occas etaveshideieegussuantisasetee tien BO SWBRAMIEY) PODOPHTHALMINAE \cs-asascacssseseecsoecascosssCsssds octave csegeasesosscoscasesaseocossaecscasassarasacaseansesueabe 37 FAMILY XANTHIDAE SUBFAMILY SUBFAMILY SUBFAMILY SUBFAMILY SUBFAMILY SUBFAMILY SUBFAMILY SUBFAMILY SUBFAMILY SUBFAMILY S WERE AMIIBY@IE TIISINA Eleteer ec teee sectc eee oee renee tte rl ect rena sacecuel ahovcunscsbaasusadeasacoute odesveseenstbee Seats 51 CONTENTS SUBBAMIE Yel GHIEORODIINAE trnrccerssseantactorsscuceres-uvacvensserenvosssecercesstsseeceesecuestensestcecrecdecersecssbeacrncreeeceecs 53 il IPAMILY: RAPE ZIMDAW 22858-0055 ss chores soteeselobeanss eased ssnhe eel Se swan cuneenn ast commu tence taneas Tory an Senet ea 57 FAMED Y: (PIEWMNEDAB: 2 .coc.iccoctvcdlelcod vous caatensecestacoruuousees Goauueaedcauacaute antec ueetinsusuracuaasandevarssueteateraete 61 PAMMEY CARPIUIIDAR sscpcassvasssivass eSeukvecanaseacnsuasvezsaceassceuh ues cces cea aieiatar draunesycunaconuatonne i aeranee eee 62 ISITE: | MIENIPPIDDA Coca ei -ciessccccccsvatewecsseeseneass couees stcteeesis asmeeen cs Resa n oe Mae ceasa ae satan cae a 63 IPAMUEY "GEGARGINIDAR ie tetrtorescccncorecaccereocsea nt oriatce cuvsons sn ecececuaitis ere niean arc cenabea santo none eee 65 IPATTEY eG RAPSTD AE tet rudters seein Manes leverantnntanse ton tales Bee eye eviese i oNwn: WOMEN NTE Ne. teat cent a t eeee 66 SUBFAMILY. |GRAPSINAR, sccatcssceshavestousaae esceioaunareatans desea hestencas A eeeeaes ee ee ee 66 SUBFAMILY. ‘ VARUNINAE::).fsscssitsccctsstescstactesssons ses sovassscusessieaes ses a Ee TO ea 69 SUBFAMILY. ‘SESARMINAE 3 )o5 cco hbo tela ie cal ae ee ae eves aces ee 69 SUBRAMIEY” PICA GUSTINAE is. csesscscdsectecessiocasesectensc sua osreteosscoasccasey Ciasae cep eaeee Ivan ean Ce 71 PAMILY IPINNOTHERIDAE csv stvcisssvsszcactecssaeeecoovssaovesisied van casaucea ares sussaevced tae ceueetneccannsees See eaee re eee Te FAMILY: OC YPODIDAE © tecccce cate scauterscacac us sauna eee cae eee cea sean ae as Se se gee e nea ae oe 2 PAMILY. CRYPTOCHIRIDAE) hiscccecatecaos ces scar ssabecuederennoe cave suse sou cee cue Season ae cea cane thee etek see ee eee 74 PAMIEY LY MENOSOMATIDAE si.s-ccas;csecccesssceusssaurervszeesunacssvsnsdensasese seus soessaduavecessevaeesere corenenemteneee 75 FAMILY INCERTAE/SEDIS :ccss.csstcadacevscscectaceveseucruna sis oveuccanasovasevaesoucassvatere stents tasceseasy sete eee 75 Leaves sucteresane losers seo esneeeu ee OSC ein Sere TRE ESOS CRC HAASE GUA eS ES Sn so eRe 76 ICME RAT WIRE! CITED). ccocescclacsuesscessss cuss coors ar cur eas Cae e a eG ee Bee See ee cae eNO Sau ee Rac ee gee 81 ACKNOWLEDGEMENT | cccccctusssccoseccuuicsvocsccoudsstsescssecccnssetecads tsua se usa eee eo ae eee eee eee 95 sas vliesia vai’conuvjiivenosuge douanp oad bGeded exeees sugen ee lebccsu ciate ee aceon MAL: RSI RII CsI Bice ane 95 This work is dedicated to MONIQUE DALLE and JOSETTE SEMBLAT Librarians at the Service Mixte de Surveillance Radiologique et Biologique and Laboratoire de Zoologie des Arthropodes, respectively. 155° W 150° 145° Hatitaa oe eas 135¢ Eiao ~ * Clark Bank H i an i . Ua Huka Nuku Hiva Fatu Huku Ua Baus ~ Hiva Oa a : f MARQUESAS Tahuata’ * Motane 10°S s° Thomasset Fatu Hiva TUAMOTU sen Manihi a Tepoto Nom a Mataiva Zousien ae a “Napuka , Pukapuka 15° ‘ PBR. ASE Takapoto , Tikei 15° i Tigian (Ss a Apataki Bellingshausen Aratika Taiaro Takume Fangatau ° T Makateasaukurats oat o Katehi , Fakahina Scill Hee » Bora Bora Niau - Roce @ Raraka Taenga Pc of Maupitie @ F ae ergs. ene Rekareka ° Tahaa 4 “Huahine, Tetiaroa aaite inf TT tak Mopelia Rata Mc Tahanea™ 0 Tepao \ Marutea North alatakoto COTES» ae ahiti rete AnaaS sete ees are Tauere ue SOCIETY Maio e Reitoru 2 Amanu Pukarua Marokatp Akiaki & Ravahere | Hao | eNahitahi dso Nengonengo _ = Paraoa _Nukutavake Manuhangi Pyaar Pinaki : Hereheretue » Ahunui a 20 20° or as ° . Nukutepipi Tureia uanurunga Vanavana Tenararo Tenarunga : Vahanga » e . sien Tematangi ° Monues Matureivavao Marutea South Rurutu ° Maria Minerve Bank _ ie Fangataufa GAMBIER s Rimatara Tubuai : 4 Temoe ee een ay ane ee wr Few Raevavae.” aioe ty Ft) ua) fe) nn 0 AUSTRAL pe 25 Thiers Bank GAMBIER Mangareva Totegegie Neilson Bank ., Rapa S : a Marotiri - a, 4. . Mekiro =f Macdonald Bank <> uae yea tem) Ne Teiku riteee 155° W 150° 145° ae Makaroa eManui Kamaka FRENCH POLYNESIA CRUSTACEA DECAPODA OF FRENCH POLYNESIA (ASTACIDEA, PALINURIDEA, ANOMURA, BRACHYURA) BY JOSEPH POUPIN SUMMARY From a bibliographic compilation and, to a lesser extent, from material collected in the field, 401 littoral and sublittoral decapods (Palinura, Anomura, Brachyura), are reported from French Polynesia. The Brachyura prevail, with 313 species, mainly Xanthidae (123 species), Portunidae (54 species), and Grapsidae (35 species). The Anomura are represented by 74 species, and the Palinura by only 14 species. The list of the deep species, ie living in depths of 100m or more, is updated. Ninety-two species are listed, making a total of 493 Polynesian species. Amongst the material recently collected, 16 species are recorded for the first time in the area: Calcinus guamensis, Calcinus imperialis, Dardanus australis, Dardanus brachyops, Albunea speciosa, Parthenope contrarius, Portunus macrophthalmus, Portunus orbitosinus, Thalamita danae, Thalamita macropus, Thalamita mitsiensis, Thalamita philippinensis, Quadrella maculosa, Planes cyaneus, Percnon guinotae, and Macrophthalmus serenei. Moreover, after the examination of the type material, Ruppelia granulosa A. Milne Edwards, 1867, originally describe from the Marquesas, is here proposed as a junior synonym of Lydia annulipes (H. Milne Edwards, 1834). ; Only 8 species, related to well defined species, are known solely from French Polynesia: Parribacus holthuisi, Micropagurus polynesiensis, Nucia rosea, Nursia mimetica, Acanthophrys cristimanus, Lissocarcinus elegans, Ozius tricarinatus, and Macrophthalmus consobrinus. For some of them, however, it is probable that their distributions extend at least to western Polynesia. The French Polynesian fauna is typically Indo-West Pacific in its composition, with few endemic forms, and a low diversity compared to the Indo-Malaysian area. It includes, however, many more species than the Hawaiian fauna, possibly because the Polynesian islands are less isolated than the Hawaiian islands. The Society, Tuamotu, and Gambier archipelagos have been well investigated, with numerous expeditions organised in these areas. In contrast, the Austral and Marquesas Islands, still remain poorly known. The French Polynesian fauna is more or less homogenous, with few regionally distinctive features. The single obvious exception is for the isolated southernmost islands, Rapa and Marotiri, subjected to a subtropical climate. In these islands, species that are very common elsewhere, are missing (Coenobita Service Mixte de Surveillance Radiologique et Biologique, SMSRB, B.P. 208, 91311 Montlhéry Cedex, France, and Muséum national d'Histoire naturelle, Laboratoire de Zoologie des Arthropodes, 61 rue Buffon, 75005 Paris. Manuscript received 8 December 1995; revised 28 March 1996 perlatus, Birgus latro, Cardisoma carnifex), and, on the contrary, at least one common species is still unknown in the northern part of French Polynesia (Panulirus pascuensis). INTRODUCTION What are the decapod crustacea known from French Polynesia? The answer to this, apparently simple question, would be very helpful for determinating the species collected during ecological studies. Moreover, from a biogeographical point of view, a check list of the species reaching this area, at the eastern limit of the Indo-West Pacific province, would be very interesting. The aim of this work therefore, is to answer this question by drawing up, mainly from a compilation of systematic and ecological studies, a list of the French Polynesian crustacea, the scope of the subject being restricted to littoral and sublittoral, Palinura (Astacidea and Palinuridae), Anomura, and Brachyura. The check list given here has been mostly compiled from bibliographical records. In a first step, the most important works dealing with the French Polynesian crustacea have been consulted. They are the works by DANA (1852b, 1855), HELLER (1865), NOBILI (1907), RATHBUN (1907), BOONE (1934, 1935), HOLTHUIS (1953), FOREST & GUINOT (1961), and more recently, those by ODINETZ (1983), MONTEFORTE (1984), GUINOT (1985), MARQUET (1988), and PEYROT-CLAUSADE (1989). In a second step, the names of the species have been updated, for changes in the generic classification, or species reduced to synonymy, by looking through more general works, like the ones by HOLTHUIS (1991), SAKAI (1976), or SERENE (1984). This research has been completed by additional consultation of two bibliographical journals, the Zoological Records and the Current Contents, and by randomly looking through reprints available at the Muséum national d'Histoire naturelle, Paris. This last step was sometimes very fruitful, with additonal species mentioned in the area, often very discreetly. For the most important families, the main works consulted during this research are the following: The Palinuridae have been found in the recent catalogue of the Marine lobsters of the world by HOLTHUIS (1991), and in his revision of the Scyllaridae (HOLTHUIS, 1985). The pagurids (Coenobitidae, Diogenidae, Paguridae) have been first searched through the work by NOBILI (1907) and the studies by FOREST, published between 1951 and 1956. Additional information has been found in the work of LEWINSOHN (1969), the report of RAHAYU (1988), and revisions of the genera, Aniculus (FOREST, 1984), Calcinus (MORGAN, 1991), Catapaguroides (DE SAINT LAURENT, 1968, 1970), Clibanarius (RAHAYU & FOREST, 1992), Pagurixus (MCLAUGHLIN & HAIG, 1984), and Trizopagurus (FOREST, 1995). Except for NOBILI's (1907) work and, for a single species, BOONE's (1935) work, the few Galatheidae known from French Polynesia come from the ecological works by PEYROT-CLAUSADE (1977a, b, 1989), KROPP & BIRKELAND (1981), and ODINETZ (1983). Almost all the Porcellanidae have been found in the works published by HAIG, between 1964 and 1992, HAIG & KROPP (1987), and KROPP (1983, 1986). For the Brachyura, the beginning of the research has been greatly facilitated by the important studies of FOREST & GUINOT (1961), MONTEFORTE (1984), and GUINOT (1985). More information has been found in: MCLAY (1991, 1993), for the Dromiidae; GALIL & CLARK (1994), for the Calappidae of the genus Matuta; GRIFFIN & TRANTER (1986), for the Majidae; STEPHENSON (1972, 1976), STEPHENSON & REES (1961, 1967), and MOOSA (1979), for the Portunidae; ODINETZ (1983, 1984a), and the works by GALIL, and co-authors, published between 1985 and 1990, for the Trapeziidae; SERENE (1984), for the Xanthidae, and CLARK & GALIL (1993) for the Pilodius xanthids; CROSNIER (1984), for the Carpiliidae and Menippidae; TURKAY (1973, 1974), for the Gecarcinidae; and SAKAI & TURKAY (1976), CRANE (1975), and BARNES (1977), for the Ocypodidae of the genera, Ocypode, Uca, and Macrophthalmus, respectively. This bibliographical compilation has been completed, in a much more limited way, by the study of some specimens collected on the field, during the last few years. From the addition of this material, 16 species are recorded for the first time in French Polynesia. The deep-water crustacea, ie collected from 100m and beyond, have already been listed in POUPIN (1996), with full references on origins of the collections, and depth ranges. A simple list is produced here, updated by inclusion of species described after the first compilation, or recently collected (cf. Appendices). HISTORICAL OLD VOYAGES: 1820-1900 At that time, the crustacea were collected during the exploring expeditions made around the world by large sailing vessels. DUPERREY, on board the Coquille (1822-1825), is one of the first to bring back some species from Tahiti and Bora Bora. They were studied by GUERIN-MENEVILLE (1829, 1838) who dedicated to DUPERREY a small ocypodid crab from Bora Bora, Gelasimus Duperreyi (now Uca tetragonon). About 10 years later (1837-1840), DUMONT D'URVILLE, chief officier of DUPERREY on the Coquille, sailed again in French Polynesia, commanding the Astrolabe and the Zélée. His vessels visited the Gambier, Marquesas (Nuku Hiva), and Society Islands. JACQUINOT (1852), naturalist, commanding the Zélée, mentioned a dozen of species from the area, and described some from the Gambier Islands, including the small Ocypode pallidula, common on the white sandy beaches of Aukena island. At the same time, the Americans, worried about participating, like the Europeans, in the discovery of remote marine areas, launched their first round the world campaign, the great U.S. Exploring Expedition (1838-1842). The squadron of 6 vessels, commanded by WILKES, left Norfolk in August, 1838. At least four vessels cruised in French Polynesia: the Flying Fish, Peacock, Porpoise, and the Vincennes. A great part of the collections from the Tuamotu Islands was lost during the wreck of the Peacock, on the banks of the Columbia river, however, the Polynesian material, about 10 Anomura and 50 Brachyura, studied by DANA (1851, 1852a-b, 1855), represents the most important collection from that area. DANA describes several new species, from Tahiti (Phymodius monticulosus, Trapezia areolata), and the Tuamotu Islands (Globopilumnus globosus, Liomera tristis, Plagusia speciosa, Thalamita integra, Trapezia bella). Between 1857 and 1859, the Austrian frigate, Novara, put in at Tahiti, during her sea voyage around the world. HELLER (1862, 1865) studied the Crustacea of this campaign. He recorded 54 species from Tahiti, and described several, for example the colourful Calcinus nitidus, and the small gecarcinid, Epigrapsus politus. Limited collections were also made at Tahiti by the famous British H.M.S. Challenger (1873-1876). They are discreetly mentioned in the works of HENDERSON (1888), for the Anomura, MIERS (1886), for the Brachyura, and BANERJEE (1960), for the grapsid crabs. This era ends with the voyages of the American ship Albatross (1899-1900, and 1900-1905), and new collections in the Society, Tuamotu, Gambier, and Marquesas Islands. RATHBUN (1907) studied the Brachyura collected by this vessel. She recorded 85 Polynesian species and described, for example, Pachygrapsus fakaravensis, a grapsid very common in the Tuamotu Islands, named after the large atoll of Fakarava. French frigate La Coquille at anchor in Matavai bay, Tahiti (1823) (Drawing by Jules-Louis LEJEUNE. Courtesy of HORIZON Magazine) BEGINNING OF THE 20TH CENTURY: 1900-1967 The voyages around the world have ended and the collections are now made by people living in French Polynesia. The most striking in that respect is certainly SEURAT, the head of a small Zoological laboratory once established at Rikitea, Gambier Islands. Between 1902 and 1905 he gathered an important collection from the Gambier Islands, and also from the Tuamotu Islands, at Hao and Marutea South. This material was studied by NOBILI (1906, 1907) with more than 130 Polynesian species, belonging to the groups here concerned, and with the description of a score of new species, for example Thalamita gatavakensis, or Thalamita seurati. FOREST (1951), for Calcinus seurati and Calcinus spicatus spp. nov., STEPHENSON & REES (1961), for Portunus guinotae sp. nov., and FOREST & GUINOT (1961), in their study on the Polynesian Brachyura, have mentioned again the material collected by SEURAT. Gilbert RANSON, of the Malacology department, Muséum national d'Histoire naturelle, Paris, also collected many specimens. In 1952, during a stay of several months, especially on the atoll of Hikueru, he gathered numerous scyllarids, pagurids, and crabs. The first two groups have been studied by FOREST (1953, 1954), with description of 5 new species, including Parribacus holthuisi and Clibanarius ransoni. The third is studied by FOREST & GUINOT (1961), who, in grouping RANSON and SEURAT material, and some smaller collections, such as the one made by CHABOUIS, a teacher at the Paul Gauguin school, Papeete, have registered about 100 crabs, 21 as new records, with some new species, such as Pilumnus ransoni. The same year, MORRISSON, as a part of the Pacific Sciences Board's Coral Atoll Program, sampled the most common Crustacea of Raroia atoll, and, to a less extent, of Pukapuka, Takume, and Tahiti. From that material HOLTHUIS (1953) produced a list of 70 species, and mentioned for the first time Hippa ovalis, from Tahiti. During these years, a few expeditions, even if they no longer have the nature of great campaigns around the world, still visited the Polynesian Islands. For example, SENDLER (1923) recorded about 30 species from Makatea, Rimatara, and Tahiti, from the collections made during the Hanseatischen Siidsee- Expedition. Some of them, like Coenobita cavipes or the gecarcinid Discoplax longipes, have never been collected since. In 1931, the yacht Alva explored the Marquesas (Nuku Hiva) and the Society Islands (Bora Bora, Raiatea, Tahiti). BOONE (1934, 1935) mentioned about 40 species collected during this cruise, and described two crabs, Actaeomorpha alvae and Lissocarcinus elegans. Two important expeditions, at an interval of 10 years, mark the end of this period. In 1957, the Americans organised the Smithsonian Bredin Expedition, which visited the Society and the Tuamotu Islands. The portunids were studied by STEPHENSON & REES (1967) and STEPHENSON (1976), with about 30 species, including some new records like Portunus iranjae, Thalamita corrugata, or Thalamita quadrilobata. GALIL (1985) and, more recently, FOREST (1995), in their works on the genera Tetraloides and Trizopagurus, respectively, also studied the material of this expedition. In 1967, the boat Pele, during the Marquesas Expedition, visited the Marquesas, Tuamotu, Society, Gambier Islands, and, in the neighbourhood, the small island of Pitcairn (HARALD, 1967). The crustacea of this campaign, deposited in Washington and Perth Museums, have been studied by STEPHENSON (1976), with some fifteen portunids, and appear, more discreetly, in the studies by SERENE (1972), for Palapedia marquesas sp. nov., GALIL & LEWINSOHN (1985), for Trapezia tigrina, or HOLTHUIS (1985), for Parribacus holthuisi. MODERN PERIOD. In 1966, with the installation of the Centre d’Expérimentation du Pacifique, several scientific investigations were made, mainly on the atoll of Moruroa. Crustacea collected during these investigations are mentioned in some systematic works, such as DE SAINT LAURENT (1967), for Catapaguroides fragilis, or GUINOT (1979) for Lophozozymus superbus, and in ecological studies, for example in CHEVALIER et al. (1968), SALVAT & RENAUD-MORAND (1969), and LABOUTE & RICHER DE FORGES (1986). These latter, during the expedition of the old minesweepper Paimpolaise, in the south of French Polynesia (MacDonald bank), have made the first Polynesian record of Panulirus pascuensis, originally described from Easter island. From these different campaigns, unstudied collections are still deposited at the Muséum national d'Histoire naturelle, Paris, for example the crustacea collected by PLESSIS, from which we give here the first record of Thalamita danae. Others collections were made during the campaigns of the fishing boat Marara, used by the Direction des Centres d'Expérimentations Nucléaires for monitoring the marine environment, as a part of the radiological safety program in French Polynesia. Although they mainly concerned the deep fauna (POUPIN, 1996), some of the most common littoral and sublittoral species were also collected, and were presented by POUPIN (1994a), in a small illustrated document. In 1971, the French Ecole Pratique des Hautes Etudes, established its research center in French Polynesia. First located at Tahiti, it was afterwards transfered to Opunohu Bay, Moorea Island, and is now denominated Centre de Recherche Insulaire et Observatoire de l'Environnement (CRIOBE). Several important works have been done by the students or researchers of this center. They are mostly ecological studies with, however, about 30 new records for the area. The cryptofauna was studied by PEYROT- CLAUSADE, at Moorea (1977, 1985), and Tikehau (1989). The study of this small fauna was completed by NAIM (1980) with a dozen of species associated with the algae, at Tiahura, Moorea. The crustacean associates of the coral Pocillopora, were studied by KROPP & BIRKELAND (1981), and by ODINETZ (1983, 1984a, b) who described two new Trapezia species, Trapezia serenei and T. punctimanus. MONTEFORTE (1984) in his Contribution a la connaissance de la faune carcinologique de Polynésie francaise, collected and studied more than 110 species, some of them, like Calcinus minutus, Calappa calappa, or Etisus anaglyptus, being new records. More recently, a dozen common species were recorded from the atoll of Nukutipipi by MERSCHARDT-SALVAT (1991), and the freshwater collections made by MARQUET (1988, 1991, 1993), have given two new grapsid records, Varuna litterata, from the Society Islands, and Ptychognathus easteranus, from the Austral and Marquesas Islands. CONVENTIONS STUDIES INCLUDED We have included only studies in which material from French Polynesia has been actually examined, excluding works where "French Polynesia" appears only in the "Distribution". These are followed by the indication, in parenthesis, of the island(s) where the material originated. Recent revisions and general syntheses, from which the names of the species are updated (changes in generic classification; synonymies), or useful in different aspects, have been added; these are followed by mention such as "Syn." or "Key", in parenthesis. Some works have been included, that do not mention new collections. These are: the important syntheses about French Polynesia, like FOREST & GUINOT (1962), followed by "Biogeography", or GUINOT (1985), followed by "List"; the ecological studies focusing on a particular island, such as DELESALLE (1985), for the atoll of Mataiva, or SALVAT & RICHARD (1985), for the atoll of Takapoto; general works dedicated to the fauna of French Polynesia, such as SEURAT (1934), CHABOUIS L. & F. (1954), or more recently the Encyclopédie de la Polynésie (cf. CHARLEUX, 1986 and SALVAT, 1986a-c) and the books by PARDON (1992) and BONVALLOT et al. (1994). For these latter, however, only the species illustrated, usually in colour, have been cited. No distinction has been made between systematic and ecological works. In her list of the Brachyura of French Polynesia, GUINOT (1985) has sometimes considered the record of a species doubtful (name followed by a ?), when it was known only from an ecological paper. As these works can easily be identified from the references, the reader will be able to judge for himself. It is clear, however, that the revision of these collections would be important, but it is often difficult, or even impossible, to retrieve the material. LOCATIONS The unit of location is the island (see map). When the name of a village, a particular locality, or a small islet on the recifal crown (Motu in Polynesian language), was indicated, the name of the corresponding island is mentioned with the following presentation: "Gatavake" = Mangareva, "Ohura" = Hao, or "Taiohae" = Nuku Hiva. The same presentation is adopted for corrections of obvious mistakes: "Tickahau" = Tikehau, "Timoe" = Temoe, or "Fakaina" = Fakahina. The Gambier Islands have a particular configuration, with 8 mountainous islands surrounded by a common recifal crown. In that particular case, we have considered as real islands three Motu of the external crown: Puaumu, Tarauru-Roa, and Vaiatekeue. Sometimes, in the oldest works, the names of the islands were old names, no longer now in use. They are translated into modern names by using MOTTLER's (1986) work, and the following presentation: "Carlshoff' = Aratika, "Clermont-Tonnerre" = Reao, or "Eimeo" = Moorea. When no particular location was specified, the reference is just followed by "French Polynesia". CLASSIFICATION The classification approximately follows BOWMAN & ABELE (1982), and, for the Xanthoidea, SERENE (1984). Subfamily ranks has been indicated only within the most important families: Portunidae, Xanthidae, and Grapsidae. The presentation has been clarified by ignoring subgeneric names in the check list. However, if they were used in the works cited, they appear in the references. LITTORAL, SUBLITTORAL AND DEEP SPECIES These three groups are here defined in the following way: littoral species are commonly collected on the reef, and in depth of few meters only; sublittoral species are collected from about 10m to 100m; and deep species are collected from 100m and deeper. It is sometimes difficult to classify the species according to these three goups, especially because our knowledge about the deep distribution of numerous species is often very limited, and will have to be revised in the future. As an example, the maximal depth known for some species has been increased, here, sometimes considerably, from collections made by traps. Some littoral forms have been found unusually deep (Carpilius convexus, 60m; Charybdis paucidentata, 100m). They are qualified as "Littoral to sublittoral" species. Sublittoral forms have been sometimes collected far beyond 100m (Dardanus brachyops, 110-300m; Dromia wilsoni, 190-350m; Thalamita spinifera, 42- 200m), and, in contrast, deep forms have been found in less than 100m (Palibythus magnificus, 70-240m; Scyllarus aurora, 90-300m; Alainodaeus rimatara, 90-350m). They are qualified as "Sublittoral to deep" species. Because of these difficulties, 12 species included in this work were also listed with the deep species (POUPIN, 1996; see Appendices 1, species with a *). It is important to realise that, in several cases, these classifications are questionable, and often only reflect the poor information that we have on that subject. For example, a species like Oreotlos potanus, known by a single specimen, has been included with the deep species, to within 1m (101m). Considering the limited accuracy of the measures at sea, O. potanus could have been reasonably considered as a sublittoral species. OTHERS CONVENTIONS When a "?” appears in front of the name of a species, it always means that the doubt on that name is ours. When it is an hesitation expressed in the work consulted, it is mentioned after the reference by "with a 9" When a species has been reduced to synonymy, two cases are considered. If the species was not originally described from French Polynesia, the full name (author and date) appears at the end of the references, after "SYNONYMS". Otherwise, this information appears clearly with the reference, and is not repeated again. In both cases the origin of the synonymy is to be found in the work followed by "Syn.". The synonymies are restricted to French Polynesian species only. As far as possible we have tried to avoid partial identifications. When a generic name is only available, the reference is not considered in the main list, but appears separately, in Appendices 2. Preliminaries identifications (aff. or cf. ) have been retained only when the species refered to is not yet recorded from French Polynesia (for example, Actaea aff. glandifera in PEYROT-CLAUSADE, 1989: 111). Otherwise, they appear under the species refered to, after "RELEVANT MATERIAL" (see for example, Lybia cf. caestifera in MONTEFORTE, 1984: 171, under Lybia caestifera). The following abbreviations are used: BM (Natural History Museum, London); CRIOBE (Centre de Recherche Insulaire et Observatoire de l'Environnement, Moorea); MNHN (Muséum national d'Histoire naturelle, Paris); USNM (National Museum of Natural History, Washington). LIST OF THE SPECIES INFRA-ORDER ASTACIDEA FAMILY ENOPLOMETOPIDAE Enoplometopus holthuisi Gordon, 1968 DISTRIBUTION. — Tuamotu - Sublittoral. REFERENCES. — Enoplometopus holthuisi - BONVALLOT et al., 1994: 144-145, photograph (Tuamotu). REMARK. — At least another Enoplometopus is present in French Polynesia (cf. Enoplometopus sp. nov. in POUPIN ef al., 1990: 16, pl. 3c). INFRA-ORDER PALINURIDEA FAMILY PALINURIDAE Justitia longimanus (H. Milne Edwards, 1837) DISTRIBUTION. — Austral (Rurutu); Society (Bora Bora, Raiatea, Tahiti); Tuamotu (Makemo, Maria, Tenarunga) - Sublittoral to deep. REFERENCES. — Justitia longimana - POUPIN et al., 1990: 16 (French Polynesia). — Justitia longimanus - POUPIN, 1994b: 46, fig. 3e’, pl. 1d, 2d (Bora Bora, Tahiti, Tenarunga, Rurutu; 62-160m); 1996: in press (Bora Bora, Makemo, Maria, Raiatea, Rurutu, Tenarunga; 80-190m). Panulirus homarus (Linné, 1758) DISTRIBUTION. — Marquesas (Fatu Hiva, Nuku Hiva); Society (Tahit1). REFERENCES. — Panulirus homarus - GORDON, 1953: 29, fig. 2b-d, 6, 7b (Marquesas "Hana Hevané" = Hanavave bay?, Fatu Hiva; Puerulus larvae only). — MICHEL, 1971: 467 (Marquesas; Phyllosom larvae only). — HOLTHUIs, 1991: 139, fig. 267-268 (Marquesas, with a ?; Syn.). — ? Panulirus (sic) spinosus (Edwards) - CANO, 1888: 179 (Tahiti) - NEW MATERIAL - Frebruary 1996, Coll. & det. J. POUPIN (Nuku Hiva) - SYNONYMS - Palinurus spinosus H. Milne Edwards, 1837 (with a ?, in HOLTHUIS, 1991: 139). Panulirus longipes (A. Milne Edwards, 1868) DISTRIBUTION. — Marquesas; Society (Tahiti); Tuamotu. REFERENCES. — Senex femoristriga - ORTMANN, 1891: 23 (Tahiti). — Panulirus longipes - MICHEL, 1971: 467 (Marquesas, Tuamotu; Phyllosom larvae only). — Panulirus longipes femoristriga - HOLTHUIS, 1991: 146, fig. 277b, 278 (Syn.). REMARK. — In the Indo-West Pacific HOLTHUIS (1991) recognises two subspecies: Panulirus longipes, the western form, distributed from Africa to Thailand, Taiwan, Indonesia, and Philippines; and P. longipes femoristriga, the eastern form, known from Japan, the Moluccas, New Guinea, New Caledonia, eastern Australia, and French Polynesia. Panulirus pascuensis Reed, 1954 DISTRIBUTION. — Austral (MacDonald bank, Marotiri, Rapa) - Littoral to sublittoral. REFERENCES. — Panulirus pascuensis - LABOUTE & RICHER DE FORGES, 1986: 7, 21, pl. 2c (MacDonald bank, Marotiri, Rapa; 40m). — SALVAT, 1986b: 70, photograph (MacDonald bank). — HOLTHUIS, 1991: 149, fig. 283-284 (Pitcairn, 500km south-east off the Gambier). — POUPIN, 1994a: 8 (after LABOUTE & RICHER DE FORGES). REMARK. — LABOUTE & RICHER DE FORGES (1986: 18) also record Panulirus polyphagus (Herbst, 1793) in French Polynesia ("Iles hautes et atolls"). This species, which is only listed without material examined, is not reported from the area by HOLTHUIS (1991: 152). We therefore consider that this record is not valid. Panulirus penicillatus (Olivier, 1791) DISTRIBUTION. — Gambier; Marquesas; Society (Tahiti); Tuamotu (Makatea, Mataiva, Moruroa, Fakarava, Hao, Raroia, Taiaro, Takapoto). REFERENCES. — Panulirus penicillatus - STIMPSON, 1860: 23 [92] (Tahiti). — BATE, 1888: 82, pl. 12-fig. 2 (Tahiti). — NoBILI, 1907: 366 (Hao). — BOONE, 1935: 67, pl. 17 (Tahiti). — SEURAT, 1934: 60 (Gambier, Tuamotu). — HOLTHUIS, 1953: 50 (Raroia). — CHABOUIS L. & F., 1954: 89 (Tahiti). — MorRIson, 1954: 16 (Raroia). — MICHEL, 1971: 467 (Marquesas, Tuamotu; Phyllosom larvae). — CHEVALIER et al., 1968: 92, 137 (Moruroa). — MONTEFORTE, 1984: 173, annex 1, tab. a (Makatea, Mataiva, Tahiti, Takapoto). — DELESALLE, 1985: 289, 293 (Mataiva). — SALVAT, 1986b: 70, photograph (French Polynesia). — BAGNIS & CHRISTIAN, 1983: 108 (Tuamotu). — HOLTHUIS, 1991: 151, fig. 285-286 10 (Tuamotu; Syn.). — PARDON, 1992: 83, photograph (Fakarava). — POUPIN, 1994a: 8, fig. 4 (Taiaro). — Cancer theresae Curtiss, 1938 (""Tautira" = Tahiti). Panulirus versicolor (Latreille, 1804) DISTRIBUTION. — Society (Tahiti). REFERENCES. — ? Panulirus fasciatus - CANO, 1888: 179 (Tahiti; cf. Remark). — Panulirus ornatus - BOONE, 1935: 63, pl. 16 (Tahiti). — POUPIN, 1994a: 8 (French Polynesia; after BOONE, and erroneously after NOBILI, 1907 and GRUVEL, 1911) - Not Palinurus ornatus (Fabricius, 1798) (cf. Remark). — Panulirus versicolor - HOLTHUIS, 1946: 142, pl. 6-j, pl. 9-b, pl. 11-e,f,m (Tahiti); 1991: 156, fig. 293-294 (French Polynesia). REMARK. — In HOLTHUIS (1991: 152) Panulirus fasciatus Fabricius, 1798 is a synonym of P. polyphagus (Herbst, 1793). However, CANO's (1888) reference to P. fasciatus would rather be P. versicolor, often recorded under P. fasciatus (HOLTHUIS, 1991: 152). Moreover, we observe that, in his catalogue, HOLTHUIS (1991) does not mention P. polyphagus from French Polynesia. According HOLTHUIS (1946: 140, 142), BOONE's (1935) Tahitian record of Panulirus ornatus (Fabricius, 1798) would be erroneous, but it is not certain that his material really belongs to P. versicolor. FAMILY SYNAXIDAE Palibythus magnificus Davie, 1990 DISTRIBUTION. — Society (Tahiti) - Sublittoral to deep. REFERENCES. — Palinurellus wienechi (sic) - ANONYMOUS, 1979: 6, 8, 11, not Palinurellus wieneckii (de Man, 1881) = Palibythus magnificus (Tahiti, 70-240m; material corresponding to the photographs examined and considered by DAVIE (1990: 686) as "almost definitely of this species"). — Palibythus magnificus Davie, 1990: 686, fig. 1a-b, 3a, c, 4a, 5a (Tahiti; but not Tuamotu). — POUPIN, 1996: in press (Tahiti, Tuamotu?). REMARK. — The beautiful set of 10 specimens, collected in 1978 in front of Taravao, Tahiti, has disappeared. It remains only the photographs examined by DAVIE (1990). Palinurellus wieneckii (De Man, 1881) DISTRIBUTION. — Tuamotu - Sublittoral. REFERENCES. — Palinurellus wieneckii - MICHEL, 1971: 460, fig. 1a-j, tab. 1 (Tuamotu; Puerulus larvae only). — HOLTHUIS, 1991: 170, fig. 315-316 (Tuamotu: larvae and juveniles; 9-27m). REMARK. — The larvae, once attributed to this species, could in fact belong to Palibythus magnificus, afterwards collected in the area (cf. previous species). FAMILY SCYLLARIDAE Arctides regalis Holthuis, 1963 DISTRIBUTION. — Tuamotu. REFERENCES. — Arctides antipodarum - MICHEL, 1971: 467 (Tuamotu; Phyllosom larvae) not A. antipodarum Holthuis, 1960 = A. regalis, with a doubt, fide HOLTHUIS (1991: 177). — Arctides regalis. — HOLTHUIS, 1991: 177, fig. 331-332 (Tuamotu; Syn.). Parribacus antarcticus (Lund, 1793) DISTRIBUTION. — Society (Maupiti, Tahiti); Tuamotu (Anaa, Manihi?, Moruroa, Raroia, Takapoto, Tureia). REFERENCES. — Scyllarus antarcticus - OWEN, 1839: 86 ("Carysfort" = Tureia). — Parribacus antarcticus - SEURAT, 1934: 60 (Tuamotu). — HOLTHUIS, 1953: 50 (Raroia) pro parte fide HOLTHUIS (1985: 74); 1985: 73, fig. 21, 25a (Anaa, 11 Maupiti, Tahiti, Takapoto; Syn.). — CHABOUIS L. & F., 1954: 89, unnumbered fig. (French Polynesia). — FOREST, 1954b: 345, fig. 26a (Tahiti). — CHEVALIER et al., 1968: 92, 137 (Moruroa). — ? BABLET, 1972: 32, pl. 10 (French Polynesia). — Parribacus ursus-major - BOONE, 1935: 54, pl. 13 (Tahiti). — Cancer barffi Curtiss, 1938: 164 ("Tautira” = Tahiti). — ? "Tiane” - PARDON, 1992: 83, photograph (Manihi) (det. according to the photograph). — Not Parribacus antarcticus - STIMPSON, 1860: 92 [23] (Tahiti). — NoBILI, 1907: 366 (Hao, "Rikitea” = Mangareva). — HOLTHUIS, 1953: 50 (Raroia), pro parte. — MorRISON, 1954: 50 (Raroia) - All = Parribacus holthuisi Forest, 1954 fide HOLTHUIS (1985: 75) - SYNONYMS - Parribacus ursus-major (Herbst, 1793). Parribacus holthuisi Forest, 1954 DISTRIBUTION. — Gambier (Mangareva); Society (Moorea, Tahiti); Tuamotu (Hao, Hikueru, Mataiva, Raroia, Takapoto). REFERENCES. — Parribacus antarcticus - STIMPSON, 1860: 92 [23] (Tahiti). — NOBILI, 1907: 366 (Hao, "Rikitea" = Mangareva). — HOLTHUIS, 1953: 50 (Raroia) pro parte. — MORRISON, 1954: 50 (Raroia) - All, not P. antarcticus (Lund, 1793) = P. holthuisi fide HOLTHUIS (1985: 98). — Parribacus holthuisi Forest, 1954b: 346, fig. 25, 26b (Hikueru, Tahiti). — MONTEFORTE, 1984: 173, annex 1, tab. a (Mataiva, Moorea, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — HOLTHUIs, 1985: 98 (Hao, Hikueru, Mangareva, Tahiti, Raroia). — SALVAT, 1986b: 70, 71, photograph (French Polynesia). Parribacus scarlatinus Holthuis, 1960 DISTRIBUTION. — Marquesas (Fatu Hiva). REFERENCES. — Parribacus scarlatinus - MICHEL, 1971: 472 (Marquesas, Omoa bay = Fatu Hiva). — HOLTHUIS, 1985: 102, fig. 26; 1991: 215, fig. 411-412 (Marquesas). Scyllarus aurora Holthuis, 1981 DISTRIBUTION. — Austral (Maria, Rurutu, Tubuai); Gambier; Marquesas (Fatu Hiva, Tahuata); Society (Maupiti, Moorea, Raiatea, Tupai); Tuamotu (Akiaki, Fangataufa, Hao, Makemo, Marutea South, Maria, Moruroa, Tuanake, Tureia, Vanavana) - Sublittoral to deep. REFERENCES. — ? Scyllarus sp. TV & V - MICHEL, 1971: 467, tab. 3 (Marquesas, Tuamotu; larvae only). — Scyllarus aurora Holthuis, 1981: 847, fig. 1-2 (Tubuai; 200m). — MANAC'H & CarsIN, 1985: 473 (Moruroa and/or Fangataufa). — POUPIN, 1996: in press (Common, 90-300m"; in the distribution, most of the islands are mentioned here for the first time). INFRA-ORDER ANOMURA FAMILY COENOBITIDAE Birgus latro (Linné, 1767) DISTRIBUTION. — Gambier (Temoe); Tuamotu (Amanu, Makatea, Marutea South, Matureivavao, Morane, Niau, Pukapuka, Raroia, Taiaro, Takapoto) - Terrestrial. REFERENCES. — Birgus latro - DANA, 1852b: 474; 1855, pl. 30, fig. Sa-b (several islands in the Tuamotu; drawing of a specimen from "Honden" = Pukapuka). — NoBILI, 1907: 375 (Amanu). — SEURAT, 1904a: 242 (Marutea South, Temoe, "Moture-vavao" = Matureivavao); 1934: 51 (French Polynesia). — SENDLER, 1923: 44 (Makatea). — HOLTHUIS, 1953: 36 (Raroia). — MOorRISON, 1954: 10 (Raroia). — FOREST, 1954a: 79; 1956a: 1073 (Niau). — CHABOUIS L. & F., 1954: 92, unnumbered fig. (Makatea). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Takapoto). — DELESALLE, 1985: 288 (Mataiva). — CHARLEUX, 1986: 80, photograph (French Polynesia). — SALVAT & 12 RICHARD, 1985: 356 (Takapoto). — SALVAT, 1986b: 71; 1986c: 8-9, photograph (French Polynesia). — BONVALLOT et al., 1994: 76, photograph (Tuamotu). — POUPIN, 1994a: 10, fig. 6, pl. 1h (Taiaro) - NEW MATERIAL - Coll. and det. J. POUPIN (Morane). REMARK. — GIBSON-HILL (1948: 10) mentions this species from the Marquesas Islands, but it is doubtful that it really occurs in these Islands, where we have made several unsuccessful investigations. Coenobita brevimanus Dana, 1852 DISTRIBUTION. — Society (Bora Bora, Tahiti); Tuamotu (Amanu, Hao, Hikueru, Makatea, Mataiva, Niau, Nukutipipi, Raroia, Takapoto, Taiaro) - Terrestrial. REFERENCES. — Cenobita clypeata Latr. - HELLER, 1865: 82 (Tahiti). — SEURAT, 1934: 52 (Amanu, Hao). — Coenobita clypeatus (Herbst) - ORTMANN, 1892a: 316, pl. 12, fig 20 (Tahiti) not C. clypeatus (Herbst, 1794) = C. hilgendorfi Terao in TERAO (1913: 388). — Coenobita clypeatus Latr. - NOBILI, 1907: 373 (Amanu, "Ohura" = Hao). — SENDLER, 1923: 42 (Makatea, "Nian" = Niau). — Coenobita hilgendorfi Terao, 1913: 388 (Syn.; cf. Remark). — FOREST, 1954a: 77 (Hikueru; Syn.); 1956a: 1072 (Hikueru, Tahiti). — Cenobita clypeatus Latreille - BOONE, 1935: 40, pl. 9 (Bora Bora). — Coenobita brevimanus - HOLTHUIS, 1953: 36 (Raroia). — MORRISON, 1954: 10 (Raroia). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Takapoto). — SALVAT, 1986b: 72 (French Polynesia). — NAKASONE, 1988: 174 (Syn.). — MERSCHARDT-SALVAT, 1991: 40 (Nukutipipi). — SALVAT F. & B., 1992: 5 (Nukutipipi). — PoOUPIN, 1994a: 11, fig. 7, pl. 1c (Hikueru, Tahiti, Taiaro). — Coenobita ollivieri (sic) - CHARLEUX, 1986: 80-81, photograph (French Polynesia) not C. olivieri (Owen, 1839) = C. brevimanus (correction according to the photograph). REMARK. — In TERAO (1913: 389) Coenobita clypeatus (Herbst, 1794) is different from Coenobita clypeatus (Latreille, 1826), and the name Coenobita hilgendorfi is proposed for LATREILLE's material. More recently, NAKASONE (1988) considers that TERAO's (1913) C. hilgendorfi is the same than C. brevimanus Dana, 1852, and states that, until 1955, DANA's species has been often referred to as, either C. clypeatus, or C. hilgendorfi. Coenobita carnescens Dana 1851 DISTRIBUTION. — Tuamotu (Ahe and/or Manihi, Aratika, Kauehi, Pukapuka, Raraka) - Terrestrial. REFERENCES. — Cenobita carnescens Dana, 1851: 272 (Paumotu); 1852b: 472; 1855, pl. 30, fig. 3a-b ("Carlshoff' = Aratika, "Honden" = Pukapuka, Raraka, "Vincennes" = Kauehi, "Waterland" = Ahe and/or Manihi). — PouPIN, 1994a: 9, fig. 5 (Text). REMARK. — In NAKASONE (1988: 165) this species would be valid, although it was considered doubtful by BOUVIER (in ALCOCK, 1905: 193). According to the drawings provided by DANA, Coenobita carnescens could be in fact the juvenile form of C. perlatus (cf. POUPIN, 1994a: 12, pl. 1d-f). Coenobita cavipes Stimpson, 1858 DISTRIBUTION. — Austral (Rimatara) - Terrestrial. REFERENCES. — Coenobita cavipes - SENDLER, 1923: 43 (Rimatara). — MIYAKE, 1991: 116, fig. 3 (cited only for the illustration). — POUPIN, 1994a: 9 (Text). Coenobita olivieri (Owen, 1839) DISTRIBUTION. — Gambier (Tarauru-Roa); Society (Tahiti) - Terrestrial. REFERENCES. — Coenobita olivieri - NOBILI, 1907: 374 ("Tarawao, Papenoo" = Tahiti, Tarauru-Roa; cf. Remark). — SEURAT, 1934: 52 (Tahiti, Gambier). — FOREST, 1956a: 1056 (French Polynesia). — POUPIN, 1994a: 14 (Tahiti; ef. Remark). — Not Coenobita olivieri - DANA, 1852b: 470 = C. spinosus H. Milne Edwards fide ORTMANN (1892a: 318). — BAGNIS & CHRISTIAN, 1983: 108, photograph (Tuamotu) = C. perlatus H. Milne Edwards fide POUPIN (1994a: 12). REMARK. — A specimen attributed by NoBILI to Coenobita olivieri (Owen, 1839) has been examined previously (PouPIN, 1994a; Tahiti "Papenoo", MNHN Pg2111). It is very close to Coenobita spinosus, and ORTMANN's (1892a) opinion that Coenobita olivieri is only a variety of C. spinosus, could be justified. Examination of the type material is required to clarify that point. 13 Coenobita perlatus H. Milne Edwards, 1837 DISTRIBUTION. — Gambier (Mangareva, Tarauru-Roa, Temoe); Society (Moorea, Tahiti); Tuamotu (Amanu, Hao, Hikueru, Kaukura, Makatea, Marutea South, Mataiva, Moruroa, Nukutipipi, Raroia, Taiaro, Takapoto, Takume) - Terrestrial. REFERENCES. — Coenobita perlata - SEURAT, 1904a: 238 (Mangareva, Tarauru-Roa, Temoe, Marutea South); 1904b: 95 (Marutea South); 1934: 51 (French Polynesia). — CHEVALIER ef al., 1968: 85, 137 (Moruroa). — Coenobita perlatus - NoBILI, 1907: 373 (Amanu, Hao, Kaukura, Marutea, Tarauru-Roa). — SENDLER, 1923: 43 (Makatea, "Uusuroa, Paumotu" = ?). — HOLTHUIS, 1953: 37 (Raroia). — MORRISON, 1954: 7 (Raroia). — CHABOUIS L. & F., 1954: 93 ("Mataia" = Tahiti). — FOREST, 1954a: 78; 1956a: 1072 (Hikueru, Takume). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 6 (Moorea). — DELESALLE, 1985: 288, 289 (Mataiva). — SALVAT & RICHARD, 1985: 359, 360 (Takapoto). — CHARLEUX, 1986: 80-81, photograph (French Polynesia). — SALVAT, 1986b: 71 (French Polynesia). — MERSCHARDT-SALVAT, 1991: 40 (Nukutipipi). — PARDON, 1992: 83, photograph (Tuamotu). — SALVAT F. & B., 1992: 5 (Nukutipipi). — BONVALLOT et al., 1994: 77, photograph (Tuamotu). — POUuPIN, 1994a: 12, fig. 8, pl. 1d,f (Hao, Mangareva, Marutea, Taiaro). — Coenobita rugosus vat. granulatus Bouvier - NOBILI 1907: 373 (Marutea, "Ohura" = Hao) not C. rugosus H. Milne Edwards, 1837 = C. perlatus fide FOREST (1954a: 78). — Coenobita rugosus - NoBILI, 1907: 373 (Hao, Kaukura) — SEURAT, 1934: 52 (NOBILI's material) - These two references, pro parte not C. rugosus H. Milne Edwards, 1837 = C. perlatus fide FOREST (1954a: 78). — Coenobita olivieri - BAGNIS & CHRISTIAN, 1983: 108, photograph (Tuamotu) not C. olivieri (Owen, 1839) = C. perlatus fide POUPIN (1994a: 12). Coenobita rugosus H. Milne Edwards, 1837 DISTRIBUTION. — Austral (Raevavae); Society (Tahiti); Tuamotu (Hikueru, Kaukura, Raraka, Raroia, Takume) - Terrestrial. REFERENCES. — Cenobita rugosa - DANA, 1852b: 471; 1855, pl. 30, fig. 1 (Raraka). — STIMPSON, 1858c: 245 [83]; 1907: 199 (Tahiti). — HELLER, 1865: 82 (Tahiti). — HENDERSON, 1888: 51 (Tahiti). — SEURAT, 1934: 52 (Kaukura) — Coenobita rugosus - ORTMANN, 1892a: 317, pl. 12, fig. 22 (Tahiti). — NoBiL, 1907: 373 (Kaukura, "Ohura" = Hao) pro parte fide FOREST (1954a: 78; some specimens are C. perlatus). — SENDLER, 1923: 42 (Tuamotu). — FOREST, 1954a: 78; 1956a: 1073 (Hikueru, Tahiti, Takume). — HOLTHUIS, 1953: 40 (Raroia). — MORRISON, 1954: 10 (Raroia). — POUPIN, 1994a: 13, fig. 9, pl. le (Raevavae, Tahiti). — Not C. rugosus (pro parte) and C. rugosus var. granulosa Bouvier - NoBILI, 1907: 373 = C. perlatus fide FOREST (1954a: 78). Coenobita spinosus H. Milne Edwards, 1837 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Ahe and/or Manihi, Amanu, Niau, Nukutipipi, Reao) - Terrestrial. REFERENCES. — Cenobita olivieri - DANA, 1852b: 470 ("Clermont Tonnerre" = Reao, Tahiti, "Waterland" = Ahe and/or Manihi) not C. olivieri (Owen, 1839) = C. spinosus fide ORTMANN (1892a: 318). — Coenobita spinosus - NOBILI, 1907: 374 (Amanu). — SENDLER, 1923: 43 ("Nian"=Niau). — SEURAT, 1934: 52 (Amanu). — FOREST, 1956a: 1056 (French Polynesia). — MERSCHARDT-SALVAT, 1991: 40 (Nukutipipi). — SALVAT F. & B., 1992: 5 (Nukutipipi). — POUPIN, 1994a: 14, fig. 10, pl. 1g (Amanu, Moorea, Tahiti). FAMILY DIOGENIDAE Aniculus aniculus (Fabricius, 1787) DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Moorea, Tahiti); Tuamotu (Ahe and/or Manihi, Aratika, Fakahina, Hikueru, Makatea, Marokau, Marutea South, Mataiva, Moruroa, Nukutipipi, Rangiroa, Raraka, Raroia, Takapoto, Tikehau). REFERENCES. — Aniculus typicus - DANA, 1852b: 461; 1855, pl. 29, fig. 1 ("Carlshoff' = Aratika, Raraka, "Waterland" = Ahe and/or Manihi). — CANO, 1888: 178 (Tahiti). — Aniculus aniculus - NOBILI, 1907: 371 (Marokau). — SEURAT, 1934: 61(Tuamotu, Gambier). — BOONE, 1935: 36, pl. 8 (Tahiti). — HOLTHUIS, 1953: 41 (Raroia). — MORRISON, 14 1954: 13 (Raroia). — FOREST, 1953b: 561; 1956a: 1067 (Hikueru, Tahiti); 1984: 21, fig. 8, 16, 28-30, 35, 66, 68, 74, 76-85 (Fakahina, Hikueru, Mangareva, Marokau, Marutea South, "Matahiva and Tiahura" = Moorea, Moruroa, Rangiroa, Raroia, Tahiti, "Tawhae, Marquesas" = Taiohae at Nuku Hiva, "Tikahau atoll"=Tikehau; Syn.). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Moorea, Takapoto); 1987: 8 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — RAHAYU, 1988: 40 (French Polynesia). — MERSCHARDT-SALVAT, 1991: 40 (Nukutipipi). — SALVAT F. & B., 1992: 5 (Nukutipipi). REMARK. — Aniculus typicus, proposed by DANA (1852c) in the place of Pagurus aniculus Fabricius, is no more a valid name (cf. FOREST, 1984: 21). Aniculus maximus Edmonson, 1952 DISTRIBUTION. — Marquesas (Fatu Hiva, Nuku Hiva). REFERENCES. — Aniculus maximus - FOREST, 1984: 61, fig. 14, 22, 59-61 (Fatu Hiva, Nuku Hiva). — SALVAT, 1986a: 6, 7, photograph (French Polynesia). — ? Aniculus sp. - SALVAT, 1986b: 71, photograph (French Polynesia; det. according to the photograph). Calcinus elegans (H. Milne Edwards, 1836) DISTRIBUTION. — Gambier (Kamaka, Makaroa, Mangareva); Society (Bora Bora, Tahiti); Tuamotu (Ahe and/or Manihi, Amanu, Apataki, Aratika, Hao, Hikueru, Marutea South, Mataiva, Raroia, Reao, Tagatau, Taiaro, Takume). REFERENCES. — Calcinus elegans - DANA, 1852b: 458; 1855, pl. 28, fig. 10a-c (‘Carlshoff' = Aratika, "Clermont Tonnerre" = Reao, "Waterland" = Ahe and/or Manihi). — HELLER, 1865: 88 (Tahiti). — NOBILI, 1907: 368 (Amanu, Hao, Kamaka, Makaroa, Marutea South, "Rikitea" = Mangareva, Tagatau, "Wakatihi" = ?). — BOONE, 1935: 23, pl. 3 (Bora Bora, Tahiti). — HOLTHUIS, 1953: 41 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST, 1953b: 555; 1956a: 1062 (Hikueru, Tahiti, Takume). — MONTEFORTE, 1984: 172, annex 1, tab. a (Mataiva). — RAHAYU, 1988: 10, 17 (French Polynesia). — POUPIN, 1994a: 15, fig. 11, pl. 2a (Hao, Taiaro). Calcinus gaimardi (H. Milne Edwards, 1848) DISTRIBUTION. — Society (Moorea, Tahiti). REFERENCES. — Calcinus gaimardi - HELLER, 1865: 87 (Tahiti). — FOREST, 1953b: 555; 1956a: 1062 (Tahiti). — Kropp & BIRKELAND, 1981: 630, tab. 5 (Moorea). — MONTEFORTE, 1984: 172, annex 1, tab. a; 1987: 8 (Moorea). — RAHAYU, 1988: 20 (Tahiti). Calcinus guamensis Wooster, 1984 DISTRIBUTION. — Marquesas (Fatu Hiva, Hiva Oa). REFERENCES. — Calcinus guamensis - NEW MATERIAL - February 1996, Coll. J. POUPIN, det. J. POUPIN & J. FOREST (Fatu Hiva, Hiva Oa). Calcinus imperialis Whitelegge, 1901 DISTRIBUTION. — Society (Tahiti); Tuamotu (Moruroa). REFERENCES. — Calcinus imperialis - NEW MATERIAL - October 1995, Coll. (in coral Pocillopora) & det. J. POUPIN (Moruroa, Tahiti "Taravao"). — MORGAN, 1991: 882, fig. 21-23 (Syn.; Key). Calcinus laevimanus (Randall, 1840) DISTRIBUTION. — Gambier (Kamaka, Mangareva, Puaumu); Society (Moorea, Tahiti); Tuamotu (Ahe and/or Manihi, Aratika, Hao, Hikueru, Kauehi, Makatea, Marutea South, Mataiva, Raraka, Raroia, Tagatau, Taiaro, Takapoto, Takume). REFERENCES. — Calcinus tibicen - DANA, 1852b: 457 ("Carlshoff' = Aratika, Raraka, "Vincennes" = Kauehi, "Waterland" = Ahe and/or Manihi) not C. tibicen (Herbst, 1791) = C. laevimanus fide MORGAN (1991: 888). — HELLER, 1865: 87 (Tahiti; cf. Remark). — HENDERSON, 1888: 61 (Tahiti; of. Remark). — Calcinus herbstii - NOBILI, 1907: 368 pro parte fide FOREST (1951: 84) (Hao, Hikueru, Kamaka, Marutea, "Puamu" = Puaumu, "Rikitea” = 15 Mangareva, Tagatau, Tahiti). — SENDLER, 1923: 42 (Makatea). — FOREST, 1951: 84 (NOBILI's material); 1953: 555; 1956a: 1062 (Hikueru, Tahiti, Takume). — Calcinus herbstii var. lividus Edw. - NoBILi, 1907: 369 (Marutea; cf. Remark). — Calcinus laevimanus - HOLTHUIS, 1953: 43 (Raroia). — MORRISON, 1954: 7 (Raroia). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Moorea, Takapoto); 1987: 8 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — RAHAYU, 1988: 10, 18, fig. 1-3 (French Polynesia). — POUPIN, 1994a: 16, fig. 12 (Hikueru, Taiaro) - SYNONYMS - Calcinus herbstii de Man, 1888; Pagurus lividus H. Milne Edwards, 1848. REMARK. — Calcinus tibicen (Herbst, 1791) is an Atlantic species. MORGAN (1991: 888) has cited several works in which C. laevimanus is referred to as C. tibicen, C. herbstii, or C. herbstii var. lividus, but without the works by HELLER, HENDERSON or NOBILI. Although not verified, this material is here attributed to C. laevimanus. Calcinus latens (Randall, 1840) DISTRIBUTION. — Gambier (Vaiatekeue); Society (Moorea, Tahiti); Tuamotu (Fakahina, Hao, Hikueru, Makatea, Mataiva, Raroia, Taiaro, Takapoto, Takume). REFERENCES. — Calcinus latens - HELLER, 1865: 88 (Tahiti). — NOBILI, 1907: 369 (Hao). — FOREST, 1951: 84, fig. 14-18 (French Polynesia); 1953b: 556 (Syn.); 1956a: 1062 (Hikueru, Tahiti, Takume). — HOLTHUIS, 1953: 44 (Raroia). — Morrison, 1954: 7 (Raroia). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — KRopp & BIRKELAND, 1981: 630, tab. 5 (Takapoto). — MONTEFORTE, 1984: 172, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 8 (Moorea). — GALZIN & POINTIER, 1985: 100 (Moorea). — SALVAT & RICHARD, 1985: 358 (Takapoto). — RAHAYU, 1988: 10, 19 (French Polynesia). — POUPIN, 1994a: 17, fig. 13, pl. 2b (Takume, Taiaro). — Calcinus herbstii - NOBILI, 1907: 368 (Hao, "canal Waiatekene” = Vaiatekeue) pro parte not C. herbstii de Man, synonym of C. laevimanus = C. latens fide FOREST (1951: 84). — Calcinus terrae-reginae - NOBILI, 1907: 369 (Fakahina, Hao, Mangareva) - SYNONYMS - Calcinus terrae-reginae Haswell, 1882. Calcinus minutus Buitendijk, 1937 DISTRIBUTION. — Society (Moorea). REFERENCES. — Calcinus minutus - MONTEFORTE, 1984: 172, annex 1, tab. a (Moorea). Calcinus nitidus Heller, 1865 DISTRIBUTION. — Society (Tahiti); Tuamotu (Takapoto). REFERENCES. — Calcinus nitidus Heller, 1865: 89, pl. 7, fig. 4 (Tahiti). — DE MAN, 1890: 111 (Tahiti). — ORTMANN, 1892a: 293 (Tahiti). — FOREST, 1956b: 218, fig. 14 (Tahiti). — RAHAYU, 1988: 10 (French Polynesia). — POUPIN, 1994a: 18, fig. 14, pl. 2c (Tahiti, Takapoto). Calcinus seurati Forest, 1951 DISTRIBUTION. — Gambier (Tarauru-Roa, Vaiatekeue); Society (Tahiti); Tuamotu (Hao, Hikueru, Fakahina, Raroia, Taiaro, Takume). REFERENCES. — Calcinus herbstii - NOBILI, 1907: 368 (Hao, Fakahina, "Taraourouroa"” = Tarauru-Roa, "chenal Waiatekene" = Vaiatekeue) pro parte not C. herbstii de Man, 1888, synonym of C. laevimanus = C. seurati fide FOREST (1951: 86). — Calcinus seurati Forest, 1951: 84, fig. 1, 3-4, 7-8 (NOBILI's material); 1953b: 556; 1956a: 1062 (Hikueru, Tahiti, Takume). — HOLTHUIS, 1953: 44 (Raroia, Takume). — MORRISON, 1954: 7 (Raroia). — RAHAYU, 1988: 19 (Hikueru, Tahiti, Takume). — POUPIN, 1994a: 19, fig. 15, pl. 2d (Hao, Taiaro). Calcinus spicatus Forest, 1951 DISTRIBUTION. — Gambier (Vaiatekeue). REFERENCES. — Calcinus herbstii - NOBILI, 1907: 368 ("chenal Waiatekene" = Vaiatekeue) pro parte not C. herbstii de Man, 1888, synonym of C. laevimanus = C. spicatus fide FOREST (1951: 84). — Calcinus spicatus Forest, 1951: 90, fig. 10-13 ("chenal Waiatekene" = Vaiatekeue). — RAHAYU, 1988: 21 (French Polynesia). 16 Ciliopagurus krempfi (Forest, 1952) DISTRIBUTION. — Marquesas (Tahuata) - Sublittoral. REFERENCES. — Ciliopagurus krempfi - FOREST, 1995: 59 fig. 10c, 11, 12c, 31d, 37g-h (Tahuata; 48m). Ciliopagurus strigatus (Herbst, 1804) DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Moorea, Tahiti). REFERENCES. — Pagurus strigatus - ORTMANN, 1892a: 285 (Tahiti). — Trizopagurus strigatus - MONTEFORTE, 1984: 172, annex 1, tab. a (Moorea, Tahiti); 1987: 8 (Moorea). — Ciliopagurus strigatus - FOREST, 1995: 49, fig. 8a, 9, 10a, 12a, 31a-b, 37d (Moorea, Nuku Hiva, Tahiti). Clibanarius corallinus (H. Milne Edwards, 1848) DISTRIBUTION. — Society (Bora Bora, Tahiti); Tuamotu (Apataki, Hao, Hikueru, Mataiva, Moruroa, Raroia, Taiaro, Takapoto, Takume). REFERENCES. — Clibanarius corallinus - HELLER, 1865: 89 (Tahiti; cf. Remark). — NOBILI, 1907: 367 (Apataki, Hao). — Boone, 1935: 17, pl. 1 (Bora Bora). — HOLTHUIS, 1953: 45 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST, 1953a: 442; 1956a: 1057 (Hikueru, Tahiti, Takume). — MONTEFORTE, 1984: 172, annex 1, tab. a (Mataiva, Takapoto). — RAHAYU, 1988: 26, fig. 4-6 (Tahiti). — POUPIN, 1994a: 20, fig. 16 (Apataki, Moruroa, Taiaro). REMARK. — HELLER (1862: 527) has also described a new Clibanarius from Tahiti, Clibanarius semistriatus. According to J. FOREST (Personal communication) this species is very doubtful and could be a Pagurus or a Paguristes. Clibanarius eurysternus Hilgendorf, 1878 DISTRIBUTION. — French Polynesia. REFERENCES. — Clibanarius eurysternus - RAHAYU, 1988: 10, 28 (French Polynesia). — RAHAYU & FOREST, 1992: 750 (Distribution only "Polynésie"). Clibanarius humilis (Dana, 1851) DISTRIBUTION. — Gambier (Mangareva, Tarauru-Roa); Society (Tahiti); Tuamotu (Hikueru, Mataiva, Moruroa, Takume). REFERENCES. — Clibanarius aequabilis - ? DANA, 1852b: 464; 1855, pl. 29, fig. 4a-f (Tahiti; cf. Remark). — NOBILI, 1907: 367. (""Rikitea" = Mangareva, Tarauru-Roa) not C. aequabilis Dana, 1852 = C. humilis fide FOREST (1953a: 443). — Clibanarius humilis - FOREST, 1953a: 443, fig. 1, 5; 1956a: 1057 (Hikueru, Tahiti, Takume). — MONTEFORTE, 1984: 172, annex 1, tab. a (Mataiva). — RAHAYU, 1988: 10, 27 (French Polynesia). — POUPIN, 1994a: 21, fig. 17 (Moruroa, Tahiti). REMARK. — Clibanarius aequabilis Dana, 1852 is an Atlantic species. Concerning DANA's material, FOREST (1953a: 446) writes: "A quoi correspond le sp. C. aequabilis de Tahiti, figuré par Dana ? II] est souhaitable que le terme d'aequabilis ne soit plus utilisé pour les espéces de ]'Indo-Pacifique." Clibanarius ransoni Forest, 1953 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Clibanarius ransoni Forest, 1953a: 446, fig. 2, 6; 1956a: 1059 (Tahiti). — RAHAYU, 1988: 30 (Tahiti). Clibanarius rhabdodactylus Forest, 1953 DISTRIBUTION. — Tuamotu (Hao, Hikueru). REFERENCES. — Clibanarius zebra - NoBILI, 1907: 367 (Hao), not C. zebra Dana, 1852 = C. rhabdodactylus fide RAHAYU & FOREST (1992: 777). — Clibanarius zebra var. rhabdodactylus Forest, 1953a: 448, fig. 3, 8; 1956a: 1059 (Hikueru). — RAHAYU, 1988: 29 (French Polynesia). — Clibanarius rhabdodactylus - RAHAYU & FOREST, 1992: 777 (Syn.). 17 Clibanarius striolatus Dana, 1852 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Clibanarius striolatus - HELLER, 1865: 89 (Tahiti). — FOREST, 1953a: 448; 1956a: 1059 (Tahiti). — RAHAYU, 1988: 10, 27 (Tahiti). Clibanarius zebra (Dana, 1852) DISTRIBUTION. — Marquesas (Fatu Hiva, Nuku Hiva, Ua Pou). REFERENCES. — Clibanarius zebra - FOREST, 1953a: 449, fig. 4 ("Taiohae" = Nuku Hiva) - NEW MATERIAL - Frebruary 1996, Coll. J. POUPIN, det. J. POUPIN & J. FOREST (Fatu Hiva, Nuku Hiva, Ua Pou). REMARK. — FOREST (1953a) states that the specimens from Marquesas (Coll. Pére Simon DELMAS) are typical of C. zebra, and distinctly larger than the rhabdodactylus variety. Dardanus australis Forest & Morgan, 1991 DISTRIBUTION. — Austral (Rapa) - Sublittoral to deep. REFERENCES. — Dardanus australis - NEW MATERIAL - March 1995, Coll. J. POUPIN, det. J. FOREST (Rapa, 70-115m). Dardanus brachyops Forest, 1962 DISTRIBUTION. — Marquesas (Tahuata); Society (Bora Bora, Maupiti) - Sublittoral to deep. REFERENCES. — Dardanus brachyops - POUPIN, 1996: in press (Bora Bora, Maupiti, Tahuata; 110-300m cf. Remark). REMARK. — This species is still known only beyond 100m in French Polynesia, but it is merely sublittoral in FOREST (1962: 365; more than 33m to 80m). Dardanus deformis (H. Milne Edwards, 1836) DISTRIBUTION. — Gambier (Mangareva?); Society (Tahiti); Tuamotu (Hao, Hikueru, Matureivavao). REFERENCES. — Pagurus difformis - HELLER, 1865: 86 (Tahiti). — HENDERSON, 1888: 57 (Tahiti). — Pagurus deformis - NOBILI, 1907: 370 (Hao, and Mangareva with a ?). — BOONE, 1935: 28, pl. 5 (Tahiti). — FOREST, 1953b: 556; 1956a: 1063 (Hikueru, Tahiti). — Dardanus deformis - HOLTHUIS, 1953: 47 (Raroia, Tahiti). — MORRISON, 1954: 7 (Raroia). — RAHAYU, 1988: 35 (French Polynesia) - NEW MATERIAL - Coll. PLESSIS (Matureivavao), coll. C. HILY (Tahiti), det. J. POUPIN. Dardanus gemmatus (H. Milne Edwards, 1848) DISTRIBUTION. — Austral (Maria); Marquesas (Ua Huka); Society (Moorea, Tahiti); Tuamotu (Manihi?, Taiaro) - Littoral to sublittoral (10-20m). REFERENCES. — Pagurus gemmatus H. Milne Edwards, 1848: 60 (Marquesas). — STIMPSON, 1858c: 234 [72] (Marquesas; new material ?). — FOREST, 1953b: 557, fig. 10-11; 1956a: 1063 (Tahiti). — Dardanus gemmatus - MONTEFORTE, 1984: 172, annex 1, tab. a; 1987: 8 (Moorea). — RAHAYU, 1988: 36 (Tahiti). — POUPIN, 1994a: 22, fig. 18, pl. 2e (Maria, Tahiti, Taiaro, Ua Huka; 10-20m). — Without name - ? SALVAT, 1986a: 23, photograph (Tahiti). — ? PARDON, 1992: 80, photograph (Manihi) - Det. after the photographs. Dardanus guttatus (Olivier, 1812) DISTRIBUTION. — Tuamotu (Tureia). REFERENCES. — Pagurus guttatus - OWEN, 1839: 82 ("Carysfort" = Tureia). — Dardanus guttatus - HAIG & BALL, 1988: 165 (Syn.). Dardanus lagopodes (Forskal, 1775) DISTRIBUTION. — Society (Maupiti?, Moorea, Tahiti, Tupai); Tuamotu (Hao, Hikueru, Moruroa). 18 REFERENCES. — Pagurus euopsis - NOBILI, 1907: 370 (Hao). — Pagurus sanguinolentus - FOREST, 1953b: 559, fig. 12-14 (Syn.); 1956a: 1064 (Hikueru, Tahiti). — Dardanus lagopodes - LEWINSOHN, 1969: 32 (Syn.). — MONTEFORTE, 1984: 172, annex 1, tab. a; 1987: 8 (Moorea). — RAHAYU, 1988: 32 (Tahiti, Tuamotu). — "Bernard I'hermite bigaré" - ? PARDON, 1992: 20, 21, double page photograph (Maupiti; det. according to the photograph) - NEW MATERIAL - Coll. and det. J. POUPIN (Moruroa, Tahiti, Tupai) - RELEVANT MATERIAL - Dardanus aff. sanguinolentus - SALVAT & RENAUD-MOoRNANT, 1969: 165 (Moruroa) - SYNONYMS - Pagurus euopsis Dana, 1852; P. sanguinolentus Quoy & Gaimard, 1824. Dardanus megistos (Herbst, 1804) DISTRIBUTION. — Society (Tahiti); Tuamotu (Hao, Raroia, Tagatau, Taiaro). REFERENCES. — Pagurus spinimanus Edw. - DANA, 1852b: 452 (with a ?); 1855, pl. 28, fig. Sa-c (Tuamotu). — BOONE, 1935: 34, pl. 7 (Tahiti). — Pagurus punctulatus Olivier - HELLER, 1865: 87 (Tahiti). — NoBILI, 1907: 370 ("Ohura" = Hao, Tagatau). — Pagurus megistos - FOREST, 1953b: 559; 1956a: 1064 (Tahiti). — Dardanus megistos - HOLTHUIS, 1953: 49 (Raroia). — MORRISON, 1954: 7 (Raroia). — RAHAYU, 1988: 33 (French Polynesia). — POUPIN, 1994a: 23, fig. 19, pl. 2g (Tahiti, Taiaro). — Without name - ? SALVAT & RIVES, 1975: 57, full page photograph (French Polynesia; det. according to the photograph) - SYNONYMS - Pagurus punctulatus Olivier, 1811; P. spinimanus H. Milne Edwards, 1848. Dardanus pedunculatus (Herbst, 1804) DISTRIBUTION. — Austral (Tubuai); Tuamotu (Moruroa) - Littoral to sublittoral. REFERENCES. — Dardanus haani - CHEVALIER et al., 1968: 119 (Moruroa). — SALVAT & RENAUD-MORNANT, 1969: 165, 176 (Moruroa). — Dardanus pedunculatus - RAHAYU, 1988: 10, 34 (French Polynesia). — POUPIN, 1994a: 24, fig. 20, pl. 2f (Tubuai; 65m) - SYNONYMS - Dardanus haani (Rathbun, 1902). Dardanus scutellatus (H. Milne Edwards, 1848) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Pagurus scutellatus - FOREST, 1953b: 560; 1956a: 1066 (Tahiti). — Dardanus scutellatus - RAHAYU, 1988: 36 (Tahiti). Diogenes gardineri Alcock, 1905 DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Marutea South). REFERENCES. — Diogenes gardineri - NOBILI, 1907: 366 (Marutea South; "Rikitea" = Mangareva). — SEURAT, 1934: 61 (Mangareva, Marutea South). — FOREST, 1956a: 1056 (French Polynesia); 1957: 530 (NOBILI's material). FAMILY PAGURIDAE Catapaguroides fragilis (Melin, 1939) DISTRIBUTION. — Tuamotu (Moruroa) - Littoral to sublittoral. REFERENCES. — Catapaguroides fragilis - DE SAINT LAURENT, 1968: 940, fig. 26 with hesitations (cf. p. 941, note 1) (Moruroa; 40m). — CHEVALIER et al., 1968: 119, 138 (Moruroa). Micropagurus polynesiensis (Nobili, 1906) DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Moruroa). REFERENCES. — Anapagurus polynesiensis Nobili, 1906a: 260; 1907: 372, pl. 1, fig. 10. ("Rikitea" = Mangareva). — FOREST, 1956a: 1056 (French Polynesia). — CHEVALIER ef al., 1968: 119 (Moruroa). — Micropagurus polynesiensis - HAIG & BALL, 1988: 184 (Syn.). 19 Pagurixus anceps (Forest, 1954) DISTRIBUTION. — Society (Tahiti); Tuamotu (Hikueru). REFERENCES. — Eupagurus anceps Forest, 1954a: 71, fig. 15-19; 1956a: 1067 (Hikueru, Tahiti). — ELDREDGE, 1967: 13 (Hikueru). — Pagurixus anceps - MCLAUGHLIN & HAIG, 1984: 135, fig. 5 (Hikueru; Syn.). Pagurixus laevimanus (Ortmann, 1892) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Eupagurus laevimanus Ortmann, 1892a: 302, pl. 12, fig. 13 (Tahiti). — Pagurixus laevimanus - MCLAUGHLIN & HAIG, 1984: 142, fig. 7 (Syn.). Pagurixus maorus (Nobili, 1906) DISTRIBUTION. — Gambier (Mangareva). REFERENCES. — Eupagurus maorus Nobili, 1906a: 259; 1907: 371, pl. 1, fig. 9 (Mangareva). — FOREST, 1954a: 73; 1956a: 1056 (French Polynesia). — Pagurixus maorus - MCLAUGHLIN & HAIG, 1984: 126, fig. 2 (Syn.). — KOMAI & ASAKURA, 1995: 341, 353 (Key). Trichopagurus trichophthalmus (Forest, 1954) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Catapaguroides trichophthalmus Forest, 1954a: 74, fig. 20-24, with a ? for the genus; 1956a: 1069 (Tahiti). — Trichopagurus trichophthalmus - DE SAINT LAURENT, 1970: 212, fig. 1-16 (Tahiti). FAMILY GALATHEIDAE Coralliogalathea humilis (Nobili, 1905) DISTRIBUTION. — Society (Moorea); Tuamotu (Hao, Tikehau). REFERENCES. — Galathea megalochira Nobili, 1906a: 260; 1907: 376, pl. 1, fig. 12 (Hao). — Coralliogalathea humilis - LEWINSOHN, 1969: 117 (Syn.). — PEYROT-CLAUSADE, 1977a, annex of the species: 24 (Moorea); 1989: 113, 115 (Moorea, Tikehau). — KRopp & BIRKELAND, 1981: 629, tab. 5 (Moorea). — ? Galathea himilis (sic) - PEYROT- CLAUSADE, 1977b: 213 (Moorea). Galathea aculeata Haswell, 1882 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Takapoto). REFERENCES. — Galathea aculeata - ODINETZ, 1983: 208 (Moorea, Tahiti, Takapoto). — ODINETZ-COLLART & RICHER DE ForGES, 1985: 201 (Moorea and/or Tahiti, Takapoto). — MONTEFORTE, 1987: 8 (Moorea). Galathea aff. amamiensis Miyake & Baba, 1966 DISTRIBUTION. — Society (Moorea) - Littoral to sublittoral. REFERENCES. — Galathea aff. amamiensis - PEYROT-CLAUSADE, 1989: 115 (Moorea; 30m). Galathea affinis Ortmann, 1892 DISTRIBUTION. — Gambier (Mangareva); Society (Moorea); Tuamotu (Hao, Marutea South, Tikehau) - Littoral to sublittoral. REFERENCES. — Galathea affinis - NOBILI, 1907: 375, pl. 1, fig. 11 (Marutea South, "Ohura" = Hao, "Rikitea and Gatavake" = Mangareva). — PEYROT-CLAUSADE, 1977a, annex of the species: 24; 1977b: 213; 1985: 462 (Moorea); 1989: 112, 115 (Moorea, Tikehau; 30m). — KRopp & BIRKELAND, 1981: 630, tab. 5 (Moorea). 20 Galathea algae Baba, 1969 DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral. REFERENCES. — Galathea algae - PEYROT-CLAUSADE, 1989: 112, 115 (Moorea, Tikehau; 30m). Galathea latirostris Dana, 1852 DISTRIBUTION. — Society (Raiatea, Tahiti). REFERENCES. — Galathea latirostris - BOONE, 1935: 50, pl. 12 (Raiatea, Tahiti). REMARK. — The status of this species, originally described by DANA (1852b: 480; 1855, pl. 30, fig. 8) from the Fiji is doubtful. The examination of topotypic material, and the revision of all the references attributed to Galathea latirostris is necessary to establish its real identity (K. BABA, personal communication). Galathea latirostris Lenz, 1902, describes form Juan Fernandez must be attributed to Phylladiorhynchus pusillus (Henderson, 1885) (cf. BABA, 1991: 487), and must not be confounded with DANA's species. Phylladiorhynchus serrirostris (Melin, 1939) DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral. REFERENCES. — Galathea serrirostris - PEYROT-CLAUSADE, 1977b: 213 (Moorea). — Phylladiorhynchus serrirostris - PEYROT-CLAUSADE, 1977a, annex of the species: 24 (Moorea); 1989: 112, 115 (Moorea, Tikehau; "...extremely abundant at 30m"). — KRopP & BIRKELAND, 1981: 630, tab. 5 (Moorea). Sadayoshia miyakei Baba, 1969 DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral. REFERENCES. — Sadayoshia miyakei - KROPP & BIRKELAND, 1981: 630, tab. 5 (Moorea). — PEYROT-CLAUSADE, 1989: 112, 115 (Moorea, Tikehau; 30m). FAMILY PORCELLANIDAE Neopetrolisthes maculatus (H. Milne Edwards, 1837) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Neopetrolisthes oshimai (sic) - PARDON, 1992: 81 (Tahiti). — Neopetrolisthes maculatus - HAIG, 1979: 127 (Syn.). REMARK. — According to HAIG (1979), Neopetrolisthes ohshimai Miyake, 1937 is a synonym of this species. This small crustacea is commensal of a sea-anemone. PARDON (1992) illustrates a shrimp Stenopus captured in a sea- anemone. "Neopetrolisthes oshimai" is only indicated in the caption, but is not visible on the photograph. Pachycheles pisoides (Heller, 1865) DISTRIBUTION. — Society; Tuamotu (Tikehau). REFERENCES. — Pachycheles pisoides - HAIG, 1966: 290 (Tuamotu; with the mention that it is the first record for the area, but without details on the material examined); 1983: 284 (Distribution only "Society Islands"). — PEYROT- CLAUSADE, 1989: 113 (Tikehau). Pachycheles sculptus (H. Milne Edwards, 1837) DISTRIBUTION. — Society (Moorea); Tuamotu. REFERENCES. — Pachycheles sculptus - HAIG, 1966: 287 (Tuamotu; same remark than for the previous species); 1983: 284 (Distribution only "Tuamotu"); 1992: 310 (Syn.), — PEYROT-CLAUSADE, 1989: 115 (Moorea). Petrolisthes bispinosus Borradaile, 1900 DISTRIBUTION. — French Polynesia. REFERENCES. — Pefrolisthes bispinosus - HAIG & KRoppP, 1987: 171, 172, fig. 1-2 (French Polynesia, only in summary and discussion). Petrolisthes borradailei Kropp, 1983 DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Huahine, Moorea, Tahiti); Tuamotu (Fakarava, Makemo, Moruroa, Rangiroa, Raroia, Taiaro, Tikehau). REFERENCES. — Petrolisthes rufescens - NOBILI, 1907: 377 ("Rikitea" = Mangareva; cf. Remark under P. rufescens). — PoupPIN, 1994a: 25, fig. 21, pl. 2h (Mangareva, Moruroa, Taiaro) not P. rufescens = P. borradailei fide KROPP personal communication. — Petrolisthes borradailei Kropp, 1983: 96, 106, fig. 3 ("Fakarova" = Fakarava, Huahine, "Makeno" = Makemo, Moorea, Nuku Hiva, Rangiroa, Raroia, "Pascua Pass" = ?, Tahiti, "Tikahua" = Tikehau). Petrolisthes coccineus (Owen, 1839) DISTRIBUTION. — Society (Moorea); Tuamotu. REFERENCES. — Porcellana coccinea Owen, 1839: 87, pl. 26, fig. 1-2 ("Low Islands" = Tuamotu). — Petrolisthes coccineus - PEYROT-CLAUSADE, 1977a, annex of the species: 24; 1977b: 213 (Moorea). — HAIG, 1983: 280; 1992: 313, fig. 9 (Syn.). Petrolisthes decacanthus Ortmann, 1897 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Petrolisthes decacanthus Ortmann, 1897a: 285, pl. 17, fig. 2 (Tahiti). — Haic & KRopp, 1987: 176 (French Polynesia, in the distribution only; Syn.). Petrolisthes eldredgei Haig & Kropp, 1987 DISTRIBUTION. — Society (Tahiti); Tuamotu (Raroia). REFERENCES. — Petrolisthes eldredgei Haig & Kropp, 1987: 180, fig. 5-6 (Tahiti, Raroia). Petrolisthes elegans Haig, 1981 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava). REFERENCES. — Petrolisthes bosci - NoBILI, 1907: 377 ("“Sakarava" = Fakarava) not P. bosci (Audouin, 1826) = P. elegans sp. nov. in HAIG (1981: 266). — Petrolisthes elegans Haig, 1981: 266, fig. 2 (Tahiti, "Sakarava" = Fakarava). — PEYROT-CLAUSADE, 1989: 115 (Moorea). Petrolisthes lamarckii (Leach, 1820) DISTRIBUTION. — Tuamotu (Raraka, Takaroa). REFERENCES. — Porcellana speciosa Dana, 1852b: 417; 1855, pl. 26, fig. 8 (Raraka). — EVANS, 1967: 409 (Raraka; syntypes at the BM). — Petrolisthes lamarckii - HAIG, 1964: 362 (Takaroa); 1992: 315, fig. 11 (Syn.). — KRopp, 1983: 100, 106 (Syn.). Petrolisthes militaris (Heller, 1862) DISTRIBUTION. — Tuamotu (Marutea South). REFERENCES. — Petrolisthes militaris - NOBILI, 1907: 377, with a ? (Marutea South). REMARK. — The presence of this species in French Polynesia is doubtful. NOBILI's determination is uncertain and, moreover, it has never been reported in the area by HAIG (1979: 122; 1982: 280; 1992: 316). 22 Petrolisthes pubescens Stimpson, 1858 DISTRIBUTION. — Marquesas (Nuku Hiva). REFERENCES. — Pefrolisthes pubescens - KRopP, 1986: 456, fig. 2 ("Taiohae, Haka Paa, baie du Controleur" = Nuku Hiva). Petrolisthes rufescens (Heller, 1861) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Porcellana rufescens - HELLER, 1865: 76 (Tahiti). — Not Petrolisthes rufescens - NOBILI, 1907: 377 (cf. Remark). — PouPIN, 1994a: 25, fig. 21, pl. 2h = P. borradailei Kropp, 1983 fide KROpP personal communication. REMARK. — Petrolisthes rufescens and P. borradailei are two very close species. The main difference concerned the posterior border of the cheliped carpus, strongly toothed in P. borradailei, more smooth in P. rufescens. We have confused these two species in a previous work (POUPIN, 1994a) and re-examination of NOBILI's material in Paris (9 ov. 8x7.3, MNHN Ga96) show that it also belongs to P. borradailei. HELLER's reference remains the only record of P. rufescens in the pacific and could also belongs to P. borradailei. Petrolisthes scabriculus (Dana, 1852) DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral. REFERENCES. — Petrolisthes scabriculus - PEYROT-CLAUSADE, 1977a, annex of the species: 25; 1977b: 213 (Moorea); 1989: 112, 115 (Moorea, Tikehau; 30m). — HAIG, 1979: 120 (Syn.). Petrolisthes tomentosus (Dana, 1852) DISTRIBUTION. — Society (Tahiti); Tuamotu (Raraka). REFERENCES. — Porcellana tomentosa Dana, 1852b: 420; 1855, pl. 26, fig. 10 (Raraka). — Petrolisthes tomentosus - Kropp, 1986: 453, fig. 1 (Tahiti, Tahitian neotype at the USNM; Syn.). Porcellana mitra Dana, 1852 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Porcellana mitra - HELLER, 1865: 74, 265, ("Siidsee" and Tahiti; cf. Remark). REMARK. — In HELLER, Tahiti is not mentioned in the main text (p. 74), but only at the end of the work, in the part concerning the geographical distribution (p. 265). It is thus not certain that the location "Tahiti" is correct for this species. Porcellana monilifera Dana, 1852 DISTRIBUTION. — Tuamotu (Raraka?). REFERENCES. — Porcellana monilifera Dana, 1852b: 413; 1855, pl. 26, fig. 3 (Raraka, with a ?). FAMILY ALBUNEIDAE Albunea speciosa Dana, 1852 DISTRIBUTION. — Society (Moorea?). REFERENCES. — Albunea speciosa - NEW MATERIAL - CRIOBE collections, Moorea (without label), 1 d 11x10.5, det. J. POUPIN. REMARK. — This specimen agrees very well with DANA's species, illustrated by SERENE (1973). Formely, Albunea speciosa was considered as endemic from Hawaii. SERENE (1973) has examined specimens from the type locality but mentions that the type material has disappeared. In 1973, THOMASSIN, has described A. madagascariensis, very close to A. speciosa. By the shape of the ocular peduncle and the number of frontal spines, the specimen from Moorea do belongs to DANA's species. FAMILY HIPPIDAE Hippa adactyla Fabricius, 1787 DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti) - Brackish water. REFERENCES. — Remipes testudinarius - ? HELLER, 1865: 72 (Tahiti; reference not found in HAIG, 1970). — DE Man, 1896: 466 ("Nuka-hiwa (Mus. Paris)" = Nuku Hiva). — Hippa adactyla - HOLTHUIS, 1953: 35 (Tahiti). — HAI, 1970: 294 (Syn.); 1974: 179, fig. 2, pl. 6 (Marquesas, distribution only) - NEW MATERIAL - 1 6 28x23.5, coll. C. HILY, det. J. POUPIN ("‘Arue" = Tahiti; confronted with DE MAN's material) - SYNONYMS - Remipes testudinarius Latreille, 1806. Hippa ovalis (A. Milne Edwards, 1863) DISTRIBUTION. — Society (Tahiti) - Brackish water. REFERENCES. — Hippa ovalis - HOLTHUIS, 1953: 35 (Tahiti). — THOMASSIN, 1969: 154, fig. 7b, 8b, 9, pl. 6, fig. 1-8 (Syn.). Hippa pacifica (Dana, 1852) DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti) - Brackish water. REFERENCES. — Remipes pacificus - NOBILI, 1907: 378 (Mangareva). — SEURAT, 1934: 60 (Tahiti). — ? Hippa sp. - CHABOUIS L. & F., 1954: 92, unnumbered fig. (French Polynesia; det. according to the shape of the front margin, on the figure). — Hippa pacifica - HAIG, 1974: 181, fig. 3 (Gambier, distribution only; Syn.). REMARK. — NOBILI's material has been re-examined (MNHN Hi38, Mangareva, 1 2 ov. 22x17, coll. SEURAT 1905, det. NOBILI 1906) and is similar to other specimens attributed by DE MAN to this species (MNHN Hi107, Java sea; MNHN Hi108, Atjeh). INFRA-ORDER BRACHYURA FAMILY DROMIIDAE Cryptodromia coronata Stimpson, 1859 DISTRIBUTION. — Gambier (Mangareva). REFERENCES. — Cryptodromia coronata - NoBILI, 1907: 378 ("Rikitea" = Mangareva). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 448 (List). — Cryptodromia ? coronata - MCLAY, 1993: 199 (Syn.; with the indication that the status of the species is uncertain). Cryptodromia fallax (Lamarck, 1818) DISTRIBUTION. — Tuamotu (Raroia). REFERENCES. — Cryptodromia canaliculata - HOLTHUIS, 1953: 3 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 448 (List). — Cryptodromia fallax - McLay, 1993: 206, fig. 18e (Syn.) - SYNONYMS - Cryptodromia canaliculata Stimpson, 1858. 24 Cryptodromiopsis tridens Borradaile, 1903 DISTRIBUTION. — Society (Moorea, Tahiti). REFERENCES. — Cryptodromiopsis tridens - MCLAY, 1991: 467, fig. 5a-d (Moorea, Tahiti). Dromia wilsoni (Fulton & Grant, 1902) DISTRIBUTION. — Austral (Raevavae); Marquesas (Tahuata); Tuamotu (Makemo, Takapoto) - Sublittoral to deep. REFERENCES. — Petalomera wilsoni - MCLAY, 1991: 470, fig. 6a-d, 7a-c, 8a-c (Makemo, Raevavae, Tahuata, Takapoto; 190-350m). — Dromia wilsoni - MCLAY, 1993: 156, fig. 16e (Syn.). — POUPIN, 1996: in press (same material than MCLAY). REMARK. — Dromia wilsoni is mainly a sublittoral species, usually found within the first 100m, but it is also recorded up to 520m (cf. in McLay, 1991: 475). FAMILY DYNOMENIDAE Dynomene hispida Desmaret, 1825 DISTRIBUTION. — Society (Moorea); Tuamotu (Marutea South, Tikehau). REFERENCES. — Dynomene hispida - NoBILI, 1907: 378 (“Marutea-Vaitutaki" = Marutea South). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 25; 1977b: 212 (Moorea); 1989: 111, 114 (Moorea, Tikehau). — GUINOT, 1985: 448 (List). Dynomene praedator A. Milne Edwards, 1879 DISTRIBUTION. — Society (Moorea, Tahiti). REFERENCES. — Dynomene praedator - ORTMANN, 1892b: 534, pl. 26, fig. 3 (Tahiti). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 448 (List). — Dynomene sp. - NAIM, 1980a: 55, fide observation and personal communication of MCLAY (Moorea, MNHN B20203). — Dynomene sinense - ODINETZ, 1983: 208 (Tahiti; MNHN B17090). — Dynomene sinensis (sic) - GUINOT, 1985: 448 (List; after ODINETZ) - These two references, not D. sinense Chen, 1979 = D. praedator fide observation and personal communication of MCLAY. Dynomene spinosa Rathbun, 1911 DISTRIBUTION. — Marquesas; Tuamotu (Raroia). REFERENCES. — Dynomene spinosa - BALSS, 1935: 115 (Marquesas). — HOLTHUIS, 1953: 3 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"; Marquesas). — GUINOT, 1985: 448 (List). FAMILY RANINIDAE Notosceles chimmonis Bourne, 1922 DISTRIBUTION. — Marquesas (Eiao) - Sublittoral to deep. REFERENCES. — Notosceles chimmonis - POUPIN, 1996: in press (Eiao; 54-101m). REMARK. — Species known between 45-52 m (SERENE & UMALI, 1972), 75-90m (RIBES, 1989), and up to 450m (Monop, 1975). Ranina ranina (Linné, 1758) DISTRIBUTION. — Marquesas (Eiao, Fatu Hiva, Nuku Hiva) - Sublittoral. 25 REFERENCES. — Ranina ranina - GUINOT, 1985: 449 (List; certainly after the following dry specimen: MNHN n°223 "Nouhiva" = Nuku Hiva) - NEW MATERIAL - Coll. and det. J. POUPIN (Eiao, Fatu Hiva; juveniles, 100m). FAMILY CALAPPIDAE Ashtoret lunaris (Forskal, 1775) DISTRIBUTION. — Marquesas (Nuku Hiva). REFERENCES. — Matuta banksii - RATHBUN, 1907: 68 (Nuku Hiva). — GUINOT, 1985: 453 (List). — Ashtoret lunaris - GALIL & CLARK, 1994: 5, fig. 1a-b, pl. la-b (Syn.) - SYNONYMS - Matuta banksi Leach, 1817. Ashtoret picta (Hess, 1865) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Matuta picta - POUPIN, 1994a: 27, fig. 23, pl. 3b (Tahiti). — Ashtoret picta - GALIL & CLARK, 1994: 18, fig. 3c-d, pl. 6a-b (Tahiti). REMARK. — In the revision of the genus Matuta by GALIL & CLARK (1994), the location "Tahiti", that should appear under Ashtoret picta, is erroneously mentioned under Ashtoret granulosa (Miers, 1877), species still unknown from French Polynesia (GALIL, personal communication). Calappa calappa (Linné, 1758) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Calappa calappa - MONTEFORTE, 1984: 173, annex 1, tab. a (Tahiti; MONTEFORTE's material verified in the CRIOBE collections, Moorea). — GUINOT, 1985: 449 (List). Calappa hepatica (Linné, 1758) DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Hao, Hikueru, Marutea North?, Marutea South, Mataiva, Moruroa, Takapoto). REFERENCES. — Calappa hepatica - NoOBILI, 1907: 378 ("Gatavake" = Mangareva, Hao, Marutea = Marutea North?, Marutea South). — RATHBUN, 1907: 67 (Bora Bora). — PESTA, 1913: 37 (Tahiti). — BOONE, 1934: 32, pl. 8-10 (Nuku Hiva). — SEURAT, 1934: 59, 60 (Hao, Marutea South). — CHABOUIS L. & F., 1954: 92, unnumbered fig. (French Polynesia). — FOREST & GUINOT, 1961: 11, fig. 1a-b, 2 (Hikueru; Syn.); 1962: 56 (Biogeography "Tahiti-Tuamotu; Marquesas"). — MONTEFORTE, 1984: 173, annex 1, tab. a, photograph p. 140c (Mataiva, Moorea, Tahiti, Takapoto); 1987: 8 (Moorea). — DELESALLE, 1985: 288 (Mataiva). — GUINOT, 1985: 449 (List). — POUPIN, 1994a: 26, fig. 22, pl. 3a (Hikueru, Moruroa, Tahiti). — Calappa tuberculata - HELLER, 1865: 69 (Tahiti) - SYNONYMS - Calappa tuberculata Fabricius, 1798. Matuta victor (Fabricius, 1781) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Matuta victor - HELLER, 1865: 69 (Tahiti). — GALIL & CLARK, 1994: 39, fig. 7a-b, pl. 13a-b (cf. Remark). REMARK. — HELLER's reference is not mentioned in the revision by GALIL & CLARK (1994). These authors consider that the eastern distribution of this species is limited to the Fiji. Thus, the revision of HELLER's material would be important to confirm the presence of Matuta victor in French Polynesia. 26 FAMILY LEUCOSIIDAE Ebaliopsis erosa (A. Milne Edwards, 1874) DISTRIBUTION. — Gambier; Tuamotu (Marutea South). REFERENCES. — Ebalia erosa - NOBILI, 1907: 378 (Marutea). — Ebaliopsis erosa - FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — SERENE, 1977: 55, fig. 5-6 (Gambier). — GUINOT, 1985: 453 (List). Heteronucia venusta Nobili, 1906 DISTRIBUTION. — Society (Moorea); Tuamotu (Fakahina, Fakarava, Hao, Tikehau). REFERENCES. — Heteronucia venusta Nobili, 1906a: 260; 1907: 379, pl. 5, fig. 14 ("Ohura" = Hao). — FOREST & GUuINOT, 1961: 13, fig. 3a-b, 4, pl. 5, fig. 1-2 (Fakahina; Syn.); 1962: 56 (Biogeography "Tahiti-Tuamotu"). — PEYROT- CLAUSADE, 1977a, annex of the species: 25; 1977b: 212 (Moorea); 1989: 113 (Tikehau). — GUINOT, 1985: 453 (List). — Nucia gelida Rathbun, 1907: 68, pl. 5, fig. 4, pl. 9, fig. 2 (Fakarava). — GUINOT, 1985: 453 (List). Nucia rosea Nobili, 1906 DISTRIBUTION. — Gambier (Mangareva). REFERENCES. — Nucia rosea Nobili, 1906a: 261; 1907: 381 (Mangareva). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). Nursia mimetica Nobili, 1906 DISTRIBUTION. — Gambier (Mangareva). REFERENCES. — Nursia mimetica Nobili, 1906a: 261; 1907: 380, pl. 5, fig. 13 (Mangareva). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). FAMILY MAJIDAE Acanthophrys cristimanus A. Milne Edwards, 1865 DISTRIBUTION. — Marquesas (Nuku Hiva). REFERENCES. — Acanthophrys cristimanus A. Milne Edwards, 1865a: 141, pl. 5, fig. 3 (Nuku Hiva). — GRIFFIN & TRANTER, 1986: 105 (cf. Remark). REMARK. — GRIFFIN & TRANTER (1986) have re-established the validity of the genus Acanthophrys, formerly transferred in Hyastenus, and Acanthophrys cristimanus is the type species of the genus. Camposcia retusa Latreille, 1829 DISTRIBUTION. — Tuamotu (Fangataufa) - Littoral to deep? REFERENCES. — Camposcia retusa - GUINOT, 1985: 452, with a ? (List) - NEW MATERIAL - Coll. and det. J. POUPIN (Fangataufa; 220m, cf. Remark). REMARK. — The presence of this species in French Polynesia, only inferred by GUINOT because of its large distribution, is here confirmed by one specimen, collected at an unusual depth. Cyclax suborbicularis (Stimpson, 1858) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Cyclax suborbicularis - FOREST & GUINOT, 1961: 15, fig. 5-6, 8 bis, 10, pl. 6, fig. 1-2 (Tahiti); 1962: 56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List). 27 Huenia proteus de Haan, 1839 DISTRIBUTION. — Society (Moorea). REFERENCES. — Huenia proteus - KROpP & BIRKELAND, 1981: 630, tab. 5 (Moorea). — GUINOT, 1985: 452 (List). Hyastenus aff. borradailei (Rathbun, 1907) DISTRIBUTION. — Society (Moorea). REFERENCES. — Hyastenus aff. borradaeilli (sic) - PEYROT-CLAUSADE, 1989: 115 (Moorea). Menaethius monoceros (Latreille, 1825) DISTRIBUTION. — Marquesas; Society (Moorea, Tahiti); Tuamotu (Fakarava, Hao, Raroia, Takapoto, Tikehau) - Littoral to sublittoral (30m). REFERENCES. — Menaethius tuberculatus Dana, 1852b: 123; 1855, pl. 5, fig. 1a-c (Tuamotu). — Menaethius monoceros - NOBILI, 1907: 382 ("Ohura" = Hao). — RATHBUN, 1907: 64 (Fakarava). — HOLTHUIS, 1953: 4 (Raroia). — ForEST & GUINOT, 1961: 14, fig. 9a-b (Tahiti; Syn.); 1962: 56 (Biogeography "Tahiti-Tuamotu, Marquesas"). — PEYROT-CLAUSADE, 1977a, annex of the species: 25 (Moorea); 1977b: 212; 1989: 112, 115 (Moorea, Tikehau; 30m). — ODINETZ, 1983: 208 (Moorea, Tahiti, Takapoto). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti, Takapoto). Micippa margaritifera Henderson, 1893 DISTRIBUTION. — Tuamotu (Tikehau). REFERENCES. — Miccipa (sic) margaritifera - PEYROT-CLAUSADE, 1989: 112 (Tikehau). Micippa parca Alcock, 1895 DISTRIBUTION. — Tuamotu (Makemo). REFERENCES. — Lophomicippa limbata Rathbun, 1907: 65, pl. 5, fig. 3, pl. 6, fig. 1, 1g (Makemo). — FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List). — Micippa parca - GRIFFIN & TRANTER, 1986: 277 (Syn.). Micippoides angustifrons A. Milne Edwards, 1873 DISTRIBUTION. — Tuamotu (Raroia). REFERENCES. — Micippoides angustifrons - HOLTHUIS, 1953: 5 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List). Perinea tumida Dana, 1851 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava, Takapoto, Tikehau). REFERENCES. — Perinea tumida - RATHBUN, 1907: 65 (Fakarava). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 25; 1977b: 212 (Moorea); 1989: 112, 115 (Moorea, Tikehau). — KROPP & BIRKELAND, 1981: 630, tab. 5 (Moorea, Takapoto). — ODINETZ, 1983: 208 (Moorea, Tahiti, Takapoto). — GUINOT, 1985: 453 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti, Takapoto). Schizophrys aspera (H. Milne Edwards, 1834) DISTRIBUTION. — French Polynesia. REFERENCES. — Schizophrys aspera - GUINOT, 1985: 453, with a ? (List). REMARK. — Cited by GUINOT, only because of the large distribution of the species (see SAKAI, 1976, or DAI & YANG, 1991: Japon, Hawaii, ... Australia). 28 Simocarcinus obtusirostris (Miers, 1879) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Trigonothir obtusirostris - FOREST & GUINOT, 1961: 14 (Tahiti); 1962: 56 (Biogeography "Tahiti- Tuamotu"). — Simocarcinus obtusirostris - GUINOT, 1985: 453 (List). — GRIFFIN & TRANTER, 1986: 98 (Syn.). Tylocarcinus dumerilii (H. Milne Edwards, 1834) DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Hao). REFERENCES. — Tylocarcinus gracilis - NOBILI, 1907: 382 (Hao, Mangareva). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). — Tylocarcinus dumerilii - GRIFFIN & TRANTER, 1986: 197, fig. 67a-b (Syn.) - SYNONYMS - Tylocarcinus gracilis Miers, 1879. FAMILY PARTHENOPIDAE Actaeomorpha alvae Boone, 1934 DISTRIBUTION. — Society (Raiatea). REFERENCES. — Acfaeomorpha alvae Boone, 1934: 37 pl. 11 (Raiatea). — FOREST & GUINOT, 1962: 56 (Biogeography "Tahiti-Tuamotu”). — GUINOT, 1966b: 759 (cf. Remark); 1985: 453 (List). REMARK. — According to the drawing published by BOONE (1934), GUINOT (1966b) considers that this species could possibly be, either Actaemorpha erosa Miers, 1878, or A. punctata Edmonson, 1935. We keep here the genus Actaeomorpha in the Parthenopidae, but GUINOT (1966b, 1967), in her study of the genera Aethra (cf. hereafter A. scruposa), Osachila, Hepatus, Hepatella and Actaeomorpha, has modified this usual classification and placed these genera in a group /ncertae sedis Parthenoxystomata (cf. GUINOT's, 1985 List). Aethra scruposa (Linné, 1764) DISTRIBUTION. — Society. REFERENCES. — Aethra scruposa - GUINOT, 1985: 453, with a ? (Society; listed only according to the large distribution of the species, without material from French Polynesia). Daldorfia horrida (Linné, 1758) DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti); Tuamotu (Fangatau, Hao, Mataiva). REFERENCES. — Parthenope horrida - NOBILI, 1907: 382 ("Fagatau" = Fangatau, Hao, Mangareva). — SEURAT, 1934: 60 ("Fagatau" = Fangatau, Gambier, Hao). — CHABOUIS L. & F., 1954: 91, fig. 7 (Tahiti, Tuamotu). —FOREST & GUINOT, 1961: 26, fig. 14 (Tahiti); 1962: 58 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 174, annex 1, tab. a (Mataiva). — GUINOT, 1985: 453 (List). — Daldorfia horrida - SAKAI, 1976: 283, pl. 96, fig. 2, text-fig. 157 (Syn.). Parthenope contrarius (Herbst, 1796) DISTRIBUTION. — Marquesas (Eiao) - Sublittoral. REFERENCES. — Parthenope contrarius - NEW MATERIAL - Coll. and det. J. POUPIN (Eiao; 42m). REMARK. — This new material (1 ? and 3 juveniles, Marara st. D38) agrees very well with the description and the good photograph published by RATBHUN (1906: 885, pl. 17, fig. 1), under Parthenope (Rhinolambrus) lamelligera (White, 1847). According to SAKAI (1976: 273) WHITE's species is a synonym of P. (Rhinolambrus) pelagicus (Riippell, 1830), but RATHBUN's material belongs to P. contrarius. Parthenope hoplonotus (Adams & White, 1848) DISTRIBUTION. — Society (Tahiti). Wg) REFERENCES. — Aulacolambrus hoplonotus - FOREST & GUINOT, 1961: 26, fig. 12a-c, 13 (Tahiti); 1962: 58 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). — Parthenope (Aulacolambrus) hoplonotus - SAKAI, 1976: 280 (Syn.) FAMILY EUMEDONIDAE Echinoecus pentagonus (A. Milne Edwards, 1879) DISTRIBUTION. — Tuamotu (Hao, Raroia). REFERENCES. — Eumedon convictor Bouvier & Seurat, 1905: 629 (Hao). — NoBILI, 1907: 382 (Hao). — SEURAT, 1934: 58 (Hao). — Eumedonus convictor - HOLTHUIS, 1953: 6 (Raroia). — MORRISON, 1954: 6 (Raroia) — Echinoecus pentagonus - SERENE et al., 1958: 152 (Syn.). — GUINOT, 1985: 453 (List). — Eumedonus pentagonus - FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). FAMILY PORTUNIDAE SUBFAMILY CATOPTRINAE Carupa tenuipes Dana, 1852 DISTRIBUTION. — Gambier (Akamaru); Society (Huahine, Maiao?, Moorea, Tahiti); Tuamotu (Makatea, Makemo, Marutea South, Pukapuka, Raroia, Tikehau). REFERENCES. — Carupa tenuipes Dana, 1852a: 85 (Tuamotu); 1852b: 279; 1855, pl.17, fig. 4a-e ("Paumotu archipelago?”). — A. MILNE EDWARDS, 1861: 386 (Tuamotu; DANA's material). — STEPHENSON & REES, 1967: 5 (Huahine, Moorea, Maiai = Maiao?, "Tickahau" = Tikehau). — SAKAI, 1976: 325 (Syn.). — STEPHENSON, 1976: 12 (Pukapuka). — MONTEFORTE, 1984: 173, annex 1, tab. a (Makatea, Moorea); 1987: 8 (Moorea). — GUINOT, 1985: 449 (List). — Carupa laeviuscula Heller, 1862: 520 (Tarti" = Tahiti); 1865: 27, pl.3, fig. 2 (Tahiti). — NoBILI, 1907: 386 (Akamaru, "Marutea-Vaitutaki" = Marutea South). — RATHBUN, 1907: 64 (Makemo). — HOLTHUIS, 1953: 9 (Raroia). — Morrison, 1954: 13 (Raroia). — FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu") - NEW MATERIAL - coll. C. HiLy, det. K. MoosA (Tikehau). Catoptrus nitidus A. Milne Edwards, 1870 DISTRIBUTION. — Marquesas (Eiao); Society (Huahine); Tuamotu (Makemo, Tikehau) - Sublittoral. REFERENCES. — Catoptrus nitidus - RATHBUN, 1907: 60 (Makemo). — FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 29 (Syn.). — GUINOT, 1985: 449 (List). — Libystes truncatifrons - STEPHENSON & REES, 1967: 6 ("Maroe" = Huahine, "Tickahau" = Tikehau) - NEW MATERIAL - Coll. J. POUPIN, det. K. Moosa (Eiao; 42m) - SYNONYMS - Libystes truncatifrons (de Man, 1887). SUBFAMILY CAPHYRINAE Caphyra rotundifrons (A. Milne Edwards, 1869) DISTRIBUTION. — Society (Bora Bora, Tahiti). REFERENCES. — Caphyra rotundifrons - RATHBUN, 1907: 60, pl. 1, fig. 4 (Tahiti). — ForREsT & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu”). — STEPHENSON & REES, 1967: 7 (Bora Bora, "Mata Uta Papeete" = Tahiti). — GUINOT, 1985: 449 (List). 30 Caphyra tridens Richters, 1880 DISTRIBUTION. — Gambier (Mangareva). REFERENCES. — Caphyra rotundifrons var. tridens Richters - NOBILI, 1907: 386 (""Rikitea, Teone Kura" = Mangareva). — Caphyra tridens - CROSNIER, 1975: 747, fig. 3a-n (Mangareva). Lissocarcinus elegans Boone, 1934 DISTRIBUTION. — Society (Raiatea). REFERENCES. — Lissocarcinus elegans Boone, 1934: 50, pl. 16 (Raiatea). — FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 27 (Distribution; French Polynesia only). — GUINOT, 1985: 449 (List). Lissocarcinus laevis Miers, 1886 DISTRIBUTION. — Marquesas (Eiao, Hiva Oa, Nuku Hiva) - Sublittoral. REFERENCES. — Lissocarcinus laevis - STEPHENSON, 1976: 12 (Nuku Hiva) - NEW MATERIAL - Coll. J. POUPIN, det. K. Moosa (Eiao, Hiva Oa; 42-53m). Lissocarcinus orbicularis Dana, 1852 DISTRIBUTION. — Society (Huahine, Moorea, Raiatea, Tahiti); Tuamotu (Hikueru, Moruroa, Tikehau). REFERENCES. — Lissocarcinus orbicularis - FOREST & GUINOT, 1961: 27, fig. 15a-b, 16a-c (Hikueru); 1962: 58 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON & REES, 1967: 7 (Huahine, Raiatea, "Tikahau" = Tikehau). — STEPHENSON, 1976: 12 (Moorea). — CHEVALIER et al., 1968: 112, 137 (Moruroa). — MONTEFORTE, 1984: 173, annex 1, tab. a (Tahiti). — GUINOT, 1985: 449 (List). — PEYROT-CLAUSADE, 1989: 113 (Tikehau). SUBFAMILY PORTUNINAE Charybdis annulata (Fabricius, 1798) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Goniosoma annulatum (Fabricius) - ORTMANN, 1893a: 82 (Tahiti). — Charybdis annulata - FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). — Charybdis (Charybdis) annulata - CROSNIER, 1962: 78, fig. 136-139, pl. 5, fig. 2 (Distribution only, Tahiti). — GUINOT, 1985: 449, with a ? (List). Charybdis erythrodactyla (Lamarck, 1818) DISTRIBUTION. — Austral (Rurutu); Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fangataufa, Hikueru, Makatea, Mataiva, Moruroa, Raroia, Taiaro, Takapoto). REFERENCES. — Goniosoma erythrodactylum - A. MILNE EDWARDS, 1861: 369 (Marquesas). — DE MAN, 1889: 424 (Tahiti). — ORTMANN, 1893a: 81 (Marquesas). — Charybdis erythrodactyla - NOBILI, 1906b: 118, fig. 3 (Nuku Hiva). — SENDLER, 1923: 40 (Makatea). — BOONE, 1934: 57, pl. 18-19 (Tahiti). — LEENE, 1936: 117, fig. 1-5 (Marquesas, Makatea). — HOLTHUIS, 1953: 6 (Raroia). — MORRISON, 1954: 16 (Raroia). — FOREST & GUINOT, 1961: 30 (Hikueru); 1962: 58 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1966a: 48 (Raroia). — CHEVALIER et al., 1968: 92, 137 (Fangataufa). — MONTEFORTE, 1984: 173, annex 1, tab. a, photograph p. 140(b) (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — SALVAT, 1986b: 72, photograph (French Polynesia). — Charybdis (Goniosupradens) erythrodactyla - LEENE, 1938: 134, fig. 77-80 (Marquesas). — STEPHENSON & REES, 1967: 13 (Bora Bora, Moorea). — STEPHENSON, 1976: 15 (""Taiohae" = Nuku Hiva). — GUINOT, 1985: 449 (List). — POUPIN 1994a: 29, fig. 25, pl. 3d (Rurutu, Taiaro) - NEW MATERIAL - Coll. and det. J. POUPIN (Moruroa). Charybdis hawaiensis Edmonson, 1954 DISTRIBUTION. — Tuamotu (Taiaro). eee NNER aN NTE lala viens Mr Sil REFERENCES. — Charybdis (Charybdis) hawaiensis - POUPIN, 1994a: 28, fig. 24, pl. 3c (Taiaro). Charybdis orientalis Dana, 1852 DISTRIBUTION. — Society. REFERENCES. — Charybdis (Charybdis) orientalis - LEENE, 1938: 69 (Society; cf. Remark). REMARK. — Although LEENE (1938) does not mention clearly the location "Society", she indicates that F. CHACE has examined, for her, in the USNM collections, a male from the Society Islands attributed to C. orientalis (confronted with the type specimen). Charybdis paucidentata A. Milne Edwards, 1861 DISTRIBUTION. — Marquesas (Hiva Oa, Tahuata); Tuamotu (Taiaro) - Littoral to sublittoral. REFERENCES. — Charybdis (Gonioinfradens) paucidentata - POUPIN, 1994a: 30, fig. 26, pl. 3e (Hiva Oa, Tahuata, Taiaro; 0-100m); 1996: in press (Hiva Oa, Tahuata). Lupocyclus quinquedentatus Rathbun, 1906 DISTRIBUTION. — Austral (Maria, Rurutu); Marquesas (Nuku Hiva); Society (Bora Bora) - Sublittoral to deep. REFERENCES. — Lupocyclus quinquedentatus - STEPHENSON, 1976: 15 ("“Hatwata" = Haatuatua bay, Nuku Hiva) - NEw MATERIAL - Coll. J. POUPIN, det. K. MoosA (Bora Bora, Maria, Rurutu; 80-110m). Portunus alexandri (Rathbun, 1907) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Callinectes alexandri Rathbun, 1907: 61, pl. 2, fig. 1, pl. 9, fig. 3, 3a-b (Tahiti). — GUINOT, 1985: 449 (List). REMARK. — This portunid has been described from Tahiti, with a paratype from "Suva, Fidjis". According to STEPHENSON (1976: 13), it is in fact a non-identifiable Portunus. Portunus dubius (Laurie, 1906) DISTRIBUTION. — Marquesas (Eiao, Fatu Hiva, Hiva Oa, Nuku Hiva, Tahuata) - Sublittoral to deep. REFERENCES. — Portunus dubius - STEPHENSON, 1976: 16 (Marquesas; numerous stations without precisions) - NEW MATERIAL - Coll. J. POUPIN, det. K. MOoSA (Eiao, Fatu Hiva, Hiva Oa, Nuku Hiva, Tahuata; 42-140m). Portunus granulatus (H. Milne-Edwards, 1834) DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Raroia, Tikehau). REFERENCES. — Neptunus (Achelous) granulatus - ORTMANN, 1893a: 72 (Tahiti). — NOBILI, 1907: 383 ("Rikitea" = Mangareva). — BOONE, 1934: 60, pl. 20 (Nuku Hiva). — SEURAT, 1934: 59 (Mangareva). — Portunus (Achelous) granulatus - RATHBUN, 1907: 60 (Bora Bora, Fakarava). — SENDLER, 1923: 40 (Tahiti). — Portunus (Cycloachelous) granulatus - HOLTHUIS, 1953: 6 (Raroia). — MORRISON, 1954: 7 (Raroia). — GUINOT, 1985: 449 (List). — Portunus granulatus - FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu, Marquesas"). — STEPHENSON & REES, 1967: 25 (Moorea, Tuamotu). — TURKAY, 1971: 127 (Tahiti). — STEPHENSON, 1976: 16 (Tahiti). — MONTEFORTE, 1984: 173, annex 1, tab. a (Moorea, Tahiti); 1987: 8 (Moorea) - NEW MATERIAL - Coll. C. HILY, det. K. Moosa (Tahiti, Tikehau). Portunus guinotae Stephenson & Rees, 1961 DISTRIBUTION. — Tuamotu (Marutea South). REFERENCES. — Portunus guinotae Stephenson & Rees, 1961: 425, fig. 1b, d, g, 2d-f (Marutea South). — FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 39 (Marutea South). — Portunus (Xiphonectes) guinotae - GUINOT, 1985: 449 (List). 32 Portunus iranjae Crosnier, 1962 DISTRIBUTION. — Marquesas (Eiao, Fatu Hiva, Hiva Oa, Tahuata); Society (Moorea) - Littoral to sublittoral. REFERENCES. — Portunus iranjae - STEPHENSON & REES, 1967: 30 (""Papetoai bay" = Moorea). — STEPHENSON, 1976: 16 ("Marquesas expedition, st. THX" = Haava strait, between Tahuata and Hiva Oa, cf. HARALD, 1967). — Portunus (Xiphonectes) iranjae - GUINOT, 1985: 449 (List) - NEW MATERIAL - Coll. J. POUPIN, det. K. MOOSA (Eiao, Fatu Hiva, Hiva Oa, Tahuata; 54m). Portunus longispinosus (Dana, 1852) DISTRIBUTION. — Marquesas (Hiva Oa, Tahuata); Tuamotu (Marutea South, Raroia). REFERENCES. — Neptunus (Hellenus) longispinosus - NoBILI, 1907: 383 (Marutea South). — Portunus (Hellenus) longispinosus - HOLTHUIS, 1953: 7 (Raroia). — MORRISON, 1954: 8 (Raroia). — Portunus longispinosus - FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1976: 16 ("Marquesas expedition st. THX, haul 5" = Haava strait, between Tahuata and Hiva Oa, cf. HARALD, 1967). — Portunus (Xiphonectes) longispinosus - GUINOT, 1985: 449 (List). REMARK. — This species can be confused with all the species belonging to the Jongispinosus complex, as Portunus iranjae and P. macrophthalmus (cf. STEPHENSON & REES, 1967; STEPHENSON, 1976; and NAGAI, 1981). Portunus macrophthalmus Rathbun, 1906 DISTRIBUTION. — Marquesas (Eiao, Hiva Oa) - Sublittoral. REFERENCES. — Portunus macrophthalmus - NEW MATERIAL - Coll. J. POUPIN, det. K. Moos (Eiao, Hiva Oa; 42- 53m). Portunus nipponensis Sakai, 1938 DISTRIBUTION. — Tuamotu (Moruroa) - Sublittoral to deep. REFERENCES. — Portunus nipponensis - POUPIN et al. 1990: 17 (French Polynesia). — POUPIN, 1996: in press pro parte (Moruroa; cf. Remark). REMARK. — This species is usually found in shallow waters (15-50m) but, in French Polynesia, it has been trapped up to 130m. Except for Moruroa, the localities mentioned in POUPIN (1996) concerned in fact a new species, related to P. nipponensis, but with a distinct male pleopod (Moosa & CROSNIER, in study). Portunus orbitosinus Rathbun, 1911 DISTRIBUTION. — Marquesas (Eiao) - Sublittoral. REFERENCES. — Portunus orbitosinus - NEW MATERIAL - Coll. J. POUPIN, det. K. MoosA (Eiao; 42m). Portunus pelagicus (Linné, 1758) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Neptunus pelagicus - HELLER, 1865: 27 (Tahiti). — Portunus pelagicus - FOREST & GUINOT, 1962: 58 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 41 (Distribution only, Tahiti). — Portunus (Portunus) pelagicus - GUINOT, 1985: 449 (List). REMARK. — Although this species has often been reported from Tahiti (BOONE, 1934; STEPHENSON & CAMPBELL, 1959; CROSNIER, 1962; STEPHENSON & REES, 1967; STEPHENSON, 1972; SAKAI, 1976; DAI & YANG, 1991), it seems that the single material examined from this locality is mentioned in HELLER (1865). It would thus be very interesting to check this reference. Portunus sanguinolentus (Herbst, 1783) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Mataiva). REFERENCES. — Neptunus sanguinolentus - CHABOUIS L. & F., 1954: 91, fig. 9 (French Polynesia). — Portunus sanguinolentus - FOREST & GUINOT, 1961: 29, fig. 17a-b, 18 (Tahiti); 1962: 58 (Biogeography "Tahiti-Tuamotu"). — a RN OSES RPL TORO SU SSE ee 33 MONTEFORTE, 1984: 173, annex 1, tab. a (Moorea). — SALVAT, 1986b: 72 (French Polynesia). — POUPIN, 1994a: 31, fig. 27, pl. 3f (Mataiva, Tahiti). — Portunus sanguinolentus sanguinolentus - STEPHENSON, 1976: 19 (Tahiti). — Portunus (Portunus) sanguinolentus - GUINOT, 1985: 449 (List). Scylla serrata (Forskal, 1775) DISTRIBUTION. — Society (Huahine, Raiatea, Tahiti, Tupai) - Brackish to sea water. REFERENCES. — Scylla serrata - HELLER, 1865: 27 (Tahiti). — MIERS, 1886: 185 (Tahiti). — BOONE, 1934: 68, pl. 25- 30 (Huahine, Tahiti). — SEURAT, 1934: 58 (Tahiti). — CHABOUIS L. & F., 1954: 90, unnumbered fig. (Huahine, Raiatea). — FOREST & GUINOT, 1961: 27 (Tahiti); 1962: 58 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (Society); 1985: 449 (List). — BABLET, 1972: 32, pl. 11 (French Polynesia). — SALVAT, 1986b: 70, 72 (French Polynesia) - NEW MATERIAL - Coll. J. POUPIN, det. K. MoosA (Raiatea, Tupai). Thalamita admete (Herbst, 1803) DISTRIBUTION. — Gambier (Mangareva); Marquesas?; Society (Bora Bora, Huahine, Moorea, Raiatea, Tahiti); Tuamotu (Fakarava, Hikueru, Kaukura, Makemo, Mataiva, Takapoto, Tikehau). REFERENCES. — Thalamita admete - HELLER, 1865: 28 (Tahiti). — ORTMANN, 1893a: 83 (Tahiti). — NOBILI, 1907: 383 (Kaukura, Mangareva). — RATHBUN, 1907: 63 (Fakarava, Makemo). — FOREST & GUINOT, 1961: 30, fig. 19a-b (Hikueru, Tahiti); 1962: 58 (Biogeography "Tahiti-Tuamotu, Marquesas" with a ?). — STEPHENSON & REES, 1967: 56, fig. 20 (Bora Bora, Huahine, Moorea, Raiatea, Tahiti, "Tikahau" = Tikehau). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — MONTEFORTE, 1984: 173, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 8 (Moorea). — DELESALLE, 1985: 288 (Mataiva). — Thalamita (Thalamita) admete - GUINOT, 1985: 449 (List). Thalamita bouvieri Nobili, 1906 DISTRIBUTION. — Gambier (Mangareva). REFERENCES. — Thalamita bouvieri Nobili, 1906a: 262; 1907: 384, pl. 2, fig. 2 ("Rikitea" = Mangareva). — SEURAT, 1934: 60 (French Polynesia). — CROSNIER, 1962: 119, fig. 201-204, pl. 10, fig. 2 (Mangareva; syntypes). — FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — Thalamita (Pseudothalamitopsis) bouvieri - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 47) - RELEVANT MATERIAL - Thalamitoides (sic) aff. bouvieri - PEYROT- CLAUSADE, 1989: 113 (Tikehau). Thalamita chaptalti (Audouin, 1826) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Thalamita chaptalii - FOREST & GUINOT, 1961: 34, fig. 21a-b (Tahiti); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — Thalamita (Neothalamita) chaptalii - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 43). Thalamita coerulipes Jacquinot, 1852 DISTRIBUTION. — Gambier (Kamaka, Mangareva); Society (Bora Bora, Huahine, Tahiti); Tuamotu (Fakarava). REFERENCES. — Thalamita coerulipes Jacquinot, 1852, pl. 5, fig. 6-10 (Mangareva). — JACQUINOT & LUCAs, 1853: 53 (Mangareva). — A. MILNE EDWARDS, 1861: 363 (Mangareva). — NOBILL, 1907: 383 (Kamaka) — RATHBUN, 1907: 63 (Fakarava, Society). — BOONE, 1934: 78, pl. 35 (Bora Bora). — FOREST & GUINOT, 1961: 32 (Tahiti); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON & REES, 1967: 64 (Huahine). — POUPIN, 1994a: 32, fig. 28, pl. 3g (Kamaka, Mangareva, Tahiti, Tuamotu). — Thalamita (Thalaminella) coerulipes - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 51). Thalamita cooperi Borradaile, 1903 DISTRIBUTION. — Society (Moorea). REFERENCES. — Thalamita cooperi - NAIM, 1980a: annex 1, tab. 3 (Moorea; material not found in MNHN). 34 Thalamita corrugata Stephenson & Rees, 1961 DISTRIBUTION. — Tuamotu (Tikehau). REFERENCES. — Thalamita corrugata - STEPHENSON & REES, 1967: 65, fig. 23 ("Tickahau" = Tikehau). — GUINOT, 1985: 449 (List). Thalamita crenata (Latreille, 1829) DISTRIBUTION. — Marquesas; Society (Bora Bora, Maiao?, Moorea, Tahiti); Tuamotu (Mataiva, Takapoto, Tikehau). REFERENCES. — Thalamita crenata - A. MILNE EDWARDS, 1861: 365 (Marquesas). — ORTMANN, 1893a: 86 (Marquesas). — RATHBUN, 1907: 62 (Bora Bora). — SEURAT, 1934: 59 (Marquesas). — FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu, Marquesas"). — STEPHENSON & REES, 1967: 66 ("Maiai" = Maiao?, "Tikahau" = Tikehau). — MONTEFORTE, 1984: 173, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 8 (Moorea). — SALVAT & RICHARD, 1985: 356, 362 (Takapoto) — Thalamita (Thalaminella) crenata - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 51) - NEW MATERIAL - Coll. C. HILY, det. K. Moosa (Tahiti, Tikehau). Thalamita dakini Montgomery, 1931 DISTRIBUTION. — Society (Bora Bora, Moorea); Tuamotu (Makatea, Mataiva, Takapoto). REFERENCES. — Thalamita dakini - STEPHENSON & REES, 1967: 69 (Bora Bora, Moorea). — MONTEFORTE, 1984: 173, annex 1, tab a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — Thalamita (Thalamitopsis) dakini - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 47). Thalamita danae Stimpson, 1858 DISTRIBUTION. — French Polynesia. REFERENCES. — Thalamita danae - NEW MATERIAL - Coll. PLEssIs, det. K. MoosA (French Polynesia). Thalamita demani Nobili, 1905 DISTRIBUTION. — Marquesas (Nuku Hiva). REFERENCES. — Thalamita demani - STEPHENSON, 1976: 20 (""Hatwata" = Haatuatua bay, Nuku Hiva). Thalamita edwardsi Borradaile, 1900 DISTRIBUTION. — Gambier (Mangareva, Temoe); Society (Tahiti). REFERENCES. — Thalamita admete var. Edwardsii Bort. - NOBILI, 1907: 383 ("Rikitea" = Mangareva, "Timoe" = Temoe). — Thalamita edwardsi - FOREST & GUINOT, 1961: 32, fig. 20a-b (Tahiti); 1962: 58 (Biogeography "Tahiti- Tuamotu"). — Thalamita edwardsi - GUINOT, 1985: 449, with " =T. admete ?" (List; cf. Remark). REMARK. — This species was formerly considered as a synonym of Thalamita admete by STEPHENSON & HUDSON (1957). This assertion is not followed by FOREST & GUINOT (1961), CROSNIER (1962) and DAI & YANG (1991). Thalamita gatavakensis Nobili, 1906 DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Tahiti). REFERENCES. — Thalamita pilumnoides var. gatavakensis Nobili, 1906a: 262; 1907: 384 (""Gatavake" = Mangareva). — Thalamita pilumnoides ssp. gatavakensis - FOREST & GUINOT, 1961: 34, fig. 22a-b, 23-25 (Mangareva); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — Thalamita gatavakensis - CROSNIER, 1962: 99, fig. 156a-c, e (Mangareva; syntypes). — STEPHENSON & REES, 1967: 75 (Bora Bora, Tahiti). — Thalamita (Thalamita) gatavakensis - GUINOT, 1985: 449 (List) - NEW MATERIAL - Coll. C. HILY, det. K. Moosa (Tahiti). Thalamita gloriensis Crosnier, 1962 DISTRIBUTION. — Society (Huahine). 35 REFERENCES. — Thalamita gloriensis - STEPHENSON & REES, 1967: 76 (Huahine). — Thalamita (Thalamita) gloriensis - GUINOT, 1985: 449 (List). Thalamita gracilipes (A. Milne Edwards, 1873) DISTRIBUTION. — Society (Tahiti); Tuamotu (Moruroa?). REFERENCES. — Thalamita gracilipes - STEPHENSON, 1976: 21 (Tahiti) - RELEVANT MATERIAL - Thalamonyx aff. gracilipes - SALVAT & RENAUD-MORNANT, 1969: 165 (Moruroa). — Thalamita (Thalamonyx) aff. gracilipes - GUINOT, 1985: 449 (List). Thalamita integra Dana, 1852 DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti); Tuamotu (Takapoto, Nukutipipi). REFERENCES. — Thalamita integra Dana, 1852a: 85; 1852b: 281; 1855, pl. 17, fig. 6a-d (Tuamotu). — A. MILNE EDWARDS, 1861: 358 (Tahiti, Tuamotu). — NoBIL, 1907: 383 ("Gatavake" = Mangareva). — FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 173, annex 1, tab. a (Takapoto). — SALVAT & RICHARD, 1985: 350 (Takapoto). — MERSCHARDT-SALVAT, 1991: 89 (Nukutipipi). — Thalamita (Thalamita) integra - GUINOT, 1985: 449 (List). Thalamita macropus Montgomery, 1931 DISTRIBUTION. — Austral (Neilson bank); Marquesas (Fatu Hiva) - Sublittoral to deep. REFERENCES. — Thalamita macropus - NEW MATERIAL - Coll. B. RICHER DE FORGES and J. POUPIN, det. K. MOOSA (Neilson bank, Fatu Hiva; 49-100/130m). Thalamita macrospinifera Rathbun, 1911 DISTRIBUTION. — Austral (Rurutu); Marquesas (Eiao, Hiva Oa); Society (Raiatea); Tuamotu (Makemo, Moruroa) - Sublittoral to deep. REFERENCES. — Thalamita macrospinifera - POUPIN, 1996: in press (Makemo, Raiatea; 120-160m) - NEW MATERIAL - F Coll. J. POUPIN, det. K. Moosa (Eiao, Hiva Oa, Makemo, Moruroa, Raiatea, Rurutu; 80-160m). Thalamita minuscula Nobili, 1906 DISTRIBUTION. — Tuamotu (Kaukura, Vahitahi). REFERENCES. — Thalamita minuscula Nobili, 1906a: 262; 1907: 386, pl. 1, fig. 15 (Kaukura, Vahitahi). — FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 49 (List with "Only from Tuamotu Is."). — Thalamita (Neothalamita) minuscula - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 43). REMARK. — This very small species (maximum width, 4mm), never recorded since its description, could be the juvenile of another species. Thalamita mitsiensis Crosnier, 1962 DISTRIBUTION. — Austral (Rurutu) - Sublittoral. REFERENCES. — Thalamita mitsiensis - NEW MATERIAL - Coll. J. POUPIN, det. K. MOOSA (Rurutu; 80m). Thalamita philippinensis Stephenson & Rees, 1967 DISTRIBUTION. — Austral (Rurutu); Tuamotu (Moruroa) - Sublittoral to deep. REFERENCES. — Thalamita philippinensis - NEW MATERIAL - Coll. J. POUPIN, det. K. Moosa (Moruroa, Rurutu; 95- 130m). Thalamita picta Stimpson, 1858 DISTRIBUTION. — Marquesas (Hiva Oa, Nuku Hiva, Tahuata); Gambier?; Society (Raiatea, Tahiti); Tuamotu (Fakarava, Makatea, Mataiva, Raroia, Takapoto). oe ee ss 36 REFERENCES. — Goniosoma lineatum A. Milne Edwards, 1861: 377, pl. 35, fig. 4 (Nuku Hiva) fide CROSNIER (1962: 138). — Thalamita alcocki - NOBILI, 1907: 384 (""Tagatau" = Gambier?). — Thalamita gardineri - RATHBUN, 1907: 63 (Fakarava, Makemo). — Thalamita picta - HOLTHUIS, 1953: 8 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST & GUINOT, 1961: 33 (Tahiti; Syn.); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 50 (Syn.); 1976: 23 ("Hoava Strait” = Haava strait, between Tahuata and Hiva Oa, cf. HARALD, 1967). — MONTEFORTE, 1984: 173, annex 1, tab. a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — Thalamita (Neothalaminella) picta - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 51) - NEW MATERIAL - Coll. J. POUPIN, det. K. MOOSA (Raiatea) - SYNONYMS - Thalamita alcocki de Man, 1902; T. gardineri Borradaile, 1902. Thalamita pilumnoides Borradaile, 1903 DISTRIBUTION. — Society (Huahine, Moorea, Raiatea). REFERENCES. — Thalamita pilumnoides - STEPHENSON & REES, 1967: 87, fig. 32 (Huahine, Moorea, Raiatea). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212 (Moorea). — MONTEFORTE, 1984: 173, annex 1, tab. a; 1987: 8 (Moorea). — Thalamita (Neothalamita) pilumnoides - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 43). Thalamita prymna (Herbst, 1803) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Anaa). REFERENCES. — Thalamita prymna - STEPHENSON, 1976: 23 (Anaa). — PEYROT-CLAUSADE, 1989: 115 (Moorea) - NEW MATERIAL - Coll. J. POUPIN, det. K. MOoSA (Tahiti). Thalamita quadrilobata Miers, 1884 DISTRIBUTION. — Society (Bora Bora). REFERENCES. — Thalamita quadrilobata - STEPHENSON & REES, 1967: 92 (Bora Bora). — Thalamita (Pseudothalamitopsis) quadrilobata - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 47). Thalamita seurati Nobili, 1906 DISTRIBUTION. — Tuamotu (Marutea South). REFERENCES. — Thalamita seurati Nobili, 1906a: 262; 1907: 385, pl. 2, fig. 1 (Marutea). — SEURAT, 1934: 60 (French Polynesia). — FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — STEPHENSON, 1972: 51 (List "Tuamotu"). — Thalamita (Pseudothalamitopsis) seurati - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 47). Thalamita spinifera Borradaile, 1903 DISTRIBUTION. — Austral (Raevavae, Tubuai); Marquesas (Eiao, Fatu Hiva, Nuku Hiva); Tuamotu (Makemo) - Sublittoral to deep. REFERENCES. — Thalamita spinifera - STEPHENSON, 1976: 24 ("Marquesas Expedition, 40-80m, 18/ix/1967 to 1/x/1967" = Marquesas, cf. HARALD, 1967) - NEW MATERIAL - Coll. J. POUPIN, det. K. MOOSA (Eiao, Fatu Hiva, Makemo, Nuku Hiva, Raevavae, Tubuai; 42-200m). Thalamita woodmasoni Alcock, 1899 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Thalamita woodmasoni - FOREST & GUINOT, 1961: 33 (Tahiti); 1962: 60 (Biogeography "Tahiti- Tuamotu"). — Thalamita (Pseudothalamitopsis) woodmasoni - GUINOT, 1985: 449 (List; subgenus from Moosa, 1979: 47). Thalamitoides quadridens A. Milne Edwards, 1869 DISTRIBUTION. — Tuamotu (Moruroa, Rangiroa). EE EEE eee 37 REFERENCES. — Thalamitoides quadridens - STEPHENSON, 1976: 26 (Rangiroa) - NEW MATERIAL - Coll. J. POUPIN, det. K. Moosa (Moruroa). SUBFAMILY PODOPHTHALMINAE Podophthalmus vigil (Fabricius, 1798) DISTRIBUTION. — Society (Moorea, Tahiti). REFERENCES. — Podophthalmus vigil - FOREST & GUINOT, 1961: 36 (Tahiti); 1962: 60 (Biogeography "Tahiti Tuamotu"). — STEPHENSON & REES, 1967: 104 ("Opunohu Bay” = Moorea). — MONTEFORTE, 1984: 173, annex 1, tab. a (Moorea). — GUINOT, 1985: 449 (List). FAMILY XANTHIDAE SUBFAMILY POLYDECTINAE. Lybia caestifera (Alcock, 1897) DISTRIBUTION. — Society (Tahiti); Tuamotu (Mataiva?). REFERENCES. — Lybia caestifera - RATHBUN, 1907: 60 (Tahiti). — FoREST & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1976: 75 (Syn.; RATHBUN's reference with a ?); 1985: 452, with a ? (List) - RELEVANT MATERIAL - Lybia cf. coestifera (sic) - MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva). Lybia plumosa Barnard, 1947 DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau). REFERENCES. — Lybia leptochelis - PEYROT-CLAUSADE, 1977a, annex of the species: 27 (Moorea) not L. leptochelis (Zehntner, 1894) = L. plumosa fide SERENE (1984: 29, 31). — Lybia plumulosa (sic) - PEYROT-CLAUSADE, 1989: 113 (Tikehau). Lybia tessellata (Latreille, 1812) DISTRIBUTION. — Marquesas; Society (Moorea, Tahiti); Tuamotu (Makatea, Mataiva, Rangiroa, Raroia). REFERENCES. — Melia tesselata - FINNEGAN, 1931: 647 (Marquesas). — Lybia tessellata - HOLTHUIS, 1953: 23 (Raroia). — Morrison, 1954: 13 (Raroia). — FOREST & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu, Marquesas”). — TAKEDA & MIYAKE, 1970: 15 ("Avatoru” = Rangiroa, Tahiti). — GUINOT, 1976: 70, fig. 17d, 18e, 19c, 20e-h, 22d, pl. 2, fig. 6 (Syn.); 1985: 452 (List). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti); 1987: 9 (Moorea). Polydectus cupulifer (Latreille, 1812) DISTRIBUTION. — Tuamotu (Raraka). REFERENCES. — Polydectus villosus Dana, 1852a: 81; 1852b: 227; 1855, pl. 13, fig. 3a-e (Raraka). — Polydectus cupulifer - FOREST & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1976: 65 (Syn.); 1985: 452 (List). — SERENE, 1984: 24, fig. 1, pl. 1a (Syn.). 38 SUBFAMILY CYMOINAE Cymo andreossyi (Audouin, 1826) DISTRIBUTION. — Society (Moorea, Tahiti). REFERENCES. — Cymo andreossyi - DANA, 1852b: 225; 1855, pl. 13, fig. 2a-b (Tahiti). — HELLER, 1865: 20 (Tahiti). — Forest & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu"), — ODINETZ, 1983: 206 (Moorea, Tahiti). — GUINOT, 1985: 450 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti). Cymo deplanatus A. Milne Edwards, 1873 DISTRIBUTION. — Tuamotu (Raroia). REFERENCES. — Cymo deplanatus - HOLTHUIS, 1953: 18 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 450 (List). Cymo melanodactylus de Haan, 1833 DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Hao, Hikueru, Moruroa). REFERENCES. — Cymo Andreossyi var. melanodactyla - NOBILI, 1907: 397 ("Otepa" = Hao, Mangareva). — Cymo melanodactylus - RATHBUN, 1907: 53 (Bora Bora, Fakarava). — PESTA, 1913: 46 (Tahiti). — BOONE, 1934: 144, pl. 74 (Tahiti). — FOREST & GUINOT, 1961: 119 (Hikueru, "Rikitea" = Mangareva); 1962: 68 (Biogeography "Tahiti- Tuamotu"). — ODINETZ, 1983: 206 (Tahiti). — SERENE, 1984: 34, fig. 8, pl. 2b (Syn.). — GUINOT, 1985: 450 (List). — ODINETZ-COLLART & RICHER DE ForRGES, 1985: 201 (Tahiti). — POUPIN, 1994a: 33, fig. 29, pl. 3h (Mangareva, Moruroa). Cymo quadrilobatus Miers, 1884 DISTRIBUTION. — Society (Moorea, Tahiti?). REFERENCES. — Cymo quadrilobatus - ODINETZ, 1983: 206 (Moorea). — GUINOT, 1985: 450 (List). — ODINETZ- COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti). SUBFAMILY TRICHIINAE Banareia parvula (Krauss, 1843) DISTRIBUTION. — Marquesas. REFERENCES. — Actaea parvula - ODHNER, 1925: 35, 51, pl. 3, fig. 13 (Marquesas). — Banareia parvula - GUINOT, 1976: 179, with a ? for the genus (Syn.; see the considerations about the generic rank); 1985: 452 (List). — Banareia parvula - GARTH et al., 1987: 243 (cited only for the generic rank). SUBFAMILY LIOMERINAE Liomera bella (Dana, 1852) DISTRIBUTION. — Gambier (Mangareva, Tarauru-Roa, Vaiatekeue); Society (Moorea, Tahiti); Tuamotu (Fakareva, Hikueru, Makatea, Makemo, Marutea South, Mataiva, Takapoto, Takaroa, Tikehau). REFERENCES. — Carpiloxanthus rugipes - HELLER, 1865: 17 (Tahiti) not Liomera rugipes (Heller, 1861) = L. bella fide SERENE (1984: 65). — Carpilodes rugatus - NoBILI, 1907: 387 ("chenal Waiatekene = Vaiatekeue, "Rikitea” = Mangareva, "Waitutaki" = Marutea South). — RATHBUN, 1907: 37 (Makemo) - All, not Liomera rugata (H. Milne Edwards, 1834) = L. bella fide SERENE (1984: 61). — Carpilodes vaillantianus - NOBILI, 1907: 387. — Carpilodes bellus - ODHNER, 1925: 16, pl. 1, fig. 9 ("Eimeo" = Moorea, Makemo, Tahiti). — BUITENDIJK, 1960: 257, fig. 2b (Tahiti). — Liomera bella - FOREST & GUINOT, 1961: 38, fig. 26a-b (Hikueru, Tahiti, "Taraourou-roa" = Tarauru-Roa); eT 39 1962: 60 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212 (Moorea); 1989: 111 (Tikehau). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 450 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti). — Liomera (Liomera) bella - SERENE, 1984: 60, fig. 21, pl. 5e (Syn.) - SYNONYMS - Carpilodes vaillantianus A. Milne Edwards, 1862. Liomera cinctimana (White, 1847) DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Moorea, Tahiti). REFERENCES. — Liomera lata - HELLER, 1865: 9 (Tahiti). — Liomera cinctimana - ORTMANN, 1893b: 450, pl. 17, fig. 8 (Tahiti). — FOREST & GUINOT, 1961: 39, fig. 27a-b (Tahiti, "Taihoae” = Nuku Hiva); 1962: 60 (Biogeography "Tahiti-Tuamotu, Marquesas"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26 (Moorea). — MONTEFORTE, 1984: 171, annex 1, tab. a (Tahiti). — GUINOT, 1985: 450 (List). — Carpilodes cinctimanus - ODHNER, 1925: 14 (Tahiti). — Liomera (Liomera) cinctimana - SERENE, 1984: 57, fig. 17, pl. Sa (Syn.) - SYNONYMS - Liomera lata Dana, 1852. Liomera laevis (A. Milne Edwards, 1873) DISTRIBUTION. — Society (Moorea). REFERENCES. — Liomera laevis - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212 (Moorea). — GUINOT, 1985: 451, with a ? (List). Liomera laperousei Garth, 1985 DISTRIBUTION. — Austral (MacDonald bank) - Littoral to sublittoral . REFERENCES. — Liomera laperousei - LABOUTE & RICHER DE FORGES, 1986: 21 (MacDonald bank, 40m; with hesitation: "semble étre une femelle juvénile de Liomera laperousei Garth, 1985 décrit de lille de Paque"). Liomera monticulosa (A. Milne Edwards, 1873) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Marutea South) - Littoral to sublittoral. REFERENCES. — Carpilodes monticulosus - NOBILI, 1907: 387 (Marutea South). — ODHNER, 1925: 21, pl. 1, fig. 18 (Tahiti, Marutea South). — Liomera monticulosa - FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — GuINoT, 1964: 11 (Syn.); 1985: 451 (List). — PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212 (Moorea); 1989: 115 (Moorea; 30m). — Liomera (Liomera) monticulosa - SERENE, 1984: 64, fig. 24, pl. 6c (Syn.). — Not Carpilodes monticulosus - RATHBUN, 1907: 37 (Fakarava, Makemo) = Liomera (Liomera) rugata (H. Milne Edwards, 1834) fide SERENE (1984: 62). Liomera pallida (Borradaile, 1900) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Carpilodes pallidus - ODHNER, 1925: 20, pl. 1, fig. 17 (Tahiti). — Liomera pallida - FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 451, with a ? (List). — Liomera (Liomera) pallida - SERENE, 1984: 62, pl. 5f (Syn.). Liomera rubra (A. Milne Edwards, 1865) DISTRIBUTION. — Society (Moorea). REFERENCES. — Liomera rubra - MONTEFORTE, 1984: 171, annex 1, tab. a; 1987: 9 (Moorea). — GUINOT, 1985: 451, with a ? (List). — Liomera (Liomera) rubra - SERENE, 1984: 65, fig. 26, pl. 6e-f, pl. 9f (Syn.). Liomera rugata (H. Milne Edwards, 1834) DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Moorea, Tahiti); Tuamotu (Fakarava, Makemo, Mataiva, Raroia, Tikehau). 40 REFERENCES. — Carpilodes rugatus - ORTMANN, 1893b: 468 (Tahiti). — ODHNER, 1925: 20, pl. 1, fig. 16 (Fakarava, Tahiti). — BOONE, 1934: 91, pl. 46 (Nuku Hiva). — HOLTHUIS, 1953: 13 (Raroia). — MORRISON, 1954: 16 (Raroia). — BUITENDUK, 1960: 259, fig. 2d (Tahiti). — Carpilodes monticulosus - RATHBUN, 1907: 37 (Fakarava, Makemo) not Liomera monticulosa (A. Milne Edwards, 1873) = L. rugata fide SERENE (1984: 62). — Liomera rugata - FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva, Moorea); 1987: 9 (Moorea). — GUINOT, 1985: 451 (List). — PEYROT-CLAUSADE, 1989: 113, 115 (Moorea, Tikehau). — Liomera (Liomera) rugata - SERENE, 1984: 62, fig. 22, pl. 6b (Syn.). Liomera semigranosa De Man, 1888 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Liomera semigranulosa (sic) - ODINETZ, 1983: 209 (Tahiti). — Liomera semigranosa - GUINOT, 1985: 451 (List). — Liomera (Liomera) semigranosa - SERENE, 1984: 63, pl. 7c, f (Syn.). Liomera stimpsoni (A. Milne Edwards, 1865) DISTRIBUTION. — Society (Moorea). REFERENCES. — Liomera stimpsoni - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212; 1989: 115 (Moorea). — GUINOT, 1985: 451, with a ? (List). Liomera tristis (Dana, 1852) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava). REFERENCES. — Carpilodes tristis Dana, 1852a: 77; 1852b: 193; 1855, pl. 9, fig. 7a-d (Tuamotu with a ?). — HELLER, 1865: 17 (Tahiti). — A. MILNE EDWARDS, 1865b: 225 (Tuamotu). — DE MAN, 1890: 50 (Tahiti). — RATHBUN, 1907: 37 (Fakarava). — ODHNER, 1925: 12, pl. 1, fig. 1 ("Eimeo" = Moorea, Tahiti). — BUITENDIJK, 1960: 254, fig. lc (Tahiti). — Carpilodes granulatus Heller, 1862: 520 (Tahiti). — Liomera tristis - FOREST & GUINOT, 1961: 38 (Tahiti); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 451 (List). — Liomera (Liomera) tristis - SERENE, 1984: 59, fig. 19, pl. 5b (Syn.). Liomera venosa (H. Milne Edwards, 1834) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Carpilodes venosus - ORTMANN, 1893b: 467 (Tahiti). — ODHNER, 1925: 22, pl. 2, fig. 1 (Tahiti). — BUITENDIJK, 1960: 259, fig. 2e (Tahiti). — Liomera venosa - FOREST & GUINOT, 1962: 60 (Biogeography "Tahiti- Tuamotu"). — GUINOT, 1985: 451 (List). — Liomera (Liomera) venosa - SERENE, 1984: 58, fig. 18, pl. 7d-e (Syn.). Neoliomera demani Forest & Guinot, 1961 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru). REFERENCES. — Neoliomera demani Forest & Guinot, 1961: 80, fig. 76, 77bis, pl. 3, fig. 3-5 (Hikueru, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — SERENE, 1984: 71, fig. 31, pl. 8b (Hikueru). — GuINoT, 1985: 451 (List). — PEYROT-CLAUSADE, 1989: 115 (Moorea). — Neoliomera pubescens - ODHNER, 1925: 28, pl. 2, fig. 6, 6a-7 (Tahiti) not Neoliomera pubescens (H. Milne Edwards, 1834) = N. demani sp. nov. in FOREST & GUINOT (1961: 80). Neoliomera insularis (White, 1847) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Neoliomera insularis - SAKAI, 1976: 398 (Tahiti; cf. Remark). — GUINOT, 1985: 451 (List; after SAKAI). REMARK. — The only reference in French Polynesia seems to be in SAKAI (1976) where "Tahiti" is cited without material examined from that island, and without older references for that location. Thus, the occurence of this species in the French Polynesia still remains to be confirmed. 41 Neoliomera pubescens (H. Milne Edwards, 1834) DISTRIBUTION. — Society (Moorea). REFERENCES. — Neoliomera pubescens - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212 (Moorea). — SERENE, 1984: 71, fig. 30, pl. 8a (Syn.; cf. Remark). REMARK. — According to SERENE (1984) it is, in most of the works, necessary to check that the material attributed to Neoliomera pubescens has not been confounded with N. demani Forest & Guinot, 1961. He also mentions that N. pubescens is known, with certainty, only from Maunitius. Neoliomera richtersi (De Man, 1889) DISTRIBUTION. — Society (Tahiti); Tuamotu (Marutea South). REFERENCES. — Actaeodes richtersii de Man 1889: 412, pl. 9, fig. 2; 1890: 51 (Tahiti). — Liomera richtersi - NOBILI, 1907: 387 (""Marutea-Vaitutaki" = Marutea South). — Neoliomera richtersii - ODHNER, 1925: 33, pl. 2, fig. 13 (Tahiti). — BUITENDIJK, 1960: 262 (Tahiti). — FOREST & GUINOT, 1961: 79, fig. 74 (Tahiti); 1962: 66 (Biogeography "Tahiti- Tuamotu"). — GUINOT, 1964: 47, fig. 17 (Tahiti); 1985: 451 (List). — SERENE, 1984: 70, fig. 28, pl. 8e (Tahiti; Syn.). Neoliomera variolosa (A. Milne Edwards, 1873) DISTRIBUTION. — Society (Moorea). REFERENCES. — Neoliomera variolosa - PEYROT-CLAUSADE, 1977b: 212, 220 (Moorea). SUBFAMILY EUXANTHINAE Alainodaeus rimatara Davie, 1993 DISTRIBUTION. — Austral (Raevavae, Rimatara); Tuamotu (Akiaki, Fangataufa, Hao, Takapoto) - Sublittoral to deep. REFERENCES. — Alainodaeus rimatara Davie, 1993: 519, fig. 6, pl. 6 (Akiaki, Fangataufa, Hao, Raevavae, Rimatara, Takapoto; 90-350m). — POUPIN, 1996: in press (same material). Euxanthus exsculptus (Herbst, 1790) DISTRIBUTION. — Gambier (Mangareva); Society (Moorea, Tahiti). REFERENCES. — Euxanthus exsculptus var. rugosus - NOBILI, 1907: 389 (Mangareva) not Euxanthus rugosus Miers, 1884 = E. exsculptus fide GUINOT-DUMORTIER (1960b: 170). — Euxanthus exsculptus - GUINOT-DUMORTIER, 1960b: 169, pl. 1, fig. 4, pl. 2, fig. 10, pl. 6, fig. 36-37, pl. 8, fig. 42-47 (""Rikitea" = Mangareva). — FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, annex 1, tab. a; 1987: 9 (Moorea). — SERENE, 1984: 86, fig. 48, pl. 11b (Tahiti). — GuINOT, 1985: 450 (List). Euxanthus sculptilis Dana, 1852 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Euxanthus sculptilis - BOONE, 1934: 107, pl. 57 (Tahiti). — ForREsT & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — GUINOT-DUMORTIER, 1960b: 167, pl. 6, fig. 39, pl. 9, fig. 49 (Syn.); 1985: 450 (List). Medaeus grandis Davie, 1993 DISTRIBUTION. — Tuamotu (Hao, Moruroa) - Sublittoral to deep. REFERENCES. — Medaeus grandis Davie, 1993: 526, fig. 8, pl. 8 (Hao, Moruroa; 90-210m). — PouPIN, 1996: in press (same material). Paramedaeus noelensis (Ward, 1934) DISTRIBUTION. — Society (Moorea, Tahiti). 42 REFERENCES. — Medaeus noelensis - FOREST & GUINOT, 1961: 56, fig. 42-43, 44a-b, pl. 1, fig. 1 (Tahiti); 1962: 62 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a (Moorea, Tahiti); 1987: 9 (Moorea). — Paramedaeus noelensis - SERENE, 1984: 90, fig. 51, pl. 12f (Syn.). — GUINOT, 1985: 451 (List). Paramedaeus simplex (A. Milne Edwards, 1873) DISTRIBUTION. — Society (Moorea). REFERENCES. — Paramedaeus simplex - PEYROT-CLAUSADE, 1977b: 212 (Moorea). — GUINOT, 1985: 451, with a ? (List). SUBFAMILY ACTAEINAE Actaea aff. glandifera Rathbun, 1914 DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral. REFERENCES. — Actaea aff. glandifera - PEYROT-CLAUSADE, 1989: 111, 114 (Moorea, Tikehau; 25m). Actaea calculosa (H. Milne Edwards, 1834) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Actaea calculosa - ODHNER, 1925: 52 (Tahiti). — FOREST & GUINOT, 1962: 64 (Biogeography "Tahiti- Tuamotu"). REMARK. — ODHNER mentions a Tahitian specimen in the collections of Hamburg. However, GUINOT (1976: 215-216) states that most of the references to Actaea calculosa, especially ODHNER (1925: 52), must be re-examined. Actaea danae A. Milne Edwards, 1865 DISTRIBUTION. — Tuamotu (Raraka). REFERENCES. — Actaeodes areolatus Dana, 1852a: 77; 1852b: 194; 1855, pl. 9, fig. 8a-d (Raraka). — Actaea danae - ForEST & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — (Actaea) danae - GUINOT, 1976: 247 (Syn.); 1985: 450, with "species inquirenda" (List). Actaea polyacantha (Heller, 1861) DISTRIBUTION. — Society (Moorea). REFERENCES. — Actaea polyacantha - PEYROT-CLAUSADE, 1989: 114 (Moorea). Actaeodes consobrinus (A. Milne Edwards, 1873) DISTRIBUTION. — Marquesas; Society (Moorea) - Littoral to sublittoral. REFERENCES. — Actaea consobrina - ODHNER, 1925: 67, pl. 4, fig. 14 (Marquesas). — Actaeodes consobrinus - GUINOT, 1976: 246, pl. 15, fig. 5, 5a (Syn.); 1985: 450, with a ? (List). — Actaeodes consobrina - PEYROT-CLAUSADE, 1989: 114 (Moorea; 30m). — Not Actaea consobrina - NOBILI, 1907: 390 = Actaea ruppellioides sp. nov. in ODHNER (1925: 47; cf. under Pseudoliomera ruppellioides). Actaeodes hirsutissimus (Riippell, 1830) DISTRIBUTION. — Society (Bora Bora, Moorea, Raiatea, Tahiti); Tuamotu (Mataiva, Tikehau). REFERENCES. — Actaea hirsutissima - HELLER, 1865: 9 (Tahiti). — RATHBUN, 1907: 42 (Bora Bora, Tahiti). — ODHNER, 1925: 69, pl. 4, fig. 13 (Tahiti). — BOONE, 1934: 124, pl. 66 (Raiatea). — FOREST & GUINOT, 1961: 78 (Tahiti); 1962: 64 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1989: 111 (Tikehau). — Actaeodes hirsutissimus - GUINOT, 1976: 245, fig. 38e, pl. 15, fig. 2, 2a (Tahiti; Syn.); 1985: 450 (List). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 213; 1985: 462 (Moorea). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva, Moorea, Tahiti); 1987: 8 (Moorea). — SERENE, 1984: 135 (Syn.). eee 43 Actaeodes tomentosus (H. Milne Edwards, 1834) DISTRIBUTION. — Society (Tahiti); Tuamotu (Makatea). REFERENCES. — Actaeodes tomentosus - HELLER, 1865: 17 (Tahiti). — GUINOT, 1976: 244, fig. 38d, 41c, pl. 15, fig. 1, la (Syn.); 1985: 450, with a ? (List). — SERENE, 1984: 134, 137 (Syn.). — Actaea tomentosa - SENDLER, 1923: 37 (Makatea). — FoREST & GUINOT, 1962: 66 (Biogeography "Tahiti-Tuamotu”). Forestia depressa (White, 1847) DISTRIBUTION. — Marquesas. REFERENCES. — Actaea depressa - BALSS, 1935: 136 (Marquesas). — Forestia depressa - GUINOT, 1976: 262 (Syn.,; with a ? for BALSS' reference); 1985: 450 (List; with a ? in front of "Marquesas"). — SERENE, 1984: 106 (Syn.; with a ? in front of BALSs' reference). REMARK. — In GUINOT (1976) and SERENE (1984), at least one specimen attributed to Actaea depressa, by BALSS (1938: 54), would be in fact a Forestia scabra (Odhner, 1925). Forestia scabra (Odhner, 1925) DISTRIBUTION. — Marquesas. REFERENCES. — Actaea scabra - BALSS, 1935: 136 (Marquesas). — Forestia scabra - GUINOT, 1976: 263 (Syn.; with a ? in front of BALSS' reference); 1985: 450 (List; with a ? in front of "Marquesas"). Gaillardiellus rueppelli (Kraus, 1843) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Gaillardiellus rueppelli - GARTH & KIM, 1983: 684 (Distribution only, Tahiti; cf. Remark). — GUINOT, 1985: 450 (List; presumably after the previous work). REMARK. — The material examined by GARTH & KIM (1983: 685) was collected in the Philippines by the Albatross (1908-1909). "Tahiti", mentioned in the “Distribution” only, corresponds neither to material examined, nor to former references cited in this work. Gaillardiellus superciliaris (Odhner, 1925) DISTRIBUTION. — Tuamotu (Raroia, Taiaro). REFERENCES. — Actaea superciliaris - HOLTHUIS, 1953: 11 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — Gaillardiellus superciliaris - GUINOT, 1976: 257 (Syn.); 1985: 450 (List). — POUPIN, 1994a: 34, fig. 30, pl. 4a, with a ? (Taiaro). Paractaea excentrica Guinot, 1969 DISTRIBUTION. — Tuamotu (Marutea South). REFERENCES. — Paractaea excentrica Guinot, 1969: 263, fig. 36 (Marutea South); 1985: 451 (List). Paractaea retusa (Nobili, 1905) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Raroia). REFERENCES. — Actaea garretti Rathbun, 1906: 852, pl. 9, fig. 8 (Society). — Actaea rufopunctata - HOLTHUIS, 1953: 11 (Raroia). — Morrison, 1954: 16 (Raroia). — FOREST & GUINOT, 1961: 79, fig. 79a-b (Tahiti) - All, not Paractaea rufopunctata (H. Milne Edwards, 1834) = Paractaea retusa form hippocrepica nov. in GUINOT (1969: 256). — Paractaea retusa form hippocrepica - GUINOT, 1969: 256, fig. 30 (Tahiti, Raroia; Syn.); 1985: 451 (List). — PEYROT- CLAUSADE, 1989: 115 (Moorea). — Paractaea retusa - MONTEFORTE, 1984: 171, annex 1, tab. a; 1987: 9 (Moorea). REMARK. — GUINOT (1969: 255), when creating the new genus Paractaea, has examined the syntype of garretti from Gilbert Islands (but not the specimen from the Society Islands), and considers that RATHBUN's species belongs to Paractaea retusa (Nobili) form garretti (Rathbun). Paractaea rufopunctata H. Milne Edwards, 1834 DISTRIBUTION. — Society (Tahiti); Tuamotu (Makemo, Marutea South). REFERENCES. — Actaea rufopunctata - NOBILI, 1907: 392 (Marutea). — RATHBUN, 1907: 43 (Makemo, Tahiti). — ODHNER, 1925: 60 (Tahiti). — FOREST & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — Paractaea rufopunctata form plumosa - GUINOT, 1969: 248, fig. 21 (Marutea). — Paractaea rufopunctata - GUINOT, 1985: 451 (List). Paractaeopsis quadriareolatus (Takeda & Miyake, 1968) DISTRIBUTION. — Society (Moorea). REFERENCES. — Paractaeopsis quadriareolatus - SERENE, 1984: 127 (Syn.; gen. nov.). — Paractaea quadriareolata - PEYROT-CLAUSADE, 1989: 115 (Moorea). Paractaeopsis tumulosus (Odhner, 1925) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Actaea tumulosa Odhner, 1925: 61, pl. 4, fig. 10 (Tahiti). — ForEST & GUINOT, 1962: 66 (Biogeography "Tahiti-Tuamotu”). — Paractaeopsis tumulosus - SERENE, 1984: 127, fig. 74, pl. 17d (Syn.; gen. nov.). — Paractaea tumulosa - GUINOT, 1985: 451 (List). Psaumis cavipes (Dana, 1852) DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Hao, Rangiroa, Raroia, Tikehau). REFERENCES. — Actaea cavipes - NOBILI, 1907: 390 ("Ohura" = Hao, "Rikitea" = Mangareva). — RATHBUN, 1907: 44, pl. 1, fig. 2 (Bora Bora, Fakarava, Rangiroa). — ODHNER, 1925: 68 ("Eimeo" = Moorea, Tahiti). — BOONE, 1934: 128, pl. 68 (Tahiti). — HOLTHUIS, 1953: 10 (Raroia). — MORRISON, 1954: 16 (Raroia). — FOREST & GUINOT, 1961: 78 (Tahiti); 1962: 64 (Biogeography "Tahiti-Tuamotu”). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212 (Moorea). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a (Moorea, Tahiti); 1987: 8 (Moorea). — Psaumis cavipes - ODINETZ, 1983: 209 (Moorea, Tahiti). — SERENE, 1984: 129, fig. 76, pl. 18f (Syn.). — GUINOT, 1985: 451 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti, Takapoto). — MONTEFORTE, 1987: 9 (Moorea). — PEYROT-CLAUSADE, 1989: 113 (Tikehau). Psaumis cellulosa (Dana, 1852) DISTRIBUTION. — Society (Moorea, Tahiti). REFERENCES. — Psaumis cellulosa - ODINETZ, 1983: 209 (Moorea, Tahiti). — GUINOT, 1985: 451 (List). Pseudoliomera granosimana (A. Milne Edwards, 1865) DISTRIBUTION. — Society (Tahiti); Tuamotu (Marutea South). REFERENCES. — Liomera granosimana - ORTMANN, 1893b: 451 (Tahiti). — NOBILI, 1907: 387 ("Marutea-Vaitutaki" = Marutea South). — Pseudoliomera granosimana - ODHNER, 1925: 79, fig. 5-6 (Tahiti). — FOREST & GUINOT, 1961: 39, fig. 28a-c, pl. 7, fig. 1-2 (Tahiti); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — SERENE, 1984: 100, fig. 56, pl. 13a (Tahiti). — GuINOT, 1985: 451 (List). Pseudoliomera lata (Borradaile, 1902) DISTRIBUTION. — Society (Moorea). REFERENCES. — Pseudoliomera lata - KROpP & BIRKELAND, 1981: 630, tab. 5 (Moorea). — SERENE, 1984: 102 (Syn.). — PEYROT-CLAUSADE, 1989: 115 (Moorea). — (Pseudoliomera) lata - GUINOT, 1985: 451, with a ? (List). — Not Actaea lata - NOBILI, 1907: 392 ("Marutea, Vaitutaki" = Marutea South) = A. ruppellioides Odhner, 1925 fide GUINOT (1962: 237). 45 Pseudoliomera ruppellioides (Odhner, 1925) DISTRIBUTION. — Tuamotu (Marutea South). REFERENCES. — Actaea consobrina - NOBILI, 1907: 390 (Marutea South) not Actaea consobrina A. Milne Edwards, 1873 =A. ruppellioides sp. nov. in ODHNER (1925: 47). — Actaea lata - NoBILI, 1907: 392 ("Marutea, Vaitutaki" = Marutea South) not Actaea lata Borradaile, 1902 = A. ruppellioides Odhner fide GUINOT (1962: 237). — Actaea ruppellioides Odhner, 1925: 47, pl. 3, fig. 9 (Marutea South; NOBILI's material). — FOREST & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — (Pseudoliomera) ruppellioides - GUINOT, 1976: 203, 246 (Genus Pseudoliomera "ou a sa proximité"); 1985: 451 (List). Pseudoliomera speciosa (Dana, 1852) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Actaea speciosa - SENDLER, 1923: 38 (Tahiti). — ODHNER, 1925: 62 (Tahiti). — FOREST & GUINOT, 1962: 66 (Biogeography "Tahiti-Tuamotu"). — (Pseudoliomera) speciosa - GUINOT, 1976: 203, 243 (Genus uncertain); 1985: 451, with a ? (List). Pseudoliomera variolosa (Borradaile, 1902) DISTRIBUTION. — Society (Moorea); Tuamotu (Mataiva, Tikehau) - Littoral to sublittoral. REFERENCES. — Pseudoliomera variolosa - KROPP & BIRKELAND, 1981: 630, tab. 5 (Moorea). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva, Moorea; cf. Remark); 1987: 9 (Moorea). — PEYROT-CLAUSADE, 1989: 111, 115 (Moorea, Tikehau; 30m). — Aff. Pseudoliomera variolosa - PEYROT-CLAUSADE, 1977a, annex of the species: 27 (Moorea). — (Pseudoliomera) variolosa - GUINOT, 1985: 451 (List). REMARK. — MONTEFORTE writes, Pseudoliomera variolosa (A. Milne Edwards, 1837 sic). Then, it could be Neoliomera variolosa (A. Milne Edwards, 1873), which is different from BORRADAILE's species (cf. SERENE, 1984: 66). SUBFAMILY ZOZIMINAE Atergatis floridus (Linné, 1767) DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Hao). REFERENCES. — Atergatis floridus - DANA, 1852b: 159; 1855, pl. 7, fig. 4 (Society and Tuamotu). — HELLER, 1865: 8 (Tahiti). — NosIL, 1907: 388 ("Ohura"” = Hao). — SEURAT, 1934: 59 (Hao). — BUITENDIJK, 1960: 268 (Society). — FoREST & GUINOT, 1961: 41 (Tahiti; Syn.); 1962: 62 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212 (Moorea). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a (Moorea, Tahiti); 1987: 8 (Moorea). — GUINOT, 1985: 450 (List). — PoUPIN, 1994a: 35, fig. 31, pl. 4b (Mangareva, Tahiti). — Afergatis ocyroe - RATHBUN, 1907: 37 (Bora Bora). — SENDLER, 1923: 37 (Tahiti) - SYNONYMS - Altergatis ocyroe (Herbst, 1801). Atergatopsis cf. germaini A. Milne Edwards, 1865 DISTRIBUTION. — Society (Moorea). REFERENCES. — Atergatopsis cf. germaini - MONTEFORTE, 1984: 170, annex 1, tab. a; 1987: 9 (Moorea). — GUINOT, 1985: 450 (List). Atergatopsis signatus (Adams & White, 1848) DISTRIBUTION. — Tuamotu (Makatea, Mataiva, Raroia, Takapoto). REFERENCES. — Atergatopsis signatus - HOLTHUIS, 1953: 12 (Raroia). — MORRISON, 1954: 16 (Raroia). — FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (Raroia); 1985: 450 (List). — MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — SALVAT, 1986b: 72 (French Polynesia). 46 Lophozozymus cristatus A. Milne Edwards, 1867 DISTRIBUTION. — Austral (Maria); Society. REFERENCES. — Lophozozymus cristatus - BUITENDUK, 1960: 292, fig. 7a (Society). — FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — POUPIN, 1994a: 36, fig. 32, pl. 4c (Maria). Lophozozymus dodone (Herbst, 1801) DISTRIBUTION. — Society (Moorea, Tahiti). REFERENCES. — Atfergatis elegans Heller, 1862: 519; 1865: 7, pl. 1, fig. 3 (Tahiti). — Lophozozymus dodone - BALSS, 1938: 39 (Tahiti). — FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a (Moorea, Tahiti); 1987: 9 (Moorea). — SERENE, 1984: 171, pl. 24e (Syn.). — GUINOT, 1985: 451 (List). — Not Lophozozymus dodone - FOREST & GUINOT, 1961: 54, fig. 39a-b (Tahiti) = Lophozozymus glaber fide GUINOT (1979: 65). Lophozozymus edwardsi Odhner, 1925 DISTRIBUTION. — Gambier (Mangareva); Society (Moorea); Tuamotu (Marutea South). REFERENCES. — Lophozozymus superbus - NoBILI, 1907: 388 (Mangareva, "Marutea Vaitutaki" = Marutea South) not Lophozozymus superbus A. Milne Edwards, 1873 = L. edwardsi fide FOREST & GUINOT (1961: 56) & GuINoT (1979: 63). — Lophozozymus edwardsi - FOREST & GUINOT, 1961: 56, fig. 41 (Mangareva); 1962: 62 (Biogeography "Tahiti- Tuamotu”). — GUINOT, 1979: 63 (Mangareva, Marutea South); 1985 (List). — MONTEFORTE, 1984: 171, annex 1, tab. a; 1987: 9 (Moorea). Lophozozymus glaber Ortmann, 1893 DISTRIBUTION. — Society (Moorea, Tahiti). REFERENCES. — Lophozozymus glaber - GUINOT, 1979: 65, pl. 8, fig. 2, 2a (Tahiti). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — MONTEFORTE, 1984: 171, annex 1, tab. a; 1987: 9 (Moorea). — Lophozozymus dodone - FOREST & GUINOT, 1961: 54, fig. 39a-b (Tahiti) not Lophozozymus dodone (Herbst, 1801) = L. glaber fide GUINOT (1979: 65). Lophozozymus pictor (Fabricius, 1798) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Lophozozymus octodentatus - BOONE, 1934: 102, pl. 54-55 (Tahiti). — SAKAI, 1976: 407, pl. 146, fig. 3 (Syn., but without BOONE's reference) - SYNONYMS - Lophozozymus octodentatus (H. Milne Edwards, 1834). REMARK. — BOONE has examined and illustrated a large male and a female from Tahiti. However, it seems that this material has never been re-examined (cf. BUITENDIJK, 1960, or SAKAI, 1976). Lophozozymus superbus (Dana, 1852) DISTRIBUTION. — Tuamotu (Moruroa, Raraka, Raroia). REFERENCES. — Xantho superbus Dana, 1852a: 74; 1852b: 167; 1855, pl. 8, fig. 5a-b (Raraka). — Lophozozymus superbus - HOLTHUIS, 1953: 23 (Raroia). — MORRISON, 1954: 13 (Raroia). — GUINOT, 1979: 63 (Moruroa); 1985: 451 (List). — Lophozozymus incisus - FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu") not Lophozozymus incisus (H. Milne Edwards, 1834) = L. superbus (Dana) (cf. Remark). — Not Lophozozymus superbus - NOBILI, 1907: 388 = L. edwardsi fide FOREST & GUINOT (1961: 56) & GUINOT (1979: 63). REMARK. — FOREST & GUINOT (1962: 62) have recorded Lophozozymus incisus (H. Milne Edwards, 1834) in French Polynesia because it has formerly been considered as a synonym of L. superbus (Dana). This opinion has been changed later on (see for example HOLTHUIS, 1953, or GUINOT, 1979). Platypodia anaglypta (Heller, 1861) DISTRIBUTION. — Society (Moorea); Tuamotu (Fakarava, Tikehau). 47 REFERENCES. — Platypodia anaglypta - RATHBUN, 1907: 38 (Fakarava). — FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 451 (List). — PEYROT-CLAUSADE, 1989: 113, 115 (Moorea, Tikehau). Platypodia granulosa (Riippell, 1830) DISTRIBUTION. — Society (Tahiti?); Tuamotu (Nukutipipi). REFERENCES. — Atergatis limbatus - ? HELLER, 1865: 8 (Tahiti). — Platypodia granulosa - ? PESTA, 1913: 41 (Tahiti). — FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — SERENE, 1984: 159, 162 (Syn.). — GUINOT, 1985: 451 (List). — MERSCHARDT-SALVAT, 1991: 89 (Nukutipipi). — Not Lophactea granulosa - NOBILI, 1907: 388 ("Rikitea" = Mangareva, Marutea). — Not Platypodia granulosa - FOREST & GUINOT, 1961: 51 (Mangareva) - These two references = Platypodia pseudogranulosa sp. nov. in SERENE (1984: 159; cf. Remark) - SYNONYMS - Atergatis limbatus (H. Milne Edwards, 1834). REMARK. — SERENE (1984: 159), for the description of Platypodia pseudogranulosa, closely related to P. granulosa, does not mention HELLER's (1865) and PEsTA's (1913) references, neither under P. granulosa, nor under P. pseudogranulosa. Thus we place these two references under both species, with uncertainty. Platypodia pseudogranulosa Seréne, 1984 DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti?); Tuamotu (Marutea South). REFERENCES. — Afergatis limbatus - ? HELLER, 1865: 8 (Tahiti; cf. Remark under P. granulosa). — Lophactaea granulosa (Riipp.) - NOBILI, 1907: 388 ("Rikitea” = Mangareva, Marutea). — Platypodia granulosa - ? PESTA, 1913: 41 (Tahiti; cf. Remark under P. granulosa). — FOREST & GUINOT, 1961: 51 (Mangareva) - NOBILI's and FOREST & GUINOT's references, not Platypodia granulosa (Riippell, 1830) = P. pseudogranulosa sp. nov. in SERENE (1984: 159). — Platypodia pseudogranulosa Seréne, 1984: 159, pl. 22d ("Rikitea” = Mangareva, Marutea; Syn.). Platypodia semigranosa (Heller, 1861) DISTRIBUTION. — Society (Moorea) - Littoral to sublittoral. REFERENCES. — Platypodia semigranosa - SERENE, 1984: 160, fig. 95, pl. 22b (Syn.; with material collected by PEYROT-CLAUSADE at Madagascar). — PEYROT-CLAUSADE, 1989: 115 (Moorea; 22m). Zozimus aeneus (Linné, 1758) DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti); Tuamotu (Fakahina, Hikueru, Makatea, Makemo, Mataiva, Moruroa, Raroia, Taiaro, Takapoto). REFERENCES. — Zozymus aeneus - DANA, 1852b: 192; 1855, pl. 10, fig. 3a (Tuamotu). — STIMPSON, 1858a: 32 [30]; 1907: 42 (Tahiti). — Noi, 1907: 388 (Fakahina). — SEURAT, 1934: 59 (Fakahina, Tahiti). — MONTEFORTE, 1984: 171, annex 1, tab. a, photograph p. 136b (Makatea, Mataiva, Takapoto). —DELESALLE, 1985: 289 (Mataiva). — SALVAT & RICHARD, 1985: 360 (Takapoto). — SALVAT, 1986b: 72 (French Polynesia). — Zozimus aeneus - RATHBUN, 1907: 38 (Makemo). — BOoNng, 1934: 99, pl. 50-53 (Nuku Hiva, Tahiti). — HOLTHUIS, 1953: 27 (Raroia). — Morrison, 1954: 16 (Raroia). — BUITENDIK, 1960: 284, fig. 6a (Society). — FOREST & GUINOT, 1961: 51 (Hikueru, Tahiti); 1962: 62 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1966a: 48 (Raroia); 1985: 451.(List). — BONVALLOT et al., 1994: 140-141, photograph (Tuamotu). — PouPIN, 1994a: 37, fig. 33, pl. 4d (Tahiti, Taiaro). — ? Zozimus sp. - SALVAT, 1986a: 19, photograph (French Polynesia; det. according to the photograph). — ? Lophozozymus sp. - BAGNIS & CHRISTIAN, 1983: 110, photograph (Tuamotu; det. according to the photograph) - NEW MATERIAL - Coll. and det. J. POUPIN (Moruroa). Zozymodes pumilus (Jacquinot, 1852) DISTRIBUTION. — Gambier?; Society (Tahiti); Tuamotu (Hikueru, Makemo). REFERENCES. — Xanthodius cristatus - RATHBUN, 1907: 41 (Makemo). — Zozymodes carinipes - NOBILI, 1907: 388 ("Tagatau" = Gambier?) not Zozymodes carinipes Heller, 1861 synonym of Z. xanthoides (Krauss, 1843) = Z. pumilus fide FOREST & GUINOT (1961: 52). — Zozymodes pumilus - FOREST & GUINOT, 1961: 52, fig. 36a-b (Hikueru); 1962: 48 62 (Biogeography "Tahiti-Tuamotu”). — SERENE, 1984: 153, fig. 90, pl. 14e (Tahiti; Syn.). — GumNot, 1985: 452 (List) - SYNONYMS - Leptodius cristatus Borradaile, 1902. Zozymodes xanthoides (Krauss, 1843) DISTRIBUTION. — Tuamotu (Takapoto, Tikehau). REFERENCES. — Zozymodes xanthoides - MONTEFORTE, 1984: 171, annex 1, tab. a (Takapoto). — GUINOT, 1985: 452, with a ? (List). — Zozymoides xanthoides (sic) - PEYROT-CLAUSADE, 1989: 113 (Tikehau). SUBFAMILY XANTHINAE Lachnopodus bidentatus (A. Milne Edwards, 1867) DISTRIBUTION. — Society (Moorea, Tahiti). REFERENCES. — Xantho arcuatus Heller, 1865: 11, pl. 2, fig. 1 (Tahiti). — Lachnopodus bidentatus - FOREST & GUINOT, 1961: 42, fig. 29-30, 32bis, 33a-b, pl. 7, fig. 1-2 (Tahiti; Syn.); 1962: 62 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a; 1987: 9 (Moorea). — GUINOT, 1985: 250 (List). Lachnopodus ponapensis (Rathbun, 1907) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Xanthias ponapensis Rathbun, 1907: 44, pl. 7, fig. 5, 5a (Tahiti). — Paraxanthias ponapensis - FOREST & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 451 (List). — Lachnopodus ponapensis - SERENE, 1984: 203 (Key). Lachnopodus subacutus (Stimpson, 1858) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Mataiva). REFERENCES. — Lachnopodus subacutus - PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva, Moorea, Tahiti); 1987: 9 (Moorea). — SERENE, 1984: 204, fig. 122, pl. 29a (Syn.). — GUINOT, 1985: 450, with a ? (List). Lachnopodus tahitensis De Man, 1889 DISTRIBUTION. — Society (Tahiti); Tuamotu (Makatea, Raroia, Takapoto). REFERENCES. — Xantho (Lachnopodus) tahitensis de Man, 1889: 418, pl. 9, fig. 4, 4a; 1890: 52 (Tahiti). — Lachnopodus tahitensis - HOLTHUIS, 1953: 22 (Raroia). — MORRISON, 1954: 16 (Raroia). — FOREST & GUINOT, 1961: 49 (Tahiti); 1962: 62 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (Raroia); 1985: 450 (List). — MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Takapoto). — SERENE, 1984: 203, fig. 123, pl. 29d (Tahiti). Leptodius davaoensis Ward, 1941 DISTRIBUTION. — Tuamotu (Hao, Hikueru, Mataiva, Moruroa, Takapoto). REFERENCES. — Leptodius leptodon Forest & Guinot, 1961: 65, fig. 55-56, 59a-b, pl. 2, fig. 3 (Hikueru) fide TAKEDA (1980: 318); 1962: 64 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva, Takapoto). — GUINOT, 1985: 450 (List). — Leptodius exaratus - NOBILI, 1907: 389 (Hao) not Leptodius exaratus (H. Milne Edwards, 1834) = L. leptodon nov. in FOREST & GUINOT (1961: 65). — Leptodius davaoensis - TAKEDA, 1980: 318 (Syn.). — POUPIN, 1994a: 38, fig. 34, pl. 4e (Hikueru, Moruroa) - RELEVANT MATERIAL - Leptodius cf. davaoensis - MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva). — GUINOT, 1985: 450 (List). Leptodius exaratus (H. Milne Edwards, 1834) DISTRIBUTION. — Marquesas (Nuku Hiva); Society; Tuamotu (Raroia). neni iii ieee aaa ee 49 REFERENCES. — Leptodius exaratus - BOONE, 1934: 110, pl. 58 (Nuku Hiva). — FoREsT & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu, ?Marquesas"). — SERENE, 1984: 183, fig. 106, pl. 26a (Syn.). — GUINOT, 1985: 450 (List). — Xantho exaratus - HOLTHUIS, 1953: 27 (Raroia). — MORRISON, 1954: 7 (Raroia). — BUITENDUK, 1960: 331, fig. 9k-m (Society). — Not Leptodius exaratus (H. Milne Edwards) - NoBILI, 1907: 389 = Leptodius leptodon nov. in ForEST & GUINOT (1961: 65) synonym of L. davaoensis fide TAKEDA (1980: 318). Leptodius gracilis (Dana, 1852) DISTRIBUTION. — Gambier (Rikitea); Tuamotu (Hikueru, Moruroa, Rangiroa, Raroia). REFERENCES. — Leptodius gracilis - NOBILI, 1907: 389 (Rikitea). — FOREST & GUINOT, 1961: 64, fig. 57, 58a-b, pl. 2, fig. 4 (Hikueru); 1962: 62 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 450 (List). — PoUPIN, 1994a: 39, fig. 35, pl. 4f (Hikueru, Moruroa). — Xantho gracilis - HOLTHUIS, 1953: 27 (Raroia). — MORRISON, 1954: 7 (Raroia). — BUITENDUK, 1960: 335 (Rangiroa). Leptodius sanguineus (H. Milne Edwards, 1834) DISTRIBUTION. — Gambier (Kamaka, Mangareva, Tarauru-Roa); Marquesas (Nuku Hiva); Society (Bora Bora, Moorea,Tahiti); Tuamotu (Ahe and/or Manihi, Fakarava, Makatea, Makemo, Marutea South, Mataiva, Rangiroa, Taiaro, Takapoto). REFERENCES. — Chlorodius sanguineus - DANA, 1852a: 79; 1852b: 207; 1855, pl. 11, fig. 11a-d ("Waterland' = Ahe and/or Manihi). — Leptodius sanguineus - NoBILI, 1907: 389 (“Rikitea” = Mangareva, Kamaka, Marutea South). — RATHBUN, 1907: 39 (Bora Bora, Fakarava, Makatea, Makemo, Mangareva, Nuku Hiva, Rangiroa, Tahiti). — SENDLER, 1923: 37 (Makatea, Tahiti). — BOONE, 1934: 116, pl. 60-61 (Nuku Hiva). — FoREST & GUINOT, 1961: 63, fig. 50a-b ("Gatavake" = Mangareva, Tahiti, "Taraourou-roa” = Tarauru-Roa); 1962: 62 (Biogeography "Tahiti-Tuamotu, Marquesas"). — MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — SERENE, 1984: 185 (Syn.). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 450 (List). — POUPIN, 1994a: 40, fig. 36, pl. 4g (Nuku Hiva, Tahiti, Taiaro). — Xantho sanguineus - BUITENDUK, 1960: 323 (Nuku Hiva). Lioxanthodes alcocki Calman, 1909 DISTRIBUTION. — Society (Moorea); Tuamotu (Makatea, Mataiva, Takapoto, Tikehau). REFERENCES. — Lioxanthodes alcocki - MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Moorea, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 451 (List). — PEYROT- CLAUSADE, 1989: 113 (Tikehau). Macromedaeus crassimanus (A. Milne Edwards, 1867) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Xantho crassimanus - BUITENDLK, 1960: 318, fig. 9c-f (Tahiti). — Leptodius crassimanus - FOREST & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — Macromedaeus crassimanus - SERENE, 1984: 179, fig. 103, pl. 25b (Syn.). — GUINOT, 1985: 451 (List). Macromedaeus distinguendus (de Haan, 1835) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Macromedaeus distinguendus - KIM, 1973: 630 (Distribution only, Tahiti). — GUINOT, 1985: 451 (Society; List). REMARK. — GUINOT (1985) has listed this species after FOREST & GUINOT (1961). These authors have actually examined some Xantho distinguendus de Haan, 1835 (p. 57, under Medaeus noelensis Ward, 1934), but they are from Hong Kong. It could be that GUINOT refers to KIM (1973), who has quoted "Tahiti" in the distribution of this species. However, this location concerns neither the material examined, nor one of the references cited by KIM under Macromedaeus distinguendus. The presence of this species in French Polynesia remains thus doubtful. 50 Macromedaeus nudipes (A. Milne Edwards, 1867) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Xantho nudipes - FOREST & GUINOT, 1961: 59, fig. 47a-b. (Tahiti); 1962: 62 (Biogeography "Tahiti- Tuamotu"). — Macromedaeus nudipes - SERENE, 1984: 178, fig. 101, pl. 25a (Syn.). — GUINOT, 1985: 451 (List). Neoxanthops cavatus (Rathbun, 1907) DISTRIBUTION. — Tuamotu (Fakarava). REFERENCES. — Cycloxanthops cavatus Rathbun, 1907: 41, pl. 5, fig. 8, pl. 6, fig. 3, 3a (Fakarava). — FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — Neoxanthops cavatus - SERENE, 1984: 212, fig. 128, pl. 29f (Syn.). — GUINOT, 1985: 451 (List). Paraxanthias notatus (Dana, 1852) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava, Makatea, Makemo, Marutea North?, Marutea South, Mataiva, Takapoto, Tikehau). REFERENCES. — Xanthodes notatus Dana, 1852a: 76 (Tahiti, Tuamotu); 1852b: 178; 1855, pl. 8, fig. 12a-b. (Society or Tuamotu). — Xanthias notatus - NOBILI, 1907: 392 (Fakarava, Makatea, Marutea = Marutea North?, Marutea South). — RATHBUN, 1907: 45 (Fakarava, Makemo). — Paraxanthias notatus - FOREST & GUINOT, 1961: 76, fig. 70a-b (Tahiti); 1962: 64 (Biogeography "Tahiti-Tuamotu”). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 451 (List). — PEYROT-CLAUSADE, 1989: 113, 115 (Moorea, Tikehau). Xanthias canaliculatus Rathbun, 1907 DISTRIBUTION. — Society (Moorea); Tuamotu (Makemo). REFERENCES. — Xanthias canaliculatus Rathbun, 1907: 45 (Makemo). — FOREST & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212 (Moorea). — GUINOT, 1985: 451 (List). Xanthias lamarcki (H. Milne Edwards, 1834) DISTRIBUTION. — Gambier (Tarauru-Roa, Temoe); Society (Bora Bora, Moorea, Raiatea, Tahiti); Tuamotu (Fakarava, Hikueru, Makatea, Makemo, Marutea South, Mataiva, Raroia, Takapoto, Tikehau). REFERENCES. — Xanthodes granoso-manus Dana, 1852a: 75; 1852b: 175; 1855, pl. 8, fig. 10a-c (Society, Tuamotu). — Xanthias lamarckii - NoBILI, 1907: 393 (Hikueru, "Timoe" = Temoe). — RATHBUN, 1907: 44 (Bora Bora, Fakarava, Makemo, Tahiti). — SENDLER, 1923: 37 (Makatea). — BOONE, 1934: 131, pl. 70 (Raiatea, Tahiti). — HOLTHUIS, 1953: 26 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST & GUINOT, 1961: 70, fig. 63, 66a-b (Hikueru, Marutea, Tahiti, "Taraourou-roa” = Tarauru-Roa); 1962: 64 (Biogeography "Tahiti-Tuamotu"), — PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212 (Moorea); 1989: 113 (Tikehau). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — SERENE, 1984: 195, fig. 112, pl. 27b (Syn.). — DELESALLE, 1985: 305 (Mataiva). — GUINOT, 1985: 451 (List). Xanthias latifrons (De Man, 1888) DISTRIBUTION. — Society (Moorea, Tahiti). REFERENCES. — Xanthias latifrons - FOREST & GUINOT, 1961: 70, fig. 67a-b (Tahiti); 1962: 64 (Biogeography "Tahiti- Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a; 1987: 9 (Moorea). — GUINOT, 1985: 451 (List). Xanthias nitidulus (Dana, 1852) DISTRIBUTION. — Tuamotu (Marutea South). REFERENCES. — Xanthodes nitidulus Dana, 1852a: 76; 1852b: 177; 1855, pl. 8, fig. 1la-c (Tuamotu). — Xanthias nitidulus - NOBILI, 1907: 392 (Marutea South). i i a Ee EEE EET ——EEE— eee 51 REMARK. — FOREST & GUINOT (1961: 72) have placed these two references under Xanthias tetraodon, with a ? and this remark "Nous nous abstiendrons pour l'instant de tirer des conclusions définitives, mais il est probable que lorsqu’on disposera d'une série de Xanthia tetraodon de diverses tailles, l'on sera amené a désigner cette espéce sous le nom de Xanthias nitidilus (Dana)". Xanthias punctatus (H. Milne Edwards, 1834) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Xanthias punctatus - FOREST & GUINOT, 1961: 68, fig. 61, 65a-b (Tahiti); 1962: 64 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 451 (List). Xanthias tetraodon (Heller, 1865) DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti); Tuamotu (Hao, Hikueru, Makatea, Marutea South?, Mataiva, Raroia, Takapoto). REFERENCES. — Eudora tetraodon Heller, 1865: 14, pl. 2, fig. 3 (Auckland = ? Tahiti in FOREST & GUINOT, 1961). — Xantho (Eudora) tetraodon - NoBILI, 1907: 389 (Hao). — Juxtaxanthias tetraodon - WARD, 1942: 92 (Mangareva). — HOLTHUIS, 1953: 22 (Raroia). — MORRISON, 1954: 16 (Raroia). — Xanthias tetraodon - FOREST & GUINOT, 1961: 72, fig. 61, 68a-c, 69bis (Hikueru, Tahiti); 1962: 64 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (Raroia); 1985: 451 (List). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). REMARK. — ODHNER (1925) and FOREST & GUINOT (1961) consider that the type locality, Auckland, mentioned by HELLER (1865) for the description of Eudora tetraodon, is a mistake, and that it could very likely be Tahiti. Moreover, FOREST & GUINOT (1961) mention that this species could be a synonym of Xanthias nitidilus (Dana, 1852) (cf. above). SUBFAMILY PANOPEINAE Panopeus pacificus Edmondson, 1931 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Panopeus pacificus - FOREST & GUINOT, 1961: 116, fig. 102, 103a-b, 104, 10Sa-b, pl. 4, fig. 3 (Tahiti); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List). SUBFAMILY KRAUSSIINAE Palapedia marquesas (Seréne, 1972) DISTRIBUTION. — Tuamotu (Anaa). REFERENCES. — Kraussia marquesas Seréne, 1972: 53, fig. 14-15, 23g, k (Anaa). — GUINOT, 1985: 449 (List). — Palapedia marquesas - NG, 1993: 141 (subfamily nov. and gen. nov.). Palapedia rastripes (Miller, 1887) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Kraussia rastripes - MONTEFORTE, 1984: 171, annex 1, tab. a (Tahiti). — GUINOT, 1985: 449 (List). — Palapedia rastripes - NG, 1993: 141 (subfamily nov. and gen. nov.). SUBFAMILY ETISINAE Etisus bifrontalis (Edmonson, 1935) DISTRIBUTION. — Tuamotu (Hikueru). 52 REFERENCES. — Etisodes electra - NOBILI, 1907: 390 (Hikueru) pro parte not Etisodes electra (Herbst, 1801) = Etisus aff. bifrontalis fide GUINOT (1964: 56, 61; cf. Remark under E. electra). — Efisus aff. bifrontalis - GUINOT, 1964: 61 (Hikueru); 1985: 450 (List) = E. bifrontalis fide SERENE (1984: 230). Etisus anaglyptus H. Milne Edwards, 1834 DISTRIBUTION. — Society (Moorea). REFERENCES. — Efisus anaglyptus - MONTEFORTE, 1984: 170, annex 1, tab. a; 1987: 9 (Moorea). — GUINOT, 1985: 450 (List). Etisus demani Odhner, 1925 DISTRIBUTION. — Society (Tahiti); Tuamotu (Hikueru?). REFERENCES. — Efisus demani - MONTEFORTE, 1984: 170, annex 1, tab. a (Tahiti). — SERENE, 1984: 227, fig. 140, 143a, pl. 31f (Hikueru?; cf. Remark). — GUINOT, 1985: 450 (List). REMARK. — The location "Hikueru" in SERENE (1984: 227) is only mentioned in the observations. It is not indicated under the material examined, and was not retrieved in the references cited by SERENE. It could be an erroneous reading in GUINOT (1964: 59), where "Hikueru" is cited under Etisus frontalis Dana, just beneath E. demani Odhner. Etisus dentatus (Herbst, 1785) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Taiaro). REFERENCES. — Efisus dentatus - BOONE, 1934: 119, pl. 62-63 (Tahiti). — FOREST & GUINOT, 1961: 86, fig. 80a-b (Tahiti); 1962: 64 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, annex 1, tab. a (Moorea, Tahiti); 1987: 9 (Moorea). — GUINOT, 1985: 450 (List). — POUPIN, 1994a: 41, fig. 37, pl. 4h (Tahiti, Taiaro). Etisus electra (Herbst, 1801) DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti); Tuamotu (Fakarava, Hikueru, Marutea South). REFERENCES. — Efisus rugosus Jacquinot, 1852, pl. 4, fig. 2. — JacQUINOT & Lucas, 1853: 33 (Mangareva). — Etisodes electra - NOBILI, 1907: 390 (Hikueru, "Rikitea" = Mangareva, Marutea South) pro parte cf. Remark. — RATHBUN, 1907: 42 (Fakarava). — FOREST & GUINOT, 1961: 89, fig. 82a-b (Tahiti); 1962: 66 (Biogeography "Tahiti- Tuamotu"). — GUINOT, 1985: 450 (List). — Etisus electra - MONTEFORTE, 1984: 170, annex 1, tab. a (Tahiti). — SERENE, 1984: 228 (Syn.). REMARK. — GUINOT (1964: 54) indicates that the material from Hikueru, attributed by NOBILI to E. electra, includes in fact three species: E. electra, Etisus frontalis (Dana, 1852), and E. aff. bifrontalis Edmonson, 1935). Etisus frontalis (Dana, 1852) DISTRIBUTION. — Society (Moorea); Tuamotu (Hikueru). REFERENCES. — Etisodes electra - NOBILI, 1907: 390 (Hikueru) pro parte not Etisus electra (Herbst, 1801) = E. frontalis fide GUINOT (1964: 54; cf. Remark under E. electra). — Etisus frontalis - GUINOT, 1964: 54 (Hikueru). — ODINETZ, 1983: 209 (Moorea). — SERENE, 1984: 229, fig. 139, pl. 3le (Syn.). — GUINOT: 1985: 450 (List). — Etisodes frontalis - GUINOT, 1985: 450 (List; cf. Remark). REMARK. — In her list, GUINOT (1985: 450) mentions the two following species: "Etisodes frontalis (Dana, 1852) and Etisus frontalis Dana, 1852". It is obviously a mistake for a single species, described under Etisodes frontalis by DANA (1852b: 187). Etisus laevimanus Randall, 1839 DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora); Tuamotu (Mataiva). REFERENCES. — Etisus macrodactylus - JACQUINOT, 1852, pl. 9, fig. 2 (Mangareva). — JACQUINOT & Lucas, 1853: 30 (Mangareva). — Etisus laevimanus - NoBILI, 1907: 390 (""Rikitea" = Mangareva). — RATHBUN, 1907: 42 (Bora Bora). — Forest & GUINOT, 1961: 88 (Mangareva); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, a Te RTE rs NO 53 annex 1, tab. a (Mataiva). — SERENE, 1984: 225 (Syn.). — GUINOT, 1985: 450 (List). — DELESALLE, 1985: 289 (Mataiva). — POUPIN, 1994a: 42, fig. 38, pl. 5a (Mangareva) - SYNONYMS - Etisus macrodactylus Bianconi, 1851. Etisus punctatus Jacquinot, 1852 DISTRIBUTION. — Gambier (Mangareva). REFERENCES. — Etisus punctatus Jacquinot, 1852, pl. 3, fig. 5. — JACQUINOT & LUCAS, 1853: 31 (Mangareva). REMARK. — This Etisus has been figured by JACQUINOT, then described by JACQUINOT & LUCAS with this commentary "Cette espéce n'ayant pas été déposée au Muséum, c'est d'aprés la figure qui en a été donnée par MM. HOMBRON et JACQUINOT que nous avons fait cette description". According to JACQUINOT & LUCAS, Etisus punctatus is related to E. macrodactylus Bianconi, 1851 (= E. laevimanus Randall, 1851) and to E. anaglyptus H. Milne Edwards, 1834. Etisus splendidus Rathbun, 1906 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru, Mataiva?, Raroia, Taiaro, Takapoto). REFERENCES. — Etisus (Etisodes) splendidus - HOLTHUIS, 1953: 21 (Raroia). — MORRISON, 1954: 16 (Raroia). — Etisus splendidus - FOREST & GUINOT, 1961: 88, fig. 8la-c (Hikueru); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (Raroia); 1985: 450 (List). — MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva?, Moorea, Tahiti, Takapoto?). — SALVAT, 1986b: 72, photograph (French Polynesia). — BONVALLOT et al., 1994: 145 (Tuamotu). — POUPIN, 1994a: 43, fig. 39, pl. 5b (Taiaro, Takapoto). SUBFAMILY CHLORODIINAE Chlorodiella barbata (Borradaile, 1900) DISTRIBUTION. — Gambier (Mangareva); Society (Moorea, Tahiti); Tuamotu (Mataiva, Marutea South, Takapoto, Tikehau). REFERENCES. — Chlorodiella barbata - FOREST & GUINOT, 1961: 96, fig. 93-94, 99a-b, 100 (Mangareva, Marutea South, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu”). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 213; 1985: 462 (Moorea); 1989: 112 (Tikehau). — NAIM, 1980a, annex 1, tab. 3; 1980b: 550 (Moorea). — THOMASSIN et al., 1982: 394 (Moorea). — ODINETZ, 1983: 97 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GALZIN & POINTIER, 1985: 100 (Moorea). — GUINOT, 1985: 450 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti). — SALVAT & RICHARD, 1985: 360 (Takapoto). Chlorodiella cytherea (Dana, 1852) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru, Mataiva, Raraka, Takapoto, Tikehau). REFERENCES. — Chlorodius cytherea Dana 1852a: 79; 1852b: 213; 1855, pl. 12, fig. 2a-c (Raraka, Tahiti). — Chlorodiella cytherea - FOREST & GUINOT, 1961: 95, fig. 90-92, 98a-b (Hikueru, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu”). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 213 (Moorea); 1989: 112 (Tikehau). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 450 (List). Chlorodiella laevissima (Dana, 1852) DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Hao, Makatea, Makemo, Mataiva, Rangiroa, Takapoto, Tikehau) - Littoral to sublittoral. REFERENCES. — Chlorodius laevissimus - NOBILI, 1907: 393 (‘‘Rikitea, Gatavake" = Mangareva, "Ohura" = Hao). — Chlorodiella laevissima - RATHBUN, 1907: 46 (Fakarava, Makemo, Rangiroa, Tahiti; 46m). — FOREST & GUINOT, 1961: 95, fig. 95-96, 101a-b (Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 213; 1985: 462 (Moorea); 1989: 111, 114 (Moorea, Tikehau). — ODINETZ, 1983: 209 (Moorea, Tahiti). — MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 54 (Moorea). — SERENE, 1984: 260, fig. 171-172, pl. 36d-e (Syn.). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 450 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti). — Chlorodiella laevissimus - BOONE, 1934: 138, pl. 72 (Nuku Hiva, Tahiti). Chlorodiella nigra (Forskal, 1775) DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Hao, Makatea, Makemo, Marutea South, Mataiva, Nukutipipi, Rangiroa, Raraka, Takapoto, Tikehau). REFERENCES. — Chlorodius niger - HELLER, 1865: 18 (Tahiti). — STIMPSON, 1858a: 33; 1907: 50 (Tahiti). — NosILI, 1907: 393 (""Gatavake" = Mangareva, Hao, Marutea, "Tikahan" = Tikehau). — Chlorodiella niger - RATHBUN, 1907: 46 (Bora Bora, Fakarava, Makemo, Rangiroa). — SENDLER, 1923: 38 (Makatea). — SEURAT, 1934: 59 ("Gatavake" = Mangareva, Hao, Marutea South, "Tikahau" = Tikehau). — Chlorodiella nigra - FOREST & GUINOT, 1961: 95, fig. 87- 89, 97a-b (Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 213 (Moorea). — KROPP & BIRKELAND, 1981: 630, tab. 5 (Moorea, Takapoto). — THOMASSIN et al., 1982: 394 (Moorea). — MONTEFORTE, 1984: 170, annex 1, tab. a, photograph p. 136a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — SERENE, 1984: 258, fig. 168, pl. 36b (Syn.). — DELESALLE, 1985: 288 (Mataiva). — GUINOT, 1985: 450 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti, Takapoto). — MERSCHARDT- SALVAT, 1991: 89 (Nukutipipi). Garthiella aberrans (Rathbun, 1906) DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral. REFERENCES. — Pilodius aberrans - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 212 (Moorea); 1989: 111, 115 (Moorea, Tikehau; 30m). — GUINOT, 1985: 451 (List). — Garthiella aberrans - TITGEN, 1986: 57, fig. 1-2 (Syn.). Liocarpilodes armiger (Nobili, 1906) DISTRIBUTION. — Society (Moorea); Tuamotu (Tikehau) - Littoral to sublittoral. REFERENCES. — Liocarpilodes armiger - PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 213 (Moorea); 1989: 111, 115 (Moorea, Tikehau; 30m). — GUINOT, 1985: 450, with a ? (List). Liocarpilodes harmsi (Balss, 1934) DISTRIBUTION. — Society (Moorea). REFERENCES. — Pilodius harmsi - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 213 (Moorea). — Liocarpilodes harmsi - SERENE, 1984: 264 (Syn.). Liocarpilodes integerrimus (Dana, 1852) DISTRIBUTION. — Gambier (Mangareva); Society (Moorea, Tahiti); Tuamotu (Fakarava, Mataiva, Takapoto, Tikehau). REFERENCES. — Actumnus integerrimus - RATHBUN, 1907: 56, pl. 1, fig. 12, pl. 8, fig. 3, 3a-b (Fakarava, Tahiti). — Pilumnus margaritatus - NOBILI, 1907: 398 ("Rikitea" = Mangareva) not Pilumnus margaritatus Ortmann, 1893 = L. integerrimus fide GUINOT (1964: 63). — Liocarpilodes integerrimus - FOREST & GUINOT, 1962: 64 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1964: 63, fig. 36a-b ("Rikitea" = Mangareva). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 213; 1985: 462 (Moorea); 1989: 111, 115 (Moorea, Tikehau). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 450 (List). Phymodius granulosus (De Man, 1888) DISTRIBUTION. — Society (Moorea and/or Tahiti). REFERENCES. — Phymodius granulosus - GUINOT, 1985: 451, with a ? (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti). 55 REMARK. — ODINETZ does not mention this material in her thesis (1983), but it is recorded in the Museum of Paris (MNHN B17071, coll. ODINETZ 1981 "Moorea, Tahiti, Takapoto, associé au Pocillopora damicornis et P. elegans", det. GUINOT). Phymodius monticulosus (Dana, 1852) DISTRIBUTION. — Gambier (Mangareva); Society (Moorea, Tahiti); Tuamotu (Fakahina, Fakarava, Mataiva, Marutea South, Takapoto). REFERENCES. — Chlorodius monticulosus Dana, 1852a: 79; 1852b: 206; 1855, pl. 11, fig. 9a (Tahiti). — STIMPSON, 1858a: 31; 1907: 50 (Tahiti). — Chlorodius Dehaanii - HELLER, 1865: 19 (Tahiti) pro parte, some sp. attributed to Phymodius ungulatus in FOREST & GUINOT (1961: 106, 114), not C. Dehaani, synonym of Phymodius granulatus (Targioni Tozzetti, 1877) in SERENE (1984: 250). — Cyclodius ornatus - NOBILI, 1907: 397 (Fakahina, Marutea South). — RATHBUN, 1907: 51, pl. 5, fig. 5, pl. 7, fig. 8 (Fakarava, Tahiti). — Phymodius monticulosus - FOREST & GUINOT, 1961: 106, pl. 10, fig. 1-6 (Fakahina, Marutea South, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — GUINOT, 1985: 451 (List). — POUPIN, 1994a: 44, fig. 40, pl. Sc (Fakahina, Mangareva) - SYNONYMS - Cyclodius ornatus Dana, 1852. Phymodius nitidus (Dana, 1852) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu ( Kaukura, Tikehau). REFERENCES. — Pilodius nitidus - NOBILI, 1907: 393 (Kaukura). — Chlorodopsis scabricula - RATHBUN, 1907: 50, pl. 1, fig. 3, pl. 9, fig. 5 (Tahiti) not Pilodius scabriculus (Dana, 1852) = Phymodius nitidus fide FOREST & GUINOT (1961: 114). — Phymodius nitidus - FOREST & GUINOT, 1961: 114, pl. 15, fig. 1-4 (Kaukura, Tahiti); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a; 1987: 9 (Moorea). — GUINOT, 1985: 451 (List). — PEYROT-CLAUSADE, 1989: 113 (Tikehau). Phymodius ungulatus (H. Milne Edwards, 1834) DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Raiatea, Tahiti); Tuamotu (Fakahina, Fakarava, Hikueru, Makemo, Mataiva, Rangiroa, Raroia, Takapoto, Tikehau). REFERENCES. — Chlorodius ungulatus - DANA, 1852b: 205; 1855, pl. 11, fig. 8a-b (Tahiti). — Chlorodius dehaanii - HELLER, 1865: 19 (Tahiti) pro parte fide FOREST & GUINOT (1961: 110), not C. Dehaani, synonym of Phymodius granulatus (Targioni Tozzetti, 1877) in SERENE (1984: 250). — Cyclodius gracilis - NOBILI, 1907: 397 (Fakahina, "Rikitea" = Mangareva). — Phymodius ungulatus - RATHBUN, 1907: 46, pl. 3-4 (Bora Bora, Fakarava, Makemo, Rangiroa, Tahiti) pro parte fide FOREST & GUINOT (1961: 110; cf. Remark). — NOBILL 1907: 393 ("Rikitea" = Mangareva). — BOONE, 1934: 140, pl. 73 (Raiatea, Tahiti) pro parte fide FOREST & GUINOT (1961: 110). — HOLTHUIs, 1953: 25 (Raroia). — FOREST & GUINOT, 1961: 110, fig. 86a-b, pl. 11, fig. 1-4, pl. 12, fig. 1-4, pl. 13, fig. 1-3, pl. 14, fig. 1-3 (Fakahina, Hikueru, Mangareva, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1964: 74, fig. 38 (Tahiti); 1985: 451 (List). — PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 213 (Moorea); 1989: 113 (Tikehau). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — SERENE, 1984: 251, fig. 158, 161, pl. 35e (Tahiti; Syn.). — DELESALLE, 1985: 288 (Mataiva). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti, Takapoto). — SALVAT & RICHARD, 1985: 356 (Takapoto) - SYNONYMS - Cyclodius gracilis Dana, 1852. REMARK. — FOREST & GUINOT (1961: 111) indicate that RATHBUN's material is only partially assignable to this species, but without mentioning the localities referring to the real Phymodius ungulatus. Thus, for this species, some of the islands mentioned by RATHBUN are doubtful. Pilodius areolatus (H. Milne Edwards, 1834) DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Hikueru, Makemo, Marutea South, Mataiva, Raroia). REFERENCES. — Actaeodes affinis Dana, 1852a: 78; 1852b: 197; 1855, pl. 11, fig. 3 (Tahiti, Tuamotu). — Actaea affinis - RATHBUN, 1907: 42 (Makemo). — Chlorodopsis areolata - NOBILI, 1907: 396, pl. 2, fig. 3 (Hikueru, "Rikitea" 56 = Mangareva, Marutea South). — HOLTHUIS, 1953: 15 (Raroia). — MORRISON, 1954: 13 (Raroia). — Pilodius areolatus - FOREST & GUINOT, 1961: 90 (Hikueru, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva). — GUINOT, 1985: 451 (List). — CLARK & GALIL, 1993: 1125, fig. la-g, 31a, 40a, 44b (Bora Bora, "Maharepa, Afareaitu, Temae" = Moorea; Syn.). Pilodius flavus Rathbun, 1893 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hao, Tikehau). REFERENCES. — Pilodius pubescens - NoBILI, 1907: 395, with a ? ("Ohura" = Hao) not Pilodius pubescens Dana, 1852 = P. flavus fide CLARK & GALIL (1993: 1130, 1146). — Pilodius flavus - FOREST & GUINOT, 1961: 95 (Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 27 (Moorea); 1989: 111, 115 (Moorea, Tikehau). — GUINOT, 1985: 451 (List). — CLARK & GALIL, 1993: 1130, fig. 4a-g, 32b, 40d, 41a (Syn.). Pilodius paumotensis Rathbun, 1907 DISTRIBUTION. — Society (Bora Bora); Tuamotu (Fakarava, Makemo, Marutea South, Tikehau). REFERENCES. — Chlorodopsis granulatus - NoBILI, 1906: 396 (Marutea South) not Pilodius granulatus Stimpson, 1859 = P. paumotensis fide GUINOT (1962: 238). — Pilodius paumotensis Rathbun 1907: 52, pl. 8, fig. 2, 2a-b (Fakarava, Makemo). — FOREST & GUINOT, 1962: 66 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 451 (List). — PEYROT-CLAUSADE, 1989: 113 (Tikehau). — CLARK & GALIL, 1993: 1143, fig. 10a-g, 35b, 43a (Bora Bora, Fakarava, Makemo, Marutea South). Pilodius pubescens Dana, 1852 DISTRIBUTION. — Society (Moorea). REFERENCES. — Pilodius pubescens - GUINOT, 1985: 451 (List; probably after NOBILI, cf. under P. flavus). — CLARK & GALIL, 1993: 1146, fig. 12a-g, 36b, 43b (""Ternae" = Temae at Moorea, Society). — Not Pilodius pubescens - NOBILI, 1907: 395, with a ? = P. flavus in CLARK & GALIL (1993: 1130). Pilodius pugil Dana, 1852 DISTRIBUTION. — Gambier (Mangareva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Hikueru, Makaiea, Makemo, Mataiva, Rangiroa, Takapoto, Tikehau). REFERENCES. — Chlorodopsis pugil - NOBILI, 1907: 395 ("Rikitea" = Mangareva). — Chlorodopsis spinipes - RATHBUN, 1907: 50, pl. 2, fig. 5 (Bora Bora, Fakarava, Makemo, Rangiroa) not Pilodius spinipes Heller, 1861 = P. pugil fide CLARK & GALIL (1983: 1149). — Pilumnus globosus - BOONE, 1934: 152, pl. 78 (Tahiti) not Globopilumnus globosus (Dana, 1852) = Pilodius pugil with a ? fide SERENE & LUOM (1959: 320). — Pilodius pugil - FOREST & GUINOT, 1961: 91 (Hikueru, Mangareva); 1962: 64 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 213; 1985: 462 (Moorea); 1989: 113 (Tikehau). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289, 305 (Mataiva). — GUINOT, 1985: 451 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea, Tahiti, Takapoto). — SALVAT & RICHARD, 1985: 358, 360 (Takapoto). — CLARK & GALIL, 1993: 1149, fig. 13a-g, 37a, 43c (Hikueru, Mangareva, "Temae, Tiahura and Afareaita" = Moorea, Tahiti, Takapoto). Pilodius scabriculus Dana, 1852 DISTRIBUTION. — Gambier (Temoe); Society (Bora Bora, Huahine, Maiao?, Moorea, Tahaa?, Tahiti); Tuamotu (Fakarava, Fakahina, Hao, Hikueru, Makatea, Makemo, Marutea North?, Marutea South, Mataiva, Raraka, Takapoto, Tikehau). REFERENCES. — Pilodius scabriculus Dana, 1852a: 80; 1852b: 220; 1855, pl. 12, fig. 9 (Raraka). — NOBILI, 1907: 394 ("Fakaina" = Fakahina, Fakarava, Marutea, Marutea South, Temoe). — FOREST & GUINOT, 1961: 91, fig. 83a-b, 84, 86bis (Hikueru, Tahiti); 1962: 66 (Biogeography "Tahiti-Tuamotu"), — PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977b: 213 (Moorea); 1989: 113 (Tikehau). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289, 305 (Mataiva). — GUINOT, 1985: 451 (List). — CLARK & GALIL, 1993: 1152, fig. 14a-g, 37b, 43d (Bora Bora, Fakahina, Hao, Hikueru, Huahine, Maiai = ee ee eeeeeeeeeeeeEeEeEeEeEeEeEeEEEEEeeeeee aff Maiao?, Marutea, Moorea, Raiatea, Tahiti, Temoe, "Tickahau" = Tikehau, "Vaiorea” = Vaitoare?, Tahaa; Syn.). — Chlorodopsis venusta Rathbun, 1907: 49, pl. 1, fig. 5 (Fakarava, Makemo). Tweedieia laysani (Rathbun, 1906) DISTRIBUTION. — Tuamotu (Tikehau). REFERENCES. — Tweedieia laysani - PEYROT-CLAUSADE, 1989: 112 (Tikehau). Tweedieia odhneri (Gordon, 1934) DISTRIBUTION. — Society (Moorea). REFERENCES. — Tweedieia odhneri - PEYROT-CLAUSADE, 1977a, annex of the species: 27; 1977: 212 (Moorea). — GUINOT, 1985: 451, with a ? (List). FAMILY TRAPEZIIDAE Jonesius triunguiculatus (Borradaile, 1902) DISTRIBUTION. — Society (Raiatea). REFERENCES. — Jonesius triunguiculatus - GALIL & TAKEDA, 1986: 165, fig. 1-4 (""Tetaro” = Raiatea). Quadrella lewinsohni Galil, 1986 DISTRIBUTION. — Marquesas (Tahuata). REFERENCES. — Quadrella sp. - MONOD, 1979: 9, fig. 1-8 (Tahuata). — Quadrella cyrenae - SERENE, 1975: 510, fig. 3- 4, pl. 1b’, e' (Tahuata; MONOD's material) not Q. cyrenae Ward, 1942, synonym of Q. maculosa Alcock, 1898 = Q. lewinsohni nov. in GALIL (1986a: 285). — GUINOT, 1985: 452 (List). — Quadrella lewinsohni Galil, 1986a: 285, fig. Sa-b, 6 (Marquesas; MONOD's and SERENE's material). Quadrella maculosa Alcock, 1898 DISTRIBUTION. — Marquesas (Fatu Hiva) - Littoral to sublittoral. REFERENCES - Quadrella maculosa - NEW MATERIAL - Coll. J. POUPIN, det. B. GALIL (Fatu Hiva; 49m). Tetralia cinctipes Paulson, 1875 DISTRIBUTION. — Austral (Rapa) - Sublittoral. REFERENCES. — Tetralia cinctipes - GALIL, 1986b: 97, fig. 1-3 (Rapa; 90m). Tetralia glaberrima (Herbst, 1790) DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Raiatea, Tahiti); Tuamotu (Aratika, Fakarava, Hikueru, Makemo, Marutea South, Tikehau). REFERENCES. — Tetralia glaberrima - DANA, 1852b: 262; 1855, pl. 16, fig. 3a-h ("Carlshoff' = Aratika, Tahiti). — ORTMANN, 1893b: 485 (Tahiti). — NOBILI, 1907: 404 (Marutea). — RATHBUN, 1907: 60 (Fakarava, Makemo, Tahiti). — BOoneg, 1934: 174, pl. 89 (Raiatea, Tahiti). — SEURAT, 1934: 59 (Tahiti, Marutea South, Marquesas). — FOREST & GUINOT, 1961: 139 (Hikueru); 1962: 70 (Biogeography "Tahiti-Tuamotu, Marquesas"). — SERENE, 1984: 281 (Syn.; cf. Remark). — GUINOT, 1985: 452 (List). — PEYROT-CLAUSADE, 1989: 111 (Tikehau). — Trapezia serratifrons Jacquinot, 1852, pl. 4, fig. 20-23. — JACQUINOT & LUCAS, 1853: 47 (Nuku Hiva). — Tetralia cavimana - HELLER, 1865: 26 (Tahiti) - SYNONYMS - Tetralia cavimana Heller, 1861. REMARK. — In SERENE (1984: 281) all the above references (except BOONE, 1934, and PEYROT-CLAUSADE, 1989) can be attributed either to T. glaberrima or toT. heterodactyla Heller, 1861. However, the specimens attributed to cavimana, and the specimens identified to glaberrima with a figure of the male pleopod, are T. glaberrima without hesitation. 58 Tetraloides nigrifrons (Dana, 1852) DISTRIBUTION. — Society (Raiatea); Tuamotu (Makemo, Pukapuka, Tikehau). REFERENCES. — Tetralia nigrifrons Dana, 1852a: 83; 1852b: 262; 1865, pl. 16, fig. 2a-d (""Honden" = Pukapuka). — Tetraloides nigrifrons - GALIL, 1985: 72, fig. 1-3 (Makemo, Tikehau, "Taoru" = Raiatea). Trapezia areolata Dana, 1852 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru, Takapoto). REFERENCES. — Trapezia areolata Dana,1852a: 83; 1852b: 259; 1855, pl. 15, fig. 8a (Tahiti). — CANO, 1888: 173 (Tahiti). — ORTMANN 1893b: 485 (Tahiti; cf. Remark). — FOREST & GUINOT, 1961: 135, fig. 133 (Hikueru). — ODINETZ, 1983: 31 (French Polynesia). — ODINETZ, 1984a: 443, fig. 3c, 4c (Moorea, Tahiti, Takapoto). — GALIL & LEWINSOMN, 1985a: 286, fig. 1, 3-4 (Tahiti). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea, Tahiti, Takapoto). — ? Trapezia ferruginea areolata - SEURAT, 1934: 59 (Tahiti). — Trapezia reticulata - KROPP & BIRKELAND, 1981: 629 (Moorea, Takapoto). — ODINETZ, 1983: 31, 205 photograph 3 (Moorea, Tahiti, Takapoto). — GUINOT, 1985: 452 (List; after ODINETZ, 1983) - All, not Trapezia reticulata Stimpson, 1858, synonym of T. septata Dana, 1852 =T. areolata fide ODINETZ (1984a: 443; does not mention KRopp & BIRKELAND, but it is probably the same material). REMARK. — In GALIL & LEWINSOHN (1985a), ORTMANN's (1893b) work, in which T. aerolata is recorded from Tahiti, New Guinea, and Palau, is cited at the same time under T. aerolata and T. septata. It is probable that the two Tahitian specimens belong to T. areolata, and the other to T. septata (opt. cit.: 291). Trapezia bella Dana, 1852 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Aratika, Hikueru, Mataiva, Takapoto). REFERENCES. — Trapezia bella Dana, 1852a: 83; 1852b: 254; 1855, pl. 15, fig. 2 ("Carlshoff' = Aratika). — FOREST & GUINOT, 1961: 133, fig. 129-130, 135a-b (Hikueru); 1962: 70 (Biogeography "Tahiti-Tuamotu"). — KrRopp & BIRKELAND, 1981: 629 (Moorea, Takapoto). — ODINETZ, 1983: 206 (Tahiti, Takapoto). — SERENE, 1984: 278 (Syn.). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Tahiti?, Takapoto). — Trapezia digitalis bella - RATHBUN, 1907: 59 (Tahiti). — Not Trapezia bella Dana - NoBILI, 1907: 403 (Hao). — SEURAT, 1934: 59 (Hao; NOBILI's material) = Trapezia speciosa fide SERENE (1984: 278). Trapezia cymodoce (Herbst, 1799) DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Tahiti); Tuamotu (Fakarava, Makemo, Rangiroa, Takapoto). REFERENCES. — Trapezia dentata - DANA, 1852b: 258; 1855, pl. 15, fig. 6a-d (Tahiti). — Trapezia hirtipes Jacquinot, 1852, pl. 4, fig. 14-16 (Nuku Hiva). — JAcQuINOT & Lucas, 1853: 44 (Nuku Hiva). — Trapezia cymodoce dentata - RATHBUN, 1907: 58 (Bora Bora, Fakarava, Makemo, Rangiroa; cf. Remark). — Trapezia cymodoce ferruginea - RATHBUN, 1907: 58 (Bora Bora, Fakarava, Makemo, Rangiroa; cf. Remark). — Trapezia ferruginea dentata - SEURAT, 1934: 59 (Tahiti). — Trapezia cymodoce - ORTMANN, 1897b: 203 (Tahiti). — SEURAT, 1934: 59 (Tahiti, Marquesas). — ODINETZ, 1983: 205, photograph 2 (Moorea, Takapoto, Tahiti); 1984a: 432, fig. 1-2 (Moorea, Tahiti, Takapoto; pro parte, some specimens would belong to T. ferruginea, cf. Remark under that species). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201(Moorea?, Tahiti, Takapoto). — GALIL & CLARK, 1990: 378 (Syn.). — Trapezia cymodoce sp.1 - KRopP & BIRKELAND, 1981: 629 (Moorea, Takapoto) fide distinction in ODINETZ (1984b: 124). — ODINETZ, 1984b: 125 (Moorea, Tahiti, Takapoto). — Not Trapezia cymodoce - DANA, 1852b: 257; 1855, pl. 15, fig. 5a-i (Tahiti). — SENDLER, 1923: 39 (Tahiti) - The two references = T. ferruginea Latreille 1825 fide GALIL & CLARK (1990: 378) - SYNONYMS - Trapezia dentata (Macleay, 1838). REMARK. — The material attributed to T. cymodoce dentata by RATHBUN (1907) has been partially re-examined by GALIL & CLARK (1990; sp. of Ellice islands "Funafuti reef") and belongs to T. cymodoce. Arbitrarily we also attribute to this species the material from French Polynesia. Concerning the T. cymodoce ferruginea also recorded by this author, from French Polynesia, Ellice, and Easter island, they would belong in part to that species (?Bora Bora, Fakarava, Makemo, Rangiroa), in part to T. guttata Riippell (2 sp. from "Mohican reef" at Rangiroa) (cf. GALIL & CLARK, 1990: 380-382), and in part to T. punctimanus (sp. from Easter island; cf. ODINETZ 1984a: 446). a 2222 E2SE<2EEEEEEEETEE 59 Trapezia digitalis Latreille, 1825 DISTRIBUTION. — Marquesas (Nuku Hiva); Tuamotu (Makatea, Takapoto). REFERENCES. — Trapezia fusca Jacquinot, 1852, pl. 4, fig. 17-19. — JACQUINOT & Lucas, 1853: 45 (Nuku Hiva). — Trapezia digitalis - SENDLER, 1923: 40 (Makatea). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu, Marquesas”). — ODINETZ, 1983: 31, 206 (Takapoto; cf. Remark). — SERENE, 1984: 277 (Syn.). — GUINOT, 1985: 452 (List). REMARK. — Although ODINETZ (1983: 31) has indicated that her material was collected at Guam, one specimen in table 9 (p. 206) is recorded from Takapoto. Trapezia ferruginea Latreille, 1825 DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Bora Bora, Moorea, Raiatea, Tahiti); Tuamotu (Takapoto). REFERENCES. — Trapezia ferruginea - DANA, 1852b: 260; 1855, pl. 16, fig. 1a-b (Tahiti, pro parte, see under T. guttata; Samoan specimens are T. serenei fide ODINETZ, 1984a: 440, 442). — BOONE, 1934: 171, pl. 88 (Bora Bora, Nuku Hiva, Raiatea). — SEURAT, 1934: 59 (Tahiti, Marquesas). — FOREST & GUINOT, 1961: 136, fig. 137a-b (Tahiti). — Kropp & BIRKELAND, 1981: 629 (Moorea). — SERENE, 1984: 273 (Syn.). — GUINOT, 1985: 452 (List). — PEYROT- CLAUSADE, 1989: 115 (Moorea). — Trapezia cymodoce - DANA, 1852b: 257; 1855, pl. 15, fig. 5a-i (Tahiti). — SENDLER, 1923: 39 (Tahiti) - The two references not Trapezia cymodoce (Herbst, 1799) = T. ferruginea fide GALIL & CLARK (1990: 380). — ODINETZ, 1984a: 432 (Moorea, Tahiti, Takapoto; pro parte, cf. Remark). — Trapezia miniata Jacquinot, 1852, pl. 4, fig. 10-13. — Jacquinot & Lucas, 1853: 43 (Nuku Hiva). — Not Trapezia cymodoce ferruginea - RATHBUN, 1907: 58 (cf. Remark under T. cymodoce). REMARK. — Trapezia ferruginea Latreille, 1825 was proposed as a synonym of T. cymodoce (Herbst, 1799) in ODINETZ (1984a), but this proposition was not followed by GALIL & CLARK (1990). Trapezia flavopunctata Eydoux & Souleyet, 1842 DISTRIBUTION. — Society (Moorea,Tahiti); Tuamotu (Hikueru). REFERENCES. — Trapezia flavopunctata - ORTMANN, 1893b: 485 (Tahiti). — FOREST & GUINOT, 1961: 136, fig. 138a-b (Hikueru). — ODINETZ, 1983: 34, 205 (Tahiti; p. 205 = "T. flavomaculata" sic). — GALIL & LEWINSOHN, 1985b: 210 ("Papetoai bay" = Moorea, Tahiti). — GUINOT, 1985: 452 (List). — Trapezia rufopunctata flavopunctata - SEURAT, 1934: 59 (Tahiti). Trapezia formosa Smith, 1869 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Mataiva, Takapoto). REFERENCES. — Trapezia formosa - KROPP & BIRKELAND, 1981: 629 (Takapoto). — ODINETZ, 1983: 206, photograph 4 (Moorea, Takapoto, Tahiti). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FoRGES, 1985: 201 (Moorea?, Takapoto, Tahiti). REMARK. — According to P. CASTRO (personal communication), who has re-examinated the material of these references, it could rather belongs to a new species. Trapezia guttata Riippell, 1830 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru, Rangiroa, Takapoto, Tikehau). REFERENCES. — Trapezia guttata - HELLER, 1865: 25 (Tahiti). — FOREST & GUINOT, 1961: 136, fig. 134, 139a-b (Hikueru, Tahiti); 1962: 70 (Biogeography "Tahiti-Tuamotu"). — ODINETZ, 1983: 205, photograph 8 (Moorea, Tahiti, Takapoto); 1984a: 442 (Moorea, Tahiti). — GuINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea?, Tahiti, Takapoto). — PEYROT-CLAUSADE, 1989: 111 (Tikehau). — GALIL & CLARK, 1990: 381 (Syn.). — Trapezia cymodoce ferruginea - RATHBUN, 1907: 58 (only some specimens from Rangiroa) not T. ferruginea Latreille, 1825 = T. guttata fide GALIL & CLARK (1990: 381; cf. Remark under T. cymodoce). — Trapezia ferruginea - DANA, 1852b: 260; 1865, pl. 16, fig. 1b (Tahiti) pro parte not T. ferruginea Latreille, 1825 = T. guttata fide GALIL & CLARK (1990: 381, 382). — Trapezia ferruginea guttata - SEURAT, 1934: 59 (Tahiti). — Trapezia 60 davaoensis - KROpP & BIRKELAND, 1981: 629 (Moorea). — GUINOT, 1985: 452 (List) - SYNONYMS - Trapezia davaoensis Ward, 1941. Trapezia punctimanus Odinetz, 1984 DISTRIBUTION. — Society (Tahiti); Tuamotu (Takapoto). REFERENCES. — Trapezia punctimanus Odinetz, 1983: 35, 206 photograph 7 (Thesis; French Polynesia); 1984a: 445, fig. 3e, 4e (Tahiti, Takapoto). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Tahiti, Takapoto). Trapezia rufopunctata (Herbst, 1801) DISTRIBUTION. — Society (Moorea, Raiatea, Tahiti); Tuamotu (Makemo, Takapoto, Tikehau). REFERENCES. — Trapezia rufo-punctata - DANA, 1852b: 255; 1855, pl. 15, fig. 3a-b (Tahiti). — Trapezia rufopunctata - RATHBUN, 1907: 57 (Makemo). — BOONE, 1934: 166, pl. 86 (Raiatea). — SEURAT, 1934: 59 (Tahiti, but not the Marquesas certainly cited after JACQUINOT & LUCAS; see below). — ODINETZ, 1983: 34 (Moorea, Tahiti, Takapoto). — GUINOT, 1985: 452 (List). — PEYROT-CLAUSADE, 1989: 112 (Tikehau). — Not Trapezia rufo-punctata - JACQUINOT, 1852, pl 4, fig. 8-9. — JACQUINOT & LucAs, 1853: 41 (Nuku Hiva) = T. tigrina Eydoux & Souleyet, 1842 fide GALL & LEWINSOHN (1985b: 166). Trapezia septata Dana, 1852 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Trapezia ferruginea areolata - SENDLER, 1923: 40 (Tahiti) not T. areolata Dana, 1852 = T. septata fide GALIL & LEWINSOHN (1985a: 288; cf. Remark). REMARK. — Although SENDLER is cited under that species in GALIL & LEWINSOMN, these authors have not examined Polynesian material. They only mention that (p. 291) "T. septata seems to be more widely distributed and more common than T. areolata. Thus, specimens identified as T. aerolata, with the exception of those mention by CANO (1888), ORTMANN (1893) (part) and FOREST & GUINOT (1961), should rightly be name T. septata". Trapezia serenei Odinetz, 1984 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Mataiva, Takapoto). REFERENCES. — Trapezia serenei Odinetz, 1983: 34, 206, photograph 6; 1984a: 440, fig. 3b, 4b (Mataiva, Moorea, Tahiti, Takapoto). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea?, Tahiti, Takapoto). — SALVAT & RICHARD, 1985: 344 (Takapoto). — Trapezia cymodoce sp. 2 - KROPP & BIRKELAND, 1981: 629 (Moorea, Takapoto). — ODINETZ, 1984b: 125 (Moorea, Tahiti, Takapoto) - Fide distinction in ODINETZ (1984b: 124). Trapezia speciosa Dana, 1852 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Aratika, Fakarava, Hao, Hikueru, Makemo, Marutea South, Mataiva, Takapoto). REFERENCES. — Trapezia speciosa Dana, 1852a: 83; 1852b: 253; 1855, pl. 15, fig. 1 ("Carlshoff' = Aratika). — NosILI, 1907: 403 (Marutea). — SEURAT, 1934: 59 (Marutea South). — FoREST & GUINOT, 1961: 133, fig. 131-132, 136a-b (Hikueru); 1962: 70 (Biogeography "Tahiti-Tuamotu"). — KROpP & BIRKELAND, 1981: 629 (Moorea, Takapoto). — ODINETZ, 1983: 205 (Moorea, Tahiti, Takapoto). — SERENE, 1984: 278 (Syn.). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea, Tahiti, Takapoto). — Trapezia digitalis speciosa - RATHBUN, 1907: 59 (Fakarava, Makemo, Tahiti). — Trapezia bella - NoBILI, 1907: 403 (Hao) not T. bella Dana, 1852 = T. speciosa fide SERENE (1984: 278). Trapezia tigrina Eydoux & Souleyet, 1842 DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Moorea, Tahiti); Tuamotu (Anaa, Makemo?, Takapoto, Tikehau). 61 REFERENCES. — Trapezia maculata - DANA, 1852b: 256; 1855, pl. 15, fig. 4 (Tahiti) not T. maculata (MacLeay, 1838) = T. tigrina fide SERENE (1984: 275) and GALIL & LEWINSHON (1984: 166). — Trapezia rufo-punctata - JACQUINOT, 1852, pl 4, fig. 8-9. — JACQUINOT & LUCAS, 1853: 41 (Nuku Hiva) not T. rufopunctata (Herbst, 1799) =T. tigrina fide GALIL & LEWINSOHN (1984: 166). — Trapezia rufopunctata var. maculata - ORTMANN, 1893b: 484 (Tahiti). — Trapezia cymodoce maculata - RATHBUN, 1907: 59 (Makemo) - These two references, with a ?, not T. maculata (MacLeay, 1838) = T tigrina fide GALIL & LEWINSOHN (1984: 167). — ? Trapezia ferruginea maculata - SEURAT, 1934: 59 (Tahiti). — Trapezia wardi - KRopP & BIRKELAND, 1981: 629 (Moorea, Takapoto). — GUINOT, 1985: 452 (List). — Trapezia tigrina - ODINETZ, 1983: 205, photograph 5 (Moorea, Tahiti, Takapoto). — GALIL & LEWINSOHN, 1984: 166, fig. 1 (Anaa, "Tikehae lagoon, Tuamotu” = Tikehau, Society; Syn.). — GUINOT, 1985: 452 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea?, Tahiti, Takapoto) - SYNONYMS - Trapezia wardi Seréne, 1969. FAMILY PILUMNIDAE Actumnus asper (Rippell, 1830) DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Marutea South). REFERENCES. — Actumnus bonnieri - NOBILI, 1907: 400 ("“Rikitea” = Mangareva, Marutea South). — Actumnus asper - BALsS, 1933: 36 (Syn.). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1964: 98 (Syn.); 1985: 452 (List) - SYNONYMS - Actumnus bonnieri Nobili, 1905. Actumnus digitalis (Rathbun, 1907) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Platypodia digitalis Rathbun, 1907: 38, pl. 1, fig. 6, pl. 9, fig. 4, 4a (Tahiti). — FOREST & GUINOT, 1962: 62 (Biogeography "Tahiti-Tuamotu"). — Actumnus digitalis - GUINOT, 1969: 225 (Syn.); 1985: 452 (List). Actumnus globulus Heller, 1861 DISTRIBUTION. — Tuamotu (Hao, Hikueru). REFERENCES. — Actumnus globulus - NOBILI, 1907: 400 ("Ohura" = Hao, Hikueru; cf. thereafter). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1969: 226 (writes about NOBILI's work: "par contre il est bien possible que les "globulus" polynésiens de NOBILI (1907, p. 50 sic) soient en fait des digitalis"); 1985: 452 (List). Actumnus obesus Dana, 1852 DISTRIBUTION. — Marquesas; Society (Tahiti). REFERENCES. — Actumnus obesus - BALSS, 1933: 37 (Marquesas). — BOONE, 1934: 154, pl. 79 (Tahiti). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1985: 452 (List). Actumnus setifer (De Haan, 1835) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Actumnus tomentosus Dana, 1852a; 1852b: 243; 1855, pl. 14, fig. 2a-c (Tahiti). —Actumnus setifer - ORTMANN, 1893b: 474 (Syn.). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List). Pilumnus merodentatus Nobili, 1906 DISTRIBUTION. — Gambier (Mangareva); Marquesas; Society (Tahiti); Tuamotu (Mataiva?). REFERENCES. — Pilumnus merodentatus Nobili, 1906a: 263; 1907: 399 (""Rikitea” = Mangareva). — SEURAT, 1934: 60 (French Polynesia). — FOREST & GUINOT, 1961: 132, fig. 128 (Mangareva; Syn.); 1962: 70 (Biogeography "Tahiti- Tuamotu"). — GUINOT, 1985: 452 (List). — Pilumnus longicornis merodentatus - BALSS, 1933: 16 (Mangareva, 62 NOBILI's material and also Tahiti, Marquesas) - RELEVANT MATERIAL - Pilumnus cf. merodentatus - MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva). — GUINOT, 1985: 452 (List). Pilumnus parvulus Nobili, 1906 DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Moruroa). REFERENCES. — Pilumnus parvulus Nobili, 1906a: 263; 1907: 398 (""Gatavake, Rikitea, Tokaerero" = Mangareva; some sp. in pearl oyster, 25m). — SEURAT, 1934: 60 (French Polynesia). — FOREST & GUINOT, 1961: 130, fig. 126, pl. 27, fig. 1 (Mangareva); 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List). — TAKEDA & MIYAKE, 1968: 6 (Key) - NEW MATERIAL - Coll. & det. J. POUPIN (Moruroa; Isp. in pearl oyster Pinctada margaritifera). Pilumnus ransoni Forest & Guinot, 1961 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Pilumnus ransoni Forest & Guinot, 1961: 130, fig. 123-124, 127, pl. 4, fig. 1-2, pl. 17, fig. 2. (Tahiti); 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 452 (List). Pilumnus tahitensis De Man, 1890 DISTRIBUTION. — Society (Moorea, Raiatea, Tahiti); Tuamotu (Fakarava, Marutea South, Tikehau) - Littoral to sublittoral. REFERENCES. — Pilumnus tahitensis de Man, 1890: 61, pl. 3, fig. 4 (Tahiti). — ORTMANN, 1893b: 437 (Tahiti). — NoBILI, 1907: 399 (Marutea South). — RATHBUN, 1907: 56 (Fakarava). — BALSS, 1933: 25 (Tahiti). — SEURAT, 1934: 60 (Marutea South). — FoREST & GUINOT, 1961: 129, fig. 125 (Raiatea, Tahiti); 1962: 70 (Biogeography "Tahiti- Tuamotu"). — GUINOT, 1985: 452 (List). — PEYROT-CLAUSADE, 1989: 111, 115 (Moorea, Tikehau; 30m). FAMILY CARPILIIDAE Carpilius convexus (Forskal, 1775) DISTRIBUTION. — Austral (Rapa, Rurutu); Society (Moorea, Tahiti); Tuamotu (Hikueru, Makatea, Makemo, Mataiva, Moruroa, Taiaro, Takapoto, Takaroa, Raroia) - Littoral to sublittoral. REFERENCES. — Carpilius convexus - STIMPSON, 1858a: 32; 1907: 37 (Tahiti). — RATHBUN, 1907: 37 (Makemo). — BOONE, 1934: 89, pl. 43-45 (Tahiti). — HOLTHUIS, 1953: 12 (Raroia). — CHABOUIS L. & F., 1954: 91, fig. 2 (French Polynesia). — MORRISON, 1954: 16 (Raroia). — BUITENDIJK, 1960: 263 (Takaroa). — FOREST & GUINOT, 1961: 37 (Hikueru, Tahiti); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (French Polynesia); 1985: 449 (List). — CHEVALIER et al., 1968: 92, 138 (Moruroa). — MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Mataiva, Moorea, Takapoto). — SERENE, 1984: 303, fig. 208-209 (Mataiva). — DELESALLE, 1985: 289 (Mataiva). — SALVAT & RICHARD, 1985: 362 (Takapoto). — SALVAT, 1986b: 72 (French Polynesia). — BONVALLOT et al., 1994: 140, photograph (Tuamotu). — POUPIN, 1994a: 45, fig. 41, pl. 5d (Hikueru, Rapa, Rurutu, Tahiti, Taiaro; up to 60m). Carpilius maculatus (Linné, 1758) DISTRIBUTION. — Gambier; Society (Moorea, Tahiti); Tuamotu (Fakarava, Hao, Hikueru, Makatea, Mataiva, Moorea, Moruroa, Raraka, Raroia, Taiaro, Takapoto). REFERENCES. — Carpilius maculatus - DANA, 1852b: 160 (Raraka). — STIMPSON, 1858a: 32; 1907: 37 (Tahiti). — HELLER, 1865: 9 (Tahiti). — NoBILI, 1907: 386 ("Ohura" = Hao). — RATHBUN, 1907: 37 (Fakarava, Tahiti). — PESTA, 1913: 39, pl. 3, fig. 4 (Tahiti, with a ?). — BOONE, 1934: 86, pl. 39-42 (Tahiti). — SEURAT, 1934: 60 (Gambier, Tuamotu). — HOLTHUIS, 1953: 12 (Raroia). — CHABOUIS L. & F., 1954: 91, fig. 1 (French Polynesia). — MORRISON, 1954: 16 (Raroia). — BABLET, 1972: 32, pl. 11 (French Polynesia). — FOREST & GUINOT, 1961: 37 (Hikueru, Tahiti); 1962: 60 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (French Polynesia); 1985: 450 (List). — CHEVALIER et al., 1968: 92, 138 (Moruroa). — CROSNIER, 1984: 302, fig. 208-209, pl. 44e (Mataiva). — MONTEFORTE, 1984: 170, ne ae eee 63 annex 1, tab. a, photograph p.140a (haut) (Makatea, Mataiva, Moorea, Tahiti, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — SALVAT, 1986b: 72, photograph (French Polynesia). — PARDON, 1992: 82, photograph (Tahiti). — BONVALLOT ef al., 1994: 141, photograph (Tuamotu). — PouPIN, 1994a: 46, fig. 42, pl. Se (Mataiva, Tahiti, Taiaro). FAMILY MENIPPIDAE Dacryopilumnus eremita Nobili, 1906 DISTRIBUTION. — Gambier (Mangareva); Tuamotu (Amanu, Hao, Makatea, Marutea South, Mataiva). REFERENCES. — Dacryopilumnus eremita Nobili, 1906a: 264; 1907: 400, pl. 2, fig. 4 (Amanu, Hao, "Rikitea" = Mangareva; gen. and sp. nov.). — FOREST & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu"). — CROSNIER, 1984: 313, fig. 240-241, pl. 47e (Mangareva, Marutea). — MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva). — GUINOT, 1985: 450 (List). Domecia glabra Alcock, 1899 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hao, Tikehau) - Littoral to sublittoral. REFERENCES. — Domecia hispida - NoBILt, 1907: 404 (Hao) not Domecia hispida Eydoux & Souleyet, 1842 = D. glabra fide FOREST & GUINOT (1961: 126). — Domecia glabra - FOREST & GUINOT, 1961: 126, fig. 115-116, 120-122, 124bis (Hao); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1962: 240, fig. 13a-b (Hao). — PEYROT- CLAUSADE, 1977a, annex of the species: 26; 1977b: 213 (Moorea); 1989: 111 (Tikehau; 30m). — GUINOT, 1985: 450 (List). Domecia hispida Eydoux & Souleyet, 1842 DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Moorea, Tahiti); Tuamotu (Hikueru, Makemo, Takapoto, Tikehau). REFERENCES. — Domecia hispida - DANA, 1852b: 251 (Tahiti). — RATHBUN, 1907: 60 (Makemo). — BOONE, 1934: 162, pl. 85 (Nuku Hiva). — FOREST & GUINOT, 1961: 126, fig. 117-119, 124bis, pl. 28, fig. 1 (Hikueru); 1962: 68 (Biogeography "Tahiti-Tuamotu, Marquesas"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 213 (Moorea); 1989: 111, 114 (Moorea, Tikehau). — NAIM, 1980a, annex 1, tab. 3 (Moorea). — KROPP & BIRKELAND, 1981: 629, tab. 5 (Moorea, Takapoto). — ODINETZ, 1983: 205 (Moorea, Tahiti, Takapoto). — GUINOT, 1985: 450 (List). — ODINETZ-COLLART & RICHER DE FORGES, 1985: 201 (Moorea and/or Tahiti, Takapoto). — Not Domecia hispida - NoBILt, 1907: 404 (Hao) = Domecia glabra Alcock, 1899 fide FOREST & GUINOT (1961: 126). Eriphia scabricula Dana, 1852 DISTRIBUTION. — Society (Tahiti); Tuamotu (Fakarava, Raroia). REFERENCES. — Eriphia scabricula Dana, 1852a: 82; 1852b: 247; 1855, pl. 14, fig. Sa-b (Tahiti). — RATHBUN, 1907: 57 (Fakarava). — HOLTHUIS, 1953: 20 (Raroia). — MORRISON, 1954: 7 (Raroia). — FOREST & GUINOT, 1961: 123, fig. 113a-b, 114 (Tahiti); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, annex 1, tab. a (Tahiti). — GUINOT, 1985: 450 (List). Eriphia sebana (Shaw & Nodder, 1803) DISTRIBUTION. — Gambier (Mangareva, Tarauru-Roa); Society (Moorea, Tahiti); Tuamotu (Fakarava, Hikueru, Makatea, Makemo, Marutea South, Mataiva, Moruroa, Pukapuka, Rangiroa, Raroia, Taiaro, Takapoto, Tauere). REFERENCES. — Eriphia laevimana Latr. - DANA, 1852b: 249; 1855, pl. 14, fig. 7a-c ("Honden" = Pukapuka, Society). — CANO, 1888: 171 (Tahiti). — NoBILI, 1907: 403 (""Gatavake" = Mangareva, Tarauru-Roa, Tauere). — Eriphia sebana - RATHBUN, 1907: 57 (Fakarava, Makatea, Makemo, "Manga Reva, Motus" = Mangareva?, Rangiroa). — SENDLER, 1923: 39 (Makatea). — HOLTHUIS, 1953: 20 (Raroia). — MORRISON, 1954: 7 (Raroia). — FOREST & GUINOT, 1961: 122, fig. 11la-b, 112 (Hikueru, Tahiti; Syn.); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, annex 1, tab. a, photograph 140a (bas) (Makatea, Mataiva, Moorea, Tahiti, Takapoto). — DELESALLE, 1985: 64 289 (Mataiva). — GUINOT, 1985: 450 (List). — SALVAT, 1986b: 72 (French Polynesia). —POUPIN, 1994a: 50, fig. 46, pl. 6a (Makatea, Marutea, Moruroa, Taiaro) - SYNONYMS - Eriphia laevimana Guérin, 1829-1844 in Latreille. Globopilumnus globosus (Dana, 1852) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Ahe?, Makatea, Manihi?, Mataiva, Raraka, Raroia, Takapoto, Tikehau). REFERENCES. — Pilumnus globosus Dana, 1852a: 81; 1852b: 236; 1855, pl. 13, fig. 10 (Raraka, Tahiti, "Waterland’ = Ahe and/or Manihi). — RATHBUN, 1907: 56 (Tahiti). — Pilumnus margaritatus Ortmann, 1893b: 436 (Tahiti). — Globopilumnus globosus - BALSS, 1933: 7, pl. 1, fig. 1-2. (Tahiti) ? pro parte. — HOLTHUIS, 1953: 21 (Raroia). — GUINOT-DUMORTIER, 1960a: 99, fig. 1-2, 5-6 (Tahiti; Syn.). — ForEST & GUINOT, 1961: 121 (Tahiti); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 450 (List). — PEYROT- CLAUSADE, 1989: 113 (Tikehau). — Not Pilumnus globosus - NOBILI, 1907: 398 (""Marutea Vaitutaki") = Liocarpilodes sp. fide GUINOT-DUMORTIER (1960a: 100). — BOonE, 1934: 152, pl. 78 (Tahiti) = Pilodius pugil with a ? fide SERENE & LuoM (1959: 320). Lydia annulipes (H. Milne Edwards, 1834) DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti); Tuamotu (Hikueru, Mataiva, Moruroa, Pukapuka, Raroia, Taiaro, Takapoto). REFERENCES. — Ruppellia annulipes - DANA, 1852b: 246; 1855, pl. 14, fig. 4a-c (Tahiti). — Lydia annulipes - HOLTHUIS, 1953: 23 (Pukapuka, Raroia). — MORRISON, 1954: 7 (Raroia). — FOREST & GUINOT, 1961: 122, fig. 109a-b, 110 (Hikueru); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a (Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). —GUINOT, 1985: 450 (List). — POUPIN, 1994a: 47, fig. 43, pl. Sf (Hikueru, Moruroa, Nuku Hiva, Taiaro). — Ruppellia granulosa A. Milne Edwards, 1867: 279 (Marquesas; new synonymy, cf. Remark). REMARK. — Ruppellia granulosa has been very rarely cited after its description. It is mentioned for the genus Lydia in SAKAI (1976: 477). It has been briefly described from a single specimen: "Cette espéce se distingue de Ruppellia annulipes par la profondeur des sillons qui limitent les lobules des régions. Ces lobules sont rugueux et granuleux. Les pattes antérieures sont également couvertes de grosses granulations peu élevées. Les autres caractéres sont les mémes que chez la Ruppellia annulipes". We have re-examined the type specimen (MNHN B9344, 1 do 17x25) and, after its comparison with specimens of L. annulipes collected in the Marquesas and the Tuamotu (cf. in PouPIN, 1994a: 47), we consider that it is a junior synonym of this species. Ozius hawaiensis Rathbun, 1902 DISTRIBUTION. — Marquesas (Nuku Hiva); Tuamotu (Fakarava, Makemo, Rangiroa). REFERENCES. — Ozius hawaiensis - RATHBUN, 1907: 54 (Fakarava, Makemo, Nuku Hiva, Rangiroa). — FOREST & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1985: 450 (List). Ozius rugulosus Stimpson, 1858 DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti). REFERENCES. — Ozius rugulosus - HELLER, 1865: 22, pl. 3, fig. 1 (Tahiti). — PEsTA, 1913: 47 (Tahiti). — Forest & GUINOT, 1961: 121, fig. 107a-b, 108 (Tahiti); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 450 (List). — PouPIN, 1994a: 48, fig. 44, pl. 5g (Nuku Hiva, Tahiti). Ozius tricarinatus Rathbun, 1907 DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti). REFERENCES. — Ozius tricarinatus Rathbun, 1907: 53, pl. 2, fig. 3 (Nuku Hiva, Tahiti). — FOREST & GUINOT, 1962: 68 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1985: 450 (List). 65 Ozius truncatus A. Milne Edwards, 1834 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Ozius lobatus Heller, 1865: 21, pl. 2, fig. 4 (Tahiti) fide CHILTON & BENNETT (1929: 750). Remark. — CHILTON & BENNETT consider, with doubt, that HELLER's species is the same as Ozius truncatus, but they do not mention "Tahiti", in the distribution of O. truncatus It is possible, as often seen in HELLER's work, that this locality was mentioned by error. Ozius tuberculosus H. Milne Edwards, 1834 DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti). REFERENCES. — Ozius tuberculosus - BOONE, 1934: 150, pl. 77 (Nuku Hiva). — POUPIN, 1994a: 49, fig. 45, pl. 5h (Nuku Hiva, Tahiti). Pseudozius caystrus (Adams & White, 1848) DISTRIBUTION. — Gambier (Kamaka, Tarauru-Roa); Society (Moorea, Tahiti); Tuamotu (Ahe and/or Manihi, Makatea, Mataiva, Moruroa, Raraka, Raroia, Taiaro, Takapoto). REFERENCES. — Pseudozius planus Dana, 1852a: 81; 1852b: 233; 1855, pl. 13, fig. 6a-h (Raraka, "Waterland’ = Ahe and/or Manihi). — EVANS, 1967: 409 ("Paumotu"; BM syntypes). — Pseudozius caystrus - NOBILI, 1907: 397 (Kamaka). — SENDLER, 1923: 38 (Makatea). — HOLTHUIS, 1953: 26 (Raroia). — MORRISON, 1954: 9 (Raroia). — FOREST & GUINOT, 1961: 125 (Tahiti, “"Taraourou-roa" = Tarauru-Roa; Syn.); 1962: 68 (Biogeography "Tahiti- Tuamotu"). — MONTEFORTE, 1984: 171, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 450 (List). — PoUPIN, 1994a: 52, fig. 48, pl. 6c (Kamaka, Moruroa, Taiaro). FAMILY GECARCINIDAE Cardisoma carnifex (Herbst, 1794) DISTRIBUTION. — Society (Bora Bora, Moorea, Raiatea, Tahiti, Tupai); Tuamotu (Ahe, Hao, Makatea, Mataiva, Nukutipipi, Pukarua, Rangiroa, Raroia, Taiaro, Takapoto, Tauere) - Terrestrial. REFERENCES. — Cardisoma obesum Dana, 1851: 252; 1852b: 375; 1855, pl. 24, fig. 1 (“Peacock" = Ahe). — STIMPSON, 1858b: 100; 1907: 111 (Tahiti). — Perigrapsus excelsus Heller, 1862: 522; 1865: SO, pl. 5, fig. 1 (Tahiti). — Cardisoma carnifex - MIERS, 1886: 220 (Tahiti). — Not, 1907: 407 (Bora Bora, Hao, Tahiti). — RATHBUN, 1907: 26 (Rangiroa, Tahiti). — SENDLER, 1923: 22 (Tahiti). — BOONE, 1934: 187, pl. 97-98 (Bora Bora). — SEURAT, 1934: 52 (Moorea, Tahiti, Tuamotu). — HOLTHUIS, 1953: 34 (Raroia). — CHABOUIS L. & F., 1954: 92, unnumbered fig. (French Polynesia). — MORRISON, 1954: 2 (Raroia). — FOREST & GUINOT, 1961: 165 (Tahiti); 1962: 74 (Biogeography "Tahiti-Tuamotu"). — EDMONSON, 1962: 25 (Raiatea). — GUINOT, 1966a: 48 (French Polynesia); 1985: 454 (List). — TURKAY, 1973: 108 (Syn.). — SAKAI, 1976: 680 (Syn.). — MONTEFORTE, 1984: 174, annex 1, tab. a (Makatea, Mataiva, Moorea, Takapoto); 1987: 6 (Moorea). — DELESALLE, 1985: 288, 295 (Mataiva). — CHARLEUX, 1986: 80, photographs (French Polynesia). — BONVALLOT et al., 1994: 78, photograph (Tuamotu). — BAGNIS & CHRISTIAN, 1983: 110-111, photograph (Tuamotu). — MERSCHARDT-SALVAT, 1991: 40 (Nukutipipi). — SALVAT F. & B., 1992: 5 (Nukutipipi). — POUPIN, 1994a: 53, fig. 49, pl. 6e (Pukarua, Tahiti, Taiaro, Tauere, Tupuai). — Without name - PARDON, 1992: 78, 79, photograph, double page (Tahiti, Papeete market) (det. according to the photograph). Cardisoma hirtipes Dana, 1851 DISTRIBUTION. — Society (Tahiti) - Terrestrial. REFERENCES. — Cardisoma hirtipes - HELLER, 1865: 35 (Tahiti). — TURKAY, 1974: 229, fig. 2, 12-13 (Tahiti). — GUINOT, 1985: 454, with a ? (List). — POUPIN, 1994a: 53 (Text). 66 Cardisoma rotundum Quoy & Gaimard, 1834 DISTRIBUTION. — Marquesas (Nuku Hiva); Tuamotu (Tikehau) - Terrestrial. REFERENCES. — Cardisoma rotundum - TURKAY, 1974: 234, fig. 1, 14 (""Tickahau-Atoll" = Tikehau). — POUPIN, 1994a: 54, fig. 50, pl. 6g (Nuku Hiva). Discoplax longipes A. Milne Edwards, 1867 DISTRIBUTION. — Tuamotu (Makatea). REFERENCES. — Discoplax longipes - SENDLER, 1923: 23, pl. 20, 1a-b (Makatea). — FOREST & GUINOT, 1962: 74 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 454 (List). Epigrapsus politus Heller, 1862 DISTRIBUTION. — Society (Tahiti); Tuamotu (Hikueru, Taiaro). REFERENCES. — Epigrapsus politus Heller, 1862: 522 (Tahiti). — HOLTHUIS, 1953: 34 ("Taravao" = Tahiti). — FOREST & GUINOT, 1961: 162, fig. 176a-b (Hikueru; Syn.); 1962: 74 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Tahiti). — GUINOT, 1985: 454 (List). — POUPIN, 1994a: 55, fig. 51, pl. 6d (Hikueru). — Nectograpsus politus Heller, 1865: 57, pl. 5, fig. 3 (Tahiti). — Not Epigrapsus politus - NOBILI, 1907: 407 (Hikueru) = Cyclograpsus integer H. Milne Edwards, 1837 fide FOREST & GUINOT (1961: 162). — SEURAT, 1934: 58 (NOBILI's material). FAMILY GRAPSIDAE SUBFAMILY GRAPSINAE Geograpsus crinipes (Dana, 1851) DISTRIBUTION. — Marquesas (Nuku Hiva); Society (Tahiti); Tuamotu (Makatea, Makemo, Mataiva, Moruroa, Pukapuka, Raroia, Taiaro, Takapoto). REFERENCES. — Geograpsus crinipes - HELLER, 1865: 48 (Tahiti). — NOBILI, 1907: 404 (Pukapuka). — RATHBUN, 1907: 28 (Makemo). — SENDLER, 1923: 32 (Makatea). — SEURAT, 1934: 52 (Pukapuka). — HOLTHUIS, 1953: 29 (Raroia). — MORRISON, 1954: 9 (Raroia). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 453 (List). — SALVAT & RICHARD, 1985: 359, 360 (Takapoto). — POUPIN, 1994a: 57, fig. 53, pl. 6h (Moruroa, Nuku Hiva, Taiaro, Takapoto). Geograpsus grayi (H. Milne Edwards, 1853) DISTRIBUTION. — Society (Tahiti); Tuamotu (Makatea, Nukutipipi, Raroia). REFERENCES. — Geograpsus grayi - KINGSLEY, 1880c: 196 (Tahiti). — ORTMANN, 1894: 707 (Tahiti). — SENDLER, 1923: 32, pl. 21, fig. 6 (Makatea). — SEURAT, 1934: 52 (Tahiti). — HOLTHUIS, 1953: 30 (Raroia). — MORRISON, 1954: 11 (Raroia). — BANERDJEE, 1960: 159 (Tahiti). — FoREsT & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea). — GUINOT, 1985: 453 (List). — MERSCHARDT-SALVAT, 1991: 40 (Nukutipipi). — SALVAT F. & B., 1992: 5 (Nukutipipi). Geograpsus stormi De Man, 1895 DISTRIBUTION. — Marquesas (Nuku Hiva). REFERENCES. — Geograpsus lividus stormi de Man - RATHBUN, 1907: 29 (Nuku Hiva). — Geograpsus stormi - BANERDIJEE, 1960: 167 (Syn.). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 58, fig. 54, pl. 7a (Nuku Hiva). EE eee 67 Grapsus depressus Heller, 1862 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Grapsus depressus Heller 1862: 521 (Tahiti; to our knowlege this species has never been mentioned after its description). Grapsus longitarsis Dana, 1851 DISTRIBUTION. — Society; Tuamotu (Fakarava, Hao, Hikueru, Makatea, Mataiva, Rangiroa, Raroia, Taiaro, Takapoto, Tike). REFERENCES. — Grapsus longitarsis Dana, 1851: 249; 1852b: 339; 1855, pl. 21, fig. 4a-d (Tuamotu). — RATHBUN, 1907: 28 (Fakarava, Rangiroa, Tikei). — HOLTHUIS, 1953: 31 (Raroia). — MORRISON, 1954: 7 (Raroia). — BANERJEE, 1960: 144, fig. 1b, 2h-n (Society, Raroia). — FOREST & GUINOT, 1961: 152, fig. 160a-b, 161, pl. 18, fig. 2 (Hikueru); 1962: 72 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 59, fig. 55, pl. 7b (Hao, Hikueru, Taiaro). — Grapsus strigosus - NOBILI, 1907: 404 (Hao) not G. strigosus (Herbst, 1799) synonym of Grapsus albolineatus Lamarck, 1818 in BANERJEE (1960: 147) = G. longitarsis fide FOREST & GUINOT (1961: 152). Grapsus tenuicrustatus (Herbst, 1783) DISTRIBUTION. — Gambier (Kamaka); Marquesas (Nuku Hiva); Society (Moorea, Tahiti); Tuamotu (Fakarava, Hao, Hikueru, Makatea, Makemo, Mataiva, Moruroa, Rangiroa, Raroia, Taiaro, Takapoto). REFERENCES. — Grapsus pictus - DANA, 1852b: 336; 1855, pl. 21, fig.1 (Tuamotu). — Grapsus grapsus - NOBILI, 1907: 404 (Hao, Kamaka) not Grapsus grapsus Linné, 1758 = G. tenuicrustatus fide FOREST & GUINOT (1961: 154). — Grapsus grapsus tenuicrustatus - RATHBUN, 1907: 27 (Fakarava, Makemo, Rangiroa). — Grapsus gracilipes - SENDLER, 1923: 31(Makatea). — Grapsus gracillimus Sendler, 1923: 32, pl. 21, fig. 5 (Makatea). — Grapsus tenuicrustatus - HOLTHUIS, 1953: 31 (Raroia). — CHABOUIS L. & F., 1954: 91, fig. 6 (French Polynesia). — Morrison, 1954: 9 (Raroia). — BANERJEE, 1960: 134, fig. 1a, 2a, c-g (Raroia; Syn.). — FOREST & GUINOT, 1961: 154 (Hikueru; Syn.); 1962: 72 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 453 (List). — SALVAT & RICHARD, 1985: 360 (Takapoto). — PARDON, 1992: 82, photograph (Tuamotu). — POUPIN, 1994a: 60, fig. 56, pl. 7c (Nuku Hiva, Tahiti, Taiaro, Takapoto). — The following works refer also probably to this species: Grapsus maculatus - KINGSLEY, 1880c: 192 (Tahiti) = G. grapsus fide ORTMANN (1894: 703). — Grapsus grapsus - SENDLER, 1923: 30 (Makatea). —BOONE, 1934: 178, pl. 90 ("Anaho Bay” = Nuku Hiva). — CHEVALIER et al., 1968: 95, 138 (Moruroa). — BAGNIS & CHRISTIAN, 1983: 112-113, photograph (Tuamotu). —Grapsus albolineatus - BONVALLOT et al., 1994: 137, photograph, (Tuamotu; det. according to the photograph) - SYNONYMS - Grapsus pictus Latreille, 1802-1803; Grapsus gracilipes H. Milne Edwards, 1853; Grapsus gracillimus Sendler, 1923. REMARK. — BANERDIEE (1960: 139) mentions that Grapsus grapsus (Linné, 1758) is solely Atlantic. More recently, MANNING & HOLTHUIS (1981: 233) have also mentioned it from the Eastern Pacific. Leptograpsus variegatus (Fabricius, 1793) DISTRIBUTION. — Austral (Rapa); Marquesas. REFERENCES. — Leptograpsus variegatus - DE MAN, 1890: 84 (Marquesas; with regard of L. ansoni H. Milne Edwards, 1853). — GRIFFIN, 1973: 461, fig. 1-6 (Syn.) - SYNONYMS - Leptograpsus ansoni H. Milne Edwards, 1853 - NEW MATERIAL - March 1995, Coll. and det. J. POUPIN (Rapa, Haurei bay, very common). Metopograpsus messor (Forskal, 1775) DISTRIBUTION. — Society (Bora Bora, Tahiti). REFERENCES. — Metopograpsus messor - KINGSLEY, 1880c: 190 (Tahiti). — MIERS, 1886: 258 (Tahiti). — RATHBUN, 1907: 29 (Bora Bora). — PESTA, 1913: 61 (Tahiti). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 61 (Text). 68 Metopograpsus thukuhar (Owen, 1839) DISTRIBUTION. — Austral (Tubuai); Gambier (Mangareva); Society (Moorea, Tahiti); Tuamotu (Hao, Taiaro). REFERENCES. — Metopograpsus thukuar - STIMPSON, 1858b: 101; 1907: 114 [47] (Tahiti). — HELLER, 1865: 43 (Tahiti). — Nosmul, 1907: 404 (Hao, "Rikitea" = Mangareva). — SEURAT, 1934: 59 (Mangareva). — BANERJEE, 1960: 186, fig. 6f-g (Tahiti). — FOREST & GUINOT, 1961: 155, fig. 162, 167 (Tahiti); 1962: 72 (Biogeography "Tahiti- Tuamotu"). — GUINOT, 1985: 453 (List). — MARQUET, 1988: 90, fig. 48, tab. 23; 1991: 130, tab. 1-2; 1993: tab. 1, 3 (Mangareva, Moorea, Tahiti, Tubuai). — POUPIN, 1994a: 61, fig. 57, pl. 7d (Hao, Mangareva, Tahiti, Taiaro). Pachygrapsus fakaravensis Rathbun, 1907 DISTRIBUTION. — Tuamotu (Fakarava, Makatea, Mataiva, Taiaro, Takapoto). REFERENCES. — Pachygrapsus fakaravensis Rathbun, 1907: 29, pl. 5, fig. 1, pl. 9, fig. 6, 6a (Fakarava). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Takapoto). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 62, fig. 58, pl. 7e (Taiaro). Pachygrapsus minutus A. Milne Edwards, 1873 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Hikueru, Makatea, Mataiva, Takapoto, Tikehau). REFERENCES. — Pachygrapsus minutus - FOREST & GUINOT, 1961: 155 (Hikueru, Tahiti); 1962: 72 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 25; 1977b: 213; 1985: 462 (Moorea); 1989: 113 (Tikehau). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 453 (List) - RELEVANT MATERIAL - Pachygrapsus aff. minutus - NAIM, 1980a: 55, annex 1, tab. 3 (Moorea; very small adult specimens, possibly of a new species). — GUINOT, 1985: 453 (List). Pachygrapsus planifrons De Man, 1888 DISTRIBUTION. — Tuamotu (Fakarava, Raroia). REFERENCES. — Pachygrapsus planifrons - HOLTHUIS, 1953: 31 (Raroia). — MORRISON, 1954: 7 (Raroia). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). — Pachygrapsus longipes - RATHBUN, 1907: 30 (Fakarava) - SYNONYMS - Pachygrapsus longipes Rathbun, 1893 (in TESCH, 1918, p. 78; with uncertainty). Pachygrapsus plicatus (H. Milne Edwards, 1837) DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava, Hikueru, Makatea, Makemo, Mataiva, Raroia, Taiaro, Takapoto). REFERENCES. — Pachygrapsus plicatus - KINGLEY, 1880c: 200 (Tahiti). — RATHBUN, 1907: 29 (Fakarava, Makemo). — HOLTHUIS, 1953: 32 (Raroia). — MORRISON, 1954: 13 (Raroia). — FOREST & GUINOT, 1961: 154 (Hikueru); 1962: 72 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Moorea, Tahiti, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 453 (List). — SALVAT & RICHARD, 1985: 359 (Takapoto). — POUPIN, 1994a: 63, fig. 59, pl 7f (Hikueru, Taiaro). Planes cyaneus Dana, 1851 DISTRIBUTION. — Austral (Neilson bank). REFERENCES. — Planes cyaneus - NEW MATERIAL - April 1995, coll. J. POUPIN, det. A. CROSNIER (Neilson bank; on a drifting buoy with cirripeds). 69 SUBFAMILY VARUNINAE Hemigrapsus crenulatus (H. Milne Edwards, 1837) DISTRIBUTION. — Society (Tahiti). REFERENCES. — Heterograpsus crenulatus Guérin - NOBILI, 1907: 405 (Tahiti; in Paris, NOBILI's material is well recorded under Hemigrapsus crenulatus MNHN B12830). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti- Tuamotu"). — Hemigrapsus crenulatus (H. Milne Edwards) - BENNETT, 1964: 81 (Syn.). — GUINOT, 1985: 453 (List). Pseudograpsus albus Stimpson, 1858 DISTRIBUTION. — Tuamotu (Fakarava, Raroia). REFERENCES. — Pseudograpsus albus - RATHBUN, 1907: 32 (Fakarava). — HOLTHUIS, 1953: 32 (Raroia). — Morrison, 1954: 10 (Raroia). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). —GUuINOT, 1985: 453 (List). Ptychognathus crassimanus Finnegan, 1931 DISTRIBUTION. — Marquesas - Freshwater. REFERENCES. — Ptychognathus crassimanus Finnegan, 1931: 649 (Marquesas). REMARK. — It seems that this species was never recorded after its description. In particular, it does not appear in the works by MARQUET (1988, 1991, 1993), who has only collected Ptychognathus easteranus (det. HOLHTUIS), in the Marquesas rivers. Ptychognathus easteranus Rathbun, 1907 DISTRIBUTION. — Austral (Rurutu); Marquesas (Hiva Oa) - Freshwater. REFERENCES. — Ptychognathus easteranus - MARQUET, 1988: 90, fig. 48, tab. 23; 1991: 132, tab. 1-2; 1993: tab. 1, 3 (Hiva Oa, Rurutu). Ptychognathus intermedius (de Man, 1879) DISTRIBUTION. — Society (Tahiti) - Freshwater. REFERENCES. — Ptychognathus intermedius - ORTMANN, 1894: 711 (Tahiti). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). Thalassograpsus harpax (Hilgendorf, 1892) DISTRIBUTION. — French Polynesia. REFERENCES. — Thalassograpsus harpax - GUINOT, 1985: 453 (List only; origin not found). Varuna litterata (Fabricius, 1798) DISTRIBUTION. — Society (Moorea, Tahiti) - Fresh & Brackish water. REFERENCES. — Varuna litterata - MARQUET, 1988: 90, fig. 48, tab. 3; 1991: 133, tab. 1-2; 1993: tab. 1, 3 (Moorea, Tahiti). — POUPIN 1994a: 67, fig. 63, pl. 8b (Tahiti). SUBFAMILY SESARMINAE Chasmagnathus subquadratus Dana, 1851 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Chasmagnathus subquadratus - ORTMANN, 1894: 728 (Tahiti). — ForREsT & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). 70 REMARK. — Species described by par DANA (1851: 251) from an uncertain locality "Novi-Zealandiae ? Novi- Hollandiae orientalis 2", not very often cited. Cyclograpsus integer H. Milne Edwards, 1837 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Fakarava, Hikueru, Kaukura?, Mataiva, Raroia, Taiaro, Takapoto). REFERENCES. — Epigrapsus politus - NOBILI, 1907: 407 (Hikueru, Kaukura) not Epigrapus politus Heller, 1862 = C. integer fide FOREST & GUINOT (1961: 160). — Cyclograpsus parvulus - RATHBUN, 1907: 36 (Fakarava). — HOLTHUIS, 1953: 32 (Raroia). — MORRISON, 1954: 5 (Raroia). — Cyclograpsus integer - FOREST & GUINOT, 1961: 160, fig. 175a- c (Hikueru; Syn.); 1962: 74 (Biogeography "Tahiti-Tuamotu"). — MONTEFORTE, 1984: 172, annex 1, tab. a (Mataiva, Moorea, Tahiti, Takapoto). — GUINOT, 1985: 454 (List). — POUPIN, 1994a: 56, fig. 52, pl. 6f (Hikueru, Taiaro) - SYNONYMS - Cyclograpsus parvulus de Man, 1897. Cyclograpsus longipes Stimpson, 1858 DISTRIBUTION. — Society (Tahiti); Tuamotu (Makemo, Marutea South, Raroia). REFERENCES. — Cyclograpsus longipes - RATHBUN, 1907: 36 (Makemo, Tahiti). — HOLTHUIS, 1953: 32 (Raroia). — Morrison, 1954: 5 (Raroia). — FOREST & GUINOT, 1961: 160 (Marutea South); 1962: 72 (Biogeography "Tahiti- Tuamotu"). — GUINOT, 1985: 454 (List). Labuanium trapezoideum (H. Milne Edwards, 1837) DISTRIBUTION. — Society (Moorea, Raiatea, Tahiti) - Freshwater. REFERENCES. — Sesarma trapezoidea - GUERIN-MENEVILLE, 1838: 14 (Tahiti). — SEURAT, 1934: 51 (Tahiti). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu"). — MARQUET, 1988: 90, fig. 48, tab. 23; 1991: 133, tab. 1-2; 1993: tab. 1, 3 (Moorea, Tahiti). — Sesarma (Sesarma) trapezoidea - NOBILI, 1907: 405 (Tahiti). — EDMONSON, 1951: 237, fig. 33b (Raiatea, Tahiti). — FOREST & GUINOT, 1961: 157, fig. 164a-b, 165 (Tahiti). — Sesarma (Sesarma) trapezoideum - RATHBUN, 1907: 33 (Tahiti). — Labuanium trapezoideum - SERENE & SOH, 1970: 402, 406 (Syn.). — GUINOT, 1985: 454 (List). REMARK. — Labuanium rotundatum (Hess, 1865) is also recorded in Polynesia by SAKAI (1976: 663; distribution only "Micronesia, Polynesia"; cited afterwards by GUINOT, 1985: 454). SAKAI must consider the Polynesia s./., with about 10 states, including French Polynesia. It is doubtful that this species have been really collected in French Polynesia since we have not find any mention of it in TESCH (1917: 193), who gives a detailed distribution, or in SERENE & SOH (1970: 402, 406), when they have established the genus Labuanium. For the moment, it thus seems better to exclude it from the area. Metasesarma rousseauxi granularis Heller, 1862 DISTRIBUTION. — Gambier (Tarauru-Roa); Society (Tahiti ); Tuamotu (Hikueru). REFERENCES. — Metasesarma granularis Heller, 1862: 522 (Tahiti). —Metasesarma rugulosa Heller, 1865: 65 (Tahiti; cf. Remark). — Metasesarma rousseauxi H. Milne Edwards - ? ORTMANN, 1894: 717 (Tahiti). — ? HOLTHUIS, 1953: 33 ("Taravao" = Tahiti). — Sesarma (Metasesarma) rousseauxi - NOBILI, 1907: 405 (Tarauru-Roa). — Metasesarma rousseauxi granularis - FOREST & GUINOT, 1961: 158, fig. 168, 169, 174a-b (Hikueru, “Papenoo"” = Tahiti; Syn.); 1962: 72 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 454 (List). REMARK. — FOREST & GUINOT (1961: 158) writes "M. rousseauxi granularis, décrit en 1862 de Tahiti par HELLER, qui, en 1865, substituait 4 ce nom, sans raison apparente, celui de M. granulosa, a été mis en synonymie avec Metasesarma rousseauxi H. Milne Edwards, 1853, par DE MAN (1889, p. 439)". The same authors recognise differences between H. MILNE EDWARDS' species and the specimens from Tahiti and Tuamotu, which they attribute to the subspecies granularis Heller. According to the location, ORTMANN's and HOLTHUIS' references should be also attributed to this subspecies. Sarmatium crassum Dana, 1851 DISTRIBUTION. — Society (Tahiti). aS. Saeco eer al REFERENCES. — Sarmatium crassum - GUINOT, 1985: 454 (List; origin ?). — DAVIE, 1992: 81, fig. 1a, 2, 3a-c (Tahiti). Sesarma angustifrons A. Milne Edwards, 1869 DISTRIBUTION. — Society (Moorea, Tahiti). REFERENCES. — Sesarma (Sesarma) angustifrons - DEMAN, 1889: 432, pl. 10, fig. 10 (Tahiti). — NoBILI, 1907: 405 (Tahiti). — SEURAT, 1934: 51 (Tahiti). — Sesarma angustifrons - FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti- Tuamotu"). — GUINOT, 1985: 454 (List). — MARQUET, 1988: 90, fig. 48, tab. 23; 1991: 133, tab. 1-2; 1993: tab. 1, 3 (Moorea, Tahiti). Sesarma jacquinoti Ortmann, 1894 DISTRIBUTION. — Society (Tahiti). REFERENCES. — Sesarma jacquinoti Ortmann, 1894: 718 (Tahiti). — FOREST & GUINOT, 1962: 72 (Biogeography "Tahiti-Tuamotu”). — GUINOT, 1985: 454 (List). SUBFAMILY PLAGUSIINAE Percnon abbreviatum (Dana, 1851) DISTRIBUTION. — Gambier (Mangareva); Society (Tahiti); Tuamotu (Raroia). REFERENCES. — Acanthopus abbreviatus Dana, 1851: 252; 1852b: 373; 1855, pl. 23, fig. 11a-c (Tahiti). — Percnon affinis - NoBILI, 1907: 406 (Mangareva), pro parte not P. affine H. Milne Edwards, 1853 = P. abbreviatum fide FOREST & GUINOT (1961: 164). — Percnon abbreviatum - HOLTHUIS, 1953: 33 (Raroia). — MORRISON, 1954: 16 (Raroia). — EDMONSON, 1959: 195, fig. 25c, 26a-c (Syn.). — FOREST & GUINOT, 1961: 164 (Mangareva; Syn.); 1962: 74 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1966a: 48 (Raroia); 1985: 454 (List). Percnon affine (H. Milne Edwards, 1853) DISTRIBUTION. — Marquesas; Gambier (Mangareva); Society (Tahiti); Tuamotu (Makatea, Mataiva). REFERENCES. — Percnon affinis - NoBILI, 1907: 406 ("Gatavake” = Mangareva), pro parte some sp. are P. abbreviatum — FOREST & GUINOT, 1961: 164 (""Gatavake” = Mangareva; Syn.); 1962: 74 (Biogeography "Tahiti- Tuamotu"). — Percnon pilimanus - BOONE, 1934: 181, pl. 92-94 (Tahiti). — FOREST & GUINOT, 1962: 74 (Biogeography "Tahiti-Tuamotu, Marquesas”). — Percnon affine - CROSNIER, 1965: 88 (Tuamotu; Syn.). — MONTEFORTE, 1984: 172, annex 1, tab. a, photograph p. 138a-bas (Makatea, Mataiva, Tahiti). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 454 (List) - SYNONYMS - Percnon pilimanus (A. Milne Edwards, 1873). Percnon guinotae Crosnier, 1965 DISTRIBUTION. — .Marquesas (Hiva Oa). REFERENCES. — Percnon guinotae - NEW MATERIAL - February 1996, Coll. & det. J. POUPIN (Hiva Oa). REMARK. — The presence of this species in French Polynesia was assumed in POUPIN (1994a: 64). It is confirmed here with a specimen collected in the Marquesas. Percnon planissimum (Herbst, 1804) DISTRIBUTION. — Gambier (Mangareva); Marquesas (Nuku Hiva); Society (Tahiti); Tuamotu (Fakarava, Hao, Makatea, Marutea South, Mataiva, Moruroa, Raraka, Taiaro, Tikehau). REFERENCES. — Acanthopus planissimus - DANA, 1852b: 372 (Raraka, Tahiti). — HELLER, 1865: 51(Tahiti). — Acanthopus tenuifrons H. Milne Edwards, 1853: 180 (Nuku Hiva). — Percnon planissimus - NOBII, 1907: 406 (Hao, Mangareva, Marutea). — MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea, Mataiva, Tahiti). — PEYROT-CLAUSADE, 1989: 113 (Tikehau). — Percnon planissimum - RATHBUN, 1907: 37 (Fakarava). — PESTA, 1913: 64 (Tahiti). — EDMONDSON, 1959: 197, fig. 25c, 27a-c (Marquesas). — FOREST & GUINOT, 1961: 163 (Marutea South, Tahiti; Syn.); 72 1962: 74 (Biogeography "Tahiti-Tuamotu, Marquesas"). — CHEVALIER et al., 1968: 95, 138 (Moruroa). — GUINOT, 1985: 454 (List). — POUPIN, 1994a: 64, fig. 60, pl. 8a (Mangareva, Taiaro). Plagusia speciosa Dana, 1851 DISTRIBUTION. — Marquesas; Society (Tahiti); Tuamotu (Ahe and/or Manihi, Hao, Hikueru, Makatea, Makemo, Mataiva, Raroia, Taiaro, Takapoto). REFERENCES. — Plagusia speciosa Dana, 1851: 252; 1852b: 369; 1865, pl. 23, fig. 9 ("Waterland" = Ahe and/or Manihi). — KINGSLEY, 1880c: 223 (Tahiti). — DE MAN, 1890: 89 (Tuamotu). — ORTMANN, 1894: 731 (Tuamotu). — NoBILI, 1907: 406 (Hao). — RATHBUN, 1907: 36 (Makemo). — SENDLER, 1923: 35 (Makatea). — BOONE, 1934: 185, pl. 95-96 (Tahiti). — HOLTHUIS, 1953: 34 (Raroia). — MORRISON, 1954: 16 (Raroia). — FOREST & GUINOT, 1961: 162, fig. 177a-c, 178 (Hao, Hikueru); 1962: 74 (Biogeography "Tahiti-Tuamotu, Marquesas"). — GUINOT, 1966a: 48 (Raroia); 1985: 454 (List). — MONTEFORTE, 1984: 172, annex 1, tab. a, photograph p. 138a-haut (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289 (Mataiva). — SALVAT & RICHARD, 1985: 362 (Takapoto). — POUPIN, 1994a: 65, fig. 61, pl. 7g (Hao, Hikueru, Taiaro). REMARK. — Plagusia immaculata Lamarck, 1818 is erroneously cited from Tahiti and the Tuamotu by DAI & YANG (1991: 563). This error must come from a quick reading of EDMONSON (1959: 194), where "Tuamotus (type locality), Tahiti” is mentioned under P. immaculata, but for remarks concerning only P. speciosa Dana. Plagusia tuberculata Lamarck, 1818 DISTRIBUTION. — Austral (Raevavae); Gambier (Kamaka, Mangareva); Marquesas (Nuku Hiva); Tuamotu (Makatea?). REFERENCES. — Plagusia squamosa - NOBILI, 1907: 406 (Kamaka, Mangareva; inferred only: reference not retrieved in recent works, and material not found in Paris). — Plagusia depressa tuberculata - FOREST & GUINOT, 1962: 74 (Biogeography "Tahiti-Tuamotu"). — Plagusia depressa - ? MONTEFORTE, 1984: 172, annex 1, tab. a (Makatea) not P. depressa (Fabricius, 1775) = P. tuberculata cf. Remark. — Plagusia tuberculata - GUINOT, 1985: 454 (List). — POUPIN, 1994a: 66, fig. 62, pl. 7h (Nuku Hiva, Raevavae, Tuamotu) - SYNONYMS - Plagusia squamosa (Herbst, 1882) (cf. SAKAI, 1976: 676, then ALCOCK, 1900: 437). REMARK. — Plagusia depressa (Fabricius, 1775) is a species of the tropical Atlantic (cf. DAWSON, 1987: 42) and cannot be MONTEFORTE's (1984) material which is more likely P. tuberculata (One specimen of this species is actually deposited in the CRIOBE collections, Moorea, Coll. and det. MONTEFORTE). FAMILY PINNOTHERIDAE Pinnotherelia laevigata A. Milne Edwards & Lucas, 1843 DISTRIBUTION. — Marquesas (Nuku Hiva). REFERENCES. — Pinnotheralia laevigata - RATHBUN, 1918: 181, fig. 115, pl. 39, fig. 1-3, pl. 40, fig. 1-2 (Marquesas "Tawhoe" = Taiohae, Nuku Hiva). — SCHMITT et al., 1973: 125 (catalogue "Marquesas Islands"). FAMILY OCYPODIDAE Macrophthalmus consobrinus Nobili, 1906 DISTRIBUTION. — Gambier (Mangareva) - Brackish water. REFERENCES. — Macrophthalmus consobrinus Nobili, 1906a: 265; 1907: 408 (Mangareva). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). — Macrophthalmus parvimanus - BARNES, 1977: 273, pro parte, only NOBILI's consobrinus, not M. parvimanus Guérin-Méneville, 1834 (cf. Remark). REMARK. — According to BARNES (1977: 273) NoBILI's Macrophthalmus consobrinus is the same as Macrophthalmus parvimanus Guérin-Méneville, 1834. However, to check that point, a large sample of M. consobrinus has been recently 73 collected in the Gambier Islands, and it appears that NOBILI's species is valid, and closely related to M. convexus Stimpson, 1858 (POUPIN, in study). Macrophthalmus convexus Stimpson, 1858 DISTRIBUTION. — Society (Bora Bora, Tahiti) - Brackish water. REFERENCES. — Macrophthalmus convexus - ORTMANN, 1894: 745 (Tahiti). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List) - NEW MATERIAL - Coll. and det. J. POUPIN (Bora Bora). Macrophthalmus serenei (Seréne, 1983) DISTRIBUTION. — Tuamotu (Tikehau). REFERENCES. — Macrophthalmus serenei - NEW MATERIAL - Coll. C. HILY, det. J. POUPIN (Tikehau). Ocypode ceratophthalma (Pallas, 1772) DISTRIBUTION. — Gambier? (Mangareva); Society (Scilly, Tahiti); Tuamotu (Fakarava, Makatea, Makemo, Mataiva, Marutea South?, Rangiroa, Raroia, Taiaro, Takapoto, Tikehau). REFERENCES. — Ocypode urvillei Guérin-Méneville, 1829: pl. 1, fig. 1, la-b; 1838: 9 (Tahiti). — OWEN, 1839: 80 ("Low Islands" = Tuamotu). — NoBILI, 1907: 407 pro parte (Marutea?, cf. Remark sous O. pallidula). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453, with a ? (List). — Ocypode ceratophthalma - STIMPSON, 1858b: 100 [46]; 1907: 108, pl. 12, fig. 2 (Tahiti). — ORTMANN, 1897a: 364 (Syn.). — RATHBUN, 1907: 26 (Fakarava, Makemo, Rangiroa). — SENDLER, 1923: 21 (Tahiti). — HOLTHUIs, 1953: 28 (Raroia). — Morrison, 1954: 9 (Raroia). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — SAKAI & TURKAY, 1976: 86, fig. 13 (Syn.). — MONTEFORTE, 1984: 173, annex 1, tab. a (Makatea, Mataiva, Takapoto). — DELESALLE, 1985: 289, 303 (Mataiva). — GUINOT, 1985: 453 (List). — HARMELIN-VIVIEN, 1985: 239 (Tikehau). — SALVAT & RICHARD, 1985: 359 (Takapoto). — POUPIN, 1994a: 68, fig. 64, pl. 8c-d (Tahiti, Taiaro, Scilly). — Ocypode cordimana - KINGSLEY, 1880b: 186 (Tahiti) not O. cordimana Desmaret, 1825 = O. urvillei, synonym of O. ceratophthalma, fide ORTMANN (1897a: 366). — ? Oxypode (sic) - CHARLEUX, 1986: 81, photograph (French Polynesia), det. according to the photograph. Ocypode cordimana Desmaret, 1825 DISTRIBUTION. — Society (Bora Bora, Tahiti). REFERENCES. — Ocypode cordimana - BOONE, 1934: 191, pl. 99-100 (Bora Bora). — FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu"). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 69, fig. 65, pl. 8e (Tahiti). — Not Ocypode cordimana - KINGSLEY, 1880b: 186 (Tahiti) fide ORTMANN (1897a: 366) = O. urvillei, synonym of O. cerathophthalma. REMARK. — BOONE mentions this species in the Tuamotu ("Paumotus") after RATHBUN (1907: 26). In fact it is O. cerathophthalma that RATHBUN has cited from this archipelago. Ocypode pallidula Jacquinot 1852 DISTRIBUTION. — Gambier (Aukena, Mangareva); Tuamotu (Marutea South?, Moruroa). REFERENCES. — Ocypode pallidula Jacquinot, 1852, pl. 6, fig. 1a (Mangareva). — SAKAI & TURKAY, 1976: 87, fig. 14- 15 ("Rikitea" = Mangareva; type material, Syn.). — JONES, 1988: 34 (Syn.). — POUPIN, 1994a: 70, fig. 66, pl. 8f (Aukena). — Ocypode cordimana (Junior) - JACQUINOT & LUCAS, 1853: 64 (Mangareva) not O. cordimana Desmaret, 1825 = O. pallidula fide SAKAI & TURKAY (1976: 87). — Ocypode urvillei - NOBILI, 1907: 407, pro parte (“Rikitea" = Mangareva, Marutea?). — SEURAT, 1934: 52 (Mangareva; NOBILI's material) - The two references, not O. urvillei Guérin-Méneville, 1829, synonym of O. cerathophthalma = O. pallidula (cf. Remark). — Ocypode laevis - CHEVALIER et al., 1968: 109 (Moruroa) - NEW MATERIAL - Coll. B. SALVAT, 1966, det. J. POUPIN (Moruroa) - SYNONYMS - Ocypode laevis Dana, 1852. REMARK. — After ORTMANN (1897a: 366), Ocypode pallidula, was usually considered as the same as O. urvillei (= O. cerathophthalma). Its validity was re-established by SAKAI & TURKAY (1976). The material mentioned in NOBILI (1907) 74 and SEURAT (1934), under O. urvillei has been collected at Marutea South and Mangareva. The specimens from this second island are in fact O. pallidula (verification in the collections of Paris: 1 sp. labelled "Ocypoda urvillei Guér., Seurat coll. 1905, Bouvier dét., G. Nobili vérif. 1906", MNHN B11841, is a real O. pallidula). The specimens from Marutea South could reasonably be O. cerathophthalma, very common in the Tuamotu. Uca chlorophthalmus crassipes (Adams & White, 1848) DISTRIBUTION. — Austral (Raevavae); Marquesas; Society (Bora Bora, Maupiti, Raiatea, Tahiti) - Brackish water. REFERENCES. — Gelasimus latreillei H. Milne Edwards, 1852: 114, pl. 4, fig. 20, 20a (Bora Bora). — Gelasimus pulchellus Stimpson, 1858b: 100 [46]; 1907: 107, pl. 15, fig. 1 (Tahiti). — Gelasimus gaimardi - HELLER, 1865: 38 (Tahiti). — Uca chlorophthalmus - NoBILI, 1907: 408 (""Taravao" = Tahiti). — Uca gaimardi - RATHBUN, 1907: 26 (Bora Bora, Tahiti). — HOLTHUIS, 1953: 29 ("Taravao" = Tahiti). — CRANE, 1957: 74, 78 (Bora Bora, Raiatea, Tahiti). — FOREST & GUINOT, 1961: 140, fig. 140-145, 153, 156a-b (Tahiti); 1962: 70 (Biogeography "Tahiti-Tuamotu"). — Gelasimus (Uca) chlorophthalmus - SEURAT, 1934: 60 (Tahiti). — Uca (Amphiuca) chlorophthalmus crassipes - CRANE, 1975: 98, 102, 599, fig. 13-14, 26c, 31c, 37h, 39a-b, 56c, 60 I-m, 68a-b, 81g, 83a, 99, pl. 15 a-f, 46b (Bora Bora, Raiatea, Tahiti, Marquesas p. 599; Syn.). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 71, fig. 67, pl. 8g (Maupiti, Raevavae, Tahiti) - SYNONYMS - Uca gaimardi H. Milne Edwards, 1852. Uca tetragonon (Herbst, 1790) DISTRIBUTION. — Austral (Rapa); Gambier (Mangareva); Society (Bora Bora, Raiatea, Tahiti); Tuamotu (Moruroa, Napuka, Raroia). REFERENCES. — Gelasimus duperreyi Guérin-Méneville, 1829, pl. 1, fig. 2, 2a (Bora Bora). — Gelasimus tetragonon - GUERIN-MENEVILLE, 1838, pl. 1, fig. 2, 2a (Bora Bora) pro parte. — HELLER, 1865: 37 (Tahiti). — KINGSLEY, 1880a: 143, pl. 9, fig. 11 (Tahiti). — MIERS, 1886: 243 (Tahiti). — DE MAN, 1891: 24, pl. 2, fig. 6 (Tahiti). — ORTMANN, 1894: 754 (Tahiti). — Uca tetragonon - NoBILI, 1907: 408 ("Rikitea, Gatavake" = Mangareva). — RATHBUN, 1907: 26 (Bora Bora). — HOLTHUIs, 1953: 29 (Raroia). — MORRISON, 1954: 8 (Raroia). — CRANE, 1957: 79 (Bora Bora). — Gelasimus (Uca) tetragonon - SEURAT, 1934: 59 (Mangareva). — Uca tetragonum - FOREST & GUINOT, 1962: 70 (Biogeography "Tahiti-Tuamotu”). — Uca (Thalassuca) tetragonon - CRANE, 1975: 77, 81, 596, fig. 37d, 63a-b, 81f, 82e, 99, pl. 13 (Bora Bora, Raiatea, Raroia, Tahiti; Syn.). — GUINOT, 1985: 453 (List). — POUPIN, 1994a: 72, fig. 68, pl. 8h (Mangareva, Moruroa, Napuka, Rapa). REMARK. — FOREST & GUINOT (1962: 70) mention U. dussumieri (H. Milne Edwards) from French Polynesia, probably after ORTMANN (1894; Gelasimus dussumieri, Tahiti, p. 755). ORTMANN's reference is cited by CRANE (1975) for two subspecies: Uca (Deltuca) dussumieri spinata (specimens from Java and Singapore) and Uca (Deltuca) dussumieri dussumieri (specimens from the Philippines and Mindanao). ORTMANN's Tahitian U. dussumieri do not appear in CRANE, who clearly indicates (p. 437) that Uca dussumieri does not occur in French Polynesia. ORTMANN's "U. dussumieri", if they exist, must probably be, either U. chlorophthalmus, or U. tetragonon. FAMILY CRYPTOCHIRIDAE Cryptochirus coralliodytes Heller, 1861 DISTRIBUTION. — Tuamotu (Makatea, Marutea South, Marokau). REFERENCES. — Cryptochirus coralliodytes - NOBILI, 1907: 409 (Marutea South, Marokau). — SENDLER, 1923: 41 (Makatea). — FOREST & GUINOT, 1962: 74 (Biogeography "Tahiti-Tuamotu"). — SEURAT, 1934: 60 (Marokau, Marutea South). — GUINOT, 1985: 454 (List). — KRopp, 1988: 873 (Revision of this species but without mention of French Polynesia). Hapalocarcinus marsupialis Simpson, 1859 DISTRIBUTION. — Society (Moorea, Tahiti); Tuamotu (Takapoto). 75 REFERENCES. — Hapalocarcinus marsupialis - KRopp & BIRKELAND, 1981: 629, tab. 5 (Moorea, Takapoto). — ODINETZ, 1983: 29, 205 (Moorea, Tahiti, Takapoto). — GUINOT, 1985: 454 (List). — ODINETZ-COLLART & RICHER DE ForRGES, 1985: 201 (Moorea and/or Tahiti). FAMILY HYMENOSOMATIDAE Elamena mathaei (Desmaret,1825) DISTRIBUTION. — Tuamotu (Tikehau). REFERENCES. — Elamena mathaei - PEYROT-CLAUSADE, 1989: 113 (Tikehau). FAMILY INCERTAE SEDIS Daira perlata (Herbst, 1790) DISTRIBUTION. — Society (Moorea, Raiatea, Tahiti); Tuamotu (Makatea, Mataiva, Hao, Hikueru, Raroia, Taiaro, Takapoto, Tikehau). REFERENCES. — Daira perlata - NOBILI, 1907: 392 (Hao). — RATHBUN, 1907: 44 (Tahiti). — SENDLER, 1923: 38 (Makatea). — BOOng, 1934: 129, pl. 69 (Raiatea, Tahiti). — HOLTHUIS, 1953: 19 (Raroia). — CHABOUIS L. & F., 1954: 91, fig. 4 (French Polynesia). — MORRISON, 1954: 15 (Raroia). — FOREST & GUINOT, 1961: 119 (Hikueru, Tahiti); 1962: 68 (Biogeography "Tahiti-Tuamotu"). — PEYROT-CLAUSADE, 1977a, annex of the species: 26; 1977b: 212 (Moorea); 1989: 112, 115 (Moorea, Tikehau). — MONTEFORTE, 1984: 170, annex 1, tab. a, photograph p. 137a (Makatea, Mataiva, Moorea, Takapoto); 1987: 9 (Moorea). — DELESALLE, 1985: 289, 305 (Mataiva). — GUINOT, 1985: 453 (List). — SALVAT & RICHARD, 1985: 362 (Takapoto). — SALVAT, 1986b: 72, photograph (French Polynesia). — POUPIN, 1994a: 51, fig. 47, pl. 6b (Hikueru, Taiaro). REMARK. — This species has sometimes been classified in the Zalasiinae Seréne, 1968 (cf. SAKAI, 1976: 513). Parapleurophrycoides roseus Nobili, 1906 DISTRIBUTION. — Tuamotu (Marutea South). REFERENCES. — Parapleurophrycoides roseus Nobili, 1906a: 264; 1907: 402, pl. 2, fig. 5 (Marutea; gen. nov. and sp. nov. described from a very small specimen, 1.3x1.7mm). REMARK. — For this species, and the following, FOREST & GUINOT (1962: 41) wnite: "...nous les considérons comme des juvéniles difficilement identifiables." Platyozius perpusillus Nobili, 1906 DISTRIBUTION. — Tuamotu (Hao). REFERENCES. — Platyozius perpusillus Nobili, 1906a: 264; 1907: 401 (Hao; described from a very small specimen, 1.45x1.75mm). REMARK. — In SAKAI (1976: 535), Platyozius Borradaile, 1902 is the same as Eucrate de Haan, 1835 (Goneplacidae). 76 DISCUSSION NUMBER OF POLYNESIAN SPECIES A total of 401 littoral or sublittoral species are recorded in this work. The number by infra-order and family is computed on table 1. The Brachyura clearly prevail, with 78% of the species, followed by the Anomura (18%), and the Palinura (4%). Within the crabs, the Xanthidae account for 123 species, distributed in 5 main subfamilies: Liomerinae, Actaeinae, Zoziminae, Xanthinae, and Chlorodiinae. The Portunidae account for 54 species, including 6 recorded for the first time from determinations made by MOOSA and CROSNIER: Portunus macrophthalmus, P. orbitosinus, Thalamita danae, T. macropus, T. mitsiensis, and T. philippinensis; half of the sublittoral species belong to this family. Two other families are also well represented, the Grapsidae, with 35 species, and the Trapeziidae, with 20 species. From these two families, Percnon guinotae, Planes cyaneus and Quadrella maculosa, are recorded for the first time. With respect to the list presented 10 years ago by GUINOT (1985), excluding the species mentioned erroneously, or not fully determined, about 60 species are added to the Polynesian Brachyura, and approximately a hundred, if the deep species are included. Within the Anomura, the Diogenidae account for 40% of the species with three main genera: Calcinus, Clibanarius, and Dardanus. Calcinus guamensis, C. imperialis, and Dardanus australis, are now recorded in French Polynesia. The porcellanids account for about 25% of the species, the single genus Petrolisthes representing 12 species out of 17. The Albuneidae are represented by only one species, Albunea speciosa, which was previously thought to be endemic from the Hawaiian islands. Only 14 palinurids are recorded in French Polynesia, of which 2 only by larvae (Palinurellus wieneckii and Arctides regalis). SPECIES ERRONEOUSLY RECORDED IN FRENCH POLYNESIA Fourteen species have been erroneously recorded in French Polynesia. They are: Panulirus ornatus (cf. under P. versicolor), Panulirus polyphagus (cf. under P. pascuensis), Dynomene sinense (cf. under D. praedator), Ashtoret granulosa (cf. under A. picta), Lophozozymus incisus (cf. under L. superbus), Labuanium rotundatum (cf. under L. trapezoideum), Plagusia immaculata (cf. under P. speciosa), Uca dussumieri (cf. under U. tetragonon), Pachygrapsus transversus, Dotilla fenestrata, Ocypode macrocera, Ocypode platytarsis, and two freshwater crabs (Potamonidae). Pachygrapsus transversus Gibbes, 1850 is recorded from Tahiti by KINGSLEY (1880c: 199) (Tahiti). According to HOLTHUIS & GOTTLIED (1958: 102) this record is obviously false, P. transversus being an Atlantic species. This conclusion is later supported by MANNING & HOLTHUIS (1981: 235), who report however the species in the Pacific, but only along the American coasts. The ocypodid Dotilla fenestrata Hilgendorf, 1869, is listed by GUINOT (1985: 453) after KROPP & BIRKELAND (1981). It is probably a mistake, because this species is not mentioned in that work, and, to our knowledge, has never been reported, elsewhere, from French Polynesia. Two other ocypodids, with an uncertain status, are also erroneously reported from Tahiti, by HELLER (1865: 42): Ocypode macrocera (H. Milne Edwards, 1837) and Ocypode platytarsis (H. Milne Edwards, 1852) (see ORTMANN, 1897a: 362). qd HELLER has mentioned two potamonids crabs in Tahiti: Thelphusa wiillerstorfi, described as a new species in 1862 (p. 520); and Thelphusa leschenaudii (H. Milne Edwards, 1853) (in HELLER, 1865: 32). RATHBUN (1904: 287) places these two references under a single species Potamon (Potamon) hydrodromus (Herbst, 1796) and writes (p. 289): "il est douteux que cette espéce ou quelqu'autre habite Tahiti". Since the recent works by MARQUET (1988, 1991, 1993), who has intensively sampled the freshwater Polynesian fauna, it is almost certain that the Potamonidae are not represented in French Polynesia. IMPROVEMENT AND CORRECTION OF THIS LIST This bibliographic compilation is of course tentative and certainly does not account for all the species living in French Polynesia. When new collections become available, other species will undoubtedly be added to the present list. Moreover, despite a careful research, it is possible that a few works, recording additional species, have passed undetected. Right now, several species listed here deserve a particular attention, either because their presence in French Polynesia remained to be confirmed, or because their identification, or taxonomic status, are uncertain. Twelve species of this list could have been erroneously recorded from French Polynesia. They are known only by larvae (Palinurellus wieneckii, Arctides regalis), are mentioned with doubt, or in an ambiguous way (Petrolisthes militaris, Porcellana mitra, Porcellana monilifera, Ozius truncatus), appear only in a part "Distribution", the origin of the French Polynesian material remaining unknown (Neoliomera insularis, Gaillardiellus rueppelli, Macromedaeus distinguendus, Thalassograpsus harpax), or, are cited from French Polynesia only because of the large geographic distribution of the species, without real collections in the field (Schizophrys aspera, Aethra scruposa). For a score of species the revision of the material would be particularly interesting. They are Coenobita cavipes, Petrolisthes rufescens, Dardanus guttatus, Matuta victor, Charybdis annulata, Portunus pelagicus, Lophozozymus pictor, Trapezia septata, Chasmagnathus subquadratus, and Cryptochirus coralliodytes, corresponding to isolated, usually old references, never again cited in recent revisions; Enoplometopus holthuisi and Neopetrolisthes maculatus, cited only in non-taxonomic books; Neoliomera pubescens, Actaea calculosa, Forestia depressa, Forestia scabra, and Actumnus globulus, for which it is clearly indicated, in systematic studies, that the revision of this material is necessary; and Trapezia formosa, re-examined in Paris and perhaps belonging to a new species (P. CASTRO, personal communication). Furthermore, about 30 additional species, recorded in ecological works, with sometimes only provisional determinations, could be added to the above mentioned species (cf. for example, Calcinus minutus, Galathea aff. amamiensis, Liomera laperousei, Paramedaeus simplex, Actaea aff. glandifera, Zozymodes xanthoides...). The status of 15 species is doubtful and must be revised. They are: Coenobita carnescens and C. olivieri, that could respectively be synonyms of C. perlatus and C. spinosus; Galathea latirostris and Cryptodromia coronata, two species whose exact identity remains to be defined; Thalamita minuscula, Parapleurophrycoides roseus, and Platyozius perpusillus, described from very small specimens which could only be the juveniles of more common species; Xanthias tetraodon, possibly a synonym of X. nitidulus; Etisus punctatus, described only after drawings, the corresponding material being lost; and Porcellana monolifera, Actaeomorpha alvae, Portunus alexandri, Grapsus depressus, Ptychognathus crassimanus, and Sesarma jacquinoti, 6 species described from French Polynesia a long time ago, and never recorded since. 78 BIOGEOGRAPHY For the Brachyura, FOREST & GUINOT (1962) have already established that the French Polynesian fauna is a part of the Indo-West Pacific fauna. Located at the eastern limit of this area, French Polynesia is characterised by a lower diversity than in the Indo-Malaysian area, considered as the origin from where the Indo-West Pacific fauna has extended. This assumption can be verified here for other groups. For the Astacidea and Palinuridea, HOLTHUIS (1991) records 27 western-pacific species (zone 71 = Malaysia, Indonesia, Philippines, New Guinea), collected within the first 100m, against only 14 in French Polynesia. For the genus Clibanarius, RAHAYU & FOREST (1992) report 20 Indonesian species, against only 7 in this work. For other diogenids the comparaison with the Indonesian fauna (in RAHAYU, 1992) reveal the following discrepancies: Aniculus 4 vs 2 species, Calcinus, 23 vs 10 species, Dardanus 10 vs 8 species, and Diogenes 18 vs 1 species. In our list, 21 species are known only from French Polynesia. For most of them, it is doubtful that they are real endemic forms: 3 have been described from very small specimens which could be juveniles of other species (Parapleurophrycoides roseus, Platozius perpusillus, Thalamita minuscula); 2 are sublittoral to deep species recently described from material collected with difficulty by the use of deep traps (Alainodaeus rimatara, Medaeus grandis); and 8 are some of the doubtful species already mentioned (Coenobita carnescens, Etisus punctatus, Porcellana monilifera, Actaeomorpha alvae, Portunus alexandri, Grapsus depressus, Ptychognathus crassimanus, and Sesarma jacquinoti). Concerning this last group, let us recall that Ruppellia granulosa, never recorded since its description from the Marquesas, is here proposed as a junior synonym of the Lydia annulipes, widely distributed in the Indo-West Pacific. The 8 remaining species, which could be true endemic forms, are the following: Parribacus holthuisi, Micropagurus polynesiensis, Nucia rosea, Nursia mimetica, Lissocarcinus elegans, Acanthophrys cristimanus, Ozius tricarinatus, and Macrophthalmus consobrinus. However, it is likely that some of them are distributed at least as far as the Western Polynesia, and that they will be recorded there when more collections are available. As an example, Calcinus nitidus, formerly considered as endemic from Tahiti, has been recently reported in the Samoa (POUPIN, 1994a). The French Polynesian fauna could be related to the fauna of the Hawaiian islands, which are of similar origin and geomorphology. A comparison between the two areas remains difficult because no detailed list of the Hawaiian fauna is yet available. We however notice that 4 species, Albunea speciosa, Charybdis hawaiensis, Panopeus pacificus and Sesarma angustifrons, are still known only from these two areas. ELDREDGE & MILLER (1995) have recently published the number of Hawaiian species, by Infra- Order. The same calculation, made after our work, is compared with the data of these authors in table 2. The most obvious result is that the French Polynesian fauna is almost twice as rich as the Hawaiian fauna. Nevertheless, the fauna of the Hawaiian islands has been well studied, with some important works, like RATHBUN (1906) or EDMONSON (1959, 1962). This discrepancy could come from the as exhaustive as possible approach that we have adopted in our compilation. In particular we have included: about 30 species recorded only in ecological works, with sometimes incomplete or only preliminary determinations; 45 doubtful species (uncertainty about the taxonomic status or the effective presence in French Polynesia); 92 species belonging to the deep fauna, which has been intensively studied and collected recently. Yet, if these three groups are eliminated from the calculation, the result remains still clearly higher in French Polynesia (326 species vs 246). Thus, this observation would reveal a real difference between the two areas, the number of species being greater in French Polynesia. A similar result has been observed for the barnacles by NEWMAN (1986). This author explains the relatively low diversity of the Hawaiian islands by their great isolation, in particular if the low islands are excluded from the chart (see opt. cit., fig. 2), and a settlement from the southern hemisphere, in part from French Polynesia. 79 Table 1 - Number of French Polynesian species, by Infra-Order and Family. The littoral and sublittoral species come from the above compilation. The deep species have been published in a previous work, updated here in Appendices 1 (The 12 sublittoral to deep species, listed in both works, are counted with the littoral and sublittoral species). INFRA-ORDER FAMILY Littoral and Deep species TOTAL sublittoral (>100m) ASTACIDEA Enoplometopidae 1 1 2 & PALINURIDEA Palinuridae 6 3 9 Synaxidae 2 2 Scyllaridae 5 1 6 subtotal 14 3) 19 ANOMURA Coenobitidae 8 8 Diogenidae 31 6 3) Paguridae 6 1 7 Parapaguridae 10 10 Chirostylidae 2 Dy Galatheidae 8 19 Di Porcellanidae 17 17 Albuneidae 1 1 Hippidae 3 3 Lithodidae 1 1 subtotal 74 39 113 BRACHYURA Dromiidae 4 1 5 Homolidae 8 8 Latreillidae 1 1 Dynomenidae 3 1 4 Raninidae 2, 2 4 Poupinidae 1 1 Calappidae 5 2 7 Leucosiidae 4 3 7 Majidae 13 3 16 Parthenopidae 5 2 7 Eumedonidae 1 1 Cancridae 1 1 Geryonidae 2 2 Goneplacidae 3 3 Portunidae 54 1 55 Xanthidae 123 14 137 Trapeziidae 20 20 Pilumnidae 9 9 Carpiliidae 2 2 Menippidae 13 13 Gecarcinidae 5) 5 Grapsidae 35 35 Pinnotheridae 1 1 Ocypodidae 8 8 Cryptochiridae 2 2 Hymenosomatidae 1 1 Incertae Sedis 3 3 6 subtotal SUIS 48 361 TOTAL 401 92 493 80 Table 2 - Total number of species in French Polynesia (this work, deep species included) and Hawaii (data of ELDREDGE & MILLER, 1995: 7). French Polynesia Hawaii Palinuridae & Astacidae 19 14 Anomura 113 43 Brachyura 361 189 Total 493 246 Few regional distinctions are observed within the French Polynesian Islands. The differences presented in table 3, where the number of species is calculated by archipelago, mainly indicate differences in the number of explorations, and amount of collections. Table 3 - Number of species by archipelago. The number in parenthesis indicates the species known only from the corresponding islands. ARCHIPELAGO TOTAL Austral 28 ~=s (7) Gambier 79 (10) Marquesas 82 (25) Society 282 (103) Tuamotu 226 (49) The Society Islands, with Tahiti, almost inevitable during a stay in French Polynesia, is of course the best studied place. Nearly as many species are known from the Tuamotu Islands, which illustrates the importance of the collections made during the voyages of either the US Exploring Expedition (Ahe, Manihi, Rangiroa, Reao...) or the Albatross (Fakarava, Rangiroa...), and those made by SEURAT (Hao, Marutea South...), RANSON (Hikueru), or MORRISSON (Raroia). The atolls of this archipelago, without rivers, are of course not colonized by fresh or brackish water species such as: the Hippidae of the genus Hippa, the Grapsidae Varuninae (Ptychognathus, Varuna), the Grapsidae Sesarminae (Labuanium, Sesarma), and some Ocypodidae such as Uca chlorophthalmus (cf. POUPIN, 1994a: 71), Macrophthalmus convexus and M. consobrinus (POUPIN, in study). At the southeastern part of Polynesia, the small archipelago of the Gambier Islands, despite its isolation, has been relatively well sampled, thanks to the collections made by SEURAT during his several years stay at Mangareva. In comparison, the Marquesas, far more extented, in the vicinity of the equator, remain poorly known. Affected by particular hydrological conditions, at the origin of the remarkable absence of a reef barrier, they, nonetheless, do not have any regional characteristics. The 25 species that, within French Polynesia, are still known only from these islands, are almost always very common in the Indo-West Pacific. The only exception seems to be the absence of the coconut crab (Birgus latro) whose presence was never verified during our frequent visits and inquiries. With only 28 species, the Austral islands have been clearly less sampled than the others. The northern islands (Maria, Rimatara, Rurutu, Tubuai, and Raevavae) have a fauna similar to the rest of Polynesia, and a lot of common species, although not yet recorded here, have been observed during our =" 81 stays: Panulirus penicillatus, Coenobita perlatus, Calcinus laevimanus, Pachygrapsus plicatus, Cardisoma carnifex... and even, in the mouths of the rivers of Raevavae, the big portunid Scylla serrata, common in the high Society Islands. Far more south, at the southern limit of the tropical area, the island of Rapa, and the islets of Marotiri, are affected by the particular climatic conditions prevailing in that place. The perceptible decrease of the water temperature allows only a feeble growth of coral, without a barrier reef. Ashore, the vegetation is affected by a milder climate, the coconut tree being almost absent. This particular situation has an influence on the decapod fauna. Some species, very common elsewhere, have never been found after several visits and inquiries: the coenobite Coenobita perlatus, the coconut crab Birgus latro and the land crab Cardisoma carnifex. In contrast, at least one species, the lobster Panulirus pascuensis, has settled in these islands, whereas it is absent in the northern Polynesia. LITERATURE CITED ALCOCK, A., 1900. — Material for a carcinological fauna of India, 6. The Brachyura Catometopa or Grapsidae. Journal Asiatic Society of Bengal, Calcutta, 69, part 2 (3): 279-456. — 1905. — Catalogue of the indian decapod crustacea in the collection of the indian Museum. Part II, Anomura, Fasciculus I, Pagurides. Calcutta: 1-197, pl. 1-16. ANONYMOUS, 1979. — Essai de péche de fond a l'extérieur du récif. Pose de casiers par le Tainui. Association Territoire de la French Polynesia, Centre National d'Exploitation des Océans, Tahiti, Centre Océanologique du Pacifique COP/D: 1-14, fig. 1-7. 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L., MCCAIN, J. C. & E. S. DAVIDSON, 1973. — Decapoda I, Brachyura I, Fam. Pinnotheridae. Jn: GRUNER, H. E. & L. B. HOLTHUIS (eds), Crustaceorum Catalogus, Pars 3: 1- 160. nn se eee 93 SENDLER, A., 1923. — Die Decapoden und Stomatopoden der Hanseatischen Siidsee-Expedition. Abhandlungen Herausgegeben von der Senckenbergischen Naturforschenden Gesellschaft, 38: 21-47, fig. 1-3, pl. 5-6. SERENE, R., 1972. — Observations on the Indo-Pacific species of Kraussia Dana 1852 (Decapoda, Brachyura). Journal of the Royal Society of Western Australia, 55 (2): 41-61, fig. 1-24. — 1973. — A new species of Decapoda Hippidae: Albunea mariellae nov. sp. from the Banda sea. Crustaceana, 24 (3): 261-264, pl. 1-2. — 1975. — Note additionnelle sur les espéces indo-pacifiques de Quadrella Dana, 1851 (Crustacea, Decapoda, Brachyura). Bulletin de la Society Zoologique de France, 100 (4): 509-521, fig. 1-13, pl. 1-2. — 1977. — Crustacés hippidés et brachyoures des iles Seychelles (lére partie). Revue de Zoologie Africaine, 91 (1): 45-68, fig. 1-43. — 1984. — Crustacés décapodes brachyoures de l'océan Indien occidental et de la mer Rouge. Xanthoidea: Xanthidae et Trapeziidae. ORSTOM Collection Faune Tropicale, 24: 1-349, fig. 1- 243, pl. 1-47. SERENE, R. & N. V. LUOM, 1959. — Note additionnelle sur les espéces de Chlorodopsis (Brachyoures). Annales de la Faculté des Sciences, Saigon: 301-340, fig. 1-5, pl. 1-3. SERENE, R. & C. L. SOH, 1970. — New Indo-Pacific genera allied to Sesarma Say 1817 (Brachyura, Decapoda, Crustacea). Treubia, 27 (4): 387-416, pl. 1-8. SERENE, R. & A. F. UMALI, 1972. — The family Raninidae and other new and rare species of brachyuran decapods from the Philippines and adjacent regions. The Philippine Journal of Science, Manila, 99 (1-2): 21-105, fig. 1-131, pl. 1-9. SERENE, R., TRAN, V. D. & V. L. NGUYEN, 1958. — Eumedoninae du Viet-Nam (Crustacea). Treubia, 24, part 2: 135-242, fig. 1-14, pl. 4-7. SEURAT, L. G., 1904a. — Observations biologiques sur les Cénobites (Cenobita perlata, Edwards). Bulletin du Muséum d'Histoire naturelle, 5: 238-242. == 1904b. — Lettre sur les travaux du laboratoire de Rikitea. Bulletin du Muséum d'Histoire naturelle, 10 (3): 94-95. — 1934. — La faune et le peuplement de la French Polynesia. Jn: P. LECHEVALIER & fils (éd.), Contribution a l'étude du Peuplement zoologique et botanique des iles du Pacifique. Society de Biogeography, 4: 41-74. STEPHENSON, W., 1972. — An annotated check list and key to the Indo-West Pacific swimming crabs (Crustacea, Decapoda, Portunidae). Bulletin of the Royal Society of New Zealand, 10: 1-64. _— 1976. — Notes on Indo-West Pacific Portunids (Decapoda, Portunidae) in the Smithsonian Institution. Crustaceana, 31 (1): 11-26, fig. 1. STEPHENSON, W. & B. CAMPBELL, 1959. — The australian portunids (Crustacea, Portunidae). III, The genus Portunus. Australian Journal of Marine and Freshwater Research, 10 (1): 84-124, fig. 1-3, pl. 1-5. STEPHENSON, W. & J. HUDSON, 1957. — The Australian Portunids (Crustacea, Portunidae). I, The genus Thalamita. Ibid., 8 (3): 312-368, fig. 1-5, pl. 1-10. STEPHENSON, W. & M. REES, 1961. — Sur deux nouveaux crustacés Portunidae indo-pacifiques. Bulletin du Muséum national d'Histoire naturelle, 2eme série, 33 (4): 421-427, fig. 1-2. — 1967. — Some portunids crabs from the Pacific and Indian oceans in the collection of the Smithsonian Institution. Proceedings of the United States National Museum, Smithsonian Institution, Washington, 120 (3356): 1-114, fig. 1-38, pl. 1-9. 94 STIMPSON, W., 1858a. — Prodromus descriptionis animalium evertebratorum que in Expeditione ad Oceanum Pacificum Septentrionalem, a Republica Federata missa, Cadwaladara Ringgold et Johanne Rodgers Ducibus, observavit et descripsit. Pars IV, Crustacea Cancroidea et Corystoidea. Proceedings of the Academy of Natural Sciences, Philadelphia, 10: 31-40 [29-37]. — 1858b. — Ibid. Pars V, Crustacea Ocypodoidea, 10: 93-110 [39-56]. — 1858c. — Ibid. Pars VII, Crustacea Anomoura, 10: 225-252 [63-90]. — 1860. — Ibid. Pars VIII, Crustacea Macrura, 10: 91-116, [22-49]. — 1907. — Report on the Crustacea (Brachyura and Anomura) collected by the North Pacific Exploring Expedition, 1853-1856. Smithsonian Miscellaneous Collections, 49 (1717): 1-240, pl. 1-26. TAKEDA, M., 1980. — Pilumnus planus Edmonson and Leptodius leptodon Forest & Guinot as synonym of Forestia depressa (White) and Leptodius davaoensis Ward (Decapoda, Brachyura). Crustaceana, 39 (3): 318-320. TAKEDA, M. & S. MIYAKE, 1968. — Pilumnid crabs of the family Xanthidae from the West Pacific. I. Twenty-three species of the genus Pilumnus, with description of four new species. OHMU Occasional papers of Zoological Laboratory Faculty of Agriculture, Kyushu University, 1 (1): 1- 58, fig. 1-13, pl. 1-4. — 1970. — Lybia edmondsoni sp. nov., a new anemone crab from the Hawaiian islands. Proceeding of the Japanese Society of Systematic, Zoology, 6: 11-15, fig. 1-6. TERAO, A., 1913. — A catalogue of hermit-crabs found in Japan (Paguridae excluding Lithodidae), with descriptions of four new species. Annotationes Zoologicae Japonenses, 8 (2): 355-391, fig. 1-4. TESCH, J. J., 1917. — Synopsis of the genera Sesarma, Metasesarma, and Clistocoeloma with a key to the determination of the Indo-Pacific species. Zoologische Mededeelingen, Leiden, 3 (2-3): 127-260, pl. 15-17. — 1918. — The Decapoda Brachyura of the Siboga expedition. I, Hymenosomidae, Retroplumidae, Ocypodidae, Grapsidae, and Gecarcinidae. Siboga Expedition, Monographie, 39 c: 1-148, pl. 1-6. THOMASSIN, B., 1969. — Identification, variabilité et écologie des Hippidae (Crustacea, Anomura) de la région de Tuléar, S.W. de Madagascar. Recueil des Travaux de la Station Marine d’'Endoume, fascicule hors série, supplément n°9: 135-177, fig. 1-20. — 1973. — Albunea madagascariensis n. sp., nouvelle espéce d'Hippidae (Decapoda, Anomura) des sables coralliens de la région de Tuléar (S.W. de Madagascar). Crustaceana, 24 (3): 265-274, fig. 1-2, pl. 1. THOMASSIN, B. A., JOUIN, C., RENAUD-MORNANT, J., RICHARD, G. & B. SALVAT, 1982. — Macrofauna and meiofauna in the coral sediments on the Tiahura reef complex, Moorea island (French Polynesia). Téthys, 10 (4): 392-397, fig. 1-5. TITGEN, R. H., 1986. — Hawaiian Xanthidae (Decapoda, Brachyura). II, Description of Garthiella, new genus, with a redescription of G. aberrans (Rathbun, 1906). Proceedings of the Biological Society of Washington, 99 (1): 56-60, fig. 1-2. TURKAY, M., 1971. — Die Portunidae des naturhistorischen Museums Genf, mit einem anhang iiber die typen von Ovalipes ocellatus floridanus Hay & Shore 1918 (Crustacea, Decapoda). Archives des Sciences de Geneve, 24 (1): 111-143, fig. 1-3, pl. 1-6. — 1973. — Zur synonymie von Epigrapsus notatus und Cardisoma carnifex (Crustacea, Decapoda). Senckenbergiana Biologica, 54 (1-3): 105-110, fig. 1-8. — 1974. — Die Gecarcinidae Asiens und Ozeaniens (Crustacea, Decapoda). Senckenbergiana Biologica, 55 (4-6): 223-259, fig. 1-19. WARD, M., 1942. — Notes on the crustacea of the Desjardins Museum, Mauritius Institute, with descriptions of new genera and species. The Mauritius Institute Bulletin, 2 (2): 49-113, pl. 5-6. 95 WILLIAMS A. B., 1982. — Revision of the genus Latreillia Roux (Brachyura, Homoloidea). Quadreni del Laboratorio di Tecnologia della Pesca, 3 (2-5): 227-255, fig. 1-8. ACKNOWLEDGEMENT The authorities of the Service Mixte de Surveillance Radiologique et Biologique, G. MARTIN and C. PAYEN, have permitted this research through the collaboration between their Institution and the Laboratoire de Zoologie des Arthropodes (Muséum national d'Histoire naturelle, Paris), where most of the literature was found. In the Laboratoire de Zoologie des Arthropodes, A. CROSNIER, J. FOREST, and D. GUINOT, have always assisted us in our research, and have contributed to improve this work by their corrections and advice. A.J. BRUCE has helped us for the english translation. Finally, the recent collections made in French Polynesia have been greatly facilitated by the whole crew of the Marara, and her last two commanding officers, R. AUDIGIER and M. BENARD. To all of them we wish to express our thanks. APPENDICES 1 - DEEP SPECIES (from 100m, and deeper) With the exception of the shrimps (Dendrobranchiata and Caridea), this list resume the work presented at the International Senckenberg Symposium, Crustacea Decapoda, Frankfurt, October, 1993 (POUPIN, 1996). Full references on the origin of the material, location and depth of the collections are available in that work. Some species, collected or described since the first compilation are added (in bold). For the new species the reader will find more information in the following works: CROSNIER (1995) for Pleurocolpus boileaui gen. & sp. nov., DAVIE (1995) for Nanocassiope oblonga nov., FOREST (1995) for the new genera Ciliopagurus and Strigopagurus, GUINOT & RICHER DE FORGES (1995) for the revision of the homolids and the new genus Yaldwynopsis, HOLTHUIS (1993) for Scyllarus rapanus nov., LEMAITRE (1994) for the genus Sympagurus, MANNING (1993) for Chaceon australis nov., POUPIN (1994b, 1995) for the genera Justitia and Naxioides, POUPIN & MCLAUGHLIN (1996) for Solitariopagurus sp. nov., and SAINT LAURENT, de & POUPIN (1996) for Eumunida treguieri nov. Twelve sublittoral to deep species, followed by a "* " are common with the previous list (cf. Conventions). INFRA-ORDER ASTACIDEA FAMILY ENOPLOMETOPIDAE Hoplometopus gracilipes de Saint Laurent, 1988 96 INFRA-ORDER PALINURIDEA FAMILY PALINURIDAE Justitia longimanus (H. Milne Edwards, 1837) * Justitia vericeli Poupin, 1994 Palinustus unicornutus Berry, 1979 (Coll. 1995, Fangataufa, 250m, det. POUPIN & CHAN). Puerulus angulatus (Bate, 1888) FAMILY SYNAXIDAE Palibythus magnificus Davie, 1990 * FAMILY SCYLLARIDAE Scyllarus aurora Holthuis, 1981 * Scyllarus rapanus Holthuis, 1993 INFRA-ORDER ANOMURA FAMILY DIOGENIDAE Bathynarius albicinctus (Alcock, 1905) Bathynarius pacificus Forest, 1993 Ciliopagurus major Forest, 1995 Ciliopagurus pacificus Forest, 1995 Ciliopagurus plessisi Forest, 1995 Dardanus australis Forest & Morgan, 1991 * (Coll. 1995, Rapa 70-115m, det. FOREST). Dardanus brachyops Forest, 1962 « Strigopagurus poupini Forest, 1995 FAMILY PAGURIDAE Solitariopagurus sp. nov. Poupin & McLaughlin, 1996 FAMILY PARAPAGURIDAE Strobopagurus cf. gracilipes (A. Milne Edwards, 1891) = S. cf. sibogae in POUPIN 1996 fide LEMAITRE (1994: 378) Sympagurus affinis (Henderson, 1888) Sympagurus boletifer (de Saint Laurent, 1972) Sympagurus bougainvillei Lemaitre, 1994 Sympagurus dofleini (Balss, 1912) Sympagurus planimanus (de Saint Laurent, 1972) Sympagurus poupini Lemaitre, 1994 Sympagurus trispinosus (Balss, 1911) Sympagurus tuamotu Lemaitre, 1994 Sympagurus wallisi Lemaitre, 1994 FAMILY GALATHEIDAE Leiogalathea laevirostris (Balss, 1913) Munida amathea Macpherson & de Saint Laurent, 1991 Munida ducoussoi Macpherson & de Saint Laurent, 1991 Munida evarne Macpherson & de Saint Laurent, 1991 Munida hystrix Macpherson & de Saint Laurent, 1991 Munida lenticularis Macpherson & de Saint Laurent, 1991 Munida longicheles Macpherson & de Saint Laurent, 1991 Munida normani Henderson, 1885 Munida ocellata Macpherson & de Saint Laurent, 1991 of Munida pasithea Macpherson & de Saint Laurent, 1991 Munida plexaura Macpherson & de Saint Laurent, 1991 Munida polynoe Macpherson & de Saint Laurent, 1991 Munida profunda Macpherson & de Saint Laurent, 1991 Munida pulchra Macpherson & de Saint Laurent, 1991 Munida rubella Macpherson & de Saint Laurent, 1991 Munida rubrovata Macpherson & de Saint Laurent, 1991 Munida sp. cf. pilosimanus Baba, 1969 Munida sp. cf. solae Baba, 1986 Sadayoshia aff. edwardsii Miers, 1884 FAMILY CHIROSTYLIDAE Eumunida keijii de Saint Laurent & Macpherson, 1990 Eumunida treguieri de Saint Laurent & Poupin, 1996 FAMILY LITHODIDAE Lithodes megacantha Macpherson, 1991 INFRA-ORDER BRACHYURA FAMILY DROMIIDAE Dromia wilsoni (Fulton & Grant, 1902) * Sphaerodromia ducoussoi McLay, 1991 FAMILY DYNOMENIDAE Dynomene tanensis Yokoya, 1933 (Coll. 1995, Fangataufa, 310m, det. MCLAY). FAMILY HOMOLIDAE Homola ikedai Sakai, 1979 1 Homola orientalis s.1. Henderson, 1888 Homologenus broussei Guinot & Richer de Forges, 1981 Hypsophrys inflata Guinot & Richer de Forges, 1981 Hypsophrys aff. murotoensis Sakai, 1979 Hypsophrys personata Guinot & Richer de Forges, 1981 Moloha aff. majora Kubo, 1936 Yaldwynopsis aff. spinimanus Griffin, 1965 FAMILY LATREILLIIDAE Latreillia metanesa Williams, 1982. FAMILY RANINIDAE Notopoides latus Henderson, 1888 Notosceles chimmonis Bourne, 1922 « Notosceles viaderi Ward, 1942 rl FAMILY POUPINIIDAE Poupinia hirsuta Guinot, 1991 FAMILY CALAPPIDAE Calappa aff. hepatica (Linné, 1758) = Calappa sp. nov. (GALIL com. pers.) Mursia hawaiensis Rathbun, 1893 FAMILY LEUCOSIIDAE Oreotlos encymus Tan & Ng, 1993 98 Oreotlos potanus Tan & Ng, 1993 Randallia serenei Richer de Forges, 1983 FAMILY MAJIDAE Cyrtomaia ihlei Guinot & Richer de Forges, 1982 Naxioides teatui Poupin, 1995 Naxioides vaitahu Poupin, 1995 FAMILY PARTHENOPIDAE Parthenope (Platylambrus) poupini Garth, 1993 Parthenope (Platylambrus) stellata Rathbun, 1906 FAMILY CANCRIDAE Platepistoma balssii (Zarenkov, 1990) FAMILY GERYONIDAE Chaceon australis Manning, 1993 Chaceon poupini Manning, 1992 FAMILY GONEPLACIDAE Carcinoplax aff. cooki Rathbun, 1906 Carcinoplax aff. crosnieri Guinot & Richer de Forges, 1981 Carcinoplax aff. verdensis Rathbun, 1914 FAMILY PORTUNIDAE Charybdis paucidentata A. Milne Edwards, 1861 * Parathranites hexagonum Rathbun, 1906 (Coll. Eiao, Vanavana; 155-240m, det. Moosa) Portunus nipponensis Sakai, 1938 * Thalamita macrospinifera Rathbun, 1911 * FAMILY XANTHIDAE Alainodaeus akiaki Davie, 1993 Alainodaeus rimatara Davie, 1993 * Banareia fatuhiva Davie, 1993 Demania garthi Guinot & Richer de Forges, 1981 Demania mortenseni (Odhner, 1925) Epistocavea mururoa Davie, 1993 Euryozius danielae Davie, 1993 Hypocolpus mararae Crosnier, 1991 Lophozozymus bertonciniae Guinot & Richer de Forges, 1981 Medaeus grandis Davie, 1993 * Meractaea tafai Davie, 1993 Meriola rufomaculata Davie, 1993 Nanocassiope oblongaDavie, 1995 Paraxanthodes polynesiensis Davie, 1993 Rata tuamotense Davie, 1993 Pleurocolpus boileaui Crosnier, 1995 XANTHOIDEA JNCERTAE SEDIS Beuroisia manquenei Guinot & Richer de Forges, 1981 Mathildella maxima Guinot & Richer de Forges, 1981 Progeryon mararae Guinot & Richer de Forges, 1981 99 2 - PARTIAL IDENTIFICATIONS These incomplete references were generally found in ecological works. To avoid partial identifications in the main list, they are presented separately hereafter, by alphabetical order. Four genera are cited for the first time in the area: Glabropilumnus, Heteropanope, Libinia, and Tylodiplax. Actaea sp. Actumnus sp. Dromia sp. Euxanthus sp. Glabropilumnus sp. : Heteropanope sp. Libinia sp. Lybia sp. Neoliomera sp. Pachygrapsus sp. Paramedaeus sp. Parthenope sp. Petrolishtes spp. Phymodius sp. Pilodius sp. Pilumnus spp. Tylodiplax sp. Xanthias sp. MONTEFORTE, 1984: 170, annex 1, tab. a (Moorea, Tahiti); 1987: 8 (Moorea). MONTEFORTE, 1984: 170, annex 1, tab. a (Makatea). — GUINOT, 1985: 452 (List). ODINETZ, 1983: 208, with a ? (Tahiti). — GUINOT, 1985: 448 (List). PEYROT-CLAUSADE, 1977a, annex of the species: 26 (Moorea), juvenile; 1977b: 212 (Moorea). — GUINOT, 1985: 450 (List). MONTEFORTE, 1984: 170, annex 1, tab. a (Mataiva). — DELESALLE, 1985: 289 (Mataiva). — GUINOT, 1985: 452 (List). PEYROT-CLAUSADE, 1989: 113 (Tikehau). SENDLER, 1923: 40 (Tahiti). PEYROT-CLAUSADE, 1977a, annex of the species: 27, juvenile (Moorea). PEYROT-CLAUSADE, 1977a, annex of the species: 27, juvenile (Moorea). HOLTHUIS, 1953: 32 (Raroia). PEYROT-CLAUSADE, 1989: 113 (Tikehau). MONTEFORTE, 1984: 174, annex 1, tab. a (Takapoto). — SALVAT & RICHARD, 1985: 350 (Takapoto). MONTEFORTE, 1984: 173. — PEYROT-CLAUSADE: 1977: 25. PEYROT-CLAUSADE, 1977a, annex of the species: 27, juvenile (Moorea). PEYROT-CLAUSADE, 1977a, annex of the species: 27, juvenile (Moorea). MONTEFORTE, 1984: 171, annex 1, tab. a, photo p. 131a (Takapoto); 1987: 9 (Moorea). — SALVAT & RICHARD, 1985: 350 (Takapoto). THOMASSIN et al., 1982: 393 (Moorea). — GUINOT, 1985: 453 (List). PEYROT-CLAUSADE, 1989: 112, 115 (Moorea, Tikehau). 100 INDEX A Gffine: PENCHONG. cosiaso cored reece cece eee 71 GODT eVIGIUMP Cr CnOn ew neater ease 71 affinis abbreviatus, ACANIROPUS ...se.sssesssessssssessessssseesseeeses 71 DCLG sian ctssesnssteaiaasetsinvsnagecnes sussesas oosenanserueenerne 55 aberrans, Garthiella, Pilodius ....c...cc+c.+cssc+sssses00se000- 54 TNE LTT ION CA tes pee PAP eee 5)3) Acanthophrys Cristim@nu ...cc.cccscsssssssesssesssseeeees 26; 78 Galathea wanes. dais ayant athitd ae 19 Acanthopus PON CHOM ooo ee che 71 GQDDIEVIGIUS ares oo sce canee eee eases see eee 71 SYMPAQUIUS -.eossseerressnesecssaseeeernsnseceesnnseceennnsesetins 96 DIGTUSSIMUSTs FOO RA os soca tact chee emenctee coteck 71 AKiaKt, AlAiNOdAeUS ........-ssseeeresseenevneeessneecesieecnsneseenns 98 LORUUTONS sion vsesaastocticacocnessndtituns wusatiinia, antares 71 Alainodaeus Achelous, Neptunus, POrtunus ....cc...cc0.-sscs00ss0e000sse-- 31 RIOR, che Meccssnscsnnancnatons cctansn chee cee on inaten cone eae 98 Actaea VUNG Gone cere ee eee sass stacdnns exes Soees 7; 41; 78; 98 CUT LIBLS eee oes sh a ee ae ee ge 55 albicinctus, Bathynaris ........ss.0errsveerseeresseeerenveeesien 96 COICUIOSG see mic tisseet coe ete ee ee re 42;77 albolineatus, GrAapSUs......ccrieecreeerieer viernes ceenneeeens 67 EGV IPOS EO RUSS, Tete ee eee can 44 AlbUrned SPECIOSA .......eeseresvneseeernnveeercnnveeeeenns 22; 76; 78 GOMSODI INR cece co er eee ee ROO 42; 45 AIbUS, PSCUdOQIAPSUS .....2-.+s.csseeroseeevneeesaseesasesenneeeons 69 Ganges eo 42 alcocki EDI ESSENSE aie RO ee 43 LiOXAntHOdES ......seeeeerervineseeerrrenveseeecnnnneeseeseennneee 49 GQ GTTOUE a8 acon sti sce cs cactcsci cece ke RR 43 TRALAMit ....seseesseessseesssesceeeseeseseesueeseseneesansennes 36 SEITE CEPT ATO ble PESO teria Bis Shit 42;77 alexandri HUSUUSSING een er ee 42 Caine ctes wooseeeeeeesesnseseeeessnnneeceennnnneecccennnassee 31 Renee te 2 al 45 er eee ee eee, 31; 77; 78 DGRV Uae ere Ftc reer ee ne 38 Algae, GAlathed «.....eeresevsvvveeeeeeeeseensnnniisteeeeccennenninss 20 POLWACONINGR een en one e 42 alvae, ACLACOMOTPHA ......r.e1vseseeeeeeneesninns 5; 28; 77; 78 FUfOPUClAIGs WAREMAN 3s chen lig SP ena 43; 44 Amamiensis, GalAHed ..........sseerreeerreessnreerrsveevsees L977 PILDPeLIOI eS wh erat weeks Rested. SEE: 42; 45 mathe, MUnida see sietr fev scesensusenasinsosaseossontacentoeete 96 SCD Gi eer eM te recrets Mece cha sects ceete atcha 43 Amphiucd, UCO viesssssesseeseeeeeeeseececcrecerreseerennnneiesieeie 74 SPicee OO Te 99 Gnaglypia PlatypOdi@, .oerecnxcc-cnecn-necnnecvennenensoreaeeeeer 46 SUPER CUUGT Ss seat cin tee ai oes ease ence 43 Anaglyptus, ELiSUS .....000.ereeereecreerrneereerneeees 6; 52; 53 LOMENIOSG cre oer Me eee res 43 anceps, Eupagurus, PAgurixus ......--crseerrrreeenneseensees 19 tum lOSat eT AEE BONIS ho oes 44 ANA COSSY1, CYMO ....seesseesvescsesseeeseeeseesereneesnceeeenesnness 38 Actaeodes ANQUIAUST PUCTULUS?. Jc foteceess- +20 sssccatente tooeeseseeteeetere 96 OPTI S oan ee wae ee Se Ne re 55 angustifrons GREOIEES oo ate tse ee ce 42 MicipPOidess ..........-.-ssveseerssseeseesoseenesnneeseennnsceenen 27 CONSODI INS Pics Semen. ee i, em ee 42 SCSAPMA ooessservssesvesvecrseecnssecsneccnesennesnseeenees 71; 78 Hi SULSSUMUS Tht Oe ek ee 42 Aniculus FICHIERS a ein ae ce ee anne: 4] CAIAIGLULLGS YRS, Bo Reavis cc. csi socscusessanssaetcereees 13 FONIETLOSTIS cr eT ee et 43 TAAXIMUS Bascave coksouns ons oaatisot ince swan eSeeaeee Re ee 14 Actaeomorpha SPs osvdeahateeeteaeess Petese ee once ce nsoanacs neat eee 14 AVaelteeti on ees in werent enn 5; 28: 77; 78 EY PIOUS ss ic ee cue aiwccoussncxsseateuas atone inane iateettarereceeeee 13 COS octdor erates etree Nene Sa ae 28 aniculus, ANicUlUs, PAQUrUS ..........0rs.erreeceveerrseesenees 13 DUNICIAL Doe cxsan cae coe RAG 28 annulata Actumnus GRAN Y DAISK. 85 BS Sieh cake Biss ccssncasnce sbossseonsseleee ee 30; 77 SPORE ee PR A CNS Ia a ce 61 GOMLOSOMA........0eoseeevveeesnrenseesnnessneenseennnceniennneennes 30 DONUT access coe tecseon niet tests oes OTs 61 annulipes Aigitalis. cccsscvcicosesssss: A ae 61 Lid... essrsessseecoseeeshecesseseneesneeesnesnseesnnenseeennes 64; 78 GIODUIUS 3. eee ee ee 61:77 Ridp ellie xs. cccchecccct cote cecs co teectea cee tevomnene nents 64 INLEGITIINUS ecard toon ee 54 antarcticus ODOSUS Riss, Geetha ee TR ha Foster 61 PAPTiDACUS «...000sessseessseeessvensserseeessnerssseesssees LO; HT SCLC sce snsins disse De ee 61 SCYLATUS ...sesccssseeessssssseseneecnsneensnnesssnnennsneesanees 10 Sis snssanvonatiodsveasess al atsaee Meee TR CEN RR 99 Antipodar um, Ar Clides ..........s.-crsveerrseeerneernneeennesennnee 10 fOMUNIOSUS Ah ps See E PR A RO ace scisis 61 Arctides aculeata: GalaMed i ciccsnncen ie eee 19 ANU POAT UM ose esesssesecessseecsvivessnnneesnneeccnnneeenneeen 10 Qdactyla; HIDDG wnas. te. acnvteaea en eaetie nore 23 TOQAIIS .oesssssseersseseecnssneeenssnsscssnecesnnaseeennes 10; 76; 77 admete, Thdlamita (.it NALA NRW 33 GCUALUS, XANENO ..ererserresvvvrvvrrernnseseeeenseceeeeeeeneeeeecen 48 Geneus, ZOTMUS ec. ake Le 47 areolata Gequabilis, CLDANALUS ....sccsesssescsessseessessessssesseeesees 16 CHOP OG OSES cos. ci veicintiesnonscuskeconassvacosencoueceraneeeets 5S A Cth SCTUPOSG «o cscsea-tts-nireanctasetnn eee 28; 77 TrQPeZiG sssveesseeeeeveeveveeseersssnsnssnnnnnnnencscees 3; 58; 60 areolatus A CLACOdES eee Net th cscseccstzniensvies RR NS 42 ILO Shed Sots osscareoceuccsisntie nee OR 55 GVTNU CH NN OCA PUOES | <-ccsc0zc-csocsscn-vsesisteseencorsetesee 54 Ashtoret PV ATUIOS Cres meticectvaevsersevesceves scives mesesttens ee 25; 76 LUN GI IS be F ces ccke tevb hetviseesisvessesass std et ee ee 25 DICE fed Se vedateh deh varias teceasee sussa recceninn dd Peeters tees 25 GASP EN HACIUTIANRUS | occ ccccs su sesus cco cavesesussseoraceen teense ees 61 ASPET. SCHIZOPNUGYS orcs sensed acoceassscoae-Jesesaeeseecoees 27; 77 Atergatis CLE GQ GNS cssarmscsce Scien CR ee ee 46 i LONI AUS psscucoecesieeevaucvnveeeesesteis SO ee 45 LEN DGLUS x tesescscchas cssedessessstens EMO tes ase 47 OGY OC fae icv sis satan taausanteccorssnent sxsunciesbeneerostere 45 Atergatopsis PONG Eases rscatbstracatvsersrectscssessssesia See ote 45 SEDNIGLUS yecteessssis dca ssesiwsscuctcnsesstivectaseseesez ete 45 Aulacolambrus hoplonotus ......10.ccssseseveeseseseretenees 29 GUT OG YSGY UGiUS <.:ccach-css02csscseesesenssiscucesttoneees 7; 11; 96 australis (CRACE OT os eI es 95; 98 DYES sp oec see aca ee eRe 17; 76; 96 B BGISSTUSIPL AICP ISL OMe as tetera secss tenses ase tce eee 98 Banareia GUI os sho soos ssi sc sedenateaduse she nuscdis saute ee 98 DGIVILIG* Ceiein oct eee wa ssecocciemen tania 38 DQNSKESM QU ava aanene cote ee Tees 25 barbata, Chlorodiella .........ceccsceccceseescessenseteeteeteeeens 53 BON fi} CONE CF os crac stecsencsscscestsuse estas Wuseessesso 11 Bathynarius CGIDICINCLUS nextevcccevtussxassiesiseits 96 DACIPICUS here ncesis SIERO 96 bella TAO Giascehoscsccetststcsauetsesatssats Be oes 38 DY AD OD zest ewistvasvicsesvateertsxs teh eo eee 3; 58; 60 bellus, Actaeodes, Carpilodes ..........:cccsccsesseeneeeeee 38 bertonciniae, LOPhOZOZYMUS ........seeceesesesesereeeeneees 98 BeUrOiSid MANQUENEL .0.......seeeseseecesesceseneneteceenenceaeeens 98 Didentatus, LAChNOPOMUS .0.........cseseceeserseseseneceenetenees 48 DifFOntalisHELiSUS cee S152 SETI STOTT 0) Nee OSE ORCC COLOR CORO EG 11; 80; 81 DISPINOSUSN PEI Olisthes iver eres eee eee: 21 Doileaui, PIEUrOCOIPUS ......cccsccecessesseesecseeseesceseeeee 95; 98 Doletifer,, SY MPGQUIUS «0x2 scsscseesssicesentteceoeesanteestonne ete 96 OMNI er yA CLUIMNUS csvescisesteusesstesdiersivescecives Seer 61 borradailei TI YASt NUS sss sessccscsssessvzesasresseva Treen 27 BelrOlisthes xccocsctrsrtiesiti scan Sees essences QUEVAZ bose; etrolisthes: 2 ks ees 21 bougainvillei, Sy MPAQurrus .........eccscecereseseeresenseeneee 96 BOUV IEF URGIGIULG ereecsessssesencossceseceentssussssrsesth ree 33 brachyops, Dardanus .0.......cccccescesevesseseseneesene 7; 17; 96 brevimanns;| GOEnODIUG vcccccc.coocecscx tote nto eessteesiees 12 DIOUSSCI TLOMOLOBZENUS nsec. -cvecessa-oerentesderaccesnceteoetes= 97 101 C CACSTILEr ONLY DIG icsacccntascsccssscsescestscs detect ne Seo 37 Calappa COILED P GN scersescsverwssedeesvewestbeussatbehe bees teettbees 6; 25 EDUCA ie sae v asst sare cinarsscesavecesosscenscacesncevece 25; 97 SP ANOV Sacsesussaievavchswadesteceletvasades caste dssaessStieee reas 97 LUDENCUIGD sede se ese se eee Ds) Calcinus CLE GIS hese dt oso ee sevens dccsostisseitees OR Ne 14 SD GUNG. A iets weve sc srscndesel sates ssesseeseoet nase ee teete oes 14 ID MCMCNISI Se ee clos oc tec xiesress sestesssestesocesnosttes 14; 76 ER DSLR Roeder ae 14; 15 UN DONIGIS iscszstvecs sesesteeccasscsseseee casa ceeciv ROE 14; 76 IGEVIMGTIYUS od ee eee AS 14; 81 IGLCnS css cece eek IIS) TRINULUS eer s, Ee De 6; 15; 77 ALERT TTS cpooz asot oe REEL EEOC EOE eee 4; 15; 78 SOUL ate neo k Mosca aes kateatstscaseeenunee ee 4,15 SDIGQUUS 25 cvs cscwescbescicucasassceviswestiepesesuiien Meteor 4; 15 LCV AE TO DINAL foi. ccstsesosconsusestzsh dodveetsneetescse srsbeee IES) LIDICEN crore sok oot an Soci eeete osm See co 14 COIGUIOSQNACIGCG ree hh ee 42;77 G@allinectes|alexand rips rescsccsccccsasnsstoveonctescesteeesen se Sill CGMPOSCI GI CLUSA aire eieccsretteescsesoievess nes oreo eet 26 canaliculata, CryptodrOmi@.............1c1ccssersesseeseeseees 45} Canaliculatus, XANtKhiAS ......0.ccccccccescececesceeesseccesseeeeses 50 Cancer GIFT oe eesctas i sees cecsrei statis Sa 11 TARA DRS OE CEES EEC DEE EOS Ee econ octoaos 10 Caphyra ROMP ONS Sed vast teats svastsaseccveteestesesseenttees 29 UL UOIS Risse ech aie Ae See 30 Carcinoplax COOK IB rece store aa savectta ed eoad tate east eases Woaewe ne 98 CH OST ON Uta eerare eae sc cieacdsvsveuetsosec niece ss Oreo 98 WEN ACTS TS yo tere ea osc aco aewsucaasbsaedoct Sees ROMO 98 Cardisoma COMPUT Os seins bessescks tani tas tee veases tees 65; 81 TELL POS eave. vescsdet covaceesc cov dates stsatazecesets eee eset 65 OD OSU a as cides oasis eed fess ck ores Siem es eo 65 VOLUN UINE reotenaes se eo pan nace 66 CArinipes, ZOZYMOAES ...........csesecseseescsssseessesensetseees 47 CAFNESCENS, COCNODIA ........cccceseccscceescsseeeeaes MDE TTS Ths CArNIfEX, CAP AISOMGA ........esecseserseveesesceseeeecsesenees 65; 81 Carpilius COTIVOXUS iF ess ech oS SR 7; 62 TILAGUL GUUS aeons ain aoc a Sea Season 62 Carpilodes DELL SW ees eactate Ga neces ee ein es 38 CUIGLIN ANUS oS ee hs is as 3 55a 39 PT QIU ALUS ecscecrccie wateenctscessesdvwv tk cccsscbeoueteuerotieettes 40 TROMMICUIOSUS Soe occcccccece sas cassn neces des eee 39; 40 QIU AUS os tecressuvcec cass cis suscs savesscssivss sevesessssasseve stort 39 TU OCLUS Ho rxcnccse cos sacs cinctets csstntsstn.dvaxesioe: Meee MOOR 4O, LIUSLES Meericee ste aces saa aac ane Soe RUIER ate sensu ee eee 40 VEAL PILI CAILULS scat cose one seoeaestvas Soest bene ceeshessanso te es 38 VEN OS US Pesce swakcsswabt evcavesttnccivvseussesbosee ever neato ye 40 GCarpiloxanthUs TUgipes sovissscssssscisesecsecissevseeseccsonenrses, IO 102 Carupa VGOVIUSC UNG coz ascucsstssceseeveesssxetuanscistexssusnisnieosties 29 LEVIUN POS) coor iae ccteacz seostensrccotsset ee entaceh ore care 29 Catapaguroides PRO QUS 5 csc verse ssivsusvascsnstasscsstcnttnsetesstenesveseostvess 5; 18 UFICHOPHENAUNUS ees teeter eee 19 CQLOPIFUs NUIDUS: nc cssencsncceseseeissnncasinrsottheue LN? 29 cavatus, Cycloxanthops, NeoxanthopS .........:.1.01000 50 COVIMMANG, DEM GUA. oe rnecsaccatscsnstentes ctnccns- ee rat 37, cavipes A CLOL ron vaccas echlcrigatcancsasecteonees eonsocnascaneas te 44 GOCNOBUG cei cscsrase ee lo 77. TSQUINUUS iii sscsccecteceutevsseatsete cgentssseintestes Sees RE 44 GAYSUUS PSCUAOZIUS ccrtescccscsanescccnascessceaness eee 65 CELLOS PSQUITIS Orne eet ge 44 Cenobita GOI TRESCONS .o vanysas gos acs ers acstetissicsien essence ee 12 GLY CAG h.nss.n.enscosssocevarsssescecatencteysen Mey ayie 12 OUI CTU oe covaccascqeas sie cuesestgsanezarsocesees CATERED 13 TUQOSG orn rissa. sesstvessusasscesecaesessacnese seein SEES 13 cerathophthalma, OCYyPOde ........c.ccssecesersseseerevensenees 73 Chaceon CLUSU AUIS rine ere er 95, 98 OUD UL, cis sasocrncseunponsesusste changers sas test ee 98 Chaptalt, TRGIGMIUG’ cio. cccsccsccsccsscocses entre 33 Charybdis CETL LG oor cw ncrcsecscteskens etesce nce ee 30; 77 CF YUY OAACEYIG « ......ccsocaocaseo--rsrsstaaneseteeccospestactine 30 RGWICVISIS’ . 5.5 o.csccss-steccct ies oeensnee eR 30; 78 ONVENEALIS 002 Vi acecinns auteur a Ret estes 31 POUCIGENIGIA.. .. avcsectsconcnae em Cota 7; 31; 98 Chasmagnathus SUBQUGArAtUs .........ccceeceeeesenseees 69; 77 Chimmonis, NOtosceles ........00...ccss0cccceserscecesessceees 24; 97 Chlorodiella DOF DOUG e.lcbec cans teste rte asset TR 53 CYR CRED sero nevis cscivtensoseeenss sos: csceucss MORSE EOE 33) TQGVISSHIIIG .ccted Uvccaccatters seciueies ascot enet oa ERTS 53) LQEVISSIINUS.. cssssue cencsseasortccnsses hu tosee+0es SSUES 54 TGC oon scecctuactanccnien ersten sac teeehe uae eee 54 Chlorodius CYTE OG wossvecssssssoesessnsiaecssansvesspscssscesseS SUReE RNR 216) CONGAAW vcccssececsu cess Seneetxene to ceeeet tac cnie sen cc ME 55 LGC VISSIINUS). ».cscessszscssessspsadsvacesceseconssnes sate tere as 53 INONUUCUIOSUS) ic ccssesscectesssseseriaa vac cess oR S13) VU QED. cvcsscseeas caxsssises scessciansvsacsoereninsesdesces SOROOINGS 54 SGMPUINCUS) ccryspceconssasseareteenusssenssonsns MASE es 49 UP QULALUS). vc ocecccsescxeansessaryrac Meee eee eee 55 Chlorodopsis I COLUG sis scsccssnssssiconsecesnssrsis siete tent ee entaaes 55 QT ANUIGLUS oo scistesccssocceessvetore ceonsssvestaevec eos Gee 56 PUUL; 10s scossivasess sivseepsaccansauaesssecensveczssscee eg MNteyRNi 56 SCODIICUIG ccc ccxcdarxsacaanasnstanaisarerieseniis SO OES 5) SDUUDES.. cssccessacvacvactrssstscascvaccssveccecsatvccsts BRYN’ 56 VENUS senpisescisainrscestnpieabsbancncscssertoesescste tee 57 chlorophthalmus AINDHLUCG socereseersossnvssussovenaseaeevess «1040 ORO LEEERO ASE « 74 GETGSIAAUS Sooo sn, 00.00ce-csecnceusetesss ses 67 MELAS QnA eli eacenrceinneedeee ees eo 97 Metasesarma TART Led Wiemrecreccor Pecrerexceccre Cre ree eat oT 70 ROUSSE GUAM ai secehtstess seeds secossassobsasaisdeesiee.cee- teem te ses 70 NU QU LOS Gyre esi een ew Rataseas dicdeleedenncasieecuseiceveeen oases 70 Metopograpsus TESS OF, vavovasscchac icskscuiies tee eadcs ies ant oie eee 67 LRU UR OI eosecswosccs osvectvew te tistre cic doc RRR 68 Micippa TRAV CANNON Aes csasiivics Peters eT 27 GUC OMereree tare cccascs cepeasas su snoven sass nsessccwnsesceesececee ce 27 108 Micippoides Angustifrons ........sscccsecsseessesceseesesenees 27 Micropagurus polynesiensis .......0.0sceccserecsseeee 18; 78 Militaris, PetrolistheS .......ccccccccssscccsscccesseseesscenees 21;77 TIUIMELIC GH NUP SIG) Jeaivecsesscesonosssensssovsssucoos snseesteues 26; 78 MINIALG TAP CT Gi svscise ssesoncsnon tence scnsersrcesnegueaveecgaes 59 MINUSCUTA, TRALAMIIG ....0.....00cccercseesscecseeseees 35; 77, 78 minutus GCOIGINUS PRS socrsons i Mooseseseieenenc 6; 15; 77 PGCHY RT APSUS)...:.ncicencsxs.csetieerglnat hess 68 Mitra, POCELANGA ......ccccccccessesseceeesneesensssneceseenseees 22; 77 MitslenstS, ThAlAMUtA .............cscseeeseccccccecesessseceees 35; 76 Miyake, SAAAYOSHIA ......6..1..ccsccescsscssensesessnsessensnsenees 20 MOlONG NGO 0) ccsacccsscocscosscesssesesnescsnsenseuessensgsasaraiet 97 MONOCEFOS, MENAethiu .........00.ceesecceersceeessessecessconees 27 monolifera, Porcellana ..........cc1scseseceeceersene 22; 77; 78 MONLICUIOSA, LIOMETGA .......1...00cccvsscesecssseessseeseseesaseees 39 monticulosus Cr PI Od CS irra oo ia ansneseanenssasnnss omecbsabratehns 39; 40 COR OAS Pe. vcssscessuesscnsssntovsvslooevsem esa sane hes 55 AY MODUS Ciseescattscsccutiespeee Maczevececoerseenntlecs 3; 55 Mortenseni, DEMANIA,..........0cceccceeseceeseeseceececascsscennerss 98 Munida CINE C OR os oss ios sk sain siise eee saie oe 96 CU COUSS OI -osbiaassieke. A saiecsurrteen Ga Nh. caer icoras 96 EV OTOP Mi aoe ches voscscalenbce eM eet ieee 96 iy SUING ois cate oes ect ace ascasesseetesssiasssastsceasdurdecaues 96 VCTLEGUT ONS ee es sei acoc cack souss eomee eee 96 VONGICNELOS ose vcccucesssnsevisesestssnvittnetecotionManscrs elem 96 TLOVIMLGM Wisse ce seen chs sass ease ee cea a ee eee 96 CICCAN TZ SR UE GA NT Se heat 96 DASUREGY es cocsccaciusesascosst coset snnerwaaee tence aucee nde te 97 PUL OSIMANUS occ cccensicccssesessssesesas sates toneseeencece yoda 97 PV OX GUT Apes sal sid ae sect sensi ssccssctessesscuatessteietaneecaae ee 97 DOW RO Cppsreroisssietrattar iscsncsccsnnodsusasuins denaasnscaipebaagss 97 PROPUNAG -oecvsccscicccsectveies cess sisstace sevascsentteeet ties 97 DUST CII Gh ist,« sus s5s c'ssnasascncsinon = ta eopa dts wae shee 97 GUDOUG on. taccse sven dee csaucn ies Gesouenssav ss wee oe 97 WUD OV AEG Bern seas Aesdhncescs ssncies ssecsvinssuscteesscoreeesses 97 SOMO. cis osaacccvexsvondeneusesucsteesae ds soso a seen etind ees 97 MUrOtOeENSIS, HYPSOPHIYS ..1.....0c.ceccvsvrerssversersesseseees 97 MUrSia HAWGICNSIS .0.....0ccccceesccesscensescessecensscessseceaneees 97 MUFULOA, EPIStOCAVEA .......seeveeesecerserereesesesessenssessenees 98 N Nanocassiope ODIONQA ........ccsccccesesscsessessssscnssnees 95; 98 Naxioides PCCD sé. ccasssvecponacaivouavencosnedsanedecsiacesseateak eeAeaeeN Ines 98 VGILGHIU, J cacsvaccasnasesnasossssuncesnncstttr aerate eae 98 NeCtOQrAPSUS POLILUS ......cceesesessecsessesscnscseseseecaseneeeses 66 Neoliomera OMIGIL ccs sssensusvunsdasaavonss vesoces cases Sea eeptnen eacees 40; 41 PPISULGTAS sasiessancincssosscoseeeuevit sede een oe EE 40; 77 DUDOSCONS aasisnsevuscsscevosstusscststiecvescieie ees 40; 41; 77 TIGTHL GUSH... cess shes saccaasteeacicessassa etaasncols eeeeoaaeartot 4] SPs ssncw vues eosesennectosnssass socinnes dasccete tens ce eee eRORSits 99 VOM OLOS Divas ccsevascevvensveseesaveccventecssincc Rees 41; 45 Neopetrolisthes PRACUICLUS ac; cicsucansacunsuccnsevuscnss 74 VAT AITO CRO ee 3; 74 ungulatus (DREADS eee oe 5)s) TAN IIOOIUS a rons cassie cones tease issuvenevas coc agstaaNameesentec’ 55 TAI COLIULUS HE GIIPUESLUS) csscessscereensoonseeedeteeeeeeeees eae 96 MEP SUIS= IG] OFS PGI TE DACUS watever ace ner se) cece eet iH VINCI OCY DOE, ti cscccscccevesassussscxcscreceves uaceceeeeemagi: 73 V vaillantianus, Carpilodes..........ccccccecensessesensecseeeenee 38 VI ARUMIN GNI OIE) .csic:.0.002<0s0ess <0 .005-52 SAR 98 Variegatus, LEptOgrAPSUS .........cseccsercvecssvscsetessevenees 67 variolosa INC OLIOINEN, GiB scoters. sas. dose sca nessessanesee ius SEREOS 41; 45 IP SCH OMOMEN.G sox. Hoedscvoegees seakesaccessio1sesstueeeeees 45 WASTED TTR CG Mey perce e oo a Reon Re SSSOACOO ho 6; 69 VEMOSC WA OMENG ics ssicagasxsnaasacisaad ree Re 40 VENOSUS, |G At PULOd OS). .nnn:-----t8so esses ess ec Nes 40 venusta (QUO Od OP SIS 5 cio sonia ncvensosornnesctaseeae ous ae ees 5/7, CLE ONU CIO éics50653 SR A eR eS 26 Verdensis;|GarGiiOp aX a vsasaccosesteees to. eect eae 98 VETIGELIPHUTESTELE Qi tenes. See re RESEARCH BULLETIN | Issued by NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C. U.S.A. OCTOBER 1997 ATOLL RESEARCH BULLETIN NOS. 443-449 NO. 443. NO. 444. NO. 445. NO. 446. NO. 447. NO. 448. NO. 449. THE EVOLUTION OF A HOLOCENE FRINGING REEF AND ISLAND: REEFAL ENVIRONMENTAL SEQUENCE AND SEA LEVEL CHANGE IN TONAKI ISLAND, THE CENTRAL RYUKYUS BY H. KAN, N. HORI, T. KAWANA, T. KAIGARA, AND K. ICHIKAWA CHECKLIST OF THE SHOREFISHES OF OUVEA ATOLL, NEW CALEDONIA BY MICHEL KULBICKI AND JEFFREY T. WILLIAMS ON THE ORIGIN OF DRIFT MATERIALS IN THE MARSHALL ISLANDS BY D.H.R. SPENNEMANN DISTRIBUTION OF RAT SPECIES (RATTUS SPP.) ON THE ATOLLS OF THE MARSHALL ISLANDS: PAST AND PRESENT DISPERSAL BY D.H.R. SPENNEMANN A POSSIBLE LINK BETWEEN CORAL DISEASES AND A CORALLIVOROUS SNAIL (DRUPELLA CORNUS) OUTBREAK IN THE RED SEA BY ARNFRIED ANTONIUS AND BERNHARD RIEGL MARINE ALGAE FROM OCEANIC ATOLLS IN THE SOUTHWESTERN CARIBBEAN (ALBUQUERQUE CAYS, COURTOWN CAYS, SERRANA BANK, AND RONCADOR BANK) BY GUILLERMO DIAZ-PULIDO AND GERMAN BULA-MEYER SCIENTIFIC STUDIES ON DRY TORTUGAS NATIONAL PARK: AN ANNOTATED BIBLIOGRAPHY BY T.W. SCHMIDT AND L. PIKULA ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. OCTOBER 1997 ACKNOWLEDGMENT The Atoll Research Bulletin is issued by the Smithsonian Institution to provide an outlet for information on the biota of tropical islands and reefs and on the environment that supports the biota. The Bulletin is supported by the National Museum of Natural History and is produced by the Smithsonian Press. This issue is partly financed and distributed with funds from Atoll Research Bulletin readers and authors. The Bulletin was founded in 1951 and the first 117 numbers were issued by the Pacific Science Board, National Academy of Sciences, with financial support from the Office of Naval Research. Its pages were devoted largely to reports resulting from the Pacific Science Board's Coral Atoll Program. All statements made in papers published in the Atoll Research Bulletin are the sole responsibility of the authors and do not necessarily represent the views of the Smithsonian nor of the editors of the Bulletin. Articles submitted for publication in the Atoll Research Bulletin should be original papers in a format similar to that found in recent issues of the Bulletin. First drafts of manuscripts should be typewritten double spaced and can be sent to any of the editors. After the manuscript has been reviewed and accepted, the author will be provided with a page format with which to prepare a single-spaced camera-ready copy of the manuscript. COORDINATING EDITOR Ian G. Macintyre National Museum of Natural History MRC-125 ASSISTANTS Smithsonian Institution Kasandra D. Brockington Washington, D.C. 20560 William T. Boykins, Jr. Theodore E. Gram Jonathan G. Wingerath EDITORIAL BOARD Stephen D. Cairns (MRC-163) National Museum of Natural History Brian F. Kensley (MRC-163) (Insert appropriate MRC code) Mark M. Littler (MRC-166) Smithsonian Institution Wayne N. Mathis (MRC-169) Washington, D.C. 20560 Victor G. Springer (MRC-159) Joshua I. Tracey, Jr. (MRC-137) Warren L. Wagner (MRC-166) Roger B. Clapp National Museum of Natural History National Biological Survey, MRC-111 Smithsonian Institution Washington, D.C. 20560 David R. Stoddart Department of Geography 501 Earth Sciences Building University of California Berkeley, CA 94720 Bernard M. Salvat Ecole Pratique des Hautes Etudes Labo. Biologie Marine et Malacologie Université de Perpignan 66025 Perpignan Cedex, France PUBLICATIONS MANAGER A. Alan Burchell Smithsonian Institution Press ATOLL RESEARCH BULLETIN NO. 443 THE EVOLUTION OF A HOLOCENE FRINGING REEF AND ISLAND: REEFAL ENVIRONMENTAL SEQUENCE AND SEA LEVEL CHANGE IN TONAKI ISLAND, THE CENTRAL RYUKYUS BY H. KAN, N. HORI, T. KAWANA, T. KAIGARA, AND K. ICHIKAWA ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. OCTOBER 1997 35°N KYUSHU f Northern Limit of Yr _. Coral Reef Formation + 30°N | BASTERN CHINA SEA Ke # {\ Tonaki Is. Fig.2A / ° SS ’ 25°N 130°E 135°E Figure 1. The Ryukyu Islands, an island arc at high latitude for reef growth. THE EVOLUTION OF A HOLOCENE FRINGING REEF AND ISLAND: REEFAL ENVIRONMENTAL SEQUENCE AND SEA LEVEL CHANGE IN TONAKI ISLAND, THE CENTRAL RYUKYUS BY H. KAN!, N. Hori2, T. KAWANA3, T. KAIGARA4 and K. ICHIKAWA> ABSTRACT Within the Indo-Pacific region Holocene reef development over the last 6000 yBP has occurred during a near stable sea level period. In particular, development of reef flat and related features have been associated with the stillstand. This is illustrated by the Holocene evolution of a reef in the Japanese Ryukyu Islands. A continuous reef structure, 8 m in thickness and 630 m in length, was observed from a fresh excavation in a modern fringing reef in the western part of Tonaki Island of the central Ryukyu Islands. Documented by 34 radiocarbon ages, the reef first reached a relative sea level of ca. 1 m above the present level at about 5200 yBP by growth of branching Acropora thickets, and by accumulation of angular clasts of tabular Acropora at the landward side. Storm features occur within all the observed reef structure and suggest that the reef has been continuously affected by high-energy events for at least 5500 yBP. The topographic outline of the modern fringing reef was formed at an early stage of sea level stillstands affected by wind, climate, and substrate topography. Delayed closure of the ‘Holocene high energy window’ resulted in vigorous reef growth on the landward side. The reef flat accreted seaward about 400 m during the last 4500 yBP with the development of spurs and grooves. The growth environment shifted from a sheltered to a wave-affected condition during the seaward accretion of the reef flat. Reduced colony sizes of tabular Acropora and decreased upward reef growth rates also occurred at about this time. With the seaward accretion, the shoreward grooves became isolated and infilled by rounded clasts. Radiocarbon age from an early archeological feature (Touma and Oshiro 1979) indicates that the formation of a Holocene tombolo is closely tied to sea level fall around 3500 yBP. This sea level fall and sufficient development of wave resistant structures at the reef edge, provided shore protection and contributed to tombolo stability. ! Department of Physical Geography, Faculty of Education, Okayama University, Okayama 700 Japan. 2 Department of Geography, Faculty of Science, Tokyo Metropolitan University, Minami-Osawa, Hachioji 192-03 Japan. 3 Laboratory of Geography, College of Education, University of the Ryukyus, Okinawa 903-01 Japan. 4 Department of Geography, Kansai Gaikokugo University, Makikata, Osaka 573 Japan. 5 Department of Geography, Komazawa University, Setagaya 154 Japan. Manuscript received 2 September 1997; revised 24 September 1997 INTRODUCTION As coral reefs grow they can alter their own environment. In mid-Holocene times, reefs that were still catching up with sea level had minimal wave baffling effects. Neumann (1972) named this the ‘Holocene high energy window’. Subsequent reef growth produces a protective crest which in turn results in the development of the back- reef area. On fringing reefs, in particular, this may result in a shallow lagoon or moat with ponding of terrestrial run-off which may limit coral growth (Ginsburg and Shinn 1964; Schlager 1981; Neumann and Macintyre 1985; Acevedo et al. 1989). Sea level stillstands of the order of millennia can produce substantial fringing-reef flats (Buddemeier and Hopley 1988). In the Indo-Pacific region where a Holocene stillstand has occurred for approximately the last 6000 years, understanding of reef growth in response to both early Holocene sea level rise and late Holocene stillstand is needed to fully understand reef evolution. This study reports on the evolution of the modern fringing reef and sea level change in Tonaki Island, the central Ryukyus (Fig. 1), as deduced from a continuous reef section in a harbor excavation across western Tonaki Reef. Supporting evidence also comes from coastal landforms including a fossil reef surface overlain and protected by beach conglomerate, marine notches, and a tombolo (Fig. 2). Tonaki Island is a high island with two small hills consisting of late Paleozoic and Cenozoic formations (Konishi 1964) joined by a tombolo (Fig. 2B). The tombolo is 330 to 740 m wide and 330 to 1000 m long. Some of the features of the tombolo developed on Tonaki Island are similar to the cay islands of the Great Barrier Reef (Gourlay 1988), Solomon Islands and British Honduras (Stoddart 1969). Hopley (1968; 1971; 1975) described development of island spits on continental high islands of North Queensland and concluded that the many similarities between the fringing reefs and associated deposits of the high islands and the low wooded islands (complex reef islands) implied a comparable Holocene history (Hopley 1982, p371). Reef structure and radiocarbon dates observed from the western Tonaki trench and coastal features has enabled us to discuss the time-series relationship between reef environmental change, sea level change, and subaerial deposit formation by reference to Figure 2. (A) Location; (B) general topography of Tonaki Island. The contour lines are drawn at 50 m intervals for land topography. Numericals in B indicates sampling sites of dated corals and beach conglomerate shown in Table 2. Location of observed reef section is indicated in B and C. (C) Modern reef topography of the northwestern Tonaki Reef. Bold arrow shows submarine ridge extending to the northwestern small rocky island (Irisuna Island). Reef zonation: a reef slope (J furrowed platform, 2 with spurs and grooves), 6 spur and groove zone of reef edge, c reef crest, d inner part of reef crest (some area recognized as a rubble flat), e shallow lagoon (3 coral alignment), f inner reef flat, g beach (4 beach rock), h land. WONARS ISLAND i archeological ruin (Higashl shell heap) c RS, observed tombolo * reef section (Fig.3) : oy {3 .¢ Zamami soNL 1B. Vie. a te Me A om Hy Yi aoa) | ) ON WGE Oe 3 4 j y Pf 2) 0 As 3B Z NGSG8 Bae, .’ + g& observed reef section MW gs 9 4 these observations and radiocarbon dates of a fossil reef surface and the earliest archeological ruin on the tombolo of Tonaki Island as described by Touma and Oshiro (1979). This relationship between the reef and associated Holocene landforms indicates how reefs respond to external and internal environmental change. MATERIALS AND METHODS In the Ryukyu Islands, many trenches have been excavated to construct harbors and associated ship channels across modern fringing reefs. Such excavations provide the opportunity to observe the continuous structure of modern reefs. Reef structure directly observed from trench walls reveals the exact location of sedimentary facies, shape and size of coral colonies and coral clasts in contrast to observations made from reef cores (Kan and Hori 1991). In Tonaki Island, a harbor excavation reaching 8 m in thickness and 630 m in length across the western Tonaki Reef, provides a continuous reef section (Fig. 2). The southern wall of the excavation provided a fresh reef section because the survey was carried out immediately after widening of the trench in 1989. The reef profile (Fig. 3) was surveyed using measuring tapes, a 5-meter pole and depth meter to define micro-topographic features and to provide the basis for later descriptions of the section. Descriptions of the reef structure were carried out from the reef surface to the foot of the excavation at approximately 5 to 10 m intervals along the entire length of the section. The survey was carried out at closely spaced horizontal intervals especially at the boundaries of the sedimentary facies. Fossil corals were collected from the excavation by using a hammer and chisel. Twenty-five radiocarbon ages were obtained for corals in this reef section (Table 1). These dates were established by methanol liquid scintillation counting, at the Dept. of Geography, Hiroshima University. Laboratory procedures were based on Fujiwara and Nakata (1984). The geomorphological map of the northwestern Tonaki Reef (Fig. 2C) was prepared to document the zonal and the micro-topographic features. The reef topography is based on the interpretation of field observations in 1989 and the 1977 color aerial photographs scaled to 1:10,000. Coastal landforms such as fossil reef surface overlain by beach conglomerate, and marine notches were investigated to document mid- to late-Holocene sea level indicators in 1991 and 1996 (Fig. 2). The altitude data were revised with reference to tide tables. Tidal values in Tonaki Island are corrected by reference to those in Zamami Island, near Tonaki (Fig. 2). Mean high (low) water level in Tonaki is about 0.6 m above (below) mean sea level (MSL) which is 1.16 m above the tidal datum. Tidal range is approximately 2.0 m at spring tide. Nine radiocarbon ages were obtained from in situ corals in the modern reef surface and in the fossil reef surface overlain by the beach conglomerate (Table 2). These ages were dated by Prof. emeritus K. Kigoshi of Gakushuin University. X-ray diffraction was used to confirm the absence of calcite in all dated samples. Age calculations are based on the Libby half-life of 5568 + 30 years. Errors are indicated at the range of + 1 o. The dates have not been corrected for isotopic fluctuations or 5 environmental factors because we have no data on the ocean !4C reservoir effect for the Ryukyu Islands. RESULTS Topography of western Tonaki Reef On Tonaki Reef, a reef crest has developed in the north where topographic zonation is clearly defined, and flat topography with poor zonation occurs on the western reef (Fig. 2C). On the northern part of the reef, spurs and grooves on the reef edge and furrows on the submerged platform off the reef edge, at a depth between 10 to 15 m align with the prevailing northerly wind of winter seasons. These features are similar to the northern reef of Kume Island described by Takahashi and Koba (1977) and Kan (1990), which lies 30 km west of Tonaki Island. The distribution pattern of reef slope coral ridges changes abruptly on the northwestern reef slope where a submarine ridge extends westward (bold arrow in Fig. 2C). The long axes of the coral ridges and honeycombed furrows suggest that a southwesterly current dominates on the western reef slope. The observed reef section bisects a large number of spurs and grooves (Fig. 3), because the orientation of spurs and grooves is oblique to the reef edge on the western reef (Fig. 2C). The dominant coral assemblage of the reef edge is tabular or encrusting Acropora. On the northern reef, a coral rubble flat of tabular Acropora clasts has developed behind the reef crest. In the shallow lagoon, branching Porites dominate in the northern part. However, the lagoon tends to shallow towards the south where sea grass beds, with branching Montipora and Pavona cactus, are found. Internal structure beneath the reef flat Four major bio-lithofacies are identified for the upper 8 m of reef structure in the western Tonaki reef flat. Three distinct zonal structures were observed: from seaward, tabular Acropora framework facies; tabular Acropora transported rubble facies; tabular Acropora reworked facies; and branching Acropora facies. These facies are arranged vertically in the reef structure (Fig. 3). The other lithofacies described below are subordinate. 1) Tabular Acropora framework facies: Jn situ growth and accretion of tabular or plate Acropora dominates the outer half of the reef structure. This facies constitutes a large part of the convex topography of spurs or paleo-spurs (Fig. 3) and abut thick beds of tabular Acropora transported rubble facies described below (Fig. 4B). While large size in situ tabular Acropora colonies, around 2 m in horizontal dimension, are dominant on the inward end of this facies zone (Fig. 4A: 340 m point in Fig. 3), colony size decreases to 20 to 50 cm at the outer edge. Coincidentally, the upward reef growth rate gradually decreases seaward (Table 1) as follows: 8.2 m/ky (between TN-12 and 13) for the earlier spur dated around 4500 yBP; 4.1 m/ky around 4000 yBP (between TN-6 and 7); 3.6 6 m/ky (between TN-4 and 5); 3.2 m/ky (between TN-2 and 3); 1.2 m/ky (between TN-1 and present). 2) Tabular Acropora reworked facies: The unconsolidated platy clasts of tabular Acropora have accumulated (Fig. 4C) to at least 6m thickness and formed a sedimentary zone behind the tabular Acropora framework facies (Fig. 3). These Acropora clasts (Fig. 4D) are relatively well preserved and essentially overturned in situ colonies. The radiocarbon ages between TN-20 and TN-21 are reversed, despite 2.6 m difference in their depths (Table 1). 3) Tabular Acropora transported rubble facies : In the outer half of the reef structure, rounded coral clasts (Fig. 4E) have densely infilled the in situ tabular Acropora facies (Fig. 3). Some clasts are coated by calcareous algae (in the form of rhodoliths; Fig. 4F), which suggests that they had been tumbled. These are similar to those that have accumulated in the present-day grooves of the reef edge. Radiocarbon ages of rubble lag approximately 400 to 900 years behind neighboring in situ tabular Acropora facies (between TN-9, 12 and 10; Fig. 3). 4) Branching Acropora framework facies: Thickets of in situ branching Acropora (Fig. 4G) form a framework zone which exceeds 100 m in width and is more than 6 m thick (Fig. 3). These colonies were relatively intact (Fig. 4H). However, calcareous algae which covered the surface of branches makes species identification difficult. Radiocarbon ages (Table 1) show this coral thicket had been growing for over 400 years. The observed upward reef growth rates were 6.3 m/ky (between TN-14 and 15) for this thicket. Aside from the fine sediment that covers the excavation (Fig. 4G), little sediment was observed infilling the branching framework. (TN-2) (TN-4) (TN-6) (TN-9) (TN-10) — (TIN-12) 880-55 2220+60 3900+65 3890 +70 3510+65 4390+70 —=— Seaward Om MSL ---- tidal datum ----------- 3 (TN-1) -5 2340.60. | 55 pS IL a? ® || eth? yi : sai Stes 3500-60 11) 4960-65 202060 3390+65 a7) ON) (TN-13) -10 (TN-3) (TN-5) 100 200 300 = tabular Acropora 2 ©] tabular Acropora 19 framework facies transported rubble facies tabular Acropora tabular Acropora 0 [= | reworked facies reworked framework facies Figure 3. Reef structure and radiocarbon ages of fossil corals in the western Tonaki Reef. Lettered squares indicate positions of photographic sites presented in Fig. 4. 7 5) Tabular Acropora reworked framework facies: Tabular corals and calcareous algae constitute blocks | to 5 m in diameter. This bio-lithofacies is similar to the tabular Acropora framework facies of the reef margin. However, directions of individual coral growth and of accumulated coral colonies are different from the framework facies. 6) Mixed coral framework facies: This facies constitutes corals, calcareous algae and skeletal grains. No dominant coral genus was observed in this facies. Beside these bio-lithofacies, the following in situ coral colonies larger than 2 m in diameter are identified in the growth fabric: foliaceous Heliopora, stubby branching Acropora, and hemispherical Lobophyllia. No terrigenous facies or Pleistocene limestone was observed within the section. Late-Holocene sea level change On Tonaki Island, part of the Holocene reef flat is surmounted by undercut blocks (Konishi 1964). It has been suggested that the reef flat originally developed 80 cm higher than the present sometime in the late Holocene (Konishi e¢ al. 1974). Beach conglomerate which overlies the landward end of the reef has also resulted in protection of the reef surface against erosion (Fig. 5). These features, together with marine notches cut into the Paleozoic limestone cliffs are considered to be sea level indicators (Kawana 1996). Nine radiocarbon ages for corals obtained from the modern and fossil reef surface (Table 2, Fig. 5) show that the sea level achieved a maximum ca. 0.9 to 1.3 m higher than present ca. 5200 yBP. This mid-Holocene maximum level extended until around 3650 yBP, with a relatively sharp fall taking place at that time (Fig. 6). (TN-14) (TN-16) (TN-17) (TN-20) (TN-22) (TN-25) 4560+70 4600+65 4580+70 5690+75 5190+75 5180+70 -70 Om } ay my wa Ae GS Se x a , mn BA ool Aue we Ne v2 ¥ Pf er ee ee eee e 5 aN = Seek OU ae Qe A a ONT an ae ON Ae ones © ye za 5510+75 3 75 5290+70 Se EMOE) (TN-21) (TN-23) 5660 1 (TN-18) (TN-18) (TN-19) a -10 400 500 600m ao mixed coral in situ stubby r~\ invisible portion framework facies branching Acropora a (wall covered by talus) branching Acropora | He | in situ foliaceous framework facies Heliopora in situ hemispherical Lobophyllia bottom of the excavated wall dated coral Table 1 Radiocarbon ages from Tonaki Reef excavation. Sample Site ae Material Labo. ee ee oy ae m) ak (yB.P.+1o) Growth : 1 . : ao Genus Life Form T( 1/2)=5568yrs het TN-1 18 6.5 Platygyra hemispherical t HR-561 2340 + 60 - 55 AZ HF TN-2 70 2.7 Acropora tabular HR-562 880 + 55 32 TN-3 70 6.4 Acropora tabular t HR-563 2020 + 60 TN-4 153 2.1. Acropora _ tabular ft HR-564 2220 + 60 3.6 TN-5 153 6.3. Acropora tabular + HR-565 3390 + 65 TN-6 214 3.4 Acropora tabular ft HR-539 3900 + 65 4.1 TN-7 214 6.1 Acropora tabular HR-540 4560 + 65 TN-8 220 4.6 Acropora _ tabular + HR-538 3500 + 65 - 60 TN-9 280 2.1. Acropora tabular + HR-560 3890 + 70 - 65 TN-10 295 2.1. Acropora tabular %* HR-534 3510 + 65 TN-11 295 4.7 Acropora tabular * HR-535 4210+ 65 TN-12 305 1.8 Acropora _ tabular + HR-536 4390 + 70 8.2 TN-13 305 6.5 Acropora _ tabular ft HR-537 4960 + 70 - 65 TN-14 3)5)// 2.4 Acropora _ tabular t HR-531 4560 + 70 6.3 TN-15 3S 7.0 Acropora tabular + HR-532 5290-70 TN-16 365 3.7. Acropora tabular HR-533 4600 + 65 TN-17 395 2.2 Heliopora foliaceous 7 HR-559 4580 + 70 TN-18 405 5.4. Acropora tabular HR-557 5440 + 75 TN-19 434 5.5 Acropora tabular HR-558 4970 + 70 TN-20 529 3.1 Acropora tabular HR-551 5690 + 75 TN-21 529 5.7. Acropora tabular HR-552 ayo ae ve) TN-22 555 2.1 Acropora ramose ft HR-553 5190 + 75 - 70 8.3 TN-23 395 5.4 Acropora ramose HR-554 5590 + 75 TN-24 606 6.3. Acropora ramose f HT-555 5660 + 75 - 70 TN-25 625 1.6 Acropora ramose ft HR-556 5180 + 70 * Site shows the sampling location corresponding to the horizontal scale of the reef section (Fig. 2). + in situ coral %¢ rounded clast accumulated in groove +t Growth rate is calculated on the basis of the TN-1 sample and the top of the spur (present). The present is assumed, as the spur has not reached sea-level. Figure 4. Holocene reef structures and components observed from the excavated wall of the western Tonaki Reef. Sites of the photographs are shown in Fig. 3. (A) Jn situ tabular Acropora facies, (B) paleo-groove (b) infilled by rounded coral clasts. Paleo- spur (a) composed of in situ tabular Acropora. (C) angular clasts of tabular Acropora reworked facies. (D) close-up of angular clast of tabular Acropora. (E) rounded coral clasts of tabular Acropora transported rubble facies. (F) a section of a rounded coral clast showing encrustations by calcareous algae. (G) branching Acropora framework facies. (H) colony of in situ branching Acropora. 10 Table 2 Radiocarbon ages of in situ corals in coastal deposits in Tonaki Island (see also Fig. 5). Loc. No. * OM YADA HW KR Wb & Radiocarbon Age (yB.P. + 1o) T(1/2)=5568yrs 5150 + 100 4410+ 80 4790 + 140 4430+ 110 4980+ 90 3640 + 190 4130+ 80 4820 + 130 4890 + 100 Material (Coral Genus) Porites Goniastrea Goniastrea Porites Goniastrea Porites Porites Goniastrea Porites Labo. No. GakK-15825 GaK-15826 GaK-15827 Gak-15828 GaK-15829 GaKkK-15830 Gak-15831 GaK-15832 GaK-15835 Explanation for Sampling Site Elevation of specimen (m) above MSL reef flat - 0.6 reef overlain by beach conglomerate 0.1 reef in front of beach conglomerate 0.1 reef overlain by beach conglomerate 0.2 do. 0.1 do. 0.77 do. 0.4 inner edge of the reef flat - 0.4 reef in front of beach conglomerate 0.2 * Location numbers are in Fig. 2B. + The higher elevations of the specimens at Loc. 6 and Loc.7 are probably due to strong wave- affected coast. ca. 0.9~1.3m above mean sea level (marine notch) Beach conglomerate MHWL 4790 + 140 4890 + 100 MSL 4820 + 130 5150 + 100 3640 + 190 4130 + 80 4410 + 80 4430 + 110 4890 + 90 Holocene coral reef Figure 5. Schematic profile of Holocene reef, beach conglomerate and marine notch at the landward end of Tonaki Reef. Radiocarbon ages of corals indicate years BP (see also Table 2). MHWL: mean high water level, MLWL: mean low water level. 1] Reef growth and island formation The western Tonaki Reef first reached the sea level at 5200 yBP by growth of branching Acropora on the landward side (Fig. 3). The accumulation of the reworked facies of angular clasts of tabular Acropora was around 5200 yBP. Around 4500 yBP, some spurs consisting of tabular Acropora framework reached sea level at the outer edge of the facies formed of tabular Acropora reworked (Fig. 3, TN-14). The transported tabular Acropora rubble facies gradually became thicker toward the inner reef flat (Fig. 3) where paleo-grooves have been entirely filled by rounded clasts (Fig. 4B). The reef flat of the western Tonaki Reef has accreted seaward about 400 m during the last 4500 yBP by the development of spurs and grooves. With the accretion of the reef margin, the landward grooves, which have became isolated from the sea, have been filled by rounded clasts. The geomorphological evidence is supported by the history of human settlement in this area. Touma and Oshiro (1979) excavated the earliest archeological ruin located on the narrow part of the tombolo of Tonaki Island (Fig. 2B). Trench excavations show that bioclastic sand accumulated up to an elevation of 3.5 to 3.8 m. Above this sand, Initial formation of spur and groove system om Development Reef catch up of tombolo ; with sea level —+— ——_— jn situ tabular Acropora ——————_____LLLL_> SSS eee tabular tabular Acropora -10 — Acropora transported -e in situ coral reworked rubble facies ~ tabular Acropora reworked clast facies -~- tabular Acropora transported rubble oa eae TIS ERT ESS ae Sir a IEC aie CR Initial formation Secondary of reef flat progradation of reef flat Figure 6. Sea level and reef growth in Tonaki Reef. The sea level curve (solid curve) until 5200 yBP is based on Kume Island, the neighboring island of Tonaki, when the 3 meters seismic uplift at around 2000 yBP (Koba et al. 1982) is subtracted from the curve of Kan ef al. (1991). The curve after 5200 yBP is based on the coastal landforms and radiocarbon ages in Tonaki Island (see text). The dashed lines show the vertical reef growth at each site in Tonaki Reef (see also Fig. 3 and Table 1). A horizontal bar indicates the range of error in radiocarbon ages. A vertical bar indicates depth between the lowest low water level and the mean sea level for Tonaki Reef (0.9 m) to represent past sea level. The vertical bar is only for ages which are reliable sea level indicators. WZ two layers of brown humic sand were observed with a thickness of between 0.3 to 0.7 m below the present-day surface soil. The lower bed, just above the bioclastic sand, contains earthenware and shell middens. The radiocarbon age of shell material was 3510+90 yBP (laboratory code: N-3080). This age coincides with the chronological determination of excavated earthenware (Touma and Oshiro 1979). DISCUSSION Storm features in the reef development The Ryukyu Islands are located in the hurricane belt. Catastrophic storm events result in obvious degradation of coral colonies (see review of Rogers 1993) and produce large amount of coral clasts (e.g. MacNeil 1954; Hernandez-Avila et al. 1977; Scoffin 1993) during cyclic growth and destruction processes (Done 1992). The fragmentation of coral colonies appears to be a widespread and extremely important method of reproduction and distribution (Highsmith 1982). However, overturning of the colony is critical to survival (Chamberlain and Graus 1975). The morphology of tabular Acropora colonies which widen toward the top, when broken, produces the overturned accumulations of the tabular Acropora reworked facies. Storms on modern reefs can result in deposition in the back reef environment (c.f., MacNeil 1954; Newell and Bloom 1970) and mixing debris of varying ages (Johnson and Risk 1987). The observed age reversal between TN-20 and TN-21 in the tabular Acropora reworked facies is of similar magnitude to those obtained by Marshall and Davies (1982) from a drill core which penetrated into reef flat rubble facies in One Tree Reef on the Great Barrier Reef. Catastrophic redistribution and abrasion by sediment have been observed during storm conditions (Kobluk and Lysenko 1992) especially along reef-edge grooves (Kan 1995). The unstabilized rubble is subjected to tumbling (Blanchon and Jones 1995) and infills cavities (Newell 1956; Edmunds and Witman 1991). The transported tabular Acropora rubble facies accumulated between the tabular Acropora framework facies (Fig. 4B) where ‘room and pillar structures’ (Tracey et al. 1948) had been formed. Storr (1964) showed that water flow over the tops of reefs is erratic and the rate of flow is generally low, whereas passageways (e.g., grooves, reef tunnels) permit high flow rates. The initial accumulation site of fragmentated colonies (i.e., back reef or reef edge grooves) may cause the differentiation between angular or rounded rubble. The observed reef structure shows that the accumulated coral clasts form rounded rubble (transported rubble facies) after the development of a spur and groove system (Fig. 6). Storm events can also destroy reef edge spurs (Stoddart 1962) and throw them onto the reef flat (Ladd 1961; Newell and Bloom 1970; Bourrouilth-Le Jan and Talandier 1985). The buried blocks of tabular Acropora reworked framework facies, along with the accumulation of a large amount of Acropora clasts and the tightly interlaced branching colonies (as in Fig. 4H) which may also be formed during the regenerative growth process of fragments (Kawaguti 1937; Gilmore and Hall 1976), suggest that the reef has been continuously affected by high-energy events. 13 Windward-leeward contrast in the reef development In the fringing reefs of the Ryukyu Islands, several studies have demonstrated that the original growth axis, where the reef first reached sea level, forms the present reef crest (Takahashi ef al. 1988; Kan et al. 1991; Kan and Hori 1993; Yonekura et al. 1994). Results of drilling in a reef of northwestern Kume Island, located 30 km west of Tonaki Island, shows the northern reef reached sea level earlier because of shallow substrate and exposure to the prevailing northerly wind of winter seasons (Kan ef al. 1991). A similar pattern of formative time lag was described by Hopley and Barnes (1985) for a fringing reef in the Great Barrier Reef. This pattern of growth helps to understand the planimetric development of the northwestern Tonaki Reef which has a similar geographical setting. A submarine ridge extending on the northwestern reef slope of the Tonaki Reef (bold arrow in Fig. 2C) may have contributed to the development of the reef crest of the northern reef because reef development is closely tied to antecedent slope break (Hubbard 1988; Kan ef al. 1995). The northern Tonaki Reef may well have initially reached sea level and acted as a breakwater against the prevailing winter northerly winds (Fig. 7A). This allowed zonal growth of branching Acropora thickets which is usually found in sheltered environments (Geister 1977; Pichon 1978; Done 1983). However, branching coral assemblages in shallow lagoons change in response to burying by sandy sediment (Nakai 1982). Siltation and suspended sediments also reduce coral growth (e.g., Aller and Dodge 1974; Dodge et al. 1974; Loya 1976). Mayer (1918) examined 4. hebes which is a synonym of A. aspera (Veron and Wallace 1984) and same or similar species to the branching Acropora of the western Tonaki section. It was shown to be sensitive to the smothering effects of silt. The vigorous growth of branching Acropora for over 400 years, with indications of little sediment influence, may be explained by the tidal current which may have been flowing between the two islands at about 5500 to 5200 yBP (arrows in Fig. 7A) before the tombolo joined them together. The northern reef may also have provided the large amount of angular clasts of tabular Acropora to the reworked facies that accumulated around 5200 yBP, because the tabular Acropora assemblage had not formed in the western reef but is distributed abundantly down to a depth of 5 m at the reef edge and decreases in deeper water in the Ryukyu Islands (Takahashi et al. 1985). At the early stage of reef flat formation, development of leeward reefs is strongly affected by the windward reef formation. Styles of reef accretion and formation of reef zonation Framework accretion with higher accumulation rates is dominated by branching corals with a high proportion of voids (e.g., Davies and Hopley 1983; Davies et al. 1985; Hopley and Kinsey 1988) and is especially associated with monospecific coral thickets (Highsmith 1982). Detrital sedimentation by storm events also results in high reef accretion rates (Davies and Hopley 1983). In the western Tonaki Reef, the early 14 stages of reef were developed by these two bio-lithofacies within a short period around 5200 yBP. They may have also contributed to the sediments of the broad inner reef flat at the western part of Tonaki Reef quickly infilling any shallow lagoons. The duration of the ‘Holocene high energy window’ has some regional variation (Hopley 1984). The development of spurs consisting of tabular Acropora framework around 4500 yBP (Fig. 3, TN-14) intimates that the ‘high energy window’ has been progressively closed since 4500 yBP. Subsequently, the area available for active carbonate production has been reduced to a zone at the reef edge (Stoddart et al. 1978) as reported in other present-day reefs (e.g., Gladfelter et al. 1978; Kinsey 1981). Since the size of wave-swept organisms depends on wave exposure (Denny ef al. 1985), the smaller colony sizes of in situ tabular Acropora on the outer reef flat suggest that the growth setting has changed to a wave-affected condition with the outward migration of the reef flat. The decline in vertical growth rates at the outward edge of the reef flat (Table 1, Fig. 6) has accompanied this change. Contrary to reefs where the crest kept pace with sea level, the delayed closure of the ‘Holocene high energy window’ resulted in vigorous reef growth toward the island. Subsequently, however, the area of active reef growth has shifted progressively seaward with the development of reef edge spurs. This gradual process has contributed to the build up of flat topography with poor zonation. Timing of island formation and sea level fall The time-series relationship between reef island formation and reef development has been previously described (e.g., Stoddart 1969; Stoddart et al. 1978; Hopley 1982; Woodroffe 1992). The Great Barrier Reef cays formed after 6000 yBP, coincident with reef flat formation, but were essentially complete in shape and size by 3000 yBP (Stoddart et al 1978). On Tonaki Island, the vigorous reef growth around 5000 yBP appears to have provided the shallow substrate to establish the tombolo between the two islands and provided enough land for human inhabitation by about 3500 yBP (Touma and Oshiro 1979; Fig. 7B). The sea level fall sometime after 3650 yBP in Tonaki Island is quite similar to those in Kosrae Island in the eastern Carolines of Micronesia where a | m fall has taken place after 3700 yBP (Kawana et al. 1995), and in the Great Barrier Reef where the mid- Holocene maximum level may have extended until 3700 yBP (Beaman ef al. 1994; Larcombe ef al. 1995). Evidence for sea level fall around 3700 yBP appears prevalent in the western Pacific. Stoddart and Steers (1977) suggested that reef islands were formed as a result of sea level fall (see also, Pirazzoli and Montaggioni 1986; Roy and Connell 1991). Schofield (1977a, 1977b) demonstrated how reef islands formed due to sea level fall and the supply of sediment from reefs. Sea level fall resulted in reduced wave force at the shore. The seaward accretion of the reef also contributed to wave attenuation because 15 Pie eet fst Seeeminestt w+ Gar 5000 F igure 7. Schematic diagrams of the formation of the Tonaki Reef. Arrows in Fig. 7A indicate the “Holocene high energy window’ at around 5200 yBP. 16 waves break farther from the shore. At 3500 yBP, the western Tonaki reef flat accreted to approximately half way across the present-day reef flat (Fig. 7B). The sea level fall and reef maturation with development of wave resistant structures at the reef edge, provided shore protection and contributed to tombolo stability. CONCLUSION Sea level achieved a maximum ca. 0.9 to 1.3 m higher than present ca. 5200 yBP in Tonaki Island, the central Ryukyus. This Holocene maximum level extended until around 3650 yBP, with a relatively sharp fall taking place at that time. Early development of the reef in Holocene times was influenced by substrate topography that produced an initial zonation contrasting the outer windward margin, and the inner sheltered reef. Spur and groove systems were initiated with major reef accretion occurring by growth of the reef edge spurs. As inner grooves have become isolated from the sea, they have been infilled during storms by rounded coral clasts. The deposition of subaerial deposits of the tombolo of Tonaki Island is linked both to the accretion of the reef flat seaward, and to a fall in sea level about 3650 yBP. ACKNOWLEDGEMENTS The authors are indebted to: Dr. D. Hopley, Prof. L.F. Montaggioni, Dr. M.K. Gagan and Dr. J. Kleypas for their invaluable comments on this manuscript; Dr. I.G. Macintyre and Dr. J.I. Tracey for their helpful review; Prof. K. Fujiwara and Prof. T. Nakata for arrangement to use radiocarbon dating system at Hiroshima University; Prof. emeritus K. Kigoshi for radiocarbon dating at Gakushuin University; Mr. Y. Nakashima, Mr. K. Takemasa, Ms. Y. Oka and Mr. T. Toubara for their field assistance; Ms. N. Nakamura and Mr. K. Yoshihama provide us information about the harbor construction. REFERENCES ACEVEDO, R., MORELOCK, J. AND OLIVIERI, R.A. 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Ser., 6, 485pp. WOODROFFE, C.D. (1992) Morphology and evolution of reef islands in the Maldives. Proc. 7th Int. Coral Reef Symp., 2, 1217-1226. YONEKURA, N., KAYANNE, H., MATSUMOTO, E., ISHII, T., MATSUSHIMA, Y., HORI, N. AND NAKAI, T. (1994) Geomorphic development of modern fringing reefs of Yoron Island, Ryukyu Arc, Japan. The Quat. Res. (Daiyonki Kenkyu), 33, 67-79. ATOLL RESEARCH BULLETIN NO. 444 CHECKLIST OF THE SHOREFISHES OF OUVEA ATOLL, NEW CALEDONIA BY MICHEL KULBICKI AND JEFFREY T. WILLIAMS ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. OCTOBER 1997 ae: de ‘Astrolabe ? O yz Récif de la Gazelle : Beautemps-Beaupré G, x @plle des Pins lle Walpole Figure 1. Map of New Caledonia and the Loyalty Islands showing the location of Ouvéa. CHECKLIST OF THE SHOREFISHES OF OUVEA ATOLL, NEW CALEDONIA BY MUCHEL KULBICKI AND JEFFREY T. WILLIAMS | ABSTRACT The shorefishes of Ouvéa, an isolated atoll in the Loyalty Islands group of New Caledonia, had not been surveyed prior to 1990. An extensive survey was conducted by ORSTOM between 1991 and 1992 to obtain baseline information on the shorefishes. A total of 653 taxa among 72 families are now documented from this area. The most diverse families are the Labridae (69 species), Pomacentridae (58 species), Gobiidae (54 species), Serranidae (39 species), Chaetodontidae (31 species) and Apogonidae (28 species). The absence or very low diversity of some families (Clupeidae, Nemipteridae, Siganidae) or genera (Abudefduf, Neopomacentrus) is similar to findings for other isolated islands of the Coral Sea. Of the 653 species recorded from Ouvéa, 51 species have not been reported from New Caledonia, a large high island to the South. Only one endemic species, Luzonichthys williamsi, has been recognized among the shorefishes at Ouvéa. A number of Pacific Plate endemic species were recorded at Ouvéa, which is positioned on the Australasian Plate to the south of the edge of the Pacific Plate. Antennarius duescus, previously known from three specimens taken at the Hawaiian Islands, is recorded from a single specimen taken at Ouvéa. Another antitropical distribution pattern is exhibited by Dinematichthys riukiuensis, which is known to occur at Fiji, Ouvéa and Queensland in the South and from Okinawa. INTRODUCTION Our knowledge of shorefishes in the Southwest Pacific has increased significantly in the last two decades with the publication of a number of checklists. Shorefishes of the Great Barrier Reef have been reported from several regions: Russell (1983) and Lowe and Russell (1990) for the southern part, Paxton et al. (1978) for Lizard Island, Allen (1989) * ORSTOM, B.P. A5, Nouméa, New Caledonia ** National Museum of Natural History, Smithsonian Institution, Washington D.C. 20560 USA Manuscript received 4 January 1997; revised 7 July 1997 D for the Coral Sea, and Paxton et al. (1989: 1 volume, out of 3 announced, of an encyclopedia of the fishes of Australia, which covers shorefishes). The fish fauna of Lord Howe, Norfolk and Kermadec islands, the southern limit of shorefishes in the Southwest Pacific, was surveyed by Allen et al. (1976) and more recently by Francis (1993) and Francis and Randall (1993). Middleton Reef, a southern reef midway between Australia and New Caledonia, has been investigated by Hutchings (1988), and Kailola (1987 a,b; 1991) published a checklist of fishes from Papua New Guinea. Kailola’s list is being updated by Allen. For New Caledonia there are two checklists, one for the main island (Rivaton et al., 1989) and one for the Chesterfield archipelago (Kulbicki et al., 1994; LeBorgne et al., 1994), a group of islands midway between the Great Barrier Reef and New Caledonia. A number of books are now available on the shorefishes of the Southwest Pacific region (Fourmanoir and Laboute, 1976, for New Caledonia and Vanuatu; Randall et al., 1990, for the Coral Sea; Allen and Swainston, 1992, for Papua New Guinea; Allen and Swainston, 1988, for Western Australia). Despite these recent efforts, the fish fauna of many areas of the Southwest Pacific remains poorly known. One such area is the Loyalty Islands, which comprise five islands, Maré, Tiga, Lifou, Ouvéa and Beautemps-Beaupré. The first three are high islands with very sparse coral reef development. The latter two are atolls, Ouvéa (900 km?) being much larger than Beautemps-Beaupré (120 km?). These atolls are located on the edge of the Australasian plate near its boundary with the Pacific plate between New Caledonia and Vanuatu (figure 1), and are the only true atolls within a 1500 km radius. Only two scientific cruises conducted studies at Ouvéa prior to 1991. Allen, in June 1973, studied the Pomacentridae of Ouvéa. ORSTOM, a French scientific organization, organized a cruise to Ouvéa in November 1979, but issued no cruise report because a cyclone considerably limited their study. A few specimens were collected and sent to the Paris Museum and the Bishop Museum (Hawaii). We are unaware of any other samples taken from Ouvéa. Carpenter and Allen (1989) report that Lethrinus “sp.2” occurs at the Loyalty Islands, but they do not provide collection data for the material they examined. In 1991, ORSTOM was asked to evaluate the fish resources of Ouvéa Atoll. During this survey, data were also collected on the atoll’s physical characteristics (water masses, geomorphology, sedimentology), the plankton, and the benthos (Chevillon, 1994; Clavier et al., 1992; Clavier and Garrigue, 1993; Kulbicki et al., 1993a, 1993b, 1994; LeBouteiller et al., 1993). A summary of these works is also available (Kulbicki, 1995). The checklist of the shorefishes of Ouvéa Atoll presented here is compiled from data gathered during the 1973 and 1979 visits and the authors’ sampling program conducted during their 1991 survey. MATERIAL AND METHODS Fish were visually censused along transects and/or collected using rotenone and SCUBA. Some specimens were caught with handlines. Visual censuses took place on the lagoon floor down to 25m, and on the outer reefs surrounding the atoll. No censuses or 5) ») collections were performed on the outer slope of the barrier reef or on the eastern side of the main island in the atoll. The locations of the stations are plotted on the map in figure 2. Thirteen collecting stations utilized rotenone (figure 2), the amount of rotenone used at each station was between 2 and 6 liters of solution containing 8% active rotenone. 20°20’S iL OCEAN PACIFIQUE 20°25’S L 20°30°S L 20°35’S i 20°40°S L 20°45’S T —— 168°10’E 1664S’E 166°20’E 166°25E 166°30'E 166°35E 166°40°E Figure 2. Location of sampling sites (open circles = handline fishing; dark circles = dives; stars = rotenone). RESULTS AND DISCUSSION A checklist, totaling 653 taxa distributed among 72 families, is presented in Table 1. There are about 300 fewer species known from Ouvéa than from nearby New Caledonia, where more than 950 reef-fish species have been recorded (Rivaton et al., 1989). The number of taxa known from Ouvéa is closer to the 795 species recorded from the Chesterfield archipelago (Kulbicki et al., 1994), another isolated area at a similar latitude, than to the 425 species recorded from Rotuma (Zug et al., 1989), a small cluster of islands about 450 km north of the Fiji Islands. Because our sampling (only 13 rotenone stations) at Ouvéa is limited and we have not sampled the eastern side of the atoll, we believe the total number of shorefish species may actually be as high as 800-900 species. Most of the species in this list have been reported from nearby New Caledonia, however 51 taxa have not been recorded from New Caledonia. Serranocirrhitus latus has not been listed in the New Caledonian checklists, but was reported from New Caledonian reefs and the Loyalty Islands as “ Dactylanthias mcmichaeli” by Fourmanoir and Laboute (1976). Most of the new records are Anguilliformes, Scorpaenidae or Gobiidae, which are usually taken only in rotenone collections. There are specimens in the Ouvéa collections representing 33 taxa that could not be identified to species. Many of these are juveniles, but 15 of these taxa are currently known to be undescribed species. Most of the 15 undescribed species are known from other localities. Their descriptions, thus, will not greatly increase the number of endemic species known from the Loyalty Islands. The only endemic species described from Ouvéa 1s Luzonichthys williamsi (Serranidae, Anthiinae). One specimen of Antennarius duescus was taken at Ouvéa. This distinctive species 1s unique in having the opercular opening situated halfway between the base of the pectoral lobe and the origin of the anal fin. It was previously known from only three specimens from the Hawaiian Islands (Pietsch and Grobecker, 1987). Its presence in the Loyalty Islands is possibly indicative of an antitropical distribution pattern. Trimma unisquamis and Gymnothorax eurostus have a similar antitropical distribution pattern, with a northern population at Hawaii. Dinematichthys riukiuensis exhibits an antitropical distribution pattern, but has its northern population at Okinawa and its southern population at the Great Barrier Reef, Ouvéa, and Fiji. A number of Pacific plate endemic species (Springer, 1982) are recorded herein from Ouvéa. These records could be described as plate margin occurrences. Pacific plate endemics taken at Ouvéa include the following taxa: Minysynchiropus laddi, Myripristis amaenus, Schismorhynchus labialis, Brotula townsendi, Centropyge nigriocellus, and Centropyge loriculus. Two species, Alticus sertatus and Neoglyphidodon carlsoni, were previously thought to be Fijian endemics. The presence of Alticus sertatus at Ouvéa provides the first record of this species west of the Fiji Islands. An undescribed species of Alticus occurs at New Caledonia, but was not collected at Ouvéa. Likewise, A/ticus sertatus has not been collected from New Caledonia. The occurrence of Neoglyphidodon carlsoni at Ouvéa is the first record outside of Fijian waters. A comparison of the diversity of the major families within several areas of the southwestern Pacific is given in Table 2. An interesting analogy with the Chesterfield archipelago (LeBorgne et al., 1994) and Rotuma (Zug et al., 1989) is the low number or lack of Abudefduf spp., Neopomacentrus spp., Clupeidae and Siganidae (Table 2). A preliminary list of fishes from Osprey Reef, off the Great Barrier Reef, based on the fish collection at the Australian Museum (Leis, pers. communication) entirely lacks these genera and families. Similarly, Norfolk, Lord Howe and Kermadec islands also have few or no members of these taxa, but have five species of Abudefduf. A survey of Elizabeth 5 and Middleton Reefs (Hutchings, 1988), located further south in the Coral Sea, indicates the presence of 3 species of Abudefduf, one species of Siganidae, but no Neopomacentrus or Clupeidae. These genera and families are well represented elsewhere in the Coral Sea and New Caledonia (Table 2). At Ouvéa, the lack of suitable habitat is not a likely reason for the low number of these species, except possibly for the Clupeidae. Indeed, the habitats where these species are found in New Caledonia are apparently present at Ouvéa (Kulbicki et al., 1993a). The early life history traits (type of egg, length of larval life, shape of the larvae, size of the larvae at recruitment on reefs) of Abudefduf and Neopomacentrus (Table 3) do not differ significantly from those of other Pomacentridae that are present at Ouvéa, Rotuma, the Chesterfield Islands, or Osprey Reef. Victor (1991), in a review on settlement strategies and biogeography, noted that duration of the larval stage rarely accounted for the geographic range of a species. The short distance between Ouvéa and the main island of New Caledonia (60 km) and the direction of the major surface currents (Kulbicki et al., 1993a; Kulbicki, 1995) would not seem to present a major obstacle to the colonization of Ouvéa by species from New Caledonia (figure 3). Fig. 3. Major currents in the vicinity of Ouvea (from Kulbicki et al., 1993a). One possible reason for the lack of certain species at Ouvéa could be related to its being a low island and New Caledonia a high island. Causal factors leading to the absence of certain taxa at Ouvéa remain speculative. ACKNOWLEDGEMENTS We wish to thank the following persons who have helped in the collection or identification of specimens during the preparation of this checklist: G. Allen, G. Bargibant, B. Carlson, P. Dalzell, A.C. Gill, B. Hutchins, H. Larson, J. Leis, J.L. Menou, R. Mooi, G. Mou Tham, S. Poss, J.E. Randall, J. Rivaton, B. Sérét, D.G. Smith. W-F. Smith-Vaniz, P. Tirard, R. Winterbottom, and the crews of the RV ALIS, DAWA and DAR MAD. We thank V.G. Springer, A.C. Gill, and K.E. Carpenter for constructive comments on earlier versions of the manuscript. REFERENCES Allen, G.R. 1989. The Fishes of the Coral Sea. Tech. Report Australian National Parks and Wildlife Services. Canberra. 151 pp. Allen, G.R. 1991. Damselfishes of the World. Mergus Press, Melle, Germany. 272 pp. Allen, G.R., D.F. Hoese, J.R. Paxton, J.E. Randall, B.C. Russell, W.A. Starck, IJ, F.H. Talbot, and G.P. Whitley. 1976. Annotated checklist of the fishes of Lord Howe Island. Records of the Australian Museum 30 (15):365-454. Allen, G.R., and R. Swainston. 1988. The Marine Fishes of North-Western Australia. A Field Guide for Anglers and Divers. Western Australia Museum. 201 pp. Allen, G.R., and R. Swainston. 1992. Reef fishes of New Guinea. 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FAO Fisheries Synopsis No.125, Vol. 12:1-149, 8 color plates. Springer, V.G. 1982. Pacific plate biogeography, with special reference to shorefishes. Smithsonian Contributions to Zoology, number 367:1-182. Thresher, R.E., and E.B. Brothers. 1989. Evidence of intra- and inter- oceanic regional differences in the early life-history of reef- associated fishes. Marine Ecology Progress Series 57:187-205. Victor, B.C. 1991. Settlement strategies and biogeography of reef fishes. Pages 231-260 in P.F. Sale (ed.). The Ecology of Fishes on Coral Reefs. Academic Press, NY. Wellington, G.M., and Victor, B.C. 1989. Planktonic larval duration of one hundred species of Pacific and Atlantic damselfishes (Pomacentridae) Marine Biology 101(4):557-567. Whitehead, P.J.P. 1985. FAO Species catalogue, Vol. 7. Clupeoid fishes of the world, part 1, Chirocentridae, Clupeidae, Pristigasteridae. FAO Fisheries Synopsis Now Zoe Vola: l-303) 10 Whitehead, P.J.P., G.J. Nelson, and T. Wongratana. 1988. FAO Species catalogue. Vol. 7. Clupeoid fishes of the world, part 2, Engraulididae. FAO Fisheries Synopsis No.125, Vol. 7:305-575. Woodland, D.J. 1990. Revision of the fish family Siganidae with description of two new species and comments on distribution and biology. /ndo-Pacific Fishes 19:1-136. Zug, G.R., V.G. Springer, J.T. Williams, and G.D. Johnson. 1989. The vertebrates of Rotuma and surrounding waters. Afoll Research Bulletin No. 316: 1-25. Table 1. List of the shorefish taxa known from Ouvéa atoll. Taxa which were not previously known from New Caledonia are marked by * and are in bold typeface. Families are ordered according to Eschmeyer (1990).The letters used in the column "Method" mean the following: V: visual census _R: rotenone Scientific name GINGLYMOSTOMATIDAE Nebrius ferrugineus CARCHARHINIDAE Carcharhinus albimarginatus Carcharhinus amblyrhinchos Carcharhinus melanopterus Galeocerdo cuvier Triaenodon obesus DASY ATIDIDAE Dasyatis kuhlii MORINGUIDAE Moringua species CHLOPSIDAE Kaupichthys species *Kaupichthys atronasus Kaupichthys hyoproroides MURAENIDAE *Anarchias cantonensis Echidna polyzona *Echidna unicolor Gymnothorax buroensis Gymnothorax chilospilus Gymnothorax eurostus Gymnothorax fuscomaculatus Gymnothorax javanicus Gymnothorax margaritophorus *Gymnothorax marshallensis Gymnothorax melatremus Gymnothorax pindae Gymnothorax rueppelliae Gymnothorax thyrsoides Gymnothorax zonipectis Uropterygius fuscoguttatus *Uropterygius makatei OPHICHTHIDAE Leiuranus semicinctus *Muraenichthys gymnotus Muraenichthys sp. *Schismorhynchus labialis *Schultzidia johnstonensis CONGRIDAE Conger cinereus Heteroconger hassi CLUPEIDAE L: line fishing A: Allen (1975 and pers. comm.) Author and date (Lesson, 1830) (Ruppell, 1837) (Bleeker, 1856) (Quoy & Gaimard, 1824) (Peron & LeSueur, | 822) (Riippell, 1837) (Miller & Henle, 1841) Schultz, 1953 (Stromann, 1896) (Schultz, 1943) (Richardson, 1844) Schultz, 1953 (Bleeker, 1857) Bleeker, 1865 (Abbott, 1861) (Schultz, 1953) (Bleeker, 1859) Bleeker, 1865 (Schultz, 1953) (Schultz, 1953) Smith, 1962 (McClelland, 1845) (Richardson, 1845) Seale, 1906 Schultz, 1953 Gosline, 1958 (Lay & Benett, 1839) Bleeker, 1850 (Seale, 1917) (Schultz & Woods, 1949) (Riippell, 1828) (Klausewitz & Eibl-Eibesfeldt, 1959) O:1979ORSTOM cruise Method WAL O AAAAAAA AA << AADAAAA DA AA OA A AD” i Herklotsichthys quadrimaculatus Spratelloides delicatulus CHANIDAE Chanos chanos SYNODONTIDAE Saurida gracilis Synodus binotatus Synodus dermatogenys Synodus jaculum Synodus hoshinonis Synodus variegatus OPHIDIIDAE Brotula multibarbata *Brotula townsendi BY THITIDAE Brosmophyciops pautzkei *Dinematichthys randall *Dinematichthys riukiuensis ANTENNARIIDAE Antennarius coccineus *Antennarius duescus Antennarius nummifer GOBIESOCIDAE Diademichthys lineatus Discotrema crinophila *Pherallodichthys species *Pherallodus species ATHERINIDAE *Atherinomorus duodecimalis Atherinomorus lacunosus *Atherion elymus Hypoatherina barnesi HOLOCENTRIDAE Myripristis amaena Myripristis berndti Myripristis kuntee Myripristis murdjan ? Myripristis pralinia Myripristis violacea Neoniphon argenteus Neoniphon opercularis Neoniphon sammara Plectrypops lima Sargocentron caudimaculatum Sargocentron diadema Sargocentron melanospilos Sargocentron punctatissimum Sargocentron rubrum Sargocentron spiniferum Sargocentron tiere Sargocentron violaceum AULOSTOMIDAE Aulostomus chinensis FISTULARIDAE (Riippell, 1837) (Bennett, 1831) (Forsskal, 1775) (Quoy & Gaimard, 1824) Schultz, 1953 Fowler, 1912 Russell & Cressey, 1979 Tanaka, 1917 (Lacépéde, 1803) Temminck & Schlegel, 1846 Fowler, 1900 Schultz, 1960 Machida, 1994 Aoyagi, 1952 (Lesson, 1831) Snyder, 1904 (Cuvier, 1817) (Sauvage, 1883) Briggs, 1976 (Valenciennes, 1835) (Schneider, 1 801) Jordan & Starks,1901 Schultz, 1953 (Castelnau, 1873) Jordan & Evermann, 1903 Cuvier, 1831 (Forsskal, 1775) Cuvier, 1829 Bleeker, 1851 (Valenciennes, 1831) (Valenciennes, 1831) (Forsskal, 1775) (Valenciennes, 1831) (Riippell, 1835) (Lacépéde, 1801) (Bleeker, 1858) (Cuvier, 1829) (Forsskal, 1775) (Forsskal, 1775) (Cuvier, 1829) (Bleeker, 1853) (Linnaeus, | 758) VR VR VRO Zep eof maze) v2) Fistularia commersonii CENTRICIDAE Aeoliscus strigatus SYNGNATHIDAE Corythoichthys amplexus Corythoichthys nigripectus Corythoichthys schultzi Doryrhamphus dactyliophorus Doryrhamphus excisus excisus *Phoxocampus diacanthus SCORPAENIDAE Dendrochirus brachypterus Dendrochirus species Pterois antennata Pterois radiata Scorpaena species Scorpaenodes albaiensis *Scorpaenodes corallinus *Scorpaenodes hirsutus Scorpaenodes kelloggi Scorpaenodes parvipinnis Scorpaenodes scaber Scorpaenopsis species Scorpaenopsis gibbosa Scorpaenopsis neglecta Sebastapistes species Sebastapistes cyanostigma *Sebastapistes mauritiana Sebastapistes strongia Sebastapistes tinckhami Setarches species CARACANTHIDAE Caracanthus maculatus Caracanthus unipinna APLOACTINIDAE *Neoaploactis tridorsalis SERRANIDAE Anyperodon leucogrammicus Aporops bilinearis Belonoperca chabanaudi Cephalopholis argus Cephalopholis miniata Cephalopholis sonnerati Cephalopholis urodeta Epinephelus caeruleopunctatus Epinephelus coioides Epinephelus cyanopodus Epinephelus fasciatus Epinephelus hexagonatus Epinephelus macrospilos Epinephelus maculatus Epinephelus merra Epinephelus polyphekadion Epinephelus rivulatus Riippell, 1838 (Ginther, 1860) Dawson & Randall,1975 Herald, 1953 Herald, 1953 (Bleeker, 1853) Kaup, 1856 (Schultz, 1943) (Cuvier, 1829) (Bloch, 1787) Cuvier, 1829 (Evermann & Seale,1907) Smith, 1957 (Smith, 1957) (Jenkins, 1903) (Garrett, 1863) (Ramsay & Ogilby,1886) (Schneider, 1801) (Temminck & Schlegel, 1844) (Bleeker, 1856) (Cuvier, 1829) (Cuvier, 1829) (Fowler, 1946) (Gray,1831) (Gray,1831) Eschmeyer & Allen, 1978 (Valenciennes, 1828) Schultz, 1943 Fowler & Bean, 1930 Bloch and Schneider, 1801 (Forsskal, 1775) (Valenciennes, | 828) (Schneider, 1801) (Bloch, 1790) (Hamilton, 1822) (Richardson, 1846) (Forrskal, 1775) (Forster, 1801) (Bleeker, 1855) (Bloch, 1790) Bloch, 1793 (Bleeker, 1849) (Valenciennes, 1830) < AAAAAAADAAAAAAAAAAA << AAAAAA Ay wn 1oe) 14 Epinephelus tauvina Gracila albomarginata Grammistes sexlineatus Liopropoma susumi Liopropoma tonstrinum Luzonichthys waitei Luzonichthys williamsi Plectranthias longimanus *Plectranthias nanus Plectranthias winniensis Plectropomus laevis Plectropomus leopardus Pseudanthias ventralis ventralis Pseudanthias hypselosoma Pseudanthias lori Pseudanthias pascalus Pseudanthias pictilis *Pseudanthias rubrizonatus Pseudanthias squamipinnis Pseudogramma polyacantha Serranocirrhitus latus Variola louti PSEUDOCHROMIDAE Cypho purpurascens Pseudochromis species Pseudochromis cyanotaenia Pseudochromis jamesi Pseudoplesiops species Pseudoplesiops howensis Pseudoplesiops multisquamatus Pseudoplesiops rosae PLESIOPIDAE Plesiops coeruleolineatus ACANTHOCLINIDAE Belonepterygion fasciolatum KUHLIIDAE Kuhlia mugil PRIACANTHIDAE Heteropriacanthus cruentatus Priacanthus hamrur APOGONIDAE Apogon species Apogon angustatus Apogon apogonides Apogon aureus *Apogon caudicinctus Apogon coccineus Apogon cyanosoma *Apogon diversus Apogon doderleini Apogon doryssa Apogon erythrinus Apogon exostigma Apogon fraenatus (Forsskal, 1775) (Fowler & Bean, 1930) (Thiinberg, 1792) (Jordan & Seale, 1906) Randall & Taylor,1988 (Fowler, 1931) Randall & McCosker,1992 (Weber, 1913) Randall, 1980 (Tyler, 1966) (Lacépéde, 1801) (Lacépéde, 1802) (Randall, 1979) Bleeker, 1878 (Lubbock & Randall, 1976) (Jordan & Tanaka, 1927) (Randall &Allen, 1978) (Randall, 1983) (Peters, 1855) (Bleeker, 1856) Watanabe, 1949 (Forsskal, 1775) (De Vis,1884) Bleeker, 1857 Schultz, 1943 Allen,1987 Allen, 1987 Schultz, 1943 Riippell, 1835 (Ogilby, 1889) (Bloch & Schneider, 1 801) (Lacépéde, 1801) (Forsskal, 1775) (Smith & Radcliffe, 1911) (Bleeker, 1856) (Lacépéde, 1802) Randall & Smith, 1988 Rtippell, 1838 Bleeker, 1853 (Smith & Radcliffe, 1911) Jordan & Snyder, 1901 (Jordan & Seale, 1906) Snyder, 1904 (Jordan & Starks, 1906) Valenciennes 1832 o) O | IO[AR], “MA SisuaoiwuDl “Gq COT WNT Wd CTA EV [joyo1es vuafiuony Gq q WIUIeIYOS 39 SZeY XO ULNOID (BUIZINY) SisuaauINs ‘Gg saa = 15) CN ETITS eC SUE IN €T XH GAG Galy HG Ol CV. 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Yyydaq on adojs tayno pue e0edse}, Joor—et0y paemaey [X] edo[s 1eymo pue soei1e} Joo1—ai10y premputy FY sjooi—daliteq Jouul Alepuocoeg sJeer jerzeydized snonutjuoosiq a0eile}y feuocose7 f Sjoot yoyed yyIM ulseq [euoodeT [7] ( Jotieg,) jeer jeseydisod snonutyuon FY TIOLV VNVddds YOdVONOd ATOLL RESEARCH BULLETIN NO. 449 SCIENTIFIC STUDIES ON DRY TORTUGAS NATIONAL PARK: AN ANNOTATED BIBLIOGRAPHY BY T.W. SCHMIDT AND L. PIKULA ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. OCTOBER 1997 “W'S! BpUOpy ‘ ye JeuoMeN sesnyoy, Arq Jo uoyeso] oy) Suimoys dey *| oins1,j $19) 3801 11 sey = yartotd aes hay GQ" S, aa gas *: om sa ei? a ie Soe yleg |euoljyen : oo sebnyio} Aug " bing PPL 4ony Bn ons yieg ES jeuoljen 5 2 sape|b1aaq upaa9 917Un) 79 oaizay fo fing aj epsapney 1104 9agoyoaay) aruy VOIYOLA HANDS NI-WYVd TYNOLLVN SYORIYOL AYA 40 NOLLVOOT = Wee : wae s eee eee SCIENTIFIC STUDIES ON DRY TORTUGAS NATIONAL PARK: AN ANNOTATED BIBLIOGRAPHY BY Thomas W. Schmidt! and Linda Pikula” ABSTRACT Dry Tortugas National Park, located 110 km west of Key West, Florida, is an elliptical, atoll-like, coral reef formation, approximately 27 km long and 12 km wide with shallow water depths ranging from 12-20 m in channels between reefs. In 1935, the area was designated Fort Jefferson National Monument, the World’s first underwater National Park unit. Central to the area is Fort Jefferson, America’s largest coastal nineteenth century masonry fort. In 1992 it was re-designated Dry Tortugas National Park. Because of the islands’ unique location, the first tropical marine biological laboratory in the Western Hemisphere was established on Loggerhead Key by the Carnegie Institution of Washington, Washington, D. C. Following the closure of the Tortugas Laboratory in 1939, aperiodic marine biological assessments have been conducted in response to man- made and natural environmental perturbations. This annotated bibliography is an attempt to provide researchers and resource managers with access to the rapidly accumulating body of information on the park’s natural resources. A total of 424 references (published and unpublished) on scientific studies in, (and what later became) Dry Tortugas National Park were annotated and indexed according to major scientific topics. Studies from a wider area were included if they also sampled in Dry Tortugas National Park. BACKGROUND Seven small islands composed of coral reefs and sand in the eastern Gulf of Mexico, approximately 110 km west of Key West, Florida comprise Dry Tortugas National Park (Fig.1). The Tortugas, an area known for its bird and marine life and shipwrecks, are an elliptical, atoll-like, coral reef formation, approximately 27 km long and 12 km wide with water depths ranging from 12-20-m in channels between reefs. 1 South Florida Natural Resources Center, Everglades National Park, 40001 State Road 9336, Homestead, Florida 33034, U.S.A. 2 NOAA Regional Library, 4301 Rickenbacker Causeway, Miami, Florida 33149, U.S.A. Manuscript received 2 September 1997; revised 2 October 1997 ee ee eee ey ae a ee II The Dry Tortugas, discovered by the Spanish explorer Ponce de Leon in 1513 and named The Turtles, Las Tortugas, were soon read on early nautical charts as "Dry Tortugas" to indicate they lacked fresh water. Central to the area and located on Garden Key is Fort Jefferson, America's largest coastal nineteenth century masonry fort. Work was begun in 1846 and continued for thirty years but was never finished. As part of the United States coastal fortification buildup after the War of 1812, Fort Jefferson was considered critical for protecting Gulf trade and ports (Murphy, 1993). Following the Fort's use as a military prison during the Civil War (where the infamous Dr. Mudd was imprisoned after President Abraham Lincoln's assassination in 1865), and its abandonment by the Army in 1874, the area was proclaimed a wildlife refuge in 1908, to protect sooty tern rookeries from egg collectors. In 1935, the area was designated Fort Jefferson National Monument, the World's first underwater National Park unit. In 1992 it was redesigned Dry Tortugas National Park to preserve and protect both historical and natural features. Early descriptive observers of Dry Tortugas natural resources include Louis and Alexander Agassiz during the 1850's, and the research vessel Blake in 1877 and 1878. Their visits resulted in a detailed map of the islands, and a description of benthic marine communities by Agassiz in 1888. In 1903, Alfred G. Mayer, under the auspices of the Carnegie Institution of Washington, recommended that a tropical marine biological research laboratory be established at the Tortugas (as opposed to other Caribbean sites) because of their isolation from continental land masses, lack of commercial fisheries, lush reefs, clear waters and proximity to the Gulf Stream. In 1904, Mayer selected Loggerhead Key as the site for Carnegie's Tortugas Marine Laboratory, the first tropical marine laboratory in the Western Hemisphere (Fig.2). Following the closure of the Laboratory in 1939, relatively few investigations were conducted in the Tortugas until the National Park Service (NPS) began in 1975, a series of cooperative, bench-mark studies to evaluate long-term changes in Marine resources in combination with the earlier Carnegie Laboratory studies. Since the initial Tortugas Reef Atoll Continuing Transect Studies (TRACTS) work of 1975-76, aperiodic biological assessments have been conducted in response to man-made and natural environmental perturbations. PURPOSE The primary purpose of this annotated report is to provide researchers and resource managers with a readily accessible document on the rapidly accumulating body of information on the natural resources of the Dry Tortugas. With the recent implementation of the Florida Keys National Marine Sanctuary, adjacent to the Park's boundary, there is a dire need for a scientific database that is centrally located, coherently organized, and directly related to the future and ongoing management and regulation of marine resource activities. Ill No complete bibliography of the scientific studies on the park's marine and terrestrial natural resources has been undertaken. In this report we have attempted to list published and unpublished reports from many fields which we feel will be useful as a starting point for natural science studies to be conducted at the Dry Tortugas for decades to come (Figs.2&3). METHODOLOGY The present bibliographic database containing 424 references was compiled using PROCITE software, and covers the period of approximately 1878-1996, with the exception of one report dated 1820. Most 1996 papers were listed through August. Arrangement is alphabetical by senior author and title. Entries are numbered in sequence, and each includes a complete bibliographic citation with abstract or summary While some attempt has been made to achieve uniformity in style, in many cases the terminology, spelling, capitalization, and phraseology of the original author or abstractor have been retained. Abstracts obtained from the Carnegie Institution of Washington's publication citations were adapted from the author's summaries of results. This bibliography includes books, book chapters, scientific articles, theses and dissertations, workshop and conference proceedings, reports, and government publications. No attempt was made to include articles from newspapers or popular boating or sport magazines Several maps and charts are cited, however. Research citations were indexed by broad fields of study, specialty sub-headings, and by both senior and joint authors. Each citation is listed under as many subject headings as is appropriate for the cited article. This cross indexing system was constructed using PROCITE. The geographic boundaries for citations in this bibliography are that the work was done either completely or partially within the Park boundary, which is the 60' contour line (Fig.4). Recently, a few studies were undertaken adjacent to or on the Park boundary, they were included also. Although all Carnegie Institution of Washington published studies were included if they were conducted at the Dry Tortugas, those studies that were conducted and identified as solely in the "Gulf Stream" or at satellite marine laboratories in Jamaica, Trinidad, Puerto Rico, Bahamas, or in the tropical Pacific were not included A broad range of marine and terrestrial topics were found, including vegetation, marine algae, invertebrates, sea water composition, and geology. The major topics were sub- divided into those specialty areas that are shared most often among the studies examined. For example, sea water composition was sub-divided into salinity and temperature, the parameters measured most often. In many cases, inclusion or exclusion of a given reference within a major topic area or specialty sub-heading was a subjective decision. ‘(ZI6T) esnompysry Aoy peoysossoT pue sBuipying AloyesOge’] suey] sesNqOT AI *7 O1NSIJ Figure 3. (a) Research yawl “Physalia” and two dockhouses (acquarium/kitchen) on the warf (1906). (b) Dr. Mayor’s laboratory and work tables (1918). VI jaat Buibursy/saysseq yueg [yf] : aU] 4noJUoD 44 09 -- NOO*8E-97 ‘6L61 ‘WIE ‘99430 JJOYS [eJUOUNUOD JojN_C suRIJIO MON Aq poyst[qnd ‘9/6] “Ty ‘peasowoy ‘yeg jeuONeN sopelsi9Aq ‘staeq “q “D Aq pojidwios * sonrunuwoos o1yjueq SuIMOYs YIeg [eUOTRN sesnyioy, Aiq jo dey ‘p o1n31.7 ajqqni pue pues aieg[] s3)K a = = Rene =r = ES | I Wwo}}Oq pseH EY E $19] au | 1 : g pue) La 3 jao1 usoybeys Fy ; jao1 yo}ed Ka aebly ( sasseib eas [| 4 3 ) Py, pooyiobbo7 > A Dork CS hay vapiog o>. d of uoss0} jor 310 OS Ved & “Ce /u0b10} yop 3404 Ke ; eA i .—/ Boy (=F a! z | ae So fee (f | i= ji {— A ; a) ih : ‘ Ade WV) 1 HRY 1 i_\ ) —, ’ = \ =a \ =, ' (==) LEA (ER T Vd TNOLLYN SVONLHOL A¥C vil LITERATURE SEARCHED The bibliographic search was predominantly done at South Florida Natural Resources Center Library, Everglades National Park, Homestead, Florida, and at the NOAA Regional Library, Miami, Florida. No starting date was established for the references in this compilation. The senior author began assembling marine archival materials (e.g., raw data sheets, correspondence, maps, etc.) and published and unpublished research results from major scientific studies conducted by NPS scientists and contractors working at the Tortugas. Pertinent record files were also searched at Dry Tortugas National Park The Park Library contains a complete 35 volume set of the Carnegie Institution of Washington's Tortugas Laboratory Papers. Volumes 1-6 were titled "Papers from the Tortugas Laboratory of the Carnegie Institution of Washington" (1908-1914), while volumes 7-14 were "Papers from the Department of Marine Biology of the Carnegie Institution of Washington" (1915-1926), Volumes 15-29 were issued as "Papers from Tortugas laboratory of Carnegie Institution of Washington" (1928-1936) and volumes 30-35 were titled "Papers from Tortugas Laboratory" (1936-1942). Each volume was given a separate publication number by the Carnegie Institution. The Year Book Series of the Carnegie Institution, which contains annual summaries by individual investigators on observations and results obtained during their visits to the Tortugas, were searched at the University of Miami's Richter Library, Coral Gables, Florida. The Richter Library contains volumes 1-12, and 20-32. Copies of annual investigator reports for volumes 13-19 and 33-39 were obtained from the Carnegie Institute of Washington, Washington, D.C. In nearly all cases it was found that the principal investigators published their final evaluations and conclusions in "Papers", while the Year Book contained mostly duplicative, preliminary, or unsubstantiated observations. For these reasons, and due to time constraints, we decided to cite only Year Book contributions for investigators who did not complete and publish their conclusions in the "Papers" series. For example, W. H. Longley published in both series, but is only cited in this bibliography under "Papers" (however, his Year Book citations can be found in the "literature cited" section of his contributions to "Papers". S. Yamanouchi, however, only published in the Year Book, and is cited here as such. We searched documents regarding the Tortugas Laboratory during two visits to the Carnegie Institution in Washington D.C. Work conducted at the Tortugas Laboratory has been published in a wide range of journals. For example, a list of scientific writings produced by activities at the Laboratory during Mayer's directorship can be found in Papers Tortugas Laboratory 19:80-90. Many publications continued to appear in the literature following the closure of the Laboratory in 1939. On-line database searches were conducted during 1993-96 at the NOAA Miami Regional Library. Subject index terms such as coral reef, geology, vegetation, marine algae, fish, Vili etc., were used to search on a variety of DIALOG electronic databases including the following: BIOSIS PREVIEWS, Dissertation Abstracts, Oceanic Abstracts, SCISEARCH, Ei Compendex, INSPEC, and GEOBASE. These individual CD’s were also searched: Aquatic Sciences and Fisheries Abstracts, Life Sciences Collection, Earth Sciences, GeoRef, NTIS (National Technical Information Service), GPO (Government Publications Office), and the OCLC (Online Computer Library Center) on- line catalog. Pertinent theses, dissertations, and journals identified in the abstracted literature were obtained via interlibrary loans. This current compilation undoubtedly does not list all available literature that might be useful in conducting research, monitoring, and resource management of the park's natural resources. There may be as many as 100 additional scientific papers generated from the Carnegie era. We would greatly appreciate additional references to the Tortugas literature, and if a sufficient number of additional articles become available, we will produce an addendum to this report. ACKNOWLEDGMENTS This report has benefited from the help of many people over the past 6 years. The original project was prompted and supported by Dr. Michael Soukup during his tenure as Director of the South Florida Natural Resources Center. Wayne Landrum, Facility Manager and Carolyn Brown-Wiley, Chief Ranger at Dry Tortugas National Park provided logistical support and took special interest in providing guidance to the pertinent files at Fort Jefferson. Ray Bowers, John Strom, and Pat Craig of the Carnegie Institution in Washington, D.C. permitted us to search their Tortugas Laboratory files, assisted in duplicating activities, and provided insightful discussions and original photographs of the Marine Laboratory. We thank George Stepney and Maria Bello of NOAA’s Regional Library in Miami for acquiring many interlibrary loans. Special recognition goes to the staff of the South Florida Natural Resources Center including Marnie Lounsbury for photocopying and collating much of the Carnegie texts, Barry Wood who produced the map figures, and Mario Alvarado who expertly produced the author and subject indexes using PROCITE. Dr. William B. Robertson, Jr., United States Geological Survey/Biological Resources Division contributed numerous references and provided encouragement during the earliest stages of the project Valuable comments were provided by Elaine Collins of the NOAA Central Library, Silver Spring, MD and Bob Hamre, former technical editor for the US Forest Service, assigned to the Beard Center under the NPS “Volunteer-in-Parks.” program. Finally, our thanks to Carol Watts, Chief of the NOAA Libraries and Information Science Division, Janice Beattie, Chief of NOAA Libraries Public Services Division, Dr. Tom Armentano, Chief of Biological Resources, South Florida Natural Resources Center, and Dr. Caroline Rogers, United States Geological Survey/Biological Resources Division for their financial support and encouragement. ANNOTATED BIBLIOGRAPHY 1. Agassiz, A.. 1888. The Florida reefs. Three cruises of the United States Coasts and Geodetic Steamer 'Blake' in the Gulf of Mexico, in the Caribbean Sea and along the coast of the United States, from 1877 to 1880., V. 1, Chapter 3, pages 52-92. Houghton, Mifflin and Co. New York. 314 pp. While the Steamer "Blake" was mostly involved in deep water dredging operations in the Gulf of Mexico and the Caribbean Sea (1877-78), a five-week visit to Fort Jefferson (Dry Tortugas) provided the author with an opportunity to work in a laboratory-like situation, to examine carefully the topography of the different groups of corals characteristic of the Florida reefs and to give an extended account of the Florida reefs in a special chapter of this book. The Tortugas, as described by Agassiz, form the most recent of the cluster of the Florida reefs, and it is here where he begins a topographical sketch of the Florida reefs from the Tortugas to Cape Florida. . 1888. The Tortugas and Florida Reefs. Memoirs of the American Academy of Arts and Sciences, Philadelphia. Centennial ed., V.II, :107-132. VII entitled "Explorations of the surface fauna of the Gulf Stream, under the auspices of the United States Coast Survey”. Agassiz reports on the formation of the Florida Reefs, commenting on the theories of Darwin, LeConte, and Hunt on this subject. At the time of this article it was believed that the elevation of the Florida Plateau, from Cape Florida southward to the Dry Tortugas and the Yucatan Banks, was based on the accumulation of coral sands, as well as animal debris brought to it by great Atlantic equatorial oceanic currents, the Gulf Stream and prevailing winds, and at which time, reef-building corals could flourish and a reef would be formed. They speculated that corals could not thrive below 6 or 7 fathoms, because siltation ooze would sink to the bottom and choke the corals. Coral reef formations were assumed to be established near strong equatorial currents which were suppliers of food for the reef- building corals. It was assumed that corals grow towards the surface as fast as the ooze deposited has closed up the circulation of the lower levels. At the time of Darwin little was known of limestone deposits formed by the accumulation of animal decay. Thus explanations of reef formation other than elevations as a result of submerged mountains and subsidence were not investigated. The Tortugas Reefs probably are newly developed, as they have not been above the sea long enough to have received the flora and fauna characteristic of the Keys north of Key West. 3. Andres, B. A. 1991. Migration of sharp-shinned hawks in the Dry Tortugas, Florida USA. Wilson Bulletin 103, no. 3: 491-93. Some species of hawks have been found to make long water crossings during migration. One of the species, the sharp-shinned hawk Accipiter striatus, seldom undertakes water crossings of >125 km. However, large numbers of sharp-shinned hawks are observed every fall in the Florida Keys including the Dry Tortugas, where water crossings are quite common. No information, however has been gathered concerning their migration after reaching the Tortugas. Based on wind speed, wind direction, and binocular observations made on Garden Key of six species of hawks (sharp-shinned hawks dominated the observations), it was found that sharp-shinned hawks are deliberately initiating an over- flight across the Gulf of Mexico directly to Central America. 4. Austin, O. L. Jr., W. B. Robertson Jr. and G. E. Woolfenden. 1972. Mass hatching failure in Dry Tortugas sooty terns, Sterna fuscata. Proceedings of the International Ornithological Congress. 627. Netherlands. The author attributes a mass hatching failure among 50,000 pairs of sooty terns (Sterna fuscata) nesting on the Dry Tortugas to damage caused by sonic booms from low-flying military aircraft. Theoretically, eggshells and embryonic tissues should withstand pressures much greater than those generated by even the most intense sonic booms. 5. Bailey, E., G. E. Woolfenden and W. B. Robertson Jr. 1987. Abrasion and loss of bands from Dry Tortugas sooty terns. Journal of Field Ornithology 58, no. 4: 413-24. During the past 25 years more than 400,000 sooty terns (Sterna fuscata) have been banded at Dry Tortugas, Florida, with size 3 aluminum bands of several different alloys. Based on large samples of bands removed from the terns, regression lines were established for each of four alloys. Differences in the slopes of the regression lines for certain of the four alloys demonstrated differences in rates of abrasion. Band loss was evident for bands of the fastest abrading alloy (2-SO) that were carried more than 20 yrs by terns banded as chicks because all band weights fell above an extension of the regression line. For this alloy, the plots of weight loss showed that band loss becomes significant at 86% of original weight. Bands of alloy 2-SO began reaching 86% of original weight at age 14 when placed on adults and 20 when placed on chicks. The regression lines for the other 3 alloys suggest that loss is likely after 17-28 yrs for bands placed on adults and after 20-25 years for bands placed on chicks. Band loss probably occurs through abrasion of the inner surface, which increases the inner diameter until the band can slip over the toes. Any gap that develops would hasten loss. 6. Bailey, P. L. 1938. Regeneration in sabellids. Carnegie Institution of Washington, Year Book 37: 84-85. Sabellid worms collected in the moat on Garden Key proved to be suitable for fixation techniques needed to conduct various regeneration experiments to determine the effects of chemical solutions on the cells. 7. Baker, B. 1994. Partitioning the National Marine Sanctuary. Bioscience 44, no. 7: 497. A management proposal is described to establish five zones in the Florida Keys National Marine Sanctuary: replenishment reserves, sanctuary preserve areas, research-only zones, wildlife management zones, and special-use zones. The Sanctuary encircles the Florida Keys, including the Dry Tortugas, for 2800 square nautical miles. 8. Ball, S.C. 1918. Migration of insects to Rebecca Shoal Light-Station and the Tortugas Islands, with special reference to mosquitoes and flies. Papers Tortugas Laboratory 12: 193-212. Carnegie Institution of Washington Publication Number 252. The circumstances which suggested the desirability of such investigations were the repeated experiences of Dr. Mayer and other scientists at Tortugas, Florida, in connection with the occurrence there of mosquitoes. These insects were abundant on Loggerhead Key only after northerly winds of several hours' duration, under conditions favorable to their migration from the mainland of Florida. Rebecca Shoal light-station was chosen as the study site, because of its isolation from the mainland and other keys, and because of its freedom from all except easily controllable breeding-places for mosquitoes. It was found that large numbers of mosquitoes and house-flies are carried by northerly and southerly winds to Rebecca Shoal light-station and the Tortugas Islands from Florida and Cuba. Easterly winds bring a few of these, as well as smaller numbers of blow-flies, horse-flies, and gnats from islands east on the Florida Reef. Occasionally Odonata, Neuroptera, and Lepidoptera are carried by the winds to these parts of the reef. Sarcophagidae breed in land crabs at Tortugas . 9. Ballantine, D. L. 1996. New records of benthic marine algae from Florida. Gulf of Mexico Science 118 i iets), Seven species of benthic marine algae are newly reported from the Dry Tortugas, Florida. These are Halimeda hummii Ballantine (Chlorophyta), Audouinella ophioglossa Schneider, Botryocladia uynnei Ballantine, Champia viellardii Kutzing, Monosporus indicus Borgesen, Hypoglossum rhizophorum Ballantine et Wynne, and Rhodogorgon ramosissima Norris et Bucher (Rhodophyta). Monosporus indicus is reported for the first time from the Atlantic Ocean. The Dry Tortugas represents the northern distributional limit for the remaining species reported, except Audouinella ophioglossa and Botryocladia uynnel. 10. Bartsch, P. 1919. "The bird rookeries of the Tortugas." Smithsonian Institution Annual Report for 1917, 2512. Smithsonian Museum. The author states that the most interesting island of the Tortugas group is Bird Key (circa 1908). Of the 32,810 birds listed for the islands, 31,200 center about that Key. A numerical listing of the summer birds is given. These rookeries were first brought to the attention of ornithologists by John Audubon in his ornithological biographies. He gives an account of a visit in May 1832. A first list of birds observed in the Dry Tortugas is given by W.E.D. Scott in his paper on birds observed during parts of March and April 1890. Drs. John B. Watson and K.S. Lashley of Johns Hopkins University made an extensive study of the wild bird colonies there, hoping to throw light on the homing instinct. The article ends with an extensive listing of bird sightings in the Tortugas up to 1919. 11. ————. 1920. Experiments in the breeding of Cerions. Papers Tortugas Laboratory 14: 1-55. Carnegie Institution of Washington Publication Number 282. Breeding experiments were conducted to determine if various forms of Cerion colonies were fixed forms, that is, will generations yield the same mode in measurement, or will changes in the local environment from season to season affect the developing organisms to such an extent as to produce an unending series of slight variations? Introduced forms were placed where native species existed. Colonies of these land snails were planted on Keys in the Dry Tortugas in 1914, 800 in 1915 on Loggerhead Key, and a third planting in 1916 of 8,317 specimens. A comparative anatomical discussion of the five species of Cerion involved in the breeding experiments is given. 12. ————. 1915. Report on the Bahama Cerions planted on the Florida Keys. Papers Tortugas Laboratory 8: 203-12. Carnegie Institution of Washington Publication Number 212. A study is made of the two races of Bahama Cerion transplanted to the Florida Keys in 1912. The conditions of the Cerion colonies are described.. Illustrations show the extent of the changes between the first generation and the parent generation. Changes that have taken place in the second generation in shell, color and sculpture are discussed. 13. . 1919. Results in Cerion breeding. Proceedings of the Biological Society of Washington, 32. Journal Washington Academy of Science 9:657 (abstr.) A short account is given of Dr. Paul Bartsch's report on the breeding of Cerions transplanted from Andros Island in the Bahamas to the Dry Tortugas. 14. ———. 1916. Visit to the Cerion colonies in Florida. Smithsonian Explorations 66, no. 17: 41-44. The author visited the Dry Tortugas through the auspices of the Carnegie Institution and the U.S. National Museum to observe the transplanted Bahamian Cerion colonies. He reported finding many adult specimens of the first Florida generation. No adult second generation specimens were found. Four hybrid specimens between the native Cerion incanum and the transplanted Bahama stock were obtained. 15. 16. 17. 18. ID) 20. Bellow, T. and C. Winegarner. 1975. Nesting of brown pelicans Pelicanus occidentalis on the Dry Tortugas, Florida. Florida Field Naturalist 3, no. 2: 47-48. On 14 June 1974 on Bush Key, Dry Tortugas, Florida T. Bellow and C. Winegarner found 5 Brown Pelican nests about 12 feet above ground in the white mangroves (Laguncularia racemosa) along the north shore. Nineteenth-century records of pelicans breeding on the Dry Tortugas are ambiguous. It appears that a few pairs did breed on the Tortugas in the mid-1800's, but by late in the century none did so. This record is the first reported nesting of this species in the 20th century on these ornithologically well-known islands. Three of the nests found in 1974 contained 2 eggs each, one nest was empty, and the fifth was not checked. Bellow, T. H. 1979. A cardinal at the Dry Tortugas, Florida. Florida Field Naturalist 7, no. 2: 31. The southern range of the nonmigratory cardinal (Cardinalis cardinalis) extends through the Florida Keys, but is considered rare in Key West. An April observation of a cardinal on Garden Key represents the second published record of this species at the Tortugas. Bennett, F. M. Commander. 1909. A tragedy of migration. Bird-Lore 11: 110-113. On, April 14, 1909 a violent storm hit the Florida Keys, including the Dry Tortugas. An apparent bird migration was in progress at the time of the storm. On April 20th the author went to the Dry Tortugas and observed hundreds of dead birds, and tens of thousands of injured and exhausted birds. A listing of the types of birds observed is given. Berrill, N. J. 1938. Budding in polystryelid ascidians. Carnegie Institution of Washington, Year Book 37: 85. The area of the budding rudiment relative to the size of the parent zooid was closely related to the size of the parent zooid, and the general nature of the colony. Blinks, L. R. 1926-1929. Electrical conductivity in Valonia. Carnegie Institution of Washington, Year Book. Note published as follows: 1926, v. 25, p. 240; 1927, v. 26, p. 217-18; 1928, v. 27, p. 270- 71; 1929, v. 28, p. 280. This study makes use of the good supply of Valonia at Tortugas for studies on the variability of electrical resistance in protoplasm. Causes of uncertainty are discussed. Bohnsack, J. A., D. E. Harper and D. B. McClellan. 1994. Fisheries trends from Monroe County, Florida. Bulletin of Marine Science 54, no. 3: 982-1018. Fishing is an important activity in the Florida Keys National Marine Sanctuary (FKNMS). Concern exists that excessive fishing could be deleterious to individual species, disrupt marine ecosystems, and damage the overall economy of the Florida Keys. We examined data from commercial, recreational, and marine life fisheries in Monroe County, Florida. Invertebrates comprised the majority of commercial landings. In 1992, the total reported commercial landings were composed of 54% invertebrates (4.09 x 10 kg) 28% reef fishes (2.19 x 10 kg), and 21% non-reef fishes (1162 x 10 kg). In the recreational headboat fishery, reef fishes accounted for 92% of 0.107 x 10 kg average total annual landings from the Dry Tortugas and 86% of 0.201 x 10 kg landed from the Florida Keys since 1981. Average annual landings for other recreational fisheries were estimated at 1.79 x 10 kg for reef fishes (45%) and 2.17 x 10 kg for non-reef fishes (55%) from 1980 through 1992. Estimated landings from the Dry Tortugas did not show distinct trends and were highly variable. Finer resolution of catch and effort data are needed, especially for recreational fisheries. Landings for some species varied greatly over time. The most conspicuous declines were for pink shrimp, combined grouper, and king mackerel, while the most conspicuous increases were for amberjack, stone crab, blue crab, and yellowtail snapper. Landings of spiny lobster have remained constant. Fisheries closed to harvest included queen conch, Nassau grouper, jewfish, and stony corals. Effective fishing effort has increased over time with more participants and more effective fishing technology. Since 1965, the number of registered private recreational vessels has increased over six times, while the number of commercial and headboat vessels has remained stable. The number of management actions have continually increased and become more restrictive with increased fishing effort. Comparison of fisheries was complicated because different fisheries targeted different species and different sized organisms. Also, landings were sometimes reported by numbers and sometimes by weight. Measures of reproductive value and spawning potential are suggested as useful parameters for comparing effects of different fisheries. The new FKNMS provides a unique opportunity to shift management emphasis from a species approach to an ecosystem and habitat based approach. 21. Bortone, S. A., P. Rebenack and D. M. Siegel. 1981. A comparative study of Diplectrum formosum and D. bittatum (Pisces: Serranidae). Florida Scientist 44, no. 2: 97-103. Specimens of the simultaneously hermaphroditic fish species Diplectrum formosum, the sandperch, and D. bivittatum , the dwarf sandperch, were collected near the Dry Tortugas, Florida, by means of shrimp trawl during December 1976. Stomach contents of 326 D. formosum (100 empty) and 325 D. bivittatum (131 empty) revealed little or no differences in their food habits relative to number and volume of food items, size of food items or the contribution, in grams each food item makes to each fish. Both species primarily consumed amphipods, shrimp, crabs, fish, and polychaetes. Temporally, both species fed at the same 2 diurnal periods. Species were collected sympatrically but there were areas where each species dominated in relative abundance. 22. Boschma, H. 1929. On the postlarval development of the Coral Maeandra aerolata (L.). Papers Tortugas Laboratory 26: 129-47. Carnegie Institution of Washington Publication Number 391. During six weeks in July and August 1925, the author studied Maeandra areolata for researches on the food of reef-corals at the Carnegie Laboratory in the Tortugas. Many of the colonies contained ripe larvae and the author reared these for the study of their development. The author concludes the development of the endotentacles which appear constantly in two successive groups of three, resembles in some way the facts recorded by de Lacase Duthiers (1872) in Actinia mesembryanthemum, some stages of which show a marked tri-radial arrangement of the tentacles. The data in the literature on the development of other coral polyps seem to prove that this successive development of the endotentacles in two groups is an exceptional case. The bilateral arrangement of the septa in the oldest stages is in accordance with that found by Duerden (1904) in Siderastrea. As in the majority of corals in which the young stages are known, the septa in Maeandra develop in two cycles, first the six endosepta and soon afterward the six exosepta. 23. Bowles, A. E., F. T. Awbrey and J. R. Jehl. 1991. Effects of high-amplitude impulsive noise on hatching success: a reanalysis of the sooty tern incident, HSD-TR-91-0006. BBN Laboratories, Inc., Canoga Park, California. This article attempts to refute the Austin article which attributed a mass hatching failure among 50,000 pairs of sooty terns (Sterna fuscata), who had nested on the Dry Tortugas to sonic boom damage from military aircraft. Theoretically, eggshells and embryonic tissues should withstand pressures much greater than those generated by even the most intense sonic booms. An experiment was conducted to test whether impulsive noise could be responsible for the hatching failure. Four pest control devices were exploded near chick eggs in various states of development: 20 chicken and 20 quail eggs. The mean peak flat sound pressure level 177.3 db re 20 upa; mean CSEL of 139; mean frequency 620 Hz. No cracking damage similar to that of the Dry Tortugas eggs occurred. Hatch rates and weights between control and exposed embryos were not significantly different. 6 24. Bowman, H. H. M. 1918. Botanical ecology of the Dry Tortugas. Papers Tortugas Laboratory 12: 109-38. Carnegie Institution of Washington Publication Number 252. As the name of these islands indicates, their vegetation is characteristically xerophytic, although the rainfall is sufficient to assure the plants the necessary amount of water. The plants are very interesting when a close study is made of their individual characteristics. The opportunity for such study was given the writer during the summers of 1915 and 1916, while pursuing another line of botanical research at Loggerhead Key, where a marine laboratory is maintained. The Tortugas are really the westernmost of all the Florida Keys, but are more detached from them and have different geological and botanical aspects. Species distributional maps were created for each of the eight islands of the Tortugas Atoll. In this treatment of the species in the Tortugas it has been aimed to give some idea of the character of the dry-climate plants inhabiting these islands, their distribution, and particularly the changes which have occurred on the various keys since Lansing's 1904 survey, with an attempt to analyze the reasons for such changes. Notes on the marine ecology of the Tortugas also are presented, along with descriptions of dominant submergent vegetation. 25. Boyden, A. 1934-1939. Serological study of the relationships of some common invertebrata. Carnegie Institution of Washington, Year Book. Note: published as follows: 1934, v. 33, p. 248-49; 1936, v. 35, p. 82; 1939, v. 38, p. 218. Preliminary results obtained through the study of the antigens collected from various major groups of animals at Tortugas were summarized. Blood relationships within Mollusca and Crustacea were emphasized. 26. Bradbury, R. C. 1992. First Florida record of variegated flycatcher Empidonomus-varius at Garden Key, Dry Tortugas. Florida Field Naturalist 20, no. 2: 42-44. The variegated flycatcher occurs throughout most of South America east of the Andes. The species migrates northward between September and February after breeding in the middle and southern part of the continent. It winters in the Guianas, northern Brazil, Venezuela, Colombia, and eastern Peru. This article describes observations of a variegated flycatcher in Florida, representing the first record in Florida and the third in the United States. 27. Breder, C. M. Jr. 1934. On the habitats and development of certain Atlantic Synentognathi. Papers Tortugas Laboratory 28: 1-35 (issued Dec. 1932). Carnegie Institution of Washington Publication Number 413. In this paper data presented are intended to form a basis for further inquiry into the comparative development and life habits of the Synentognathi, which includes the familiar Belonidae (needlefish), Hemiramphidae (halfbeaks), and Exocoetidae (flying fish). The data on which the present paper are based represent some field studies and laboratory work on material gathered in the Dry Tortugas, Florida during May and June 1929 The feeding habits, leaping, and flight during the presence and absence of light, eye specialization, enemies, and ontogeny and phylogeny are discussed. A new species, Strongylura longleyi is described. A key to the Tortugas Synentognathi is provided, along with tables, beak measurement and eye development. The Exocoetidae form the major item of diet of a variety of sea birds, about the Tortugas at least. Young Exocoetidae pass the most dangerous part of their day when the sun is low, at which time they are unable to see their predacious enemies coming from below because of light conditions. The eyes of Belonidae are provided with elaborate equipment to protect them from the brilliance of their environment. 28. Breder, C. M. Jr. and J. E. Harris. 1936. Effect of light on orientation and stability of young plectognath fish. Papers Tortugas Laboratory 29: 23-36 (issued Nov. 1935). Carnegie Institution of Washington Publication Number 452. Under certain circumstances some plectognath species will respond to a strong beam of light by violent gyrations. This was first observed by Breder (1929) at the Tortugas Laboratory. The mechanism by which these movements are effected, their relationship to the intensity and duration of the stimulus, and the disappearance of the phenomenon with advancing age of the animal give rise to a number of interesting problems concerning the action of the receptor-effector system in these fish. This paper is an attempt to explain this feature of fish behavior. It was found that small specimens of Monocanthus and Lactophrys, if exposed to a beam of light, frequently exhibit somersaulting or rotational movements of great rapidity. Somersaulting is produced by passing the locomotor waves in opposite direction along the dorsal and anal fins. The rotational movement is accompanied by the deflection of the dorsal and anal fins to the opposite sides of the body, the direction of motion of the undulations being usually antero-posterior in both fins. Various combinations of fin and tail movements may occasionally give other twisting gyratory movements. The primary response to light is always an attempt at reorientation of the animal so that the light is incident upon the dorsal surface. In fish kept in complete darkness and "sensitized" by repeated stimuli, gyrations may continue after the light has been removed, and even mechanical stimuli may initiate similar paroxysms, the equilibrating system apparently being more or less permanently deranged. Specimens of Monocanthus over 50 mm. in length do not usually display this behavior, and species other than plectognaths show it very feebly or not at all. The integration of gravitational stimuli into the behavior pattern is apparently not perfectly attained until a comparatively late stage in development, and light is the primary orienting factor. The gyrations are apparently due to instability, consequent upon overcorrection. 29. Brinley, F. J. 1937-1938. Studies on the implantation of embryonic fish tissue, with notes on the spawning habits and development of four species of fish. Carnegie Institution of Washington, Year Book. Note: published as follows: 1937, v. 36, p. 86; 1938, v. 37, p. 86-7. Livers and spleens were transplanted from embryos of the hard head shiner to other embryos of the same age. No apparent effect on the host was noticed. Eggs of Pomacentrus and parrot fish were collected for observation, along with nurse shark embryos. Additional work was performed on the origin of muscular movement in these species. 30. Brooks, H. K. 1962. Reefs and bioclastic sediments of the Dry Tortugas (abs.). Geological Society of America. Special Paper 73: 1-2. Many miscomprehensions exist relative to origin of the Florida Reef track and, in particular, its southwestern extremity-the banks, shoals, and reefs known as the Dry Tortugas. They are not an atoll as stated by Vaughan (1914). The component physiographic features rise from a shallow limestone platform 80 to 100 feet below sea level. Relief features are banks and shoals of bioclastic sands. Their genesis and circulation distribution are related to the prevailing seasonal storm patterns. Large patches of Acropora cervicornus (Lamark) are widely distributed through the area in water less than 60 feet deep. Live coral on these patches is sparse. Proliferation of the staghorn corals is slow, but cumulative growth has produced a magnitude of skeletal remains. The coralla are preserved and are ultimately indurated into a porous rocky mass by the luxuriant growth of Lithohamnion and its cognate encrusting associates. The shallow reefs of Garden and Loggerhead Keys, populated by calcareous algae, alcyonarians, and scleractinians, etc., originate upon a foundation of the remains of these organisms. This can be seen where erosion in surge channels has exposed the underlying materials. SIE a2 33% 34. Brooks, W. K. 1908. Salpa floridian (Apstein) Part II in the Pelagic Tunicata of the Gulf Stream. Papers Tortugas Laboratory 1: 75-89. Carnegie Institution of Washington Publication Number 102. This rare Salpa about which little is known, has been noted in this paper. Mature specimens of both stages of Salpa were found, in May 1906, on the surface in the vicinity of the Marine Biological Laboratory at Tortugas, Florida; and an opportunity was afforded to study and sketch them while alive, and thus to make additions to, and some slight- corrections of, the count of the species.. Brooks, W. K and C. Kellner. 1908. On Oikopleura tortugensis, n.sp. a new appendicularian from the Tortugas, with notes on its embryology in Part IV, The Pelagic Tunicata of the Gulf Stream. Papers Tortugas Laboratory 1: 73-95. Carnegie Institution of Washington Publication Number 102. This species was found in abundance near the Marine Laboratory. The specimens are from 5 to 8 mm. long and occur in great swarms at the depth of 5 to 6 fathoms. A description of the species is provided. Brown, D. E. S. 1935. Cellular reactions to high hydrostatic pressures. Carnegie Institution of Washington, Year Book 34: 76-77. Physiological studies were carried out on the muscles of crabs and fish collected in deep (100 fathoms) and shallow water of the Tortugas. Brown, W. Y. and W. B. Robertson Jr. 1975. Longevity of the brown noddy. Bird-Banding 46, no. 3: 250-251. Despite its abundance and pantropical range, little published information exists on the longevity of the brown noddy (Anous stolidus). Woodward (Atoll Research Bull, 164: 280,1972) reported a maximum known survival of 10 years for brown noddies banded as adults on Kure Atoll , Hawaii. Brown noddies on Manana Island, Oahu, Hawaii (A. s. pileatus) and the Dry Tortugas, Florida (A. s. stolidus), are among the few populations that have been banded over a period long enough to provide quantitative data on longevity. Twelve of the brown noddies banded on Manana before 1948 were recaptured dead or alive before 1960, the longest interval from banding to recapture being 13 years. On 23 May 1972 Brown recaptured on Manana a brown noddy that had been banded there as a juvenile on 12 June 1947, 25 years earlier. 35. Bullington, W. E. 1940. Some ciliates from Tortugas. Papers Tortugas Laboratory 32: 179-221 (issued Sept. 1940). Carnegie Institution of Washington Publication Number 517. During the summers of 1930, 1931, and 1935, during a special study of spiraling in certain species of ciliates at Tortugas, there appeared in the author’s cultures from time to time many other species which seemed to be new or little known. There are now fifteen species, either new to science or little known, about which it is believed sufficient information is available to justify their description or redescription. Five of the fifteen species have previously been described, but none of them is well known. Ten were described as new. They were characterized by amazing shades of color, yellow and red predominating. The species here discussed and described constitute only a few of those which have been seen at Tortugas at one time or another, but these are all the author feels justified in discussing, at the present time, with the information at hand. 36. Burkenroad, M. 1929. Studies upon plankton and the mechanism of sound production in Haemulidae. Carnegie Institution of Washington, Year Book 28: 283-90. Daily tows were made from May 31 to August 19 near Loggerhead Key. The variety of species and numbers of individuals found disputed the notion that the Tortugas region "once noted for the variety and richness of its floating life, has gradually become in recent years an almost desert sea." 37. Caira, J. N. and M. H. Pritchard. 1986. A review of the genus Pedibothrium Linton, 1909 (Tetraphyllidea Onchobothriidae) with a description of two new species and comments on the related genera, Pachybothrium Baer and Euzet, 1962 and Balanobothrium Hornell, 1912. Journal of Parasitology 72, no. 1: 62-70. A review of the genus Pedibothrium Linton, 1909 is based on type and voucher specimens. The type species, Pedibothrium globicephalum Linton, 1909 is redescribed. Descriptions of Pedibothrium brevispine Linton, 1909 and Pedibothrium longispine Linton, 1909 are emended. Two new species are described, the generic diagnosis is emended, and a key is provided. 38. Calder, D. R. 1992. Similarity analysis of hydroid assemblages along a latitudinal gradient in the Western Atlantic. Canadian Journal of Zoology 70, no. 6: 1078-85. Shallow-water (0-100 m depth) hydroid faunas reported from 26 locations along the western North Atlantic coast between the high Canadian Arctic archipelago and the Caribbean Sea were compared. Species numbers varied widely between locations, but were highest in the tropics and subtropics, lowest in arctic and subarctic waters, and intermediate in mid-latitudes. Percentages of species producing free medusae were lowest in high latitudes, intermediate in low latitudes, and highest in mid-latitudes (especially in estuaries). In a numerical analysis, similar hydroid faunas were identified at locations (1) between the high Canadian Arctic islands and the Strait of Belle Isle off western Newfoundland; (ii) between the Gulf of St. Lawrence and Chesapeake Bay; (iii) between North Carolina and southeastern Florida (south as far as St. Lucie Inlet), and including the northern Gulf of Mexico; (iv) in the Caribbean Sea, together with Dry Tortugas and the oceanic island of Bermuda. The greatest change in hydroid species composition along the coast appeared to occur around Cape Hatteras. 39. Carrier, J. C., H. L. Pratt Jr. and L. K. Martin. 1994. Group reproductive behaviors in free-living nurse sharks, Ginglymostoma cirratum. Copeia 3: 646-56. Mating events of the nurse sharks were observed in a nine-day period in the Dry Tortugas islands. There were four stages of mating: precoupling, coupling, positioning and alignment, and insertion and copulation. Films were made of four of the mating events. Seminal fluid released into the water was obtained following one of the copulations. It showed the presence of free, nonpackaged sperm cells. Of the fifty mating events observed, ten of these involved multiple males attempting to copulate with single females. 40. Carrier, J.C. and H. L. Pratt Jr. 1997. Habitat management enclosure of a nurse shark breeding and nursery grounds. Fisheries Research (In press). Based on nurse shark breeding studies conducted at Dry Tortugas, a sanctuary for nurse shark reproductive and nursery activities is being established at Dry Tortugas National Park. 41. Cary, L. R. 1915. The Alcyonaria as a factor in reef limestone formation. Proceedings of the National Academy of Science 1: 285-89. In many areas of the Floridean-Antillean region, Gorgonaceae rather than stony corals make up the most characteristic feature of the lime-secreting organisms permanently attached to the bottom. In this paper, data are presented on the amount of material contributed to reef formation by gorgonians. Three factors were taken into consideration: spicule content (the amount of lime held as spicules in the colonies), distribution of gorgonians on the Tortugan reefs (the bulk of the gorgonians on any reef area) and 10 42. 43. 44. disintegration of the coenenchyma of the colonies and the addition of their spicules to the reef building materials. Using line surveys and the weight and percentage of spicules in the colonies, it was found that the amount of lime held as spicules in the tissue of living gorgonians per acre of reef area is 5.28 tons. Next to the destruction of the colonies by wave action (storms), the greatest mortality of the colonies is from overgrowth of tissues by other organisms. The destruction of Tortugan gorgonian colonies was nearly complete in the hurricane of October 1920. It has been estimated that nearly one-fifth of the gorgonian colonies are destroyed annually. . 1918. The Gorgonaceae as a factor in the formation of coral reefs. Papers Tortugas Laboratory 9: 341-62. Carnegie Institution of Washington Publication Number 213. An important constituent of the limestone of coral reefs is the calcium carbonate secreted in the skeletal structures of Anthozoa and marine calcareous algae. Representatives of the Hydrozoa were important reef formers in past geological epochs, but in the formation of modern reefs they constitute a minor factor. Representatives of the Anthozoa, the stony and flexible corals, are among animals the only important agents in the formation of the modern reefs. The results of this study show that over large reef areas, in the Tortugas at least, the gorgonian fauna is by far the most important element contributing to the formation of reef limestones. The amount of spicules in the tissues of gorgonian colonies would average at least 5.28 tons to the acre for all of the reefs in the Tortugas group. The figures given represent only a potential contribution to reef formation but a study of the normal cycle of changes in the gorgonian fauna of this region has shown that at least a fifth of this amount of calcium carbonate, as spicules, will be added to the reef limestones annually . . 1934. Growth of some tissues of Ptychodera bahamensis in vitro. Papers Tortugas Laboratory 28: 195-213. Carnegie Institution of Washington Publication Number 435. Nearly all refinements and expansions of the technique of tissue culture have taken place with warm-blooded animals as the experimental material. This line of development has, no doubt, been followed because of its possible medical application. Technical difficulties extending this method to invertebrates, where the necessary asepsis is more difficult to attain, have also played a part. The writer developed a technique which was successfully applied to some tissues of eleven species of marine animals belonging to seven phyla. In all cases, both migration and cell multiplication were obtained. Two organisms seemed to offer particularly favorable material for tissue culture. One was the gastropod Astroea longispina; the other was the enteropneustan Ptychodera bahamensis. This being the most convenient material with which to work, investigations in 1932 were confined to the tissues of this species alone. The technique of a method, using either hexyl-resorcenol or ultraviolet radiation in amounts harmless to the tissues, for growing in vitro the cells of marine invertebrates is described. Because of their structure, members of the Enteropneusta lend themselves especially well to the obtaining of explants composed of one or of several types of tissue. The growth and reproduction of cells from the caecal portions of the intestine are recorded in detail. The changes undergone by muscle cells when removed from the body of an animal show a characteristically reversible series of stages peculiar to this type of cell. The bearing of the observations on Ptychodera cells to broader problems of cytology is considered . . 1915. The influence of the marginal sense organs on functional activity in Cassiopea xamachana. Proceedings of the National Academy of Science 1: 611-16. The influence of sense organs (nervous system) on the rate of regeneration was examined at the Dry Tortugas using the disks of the rhizostomous medusa Cassiopea, which can be 11 separated from the oral arms and kept in dishes of seawater for an indefinite period. Pairs of disks were examined from which all of the thopalia were removed, while from the other equal amounts of tissues were removed from the bell margin between the thopalia. In all instances, the disks where the half on which the thopalia remained regenerated at a more rapid rate than the inactive half. Other experiments focused on influence of sense organs on the rate of metabolism as measured by production of carbon dioxide. Carbon dioxide produced was always greater for the normal disk containing sense organs. It was concluded by the author that in this type of experiment there is some other form of metabolic activity which is of greater importance as a source of CO) and which is more directly under the influence of the sense organs than is the activity of the muscular system. 45. ———. 1916. The influence of the marginal sense organs on the rate of regeneration in Cassiopea xamachana. Journal of Experimental Zoology 21, no. 1: 1-31. Studies were conducted on accepting the view of the direct or indirect influence of the nervous system on regeneration in Cassiopea xamachana collected from the Fort Jefferson moat at Dry Tortugas. Experiments conducted to determine the influence of sense organs on the rate of regeneration were inconclusive, when testing entire disks with sense organs removed, compared with specimens where the sense organs remained because of wide differences in physiological activity between different individuals. Half disks with sense organs regenerated more rapidly than those half disks without sense organs. Other experiments involving electrical stimulus by induction shocks on disk halfs, with and without sense organs, indicated regeneration is faster in the activated half, than from the inactive disk. These experiments indicate the rate of regeneration is simply one expression of the general metabolic activity of an animal, and as such is subject to the influence of the nerve centers, as are many of the functional activities. 46. 1914. Observations upon the growth-rate and ecology of gorgonians. Papers Tortugas Laboratory 5: 79-90. Carnegie Institution of Washington Publication Number 182. This report provides a record of observations extending over a 3-year period on the growth rate of Gorgonia flabellum and Plexaura flexuosa on the reefs around the Dry Tortugas, Florida. Ecological observations are supplemented by observations made in Jamaica. For effective attachment of the planule, the presence of depressions or cracks into which the planule could settle appears to be the most important factor. In comparison with young coral polyps the gorgonian colony has an obvious advantage, in that is most rapid growth is perpendicular to the surface, which permits its most rapidly growing part to secure food and oxygen. Wave action during very severe storms is by far the most destructive agent to which Gorgonia are subjected. It appears that the greatest destruction by storms comes from the tearing of the Gorgonia colonies from the substrate rather than laceration of tissue. 47. 1917. Studies on the physiology of the nervous system of Cassiopea xamachana. Papers Tortugas Laboratory 11: 121-70. Carnegie Institution of Washington Publication Number 251. In this paper are gathered the results of several distinct lines of experimentation. They deal with some phase of the physiology of the nervous system of Cassiopea and represent portions of a general program of research on the nervous system of the lower animals. On account of its ability to live under adverse conditions and to withstand practically any type of operation, Cassiopea is an especially favorable form for experimentation and has been used as a subject for many researches. The experiments with entire disks, when the rates of regeneration of specimens on which the sense-organs remained are compared with those of specimens from which all sense-organs are removed, are inconclusive because of wide differences in physiological activity between different individuals. When we compare the 12 insulated halves of a disk, on one of which the sense-organs remain, while all of them have been removed from the other half, it is found that the half-disk with sense-organs always regenerates most rapidly. When all the sense-organs are removed from a disk and the halves insulated, the regeneration is faster from the activated than from the inactive half- disk. These experiments indicate that the rate of regeneration is simply one expression of the general metabolic activity of an animal, and as such is subject to the influence of the nerve-centers, as are many other functional activities. Briefly summarized, the results of the observations made on the starved Cassiopea are as follows: In general the smaller Cassiopea loses relatively more in weight than does the larger Cassiopea. The percentage of water found in the entire body is nearly the same in all sizes of Cassiopea. The nitrogen-content of the entire body is higher in the small than in the larger Cassiopea. However, the absolute amount of nitrogen found in the starved Cassiopea is considerably higher than in the normal having the same bodyweight. The loss in weight of the different parts in the starved Cassiopea remains the same proportionately to those in the normal Cassiopea. 48. . 1918. A study of respiration in Alcyonaria. Papers Tortugas Laboratory 12: 185-91. Carnegie Institution of Washington Publication Number 252. Although the respiration of many species of invertebrates has been studied, the only references to that of Alcyonaria are those given by Montuori (1913), who studied two species, Alcyomeum pallidum and Gorgonia cavolinii. In these experiments the total weight of the colony was taken as the basis of comparison without taking into account the proportion of inert skeletal material- the spicules in the first species and the spicules and chitinous axis in the latter. The observations recorded were made as part of a study of the ecological factors determining the distribution of Alcyonaria on the coral reefs of southern Florida. All the species of the genus Gorgonia and the closely related Xiphigorgia, which have as a group the highest rate of respiration, are next to Briareum the most resistant to increased temperature. Taken all together these observations indicate that some other factor is the controlling agency in the ability of a marine organism to withstand high temperatures. The acidity of the water at the close of the heat experiments was always greater than in respiration experiments carried on at 27.5° C. This may be only an expression of the abnormality of their metabolism at high temperatures, or have a causal relation to the death of the organism. 49. Cate, C. N. 1978. New species of Ovulidae and reinstatement of Margovula pyrulina (A. Adams, 1854) (Gastropoda). Nautilus 92 , no. 4: 160-167. Eight species of living Ovulidae are described as new, and the species M. pyrulina is reinstated. The 8 new species are listed as follows: Prionovolva castanea from the Gulf of Oman; Aperiovula testudiana from Mukaishima, Japan; Primovula santacarolinensis from Mozambique; P. uvula from Moreton Bay, Queensland, Australia; Crenavolva periopsis from Java, Indonesia; Speculata advena from off Sand Key, Florida; Cyphoma rhomba from Fort Lauderdale Reef, Florida; and Psudocyphoma gibbulum from off the Dry Tortugas Islands, Florida. 50. Chambers, E. L. 1937. The movement of the egg nucleus in relation to the sperm aster in the sea- urchin, Lytechinus varigeatus. Carnegie Institution of Washington, Year Book 36: 86-87. With the aid of a camera, 30-second observations on the positions (rate and direction of movement) of the egg nucleus and sperm aster of Lytechinus were made. 51. Child, C. A. 1992. Shallow-water Pycnogonida of the Gulf of Mexico. Memoirs of the Hourglass Cruises 9, no. 1: 1-86. This paper treats 11 species in 8 genera of the Pycnogonida that were collected during the Hourglass Cruises, a sampling program conducted on the central West Florida Shelf for 28 13 months during 1965-1967. Five benthic stations in depths from 6 to 72 m were sampled monthly with dredges and trawls among each of two transects. Treatments of 20 more species in 6 additional genera from other shelf collections are also included to offer a comprehensive survey of species (a total of 31 species in 14 genera) known from the continental shelf of the Gulf of Mexico, excluding the Dry Tortugas and the Florida Keys. Three of these species were previously unreported from the Gulf. Two new species, Ascorhynchus crenatum and A. horologium, are described from the Hourglass material, and an additional new species, Anoplodactylus dauphinus, is described from the other material. Artificial taxonomic keys are provided for all Gulf of Mexico families and species, and checklists are provided for all species known or expected to occur in the Gulf. All species are diagnosed and illustrated, and their distributions are given. Only four species were taken during the Hourglass Cruises with sufficient frequency to allow analysis of their distributions and abundances. 52. Clapp, R. B. and W. B. Robertson Jr. 1986. Nesting of the masked booby Sula dactylatara on the Dry Tortugas, Florida. The first record for the contiguous United States. Colonial Waterbirds 9, no. 1: 113-16. In both 1984 and 1985 masked boobies (Sula dactylatra) attempted to nest on sandy islets at the Dry Tortugas, Florida. Nesting attempts failed because the nest sites were washed away by summer storms. It seems likely that this species will eventually nest there successfully and will establish a small breeding population. This is the first documented nesting by this species in the contiguous United States. 53. Clark, H. L. 1919. The distribution of the littoral echinoderms of the West Indies. Papers Tortugas Laboratory 13: 49-74. Carnegie Institution of Washington Publication Number 281. The purpose of this investigation was to determine if the distribution of littoral echinoderms varied among various northern West Indian islands bounded by Bermuda to the North, Tobago to the South, and the Tortugas to the west. Five classes of echinoderms are discussed, including Comatulidea (feather-stars), Asteroidea (sea-stars), Ophiuroidea (brittle-stars), and Echinoidea (sea urchins). The number of species and numbers of individuals of the classes are discussed. Of the island areas investigated, the Tortugas appears to be the richest in the number of species found with 76 littoral echinoderms, with 70 available to collect by hand. The sea urchins, Echinometra lacunter were reported to be excessively abundant on many reefs, actually occurring by the thousands. The number of echinoid species found throughout the region and Florida are compared with comments provided on the origin of echinoids in the West Indies. Next to brittle-stars, holothurians were considered to be the most abundant of the littoral echinoderms. 54. Clark, L. B. and W.N. Hess. 1942. Swarming of the Atlantic palolo worm, Leodice fucata (Ehlers). Papers Tortugas Laboratory 33: 21-70 (issued Oct. 1940). Various organisms show reproductive activity coinciding with the lunar cycle. At the Dry Tortugas, swarming observations were recorded during 1937-39 on the Atlantic palolo worm, in association with the quarter-moon phase. Other important factors determining the time when the worms reproduce, include the maturity of the animal and the amount of water turbulence. It was concluded that: (1) the stimulating effect of the first-quarter moon is less than that of the third-quarter, (2) the worms increase in sensitivity to the stimuli inducing the swarming as they become sexually mature, and (3) wave action and water turbulence above a certain level induced by an 8 mph wind decreases or prevents swarming at the Dry Tortugas. 14 a5: 56. 57, 58. 52: 60. 61. Cole, L. J. 1906. Ant Studies. Carnegie Institution of Washington, Year Book 5: 110. To investigate the biology of ants at the Tortugas, specimens were collected, and observations were made. Collie, M. R. 1979. A Sabine's gull at the Dry Tortugas. Florida Field Naturalist 7, no. 2: 28. A photographic observation was made by the author during August 1978 of a Sabine's gull at Garden Key, Dry Tortugas. There are four other records of this species in Florida along the Atlantic coast, however this sighting represents the first record of the Sabine's gull at the Tortugas. Colman, J. 1931. The superficial structure of coral reefs: animal succession on prepared substrata. Carnegie Institution of Washington, Yearbook 30: 395. Plant and animal successions were examined on concrete cubes planted in the water at three sites: Fort Jefferson moat, an iron wreck east of Loggerhead Key, and northwest of loggerhead Key. Also, a detailed ecological survey of Long and Bush Keys was made. Conger, P. S. 1924-1935. Diatom studies. Carnegie Institution of Washington, Year Book. Note: published as: 1924, V. 23, p. 220; 1925, v. 24, p. 221; 1926, v. 25, p. 240; 1927, v.26, p. 220; 1928, v. 27, p. 271; 1929, v.28, p. 283; 1930, v. 29, p. 323. Diatom studies were conducted in association with A. Mann. Narrative same as in reference no. 218. Conklin, E. G. 1908. The habits and early development of Linerges mercurius. Papers Tortugas Laboratory 2: 153-70. Carnegie Institution of Washington Publication Number 103. The jellyfish, Linerges mercurius, Scyphomedusa, was investigated at Tortugas and Nassau Harbor. The sudden appearance in great numbers of this species at Tortugas was noted, followed by their rapid disappearance. Normal movements of the medusae are described as well. Other phases of early development, including egg-laying, egg structure, maturation and fertilization, first cleavage, second, and later cleavages, and blastula, gastula, and planula are described. Experiments on isolation of blastomeres and centrifugalized eggs are presented. The organization of the egg of Linerges and mechanics of cell division are described. . 1908. Two peculiar actinian larvae from the Tortugas, Florida. Papers Tortugas Laboratory 2: 171-86. Carnegie Institution of Washington Publication Number 103. During middle-of-the-day sampling tows at the Tortugas in May 1905, two peculiar larvae were collected. They did not undergo metamorphasis in aquaria. Natural history notes on living larvae are provided. Based on literature description, they were probably Zoanthidae of the Order Hexactinia. A band of strikingly brilliant, locomotor cilia was noted as most peculiar for these larvae. Their size, shape, and coloration are described. Yellowish-green symbiotic algae occur in both types and are hypothesized to be associated with their metabolism and play an important role in their nutrition. The morphology and histological character of these two types are similar but minor differences are described. Although these types have only been collected a few times world-wide, they are not considered rare at Nassau and the Dry Tortugas. Coonfield, B. R. 1940. Chromatophore reactions of embryos and larvae of Pomacentrus leucostictus. Papers Tortugas Laboratory 32: 169-78 (issued Sept. 1940). Carnegie Institution of Washington Publication Number 517. Interest in the origin of the changing color patterns of fishes and certain other vertebrates, together with certain questions that have been raised by the work of investigators in their 62. 15 study of this problem in embryos of vertebrates, prompted an investigation of the color mechanism in developing fishes. Both embryos and larvae of Pomacentrus leucostictus, which is found in abundance in the Dry Tortugas, were used in this study. It was concluded that melanin granules migrate within the melanophores of Pomacentrus embryos as soon as these pigmentary bodies are completely formed. The melanophores of embryos a few hours of age contract in response to pressure applied with forceps and to a temperature of about 8°C. The melanophores of a majority of developing embryos, from their beginning up to a few hours before hatching, are found to be in a stellate state regardless of whether these young are over a white or a black background or are in total darkness. A few hours before hatching, the melanophores contract when the embryos are over a white background, expand when they are over a black background, and contract when they are in total darkness. This response continues in these young on through the hatching period and for a few hours after hatching. Larvae of two or more days after hatching do not show any conclusive response to different backgrounds or to the absence of light. The eyes of these young fish are believed to have no function in controlling their melanophore responses. The evidence is in favor of the release of a hormone within the capsule just before the embryos hatch. This agency either permits or directly causes the melanophores to respond to various environments. . 1940. The chromatophore system of larvae of Crangon armillatus. Papers Tortugas Laboratory 32: 121-26 (issued May 1940). Carnegie Institution of Washington Publication Number 517. The ability of certain animals to imitate the color in their background 1s so striking that is has received the attention of investigators for a considerable period of time. This feature has been observed principally in fishes, amphibians, and reptiles of the vertebrates, and in crustaceans of the invertebrates. This paper adds to this field of study the results of observations on the reactions of the chromatophore system of larvae of Crangon armillatus. The erythrophores of normal larvae of Crangon armillatus react as follows according to different backgrounds: the pigment is dispersed when the animals are kept in a white illuminated bowl. The erythrophores of enucleated specimens show the following conditions when subjected to different backgrounds: the pigment becomes concentrated when the specimens are kept over a white illuminated background; the pigment is dispersed when these specimens are subjected to black illuminated bowls. The time required for concentration of pigment in the erythrophores is much longer than that required for its dispersion. Ablation of the eyes permits the erythrophores to react directly to stimulations caused by different backgrounds . 63. Coutiére, H. 1910. The snapping shrimps (Alpheidae) of the Dry Tortugas, Florida. Proceedings of the United States National Museum 37: 485-87. The Alpheidae collected by Dr. McClendon at the Tortugas in 1908 are discussed. The Alpheidae are referable to eight different forms, including one new species and one new subspecies: Alpheus formosus Gibbes. Alpheus cristulifrons Rathbun and Alpheus armillatus H. Milne-Edwards. 64. Cowles, R. P. 1908. Habits, reactions, and associations in Ocypoda arernaria. Papers Tortugas Laboratory 2: 1-41. Carnegie Institution of Washington Publication Number 103. On Loggerhead Key, investigations were made on the behavior of Ocypoda arernaria. It was found that adult ghost crabs build two kinds of burrows. One consists of a single tunnel extending down in the sand for 3 to 4 feet. The other is similar, except that it is shorter and has a passage branching off from it, which is used for escape. Young ocypodas make short burrows, only a few inches long, which often extend vertically downward. Breeding in the region of Loggerhead Key probably occurs in the spring and early summer. 16 65. Ocypoda is a scavenger and a cannibal. The eyes do not seem to play an important role in the detection of food, but they undoubtedly lead individuals to objects which may be food. That Ocypoda is stimulated by odors was not conclusively shown, but certain experiments point strongly in that direction. The eyes are highly developed, so far as crustacean eyes are concerned; they are quite sensitive to large differences in the intensity of light; they do not react to different colors; they aid much in the search for food, in the detection of enemies, and in the accuracy of locomotion. Ghost crabs probably do not have vision such as that of the human eye, nor do they see the color and finer characters of the surface of an object, but they undoubtedly see its outlines and possibly some of the more evident irregularities of the surface made evident by differences in lighting. The color-pattern seen through the carapace of Ocypoda changes in intensity under different conditions of temperature and light. In the absence of light when the temperature is anywhere between 22° C. and 45° C., and undoubtedly when it is even lower or higher, a light coloration occurs. Generally in diffuse light and even direct sunlight a dark coloration appears, provided the temperature is not too high. Usually at low temperatures, not above 35° C., a light coloration is the rule, and it occurs independently of the intensity of light. At high temperatures, above 35° C., a light coloration is the rule, and it occurs independently of the intensity of light. No indication of audition was observed in Ocypoda. The so-called "auditory organs” are equilibrating organs. Ocypoda has a stridulating ridge on the palm of its large chela. . 1911. Reaction to light and other points in the behavior of the starfish. Papers Tortugas Laboratory 3: 95-110. Carnegie Institution of Washington Publication Number 132. Experiments were designed to test the reactions of starfish to light. Two species were used Echinaster crassispina and Astropecten duplicatus. Both are migratory and are found in open waters over sandy bottoms, in areas generally exposed to light. In the Tortugas laboratory, starfish were placed in aquaria wooden boxes and tested for movement up in response to light, inclines, vertical walls, and tilted floors. Every specimen reacted positively, moving toward bright light. Even with eye-spots removed, movements towards light are positive, but not as quick as in normal individuals. When tested in different degrees of water temperature the reaction to light was positive at ordinary temperatures. Quality of light was tested using various color screens (UV, violet, blue, green, yellow, orange and red) in a box with closed and open ends. The source of light was sunlight. Ina series of 10 tests with varied orientation and handling, in nearly every test starfish moved toward light without hesitation. 66. Criales, M. M. and T. N. Lee. 1995. Larval distribution and transport of penaeoid shrimps during the Tortugas Gyre in May-June 1991. Fishery Bulletin 93: 471-82. As part of the Southeast Florida and Caribbean Recruitment (SEFCAR) project, penaeoid shrimp larvae were collected during the spring and summer cruise of the RV Longhorn in the Lower Florida keys and Dry Tortugas from 29 May to 30 June 1991. Larvae of the pink shrimp, Penaeus duorarum, and the rock shrimp, Sicyonia sp., were distributed inshore close to the Dry Tortugas Grounds adjacent to the boundaries of Dry Tortugas National Park, whereas larvae of the oceanic shrimp Solenocera sp. showed mainly an offshore distribution. Significant concentrations of Solenocera sp., Sicyonia sp. and P. duorarum larvae at the Tortugas transect in early June were found within and above the seasonal thermocline, while the cold cyclonic Tortugas Gyre was intensively developed. For Solenocera sp., which spawn on the outer ridge of the gyre followed by onshore Ekman transport. Penaeus duorarum, which spawn in the shallow Tortugas Grounds, showed a mode of zoea II-III progressing to postlarvae I at the Tortugas Grounds during the 15 days in which the drifter Halley recirculated in the interior of the Tortugas Gyre. Retention of P. duorarum larvae by the internal circulation of the gyre at the spawning 1, grounds may be an important mechanism for local recruitment of these shrimp to the nursery ground of Florida Bay, Everglades National Park. 67. Cushman, J. A.. 1922. Shallow-water Foraminifera of the Tortugas region. Papers Tortugas Laboratory 17: 1-85. Carnegie Institution of Washington Publication Number 311. The paper gives the results of a study of collections made in the waters about the Tortugas Laboratory of the Carnegie Institution of Washington. Collecting was done largely from the boats, the most satisfactory method that was used with the Darwin. Collecting in the moat at Fort Jefferson in shallow water on Long Key, as well as on the reefs and flats, was done by hand. The Tortugas region presents an ideal spot for studying the shallow-water tropical Foraminifera of this particular region. It is removed from influence of shore conditions; the water is at all times warm and pure, so that ecological conditions that are present are constant. The twenty stations from which bottom samples were studied in the preparation of this paper, together with collections from reef flats and from the eel-grass, give a considerable range of conditions. The only stations at which Rotalia was found are two in the moat at Fort Jefferson and the other in a very shallow lagoon at Long Key, where the water was warm at low tide in June. By most authors, these specimens would ordinarily be referred without question to Rotalia beccarii (Linnaeus). There are differences from northern material, and in probability the Tortugas specimens belong to different species. On the banks of dead coral which become exposed at spring tides, great masses of attached Foraminifera develop. Of these, the most abundant is Homotrema, which makes an appreciable contribution to the mass of material. With it, in crevices of the dead coral, was a new species of Haliphysema. On the eel-grass (Posidonia), which forms in shallow water inside the reef, there is Jridia, Planorbulina, Discorbis, Orbitolites, with a peculiar miliolid which spreads over the surface. The mass of these must add appreciably to the amount of carbonate of lime added to the bottom. The forms are rapid in their growth, as the leaves of Posidonia are quickly covered in their growth by Foraminiferea and other encrusting animals. 68. Cutright, P. E. 1937. Studies on the development of the dorsal spine of sting rays. Carnegie Institution of Washington, Year Book 36: 90. This report describes the collection of southern sting rays for a histological examination of the stinging mechanism. 69. Dall, W. H. 1889. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico 1877-78 and in the Caribbean Sea (1879-80), by the U.S. Coast Survey Steamer "Blake":, Lieutenant Commander C.D. Sigsbee, U.S.N., and Commander J.R. Bartlett, U.S.N. Commanding. XXIX- Report on the mollusca . Part II Gastropoda and Scaphoda. Bulletin of the Museum of Comparative Zoology at Harvard College 18: 1-492, with thirty one plates. This listing of Mullusca collected by the “Blake” is supplemented by the southern dredgings of the U.S. Fish Commission Steamer "Albatross" and other material collected from the region. A systematic description and account of the gastropods and scaphopods is given and illustrated. Nomenclature is discussed and rectified in several cases. 70. Darby, H. H. 1934. The mechanisms of asymmetry in the Alpheidae. Papers Tortugas Laboratory 28: 347-61 (issued Feb. 1934). Carnegie Institution of Washington Publication Number 435. In 1901, Przibram reported a series of striking experiments on the regeneration of chelae of Alpheus dentipes, A. platyrhynchus and A. ruber. There is a pronounced quantitative and qualitative difference between the right and left chelae. One chela is several times as large as the other. Przibram showed that if the snap-claw is removed, at the next molt the pinch- 18 Hil claw is changed into a snap-claw and a new pinch-claw is regenerated on the stump of the old snap-claw. This unusual reversal of asymmetry was confirmed by Wilson (1903) and Zeleny (1905). It was also shown that if both chelae are removed at the same time, they regenerate in their original positions. These experiments seem to indicate that the final degree of morphological expression of the gene may in certain cases depend on the environment. In Crangon armillatus, an asymmetrical individual, it is symmetrical ten days later. The morphological expressions of the gene are so concrete that it is difficult to realize that the gene may also be the controlling agency in the production of definite chemical substances, whose presence is manifested only by their physiological reactions. Environment from that point of view can quite easily be thought to control the amount of a substance produced. Changes might well be induced by radiation, due to the ionization of the cell. Crangon has shown itself to be an organism in which studies on development as an expression of the activity of the gene can be undertaken with some hope of success. The nature of the regeneration of chelae in two members of the family of Alpheidae has been studied; in particular, in Crangon armillatus. It has been shown that at certain stages in the development of the chelae, a state is reached that permits the determination of which side is to have the large chela, or snap-claw. Equal chelae have been produced experimentally and are of three varieties: (1) both small (pinch-claws); (2) both large (snap- claws); (3) both intermediate . . 1940. Symmetry in normally asymmetrical crustacea. Papers Tortugas Laboratory 32: 61- 64 (issued Oct. 1939). Carnegie Institution of Washington Publication Number 517. A symmetrical specimen of Crangon armillatus was found in nature with two snap claws. These claws differed in no way from snap claws produced experimentally and reported previously. 72. Darby, H. H., E. R. F. Johnson and G. W. Barnes. 1937. Studies on the absorption and scattering of solar radiation by the sea: spectrographic and photoelectric measurements. Papers Tortugas Laboratory 31: 191-205 (issued Oct. 1936). Carnegie Institution of Washington Publication Number 475. A considerable amount of work has been done in recent times on the penetration of radiation into sea and lake water. The importance of this work, in such matters as plant and animal metabolism and under-water photography, is obvious. The amount and spectral distribution of scattering, and the penetration of the ultraviolet component, are two phases of the subject which have received scant attention. These studies were made from the yacht, Elsie Fenimore at the Tortugas Laboratory. A comparative study has been made of two methods of evaluating the transmission of various wave lengths of light through sea water: (1) photometry by means of photoelectric cells, and (2) photographic spectrophotometry. Bertel's observation that ultraviolet light penetrates a considerable distance into the sea has been confirmed. The extent of penetration is greater than would be expected from the laboratory data of Hulbert and Sawyer. A rough evaluation of the transmissive exponent from 4500A to 3250A was made, which indicates the magnitude of the disagreement. Scattering was found to be selective, becoming greater with decreasing wave length. The spectral distribution of scattered radiation is indicated. The importance of these observations for biological systems is outlined. 73. Davis, G. E. 1977. Anchor damage to a coral reef on the coast of Florida. Biological Conservation 11: 29-34. Twenty percent of an extensive staghorn coral Acropora cervicornis has recently been damaged by boat anchors in Fort Jefferson National Monument, Dry Tortugas, Florida. It is suggested in this article that this type of damage may occur in other coral reef sanctuaries 74. De 76. 19 unless anchor-sensitive areas are identified and closed to anchoring. Alternatively, mooring buoys should be provided by sanctuary managers. . 1982. A century of natural change in coral distribution at the Dry Tortugas, Florida USA. A comparison of reef maps from 1881 and 1976. Bulletin of Marine Science 32, no. 2: 608- 73. Changes in coral reef structure and composition at Dry Tortugas, Florida were compared over a 95-year interval from benthic maps prepared in 1881 and 1976. Living hermatypic corals occupied less than 4% of the 23,000-hectare area mapped, and showed little change in area during the interval between maps. However, major changes in coral species distributions and reef types were apparent. In 1976, a lush 220-hectare Acropora cervicornis reef occupied what had been octocoral dominated hard bottom in 1881. The 44-hectare swath of A. palmata on the reef crest in 1881 was reduced to two small patches totaling less than 600 m* in 1976. More than 90% of the extensive thickets of A. cervicornis at Dry Tortugas were killed during the winter of 1976-77, apparently as a result of thermal shock. These changes in coral distribution and abundance demonstrated the natural dynamic nature of coral reefs, and showed the important role occasional short-term extreme climatic events can play in shaping coral reef structure and species distribution. The importance of protecting living corals and the value of ecosystem level sanctuaries as dynamic standards are discussed. . 1977. Effects of recreational harvest on a spiny lobster, Panulirus argus population. Bulletin of Marine Science 27, no. 2: 223-36. A commercially unfished population of Panulirius argus was studied in Fort Jefferson National Monument at Dry Tortugas, Florida, from April 1971 to July 1975. For 29 months all harvest was prohibited, then an experimental sport harvest (hand caught by recreational divers) was allowed in 50% of the area for a period of 8 months, followed by 16 months of complete protection for assessment of recovery. Data on the size, abundance, and natural history of the lobsters were collected using SCUBA, and commercial trapping techniques. A total of 4,257 lobsters, with a mean carapace length of 101 mm, was tagged and released at Dry Tortugas. The existence of a resident adult P. argus population was demonstrated by the recovery of all recaptured lobsters (7.3%) with 10 km of their respective capture sites up to 104 weeks after release. Immediately following the experimental sport harvest, the population in the sport harvested area showed a 58% reduction in trap catch rate and dispersed to 42% of its pre-harvest lair occupancy density, while the population in the unharvested control area remained essentially unchanged. The catch rate in the sport harvested area recovered to 78% of its pre-harvest level after 1 year of complete protection from harvest, and the lair occupancy rate recovery was 71% after 16 months of post harvest protection. The pre-harvest standing crop was estimated at 58.3 kg/ha, wet weight. . 1977. Fishery Harvest in an Underwater Park. Proceedings, Third International Coral Reef Symposium, 605-8 no. 2. RSMAS, Univ. Miami, Coral Gables, Florida. There is a potential conflict between park management for preservation of maximum species richness and fishery harvest in parks. The recreational harvest of spiny lobster, Panulirus argus, at Ft. Jefferson National Monument, Dry Tortugas, Florida, demonstrates the nature and extent of the conflict. An eight-month-long diver harvest, limited by a daily bag limit of two lobsters, reduced the previously unfished population by 585 and significantly altered the local lobster distribution. Growth and natural recruitment did not restore the population to its 58.3 kg/ha pre-harvest level, even after 16 months with no additional harvest. The trophic status of spiny lobsters as high level carnivores and current ecological theory combined with the harvest impact observed at Dry Tortugas suggests that community structure and species richness would be significantly altered by the harvest. 20 77. 78. 19) 80. 81. . 1975. Minimum size of mature spiny lobsters, Panulirus argus, at Dry Tortugas, Florida USA. Transactions of the American Fisheries Society 104, no. 4: 675-76. Of 1,594 female spiny lobsters examined during April 1973-1975 at the Dry Tortugas, 55% were bearing eggs (berried). The specimens ranged in carapace length from 39 mm to 140 mm. No berried females were found with carapace lengths less than 78mm. Maturity was reached by one half of the females in the 86-95 mm size class. The current minimum legal size for sport and commercial lobster fishing in Florida is 76-mm carapace length. . 1974. Notes on the status of spiny lobsters, Panulirus argus, at Dry Tortugas, Florida, SUSF-SG-74-201. State University (Florida) System. Sea Grant Program. Publ.. Until mid-1971, sport harvest of spiny lobsters, primarily Panulirus argus, was permitted in the 19,000 hectares underwater preserve created in 1935 which included the Dry Tortugas atoll. At that time there was a two lobster per person per day limit. Few visitors reached the isolated atoll during the first 20 years, with an average of some 1,200 people per year. Annual visitation increased to over 21,000 in the late 1960's and early 1970's. Concern was expressed for the protection of the quality and quantity of the lobsters found in the area. The primary objective of the study was to assess the impact of human harvest on a natural unperturbed lobster population. . 1981. On the role of underwater parks and sanctuaries in the management of coastal resources in the southeastern United States. Environmental Conservation 8, no. 1: 67-70. Aquatic resources in parks and reserves are not as adequately protected as comparable terrestrial resources. Thus the values of protected aquatic ecosystems as standards for comparison, reservoirs of genetic materials, and ‘emotional’ reserves, are apt to be greatly diminished. Even seemingly static ecosystems such as coral reefs are dynamic, changing dramatically in response to natural short-term environmental variations. Such ecosystems require protected natural areas as dynamic standards that will allow distinctions to be drawn between effects of exploitation or pollution and normal variation. Furthermore, fisheries harvests may reduce the size at which exploited species mature, and reduce the amount and variability of genetic material produced by exploited populations. The seven underwater parks or sanctuaries established since 1935 (Dry Tortugas) in Florida and the U.S. Virgin Islands exhibit wide variations in the degree of protection accorded to aquatic resources, a range being apparent from nearly complete protection in the first parks to be established to virtually no protection at all in the recently established parks. The consequences of permitting consumptive uses of aquatic resources in parks and reserves need to be objectively evaluated. Unless these consumptive uses are severely curtailed or eliminated, the primary values of the parks and reserves may never by realized. . 1980. Spiny lobster series. Gary E. Davis (ed.), 27 pgs. American Fisheries Society: Bethesda, MD. This series of papers regarding spiny lobster management represents the efforts of a broad cross section of the scientific fisheries community. Not only is there a diverse array of disciplines from biochemical genetics to ecology and economics, but nearly every source of research endeavor is represented. Members of two federal agencies (National Marine Fisheries Service and National Park Service), a state agency (Florida Department of Natural Resources), a public university (University of Florida), a private university (Nova University), and a private company (Science Applications, Inc.) have combined their efforts on a common subject that has already spawned thousands of scientific papers and countless popular articles. Davis, G. E. and J. W. Dodrill. 1989. Recreational fishery and population dynamics of spiny lobsters, Panulirus argus, in Florida Bay, Everglades National Park, 1977-1980. Bulletin of Marine Science 44, no. 1: 78-88. 21 Florida spiny lobsters Panulirus argus, occupied the southern two-thirds of Florida Bay in Everglades National Park. Field studies of 3,570 tagged lobsters revealed that they pass through Florida Bay, using it for less than three years as juveniles, between their planktonic larval stages in the open ocean and adulthood on coral reefs. Lobsters from the bay support commercial and recreational fisheries outside of Everglades National Park from Dry Tortugas to Pacific Reef near Miami. Growth rates of juvenile lobsters in Florida Bay are the highest on record, which may be a reflection of optimum habitat with abundant food and shelter. 82. Davis, G. E. and J. W. Dodrill. 1980. Marine parks and sanctuaries for spiny lobster fisheries management. Proceedings of the Gulf and Caribbean Fisheries Institution, pp.194-207, 32nd Annual Session. National parks and sanctuaries with significant marine resources can play important roles in effective fisheries management. However, if fishery resources are exploited and not protected to the same extent terrestrial resources are protected in parks and sanctuaries, they may not be available to provide the dynamic standards for comparison, reproductive/genetic reserves, unique educational opportunities, and recreational escape. Observations of more than 15,000 specimens of P. argus tagged at the National Park were analyzed to provide data on migration patterns, natural mortality, reproduction and development. Main factors affecting these populations were seasonality, stress in juveniles, and sexual proportions in adults. Studies in non-exploited populations gave good estimates of natural mortality. Size at first maturation was greater in non-exploited populations than in exploited populations. Juveniles of P. argus show an extensive directional migration pattern of 200 KM, while adults exhibit a restricted pattern for about two years. Returned tags during the 1977-78 season in Florida came from sports fisherman (49%), from commercial fisherman, (51%), and commercial traps (11%). This return proves that sports catches were only 9% of the total in the northern part of the Florida Keys (if all the tags were reported). The average of lobsters escaping from traps that were never recovered was 1.2% daily, during the fourteen days that these were in operation. 83. Davis, J. H. Jr. 1940. The ecology and geologic role of mangroves in Florida. Papers Tortugas Laboratory 32: 303-412 (issued Sept. 1940). Carnegie Institution of Washington Publication Number 517. The mangrove swamps of the low-lying coasts and islands of central and southern Florida were studied during five seasons to determine the ecology of these unique littoral swamps, and to obtain some idea of their importance as geologic agents in extending the coasts and forming islands in the shoal-water regions. Five coastal and insular regions of the peninsula were selected and a number of stations established in each for observations and experimental studies. The report is divided into two parts. The "Ecology of the Mangroves" is concerned with the types of plant communities of the mangroves and associated vegetation, and the successional relationships of some of these communities. "The Geologic Role of the Mangroves" considers the accretions of sedimentary and cumulose soils in connection with the different agents that bring them about, and more significantly, the role of the different mangrove communities in forming soils at higher and higher levels. The most apparent succession of the mangrove communities consists of a pioneer Rhizophora, a mature Rhizophora consocies, an Avicennia salt-marsh associes, not always, flooded by salt or brackish water, a Conocarpus transition associes, seldom if ever flooded by water, and a tropical or semitropical forest association, which is the actual climax of the region. Besides Rhizophora, the Tortugas Keys have a young swamp of Laguncularia around a pond on Bush Key, some young plants of Avicennia on Bush Key, and a few old ones on Garden Key. Concocarpus was established on both Garden Key and Bush Key. How these species got to the Tortugas and to many of the most isolated of the Florida Keys is not certain, but should be considered. 727) 84. . 1942. The ecology of the vegetation and topography of the Sand Keys of Florida. Papers Tortugas Laboratory 33: 113-95 . Carnegie Institution of Washington Publication Number 524. This is a study of the vegetation and some of the physiographic features of about thirty islands of the Florida Keys, in an area extending west from Key West, Florida, and including the Dry Tortugas Keys. About thirty islands of the Florida Keys beyond Key West were investigated during the summer of 1940 and winter of 1942, and to some extent during the summers of 1937 and 1938. These studies were concerned with the topography and vegetation of these small, relatively isolated, and partly tropical islands. These islands are here termed the Sand Keys because most of the parts above high tide are composed of coarse calcareous sands, and also because this name was used by Willspaugh (1907). A few of the Marquesas and Tortugas Keys have changed a great deal. The strand areas on a number of islands seem to be increasing at the expense of the mangrove swamps. The mangrove swamps have spread over wide areas in some instances and seem to be aiding in building up the islands. Most of the constructional processes are, however, due to maritime factors such as the ocean currents and tides. This paper is also a part of a series of studies of the plant ecology of southern Florida. This and the author's study of mangrove vegetation together describe most of the coastal and insular vegetation of that region. 85. Davis, R. A. Jr. and C. W. O'Neill. 1979. Morphodynamics of East Key, Dry Tortugas, Florida. in Guide to Sedimentation for the Dry Tortugas, Fort Jefferson National Monument Florida Southeast Geological Society Publication 21: 7-13. East Key is comprised wholly of biogenic sand and fine gravel. It lacks beachrock or bedrock which may act as a stabilizing agent such as on Loggerhead Key. During the past two centuries, maps and charts documented the size, shape, and location of East Key. The Key moved in a generally southeasterly direction across the shallow carbonate bank. East Key was preserved, unlike some other islands, because of its easterly position with respect to the deep lagoon. Those islands west of the lagoon moved easterly and disappeared into the lagoon (O'Neill, 1976). East Key has decreased markedly in size during its southeasterly movement. In addition there appears to be a change in morphology which is related to seasonal changes in predominant wind direction. 86. de Laubenfels, M. W. 1936. A discussion of the sponge fauna of the Dry Tortugas in particular and the West Indies in general, with material for a revision of the Families and Orders of the Porifera. Papers Tortugas Laboratory 30: 1-225. Carnegie Institution of Washington Publication Number 467. Sponge specimens were collected near the Dry Tortugas by scientists affiliated with the Carnegie Institution of Washington, or working at the laboratory maintained on Loggerhead Key. These were sent to the U.S. National Museum to be studied by the author. The West Indian region has long been known as one of the richest collecting grounds for sponges in the world, and the Dry Tortugas offers a representative sample of it. The author identified several new families and in many cases proposed new names for families already in use. Representatives of each of the species discussed in this paper have been deposited in the United States National Museum. Each new species is described in detail. 87. ———. 1934. Physiology and morphology of Porifera exemplified by Jotrochota birotulata Higgin. Papers Tortugas Laboratory 28: 37-66. Carnegie Institution of Washington Publication Number 435. The experimental work upon which this article is based was carried on during the summers of 1927 and 1928 at the Tortugas Laboratory. A taxonomic description of the sponge was provided. It was found that a hyaline ground mass or slime plays a very important role in the life of Iotrochota and perhaps numerous other sponges. Judging only by items visible 23 in living Jotrochota cells, which were kept track of by conspicuously colored inclusions, new sponges resulted from disassociated cells without intermediate differentiation and respecialization. Reproductive bodies (gemmules) seem to result in Jotrochota by the migration together of cells previously specialized for the purpose. Bispecific conglomerations could be secured between Jotrochota and other species, and these remained alive for two weeks or more, but whatever cell motility occurred within them tended toward the ultimate segregation of the two species after somewhat the manner in which animal gratings finally terminate. Amebocytes of Jotrochota sometimes ingested flagellates which subsequently appeared as intracellular inclusions, and perhaps became the symbionts whereby there occurred a certain amount of photosynthesis, the existence of which was indicated by experimentation. 88. . 1953. Sponges of the Gulf of Mexico. Bulletin of Marine Science of the Gulf and Caribbean 2, no. 3: 511-77. In 1948, a collection of sponges was made by the Marine Laboratory of the University of Miami in the eastern Gulf of Mexico. Twenty-two stations were studied, at depths from 6 to 20 meters, in the area between Dry Tortugas and the northeastern part of the Gulf. The collection comprises 52 species in 41 genera, all within the class Demospongea. Of these 11 species are new. Additional description is provided for a number of species. An analysis of the sponge collection by stations is included. 89. de Renyi, G. S. 1934. Studies of nerve cells of invertebrates. Carnegie Institution of Washington, Year Book. 33: 250. The nerve tissue (neuroplasm) of gastropods (Strombus gigas), Aplysia protea, Olivia litterata, Cypraea exanthema, Casio cameo), decapods (Panulirus argus, Crangon armillatus, Ocypoda albicans), and hemichordates (Ptychodera bahamensis) were studied. The neoplasm of the Gastropoda and Hemichordata exhibited viscosity, and a certain degree of elasticity, whereas decapodean neuroplasm was liquid. 90. Deflaun, M. F. 1987. "The distribution and molecular characterization of dissolved DNA in aquatic environments." University of South Florida. Ph.D. Dissertation The distribution of dissolved DNA in oceanic, estuarine and freshwater environments in southwest Florida and the Gulf of Mexico was determined by using a method for the measurement of dissolved DNA based on the fluorescence of Hoechst 33258-DNA complexes. Oceanic concentrations of extracellular DNA ranged from 0.2 to 19 decreasing as a function of distance from the shore and depth in the water column. Samples of the mucus-rich coral surface microlayer (CSM) collected on reefs in the Dry Tortugas had dissolved DNA concentrations from 1.8 to 11.7 times that in the overlying water. Estuarine concentrations, measured at three stations in Tampa Bay, FL over a 15- month period, followed the seasonal trend in concentrations in offshore environments, while variations in the estuary were significant, with maximum concentrations in nighttime samples. Although concentrations of dissolved DNA in the eutrophic Alafia River were generally higher than those in the oligotrophic Crystal River, values as low as 1.14 were measured in the Alafia. A wide range of molecular weights (determined by agarose gel electrophoresis) was found for extracellular DNA concentrated from various aquatic environments. These results indicated that dissolved DNA is in a size range sufficient to contain gene sequences, which may be important in natural transformation of microbial populations. A model system for probing extracellular DNA from aquatic environments was developed using the plasmid containing the herpes simplex thymidine kinase (TK) gene. Plasmid DNA and the TK gene fragment added to artificial seawater were concentrated and labeled TK to establish percent recovery and detection limits for the method. The degradation of plasmid DNA added to a natural seawater sample was monitored over a 36 h period by probing with the TK gene probe. Intact plasmid was 24 detected for up to 4 h and DNA hybridizable to the TK probe was detected for up to 24 h. These methods were used to probe for the TK gene in environmental samples of extracellular DNA. Hybridization to the TK probe was detected in both freshwater and estuarine samples. 91. Dinsmore, J. J. 1972. Sooty tern behavior. Bulletin of the Florida State Museum of Biological Science 16, no. 3: 129-79. A four-year study of the breeding behavior of sooty terns (Sterna fuscata) was made at Bush Key, Dry Tortugas in the southeastern Gulf of Mexico. The results are compared with the behavior of other terns and the differences discussed, particularly in regard to the pelagic environment the sooty tern inhabits. sooty terns have a lower clutch size, longer period of development of the chick, and first breed when older than most other terns, many of which feed in marshes and coastal waters. These characteristics of sooty tern breeding biology are similar to those of many other pelagic birds. A distant food supply and high adult survivorship apparently have contributed to these differences from other terns. 92. Dinsmore, J. J. and W. B. Robertson Jr. 1972. Sooty tern feeding on moths. The Auk 89, no. 2: 93. Dole, R. 440. While banding sooty terns (Sterna fuscata) at Bush Key, Dry Tortugas, Florida on June 28, 1970, an adult tern regurgitated two moths 1.5 to 2 cm long together with several unidentified fish. The moths were identified to the family Noctuidae. Although the food of sooty terns at the Dry Tortugas has not been studied in detail, sizable collections of food regurgitated show that this population feeds on fish and squid. In 13 years of tern banding, this is the first time an insect has been found as part of the sooty tern's diet . B. 1914. Some chemical characteristics of sea water at Tortugas and around Biscayne Bay, Florida. Papers Tortugas Laboratory 5: 69-78. Carnegie Institution of Washington Publication Number 182. The chemical tests at Tortugas were performed by the writer in June 1913, in the Marine Biological Laboratory, Tortugas, Florida, for the primary purpose of ascertaining what soluble effect, if any, carbon dioxide in sea-water might have on coral and other deposits of calcium carbonate. The tests of waters from Biscayne Bay were made to ascertain the differences in concentration of sea-water in the bay and the diluting effect of Miami River. The salinities of the three samples taken outside the reefs agree closely with each other and with the salinity of Gulf water at Tortugas, Florida (36.01 ppt), which is somewhat greater than that of standard ocean water (35.02 ppt.). The water in the south part of the bay is somewhat more concentrated having salinities of 36.73, 36.64, and 36.64 ppt., respectively. This evidence that the water in this part of the bay is concentrated by evaporation during its retention in the shallows serves further to indicate that circulation there is not very rapid and that the greater bulk of the water inside the keys is not thoroughly mixed or shifted by the tides. Sample 1 has a salinity obviously higher than the pure water of Miami River alone may be expected to have, and represents admixture with bay water; carbonates are absent from it, but bicarbonates are much higher than in the normal drainage from the Everglades and may be attributed to reaction of the carbon dioxide that the river water carries. 94. Domeier, M. L. Speciation in the Serranid fish Hypoplectrus. Bulletin of Marine Science 54, no. 1: 103-41. Research was conducted to determine the species status of individual color morphs of fishes in the genus Hypoplectrus (family Serranidae). Crossing two morphs of Hypoplectrus (H. unicolor x H. gema) in the laboratory produced an F1 generation with an intermediate phenotype to that of the parental types. This intermediate morph cannot be assigned to any known morph and is thus termed a hybrid. Individuals of several 25 Hypoplectrus morphs were found to select only individuals of the same morph as a mate when provided a choice. Individual fish can sometimes be forced to mate with an individual of a different morph by not providing a choice of mates. The occurrence of hybrids was found to be low in the field, corresponding to the low occurrence of mixed matings in the field. Some differences in distribution were found between the different hamlet morphs. The new data provided by this study, which includes specimens collected from the Dry Tortugas, indicate that the different color morphs warrant full species rank. It is hypothesized that speciation in Hypoplecturs was driven by the rise and fall of sea level during the last ice age. 95. Donaldson, H. H. 1916. Experiment on the feralization of the albino rat. Carnegie Institution of Washington, Year Book 15: 200-201. Domesticated albino Norway rats were released on East Key to determine if changes in brain weight occur over successive generations in a wild state. Since the rats were unmarked, it was impossible to ascertain if differences in weight were from new breeds or from animals in the original colony. 96. Doyle, W. L. 1936. Cytology of Valonia. Papers Tortugas Laboratory 29: 13-21 (issued Nov. SHE 1935). Carnegie Institution of Washington Publication Number 452. For a number of years algal cells with large vacuoles have been the subject of research on the permeability of the plasma membrane. Prominent among forms investigated is Valonia. This paper describes the cytology of Valonia ventricosa and Valonia macrophysa with particular emphasis on structures of significance in physiological investigations. The cells were collected on Bush Key Reef and from the moat at Fort Jefferson and kept in finger bowls in the laboratory. The morphology of the various structures in the cytoplasm of Valonia macrophysa and V. ventricosa is described. The plastids produce starch and lipoid granules and are sufficiently numerous as to constitute two-thirds-of the volume of the cytoplasm. There are approximately three hundred nuclei per square millimeter of cell surface in the coenocytes. Mitosis is intranuclear. In the development of the rhizoidal hapteron cells of the aplanospores, the mitochondria arise from plastids of the coenocyte in which the aplanospores were formed. The large central vacuoles of the coenocyte arises by fusion of small vacuoles formed in the cytoplasm. Double vital staining of artifact vacuoles is noted. . 1940. The structure and composition of Valonia ventricosa. Papers Tortugas Laboratory 32: 143-52 (issued Sept. 1940). Carnegie Institution of Washington Publication Number 571. The physiology of Valonia has been dealt with extensively by numerous authors. Cells were collected from Long Key and the adjacent reef and brought to the laboratory, where they were kept in large glass jars of sea water which was changed daily. Measurements have been made of the relation of the volume and thickness of the cytoplasm and cell wall to the size of the coenocyte. The specific gravities of various cell constituents and of cells of various sizes have been measured. From a consideration of the results presented it would appear that the level of metabolic rate in Valonia is of a low order, but not necessarily of a different order of magnitude from that of the barley-root and potato-slice systems. 98. Doyle, W.L. and M. Metcalfe Doyle. 1940. The structure of zooxanthellae. Papers Tortugas Laboratory 32: 127-42 (issued May 1940). Carnegie Institution of Washington Publication Number 517. The structure of the zooxanthellae in various invertebrate reef organisms under various conditions was investigated at the Dry Tortugas in 1934, 1935 and rechecked in 1939. Ten 26 species of corals and foraminifera were studied in their living conditions, as well as after fixation. Zooxanthellae in foraminiferans, were examined for the effects of light in normal gas tensions, in increased carbon dioxide tensions, and on specimens in oxygen and hydrogen; while in corals the comparative cytology of zooxanthellae was studied. For the large heads of the Orbicella (Madrepora), the amount of light present at the top and bottom of the corals determined the natural variations in the amount of calcium oxalate crystals in zooxanthellae. Increased levels of crystals were found at the bottom in darkness, while no crystals were found at the top. Similar results were found in foraminiferans. The converse is true for the amount of starch present. The zooxanthellae in corals under the most intense natural light conditions contains little starch, but abundant oil droplets. It was concluded that, overall, for the greater part of the day, the zooxanthellae, as well as the corals, are in need of oxygen. 99. Drew, G. H.. 1914. On the precipitation of calcium carbonate in the sea by marine bacteria, and on the action of denitrifying bacteria in tropical and temperate seas. Papers Tortugas Laboratory 5: 7-45. Carnegie Institution of Washington Publication Number 182. The investigations described in this paper were made in the summers of 1911 and 1912 under the auspices of the Carnegie Institution of Washington. The intent was to study the action of marine denitrifying bacteria in tropical seas. The discovery that these denitrifying bacteria also possess the power of precipitating calcium carbonate from soluble calcium salts present in sea-water has overshadowed the primary object of the work. The observations so far available are few, and the area they cover too small, to attempt to make broad generalizations. However, it can be stated that the very extensive chalky mud flats forming in the neighborhood of the Florida Keys are now being precipitated by the action of the bacterium calcis on the calcium salts present in solution in sea-water. The investigation can at most be considered to offer a mere indication of the part played by bacterial growth in the metabolism of the sea. To obtain a real insight into the question, it would be necessary to make more extensive bacterial and chemical observations in tropical, temperate, and arctic waters, to study the bacteriology of other areas where calcium carbonate is being precipitated from the sea, and to make further investigations in the laboratory into the chemistry of the reactions that can be brought about by various species of marine bacteria. 100. Dustan, P. 1985. Community structure of reef-building corals in the Florida Keys , Carysfort Reef, Key Largo, and Long Key Reef, Dry Tortugas. Atoll Research Bulletin 282-292: 1-29. This communication is the result of two parallel studies on the distribution of reef-building corals on Carysfort Reef, Key Largo and Long Key Reef, Dry Tortugas. The aim of the projects was to characterize the species composition of reef-building corals from the northern and southernmost localities of the Keys, and through comparison attempt to identify the impact of man on the reefs in the Key Largo area of the northern Florida Keys. 101. Dustan, P., W. Jaap and J. Halas. 1976. The distribution of members of the Class Sclerospongiae. Lethaia 9, no. 4: 419-20. The Sclerospongiae play an important and sometimes major role in the construction and infilling of reefs in tropical waters. Modern sclerosponges are limited to dark, quiet, sediment-shaded habitats. This study describes the distributions of sclerosponges in the Bahamas and the Florida Reef Tract. The sponges were found in the Grand Bahamas. After extensive SCUBA diving in Pennekamp Park and the Dry Tortugas, no Sclerospongiae were found. Cold water temperature, or alternatively few, if any larvae to colonize the reef tract are possible explanations for the lack of Sclerospongiae in the Florida Reef Tract. 27 102. Edmondson, C. H. 1908. A variety of Anisonema vitrea. Papers Tortugas Laboratory 1: 191. Carnegie Institution of Washington Publication Number 102. Notes are provided on the protozoan, Anisonema. Anisonema vitrea (Dujardin) is a flagellated protozoan, elongate-oval in form, the anterior end broadly rounded, the posterior more acutely rounded. Anisonema vitrea is distinguished from other species of the genus by eight furrowed surfaces extending in a slightly spiral manner from one end of the body to the other. During the summer of 1906, while working on marine Protozoa at the Tortugas, Fla., the author studied a form considered as a variety of the above species entitled Anisonema vitrea (Duj.) var. pentagona. A description of the difference between the species and variety is presented . 103. Erseus, C. and M. R. Milligan. 1988. A new Bathydrillus oligochaeta Tubificidae from the eastern Gulf of Mexico. Bulletin of Marine Science 42, no. 2: 292-95. Bathydrilus natabilis is described from 4-58.5 meter depths off Crystal River and Dry Tortugas in the eastern Gulf of Mexico. The species is characterized by large, finely pectinate, penial setae in segment 11 and entally curved, single-pointed, spermathecal setae in segment 10 which distinguish it from all congeners. 104. Farfante, I. P. 1980. A new species of rock shrimp of the Genus Sicyonia penaeoidea, with a key to the western Atlantic species. Proceedings of the Biological Society of Washington 93, no. 3: 771-80. Sicyonia olgae, new species, ranges from Dry Tortugas Islands, Florida, to Surinam. It differs from Sicyonia typica (Boeck, 1864), its closest western Atlantic relative, in possessing sublateral carinae on the carapace, and in lacking posterior pleural sulci on the first three abdominal somites. Also distinctive are the sharply pointed, mesially directed, distomesial projections of the petasma in the male, and in the female the pair of long, slender spines on sternite XI and rounded posterolateral processes of the median plate of sternite XIII. A key to the western Atlantic species of Sicyonia is supplemented by synopses of their geographic and depth ranges which include many extensions of previously known limits. 105. Feinstein, A. A. A. R. Ceurvels R. F. Hutton and E. Snoek. 1955. "Red tide outbreaks off the Florida West Coast." Report to the Florida State Board of Conservation of Marine Laboratories . A compilation of reports of red tide on the west coast of Florida from 1844 to January, 1955 is given. Also included are two working diagrams of incidence of red tide, suggesting that red tide occurs more frequently in the months of August through January, and that individual red tide outbreaks are part of larger outbreaks, which seem to move from south to north, and summer outbreaks appear to originate mostly north of Venice, winter and spring outbreaks further south. Further data are required to give complete support. If this is substantiated, control may be exerted by action in a limited focal area or areas of origin. Otherwise, the problem of control may be of the greatest difficulty, since it will require action over a much wider area. | 106. Field, R. M. 1919. Investigations regarding the calcium carbonate oozes at Tortugas, and the beach rock at Loggerhead Key. Carnegie Institution of Washington, Year Book 18: 197-98. Calcium carbonate accumulations in the shallow lagoons and channels between the reef flats were examined to ascertain their origin. Carbonate ooze hardens rapidly when exposed to air and when flooded with saltwater, mud-cracked zones can be formed similar to those in the geologic record, as in the Stones River limestone formation. An account is given on the origin of the "beach rock" found between the high and low water marks on Loggerhead Key. 28 107. : 1920. Origin of "beach rock" (coquina) at Loggerhead Key, Tortugas (abs.). Bulletin of the Geological Society of America 31: 215. A study was made to discover the origin of the "beach-rock" or cemented shell-sands which occur between high and low tides. By means of a stand-pipe and pump, it was found that during heavy rains a shell key acts like a reservoir, and the meteoric water dissolves CaCO; on its way through the loose shell sands. The ground water was found to contain 40 per cent more CaCo; in solution, or colloidal suspension, than the normal sea water. This concentrated solution of CaCo; has a strong cementing value, and is probably an important factor in the formation of the "beach-rock" where the ground water flows out through the beach sands, between tides. 108. Fisk, E. J. 1976. Black phoebe sighted at Dry Tortugas. Florida Field Naturalist 4, no. 2: 39. An observation of a black phoebe on Loggerhead Key, Dry Tortugas on April 13, 1976 is recorded. This is the fourth sighting and only spring record for Florida of a black phoebe. 109. Gauld, G. 1820. An accurate chart of the Tortugas and Florida Keys or Martyrs, surveyed by 110. Gee, H.. George Gauld, A.M. in the years 1773, 1774, 1775. London, W. Faden. First nautical chart of the Dry Tortugas is produced. 1934. Lime deposits and the bacteria. I. Estimate of bacterial activity at the Florida Keys. Papers Tortugas Laboratory 28: 67-82 (issued Dec. 1932). Carnegie Institution of Washington Publication Number 435. Aerobic organisms were collected from the Florida Keys. Viable counts indicate that open areas are only thinly populated with these forms, but that sheltered areas may permit increased activity. Conditions in the mud are such as to favor the growth of anaerobes. There is a possibility that specific groups, such as the purple sulphur organisms are at work in addition to the conventional aerobes. 111. Gee, H. and C. B. Feltham. 1934. Lime deposition and the bacteria. II. Characteristics of aerobic 112. Gersh, I. 113. Gilmore, bacteria from the Florida Keys. Papers Tortugas Laboratory 28: 83-91 (issued Dec. 1932). Carnegie Institution of Washington Publication Number 435. General bacterial conditions at the Florida Keys during the 1930 season have been discussed by Gee (1932). There was reported a collection of 138 representative aerobic organisms recovered from the water and mud of Bird Key harbor between Bird and Garden Keys, of the Marquesas lagoon, and of one vertical one in the vicinity of the Gulf Stream. Preliminary examinations were made of them at the Tortugas laboratory. The strains were found to be Gram-negative rods, ammonia-producing, and possibly fermenting. The collection was subsequently studied exhaustively at the Scripps Institution during the winter of 1930-31. When freshly isolated, these bacteria displayed considerable variation in size, in colony features and color, and in their degree of physiological activity. 1935. Studies on the anterior pituitary gland of the nurse shark. Carnegie Institution of Washington, Year Book 34: 81. Experiments were planned on the nurse shark to determine which of the activities of the anterior pituitary gland are referable to the eosinophile cells. R. G. and R. S. Jones. 1988. Lipogramma flavescens, a new grammid fish from the Bahama Islands with descriptive and distributional notes on L. evides and L. anabantoides. Bulletin of Marine Science 42, no. 3: 435-45. In 1981, dredge collections made north of the Dry Tortugas by Continental Shelf Associates under contract with the Bureau of Land Mangement documented the first continental record of L. anabantoides. 29 114. Ginsburg, R. N. 1953. Beach rock in south Florida. Journal of Sedimentary Petrology 23: 89-92. 115. Goldfarb 116. IIY/, The rapid intertidal lithification of beach deposits in the coral seas has received the attention of numerous investigators. Study of beach rock from the Dry Tortugas shows that the aragonite cement is precipitated from the sea water remaining in the beach sands at low tide. High temperatures, rate of beach drainage, and the permanence of the beach control the localization of beachrock. The recognition of beachrock in the fossil record is briefly discussed. , A. J. 1913. Changes in concentration of sea-water and their influence upon regeneration. Proceedings of the Society for Experimental Biology and Medicine 10, no. 3. The regeneration under changed densities of sea water was observed under conditions that endured the elimination of uniformity of associated factors such as size of medusae, volume, surface and depth of solutions, extent of injury, level of amputation, temperature, crowding, aeration, etc. Dilutions were made with water containing a known quantity of sea salts, and concentrated solutions were made by slow evaporation, which corrected certain errors in previous experiments. Results were compared with those of Loeb. It was found that both the hydroid Eudendrium of Woods Hole as well as Cassiopeia of Dry Tortugas differed radically from the Loeb experiments, in respect to the range of solutions in which animals lived or regenerated, the optimum solutions, the normality of the regenerated parts and the character of the curve. It is stated that Loeb's curve probably is limited to Tubularia of Naples, and does not represent the behavior of organisms to changes of density of sea water, and that the differences in the behavior of these three organisms can hardly be correlated with the differences in concentration of the sea water in which they normally live. . 1914. Changes in salinity and their effects upon the regeneration of Cassiopea xamachana. Papers Tortugas Laboratory 6: 83-94. Carnegie Institution of Washington Publication Number 183. Cassiopea xamachana, a large scyphomedusa, is very abundant in the very shallow waters of the moat at Fort Jefferson, Dry Tortugas, Florida. The present report considers to what extent changes in salinity influence regeneration in Cassiopea, and the results of the investigation are compared with those previously obtained with the hydroid Eudendrium ramosum of Woods Hole, Massachusetts and with the observations of Loeb with the hydroid Tubularia of Serino Bay, Italy. The object of this investigation was to ascertain to what extent changes in salinity affected Cassiopea xamachana normally subject to relatively great variation in the concentration of the sea-water, and to compare the results with those of the hydroid Eudendrium and the hydroid Tubularia. The following variable factors were uniform for the series: size of medusae; volume, surface, and depth of the solutions; extent of injury; level of amputation; temperature; crowding. Injurious or other variable factors were guarded against. Cassiopea lived in solutions ranging from 40 to 153 per cent sea-water solutions. Regeneration occurred in solutions containing 50 to 133 per cent sea-water. Normal regeneration of the arms occurred within much narrower range, namely 75-105 per cent. Beyond these limits regeneration was atypic . . 1918. Effects of aging upon germ cells and upon early development. Part II. Biological Bulletin 34, no. 6: 372-409. In a previous preliminary experiment it was shown that freshly liberated eggs of different females of three different species of sea urchins (Toxopneustes and Hipponoe collected from the Tortugas, and Arbacia from Woods Hole, Mass.) varied in respect to size, jelly layer, membrane formation, and cleavage. In this paper the same technique and the same three species of sea urchins were used to determine the physiologic condition of the germ cells, and then determine the nature of the changes in the eggs as they became increasingly overripe. As eggs in good physiologic condition aged, their volume increased until they 30 118. 120. 121. became smaller than the norm. Eggs in poor condition were reduced in size., in all three species, there was a loss in jelly layer with age, depending on the condition of the egg. In all three species, as the eggs aged, the membrane appeared closer to the surface, becoming thinner until none was formed. The rate of decrease in cleavage with age was greater in Toxopneustes and Hipponoe than in Arbacia. Overall, the change in size, jelly membrane, and cleavage with aging of germ cells are accurate, convenient and corroborative indices of physio-chemical and morphologic changes in the eggs as they age, and afford convenient measures of loss in vitality, or physical deterioration. . 1914. Experimentally fused larvae of echinoderms with special reference to their skeletons. Papers Tortugas Laboratory 6: 103-21. Carnegie Institution of Washington Publication Number 183. The early work of Loeb, Morgan, and Herbst on the production of multiple embryos from a single egg suggested the reverse experiment of grafting or reuniting several fertilized eggs into on embryo. In 1912, the writer repeated these experiments with the American form Arbacia punatulata and succeeded only after slightly modifying Driesch's method. Subsequently, in the performance of other experiments, it was discovered that eggs could be agglutinated and fused quite as readily by a very different method, which was not only simpler but free of certain objections that might be urged against previously known methods. The new method consisted in using an isotonic or slightly hypotonic NaCl solution diluted with varying quantities of sea-water. . 1913. The influence of the central nervous system in regeneration of an annelid worm. Proceedings of the Society for Experimental Biology and Medicine 10, no. 3. np. (No abstract available). . 1914. Regeneration in the annelid worm Amphinsoma pacifica, after removal of the central nervous system. Papers Tortugas Laboratory 6: 95-102. Carnegie Institution of Washington Publication Number 183. In a previous publication, the writer found that the head of the earth-worm Lumbricus was regenerated in the entire and permanent absence of the nerve-cord from the amputated region. The marine annelid worm Amphinoma pacifica readily regenerated a head at all levels except the distal eighth of the worm. Regeneration may be prevented by a severe injury, either to the digestive tract or to the central nerve system; the greater the injury the more likely will regeneration be inhibited. Many pieces did not regenerate after removing the alimentary tract from five or more segments nearest the amputated level. Many pieces, about one-third, failed to regenerate after removing the nerve-cord by the forceps, i.e., with little injury to adjoining tissues. All failed to regenerate after removing the nerve-cord by the "window" method. The operated worms were examined in serial sections. In one group a regenerated nerve-cord connected the regenerated "brain" and commissures with the old intact nerve-cord. In a second group the regenerated nerve-cord approached and in instances reached the amputated level, yet no head was formed. In a third group, the nerve- cord had regenerated, but several segments nearest the amputated end were yet without any nerve-cord or ganglia. These worms nevertheless had regenerated a head with its typical brain and nerve-commissures. . 1917. Variability of eggs and sperm of sea-urchins. Papers Tortugas Laboratory 11: 71-87. Carnegie Institution of Washington Publication Number 251. A clear understanding of the variability in normal fresh eggs and sperm is necessary in order to appreciate and to evaluate the changes that take place in overripe germ-cells. This paper deals exclusively with the qualitative and quantitative differences of such freshly collected sea-urchin eggs and sperm and with the differences in their early development. Sil 122. . 1917. Variability of germ cells of sea urchins. Proceedings of the National Academy of Science 3: 241-45. Three different species of sea urchins (Toxopneustes and Hipponoe collected at the Dry Tortugas, Arbacia collected at Woods Hole, Massachusetts) were used to determine the normal variability of sea urchin germ cells. Having determined the optimum and constant conditions of germ cells, studies were conducted to examine variations in size and shape of eggs, the jelly layer of eggs, membrane formation, and cleavage among the three species. Amazingly large variations were found in fresh germ cells among species, thus suggesting that among other investigators of the varying behavior of the eggs, a large part of the variation was probably due to the physiologic conditions of the eggs which these investigators used. 123. Goodrich, H. B. 1935. Color patterns in fish. Carnegie Institution of Washington, Year Book 34: 81. Studies were carried out to investigate internal conditions which may control the development and maintenance of color patterns in fish by transplanting scales and tissues from one type of pigment area to another. 124. Gordon, M. 1933. The internal pigment systems of fishes. Carnegie Institution of Washington, Year Book 32: 268. The internal pigmentary systems of major taxonomic groups were examined. Halichores bivittatus and Lutjanus griseus showing possibly neoplastic growths were collected for study. 125. Goy, J. W. 1982. West Indian Stenopodidae. 2. Occurrence of Richardina spinicincta Crustacea, Decapoda, Stenopodidea off the Dry Tortugas. Bulletin of Marine Science 32, no. 1: 344- 47. An examination of Richardina spinicincta collected by W.L.Schmitt in August of 1932 is made. It is concluded that this specimen is truly R. spinicincta, that this is the sixth known specimen of the species, and the first record of the genus in the Western Atlantic. The occurrence suggests that the genus occurs at shallower depths than those recorded in previous literature. 126. Grave, C. 1934. The Botryllus Type of Ascidian larva. Papers Tortugas Laboratory 28: 143-56 (issued Dec. 1932). Carnegie Institution of Washington Publication Number 435. Free-swimming larvae of at least three well-defined types are found in life cycles of ascidians; one, characteristic of species of Molgula (Grave '26) and related genera, that has one sense organ only, a statolith, in its sensory vesicle. The nerve cord lies in a mid- dorsal position above the notochord, the caudal fin is expanded vertically in the median plane and adhesive papillae are lacking. In the text the structural organization of a type of larva characteristic of species of Botryllus and related genera is described. The body is egg shaped, its depth being approximately the same as its width. Three conical sensory papillae arranged in the form of an equilateral triangle are borne at the anterior smaller end of the body, two located on either side of the median plane dorsal to the central body axis, one in the median plane ventral to the central axis. The same gross parts found in the Central nervous system of larvae of other types are present. The anterior end of the visceral ganglion bends to the right and expands to form the sensory vesicle, which, in contrast with that of larvae of other types, does not project to the level of the dorsal surface of the body but retains an interior position relatively far below the surface. 127. ———. 1936. Metamorphosis of ascidian larvae. Papers Tortugas Laboratory. 29: 209-91 (issued Dec. 1935). Carnegie Institution of Washington Publication Number 452. 32 The studies of metamorphosis of larvae of ascidians were made during the summers of 1927, 1930, and 1933 at the Tortugas Laboratory with the purpose of finding methods of accelerating and controlling metamorphosis and thus of discovering something of the fundamental nature of the internal mechanism involved and the environmental conditions with which it is causally related. The observations made in the cqurse of this investigation are interpreted as follows: The ascidian larva is a dual organism, the action system of the larva being quite separate from the action system of the ascidiozooid. Metamorphosis advances by three stages; (a) changes in the adaptive responses of the larva to light and gravity; (b) the attachment of the larva to the surface of some foreign object; (c) the disruptive phase during which the entire larval action system is destroyed. Swimming activity causes the production and concentration of some metabolic product in the larval tissues that is essential to the induction of metamorphosis. The presence in the larval tissues of metabolic products resulting from swimming is not alone sufficient to induce metamorphosis, but another substance with which this metabolic product may react is equally necessary. The great variability of ascidian larvae of the same species in the duration of their free-swimming period is apparently due to variability in the time of formation of the susceptibility substance and hence to the time of differentiation of the larval organ that produces it. Metamorphosis may be induced artificially by diverse chemical and biological substances placed in sea-water with groups of larvae in lactic acid. Metamorphosis is rapidly and consistently induced in the larva of Phallusia nigra. The activating agents extracted from the fresh ascidian tissues that were so specific in their effects may also be endosymes of a highly specialized kind, each found only in a single species of ascidian. The mechanism of metamorphosis is comparable in its organization to that of development of an egg, which also may be activated by numerous and diverse chemical and physical agencies. 128. Grave, C. and P. A. Nicoll. 1940. Studies of larval life and metamorphosis in Ascidia nigra and species of Polyandrocarpa. Papers Tortugas Laboratory 32: 1-46 (issued Oct. 1939). Carnegie Institution of Washington Publication Number 517. Experimental studies made during the summer of 1933 (Grave, 1936) demonstrate that sea- water extract of pharyngeal, atrial, or mantle tissues of adult Ascidia nigra is effective in inducing 100 per cent metamorphosis in groups of Ascidia larvae within 3 hours after hatching and that similar extract of tissues of Polyandrocarpa induces 100 per cent metamorphosis in groups of Polyandrocarpa \arvae within 42 minutes after liberation from the parent colony. These observations led to a search during the summers of 1935 and 1936 for a specific chemical substance in the tissues of these ascidians having the properties required for the rapid acceleration of the process of metamorphosis. An account of the methods and results of this work is given in this paper. It was found that the amino acids |-histidine, leucine, glycine, cysteine, and d,]-alanine, in the form received from the laboratories in which they were prepared, accelerated metamorphosis in groups of larvae of both types. A sea-water extract of free-swimming larvae or of late embryonic stages of Ascidia has the same accelerating effect on metamorphosis of Ascidia larvae as an extract made from tissues of the adult ascidian. Heating adult Ascidia tissue or releasing distilled- water extracts of the tissue for several hours does not destroy the accelerating substance. Non-toxic concentrations of copper, iron, and aluminum salts induce early metamorphosis to a marked degree. The duration of the free-swimming period of Ascidia larvae is longest at the beginning of the breeding season of the species and becomes gradually shorter as the season advances. 129. Gudger, E. W. 1921. Notes on the morphology and habits of the nurse shark, Ginglymostroma cirratum. Copeia 98: 57-59. A physical description of the nurse shark as observed by the author for several summers in the southern Florida Keys and Dry Tortugas is given. 130. Wil, N32, 133), 33 . 1929. On the morphology, coloration, and behavior of seventy teleostean fishes of Tortugas, Florida. Papers Tortugas Laboratory 26: 149-204. Carnegie Institution of Washington Publication Number 391. In the course of work at the Tortugas, 70 teleosts, belonging to 28 families, have been studied. Habits have been recorded herein that stand out prominently to the taxonomist. First, basing the classification of tropical fishes on coloration is a very dangerous thing. Most of the fishes in Tortugas have from two to five color phases in life and, even when studying the fish in a state of comparative quiet in an aquarium, it is very difficult to determine which is its normal color. When a fish dies, its color changes either entirely or in its intensity, so that the coloration of the dead fish is markedly different from that of the live fish. It is equally dangerous to describe and classify a tropical fish from a single specimen, since these fishes are so very variable in the number of fin rays, in the relative proportions of the body, in scale count, and in the many details which help to distinguish one species from another.. . 1918. On the use of the diving helmet in submarine biological work. American Museum Journal 18: 135-38. The use of the diving helmet for research at the Dry Tortugas was initiated in 1915 by Longley and Carey, for fish observations and photography. Its use was declared new for underwater work. However, such is not the case. The use of the helmet alone replaced cumbersome diving suits (scaphanders) used by the commercial spongers out of Tarpon Springs, Florida, and early workers on the construction of the overseas railroad, The Florida East Coast Railway Extension from Homestead to Key West, Florida. The diving helmet in biological work dates back to around 1845, when M. Milne-Edwards conducted bottom surveys off the coast of Sicily. In 1679, pressurized air was first supplied to Borelli, who attached a simple air compressing pump to a leather diving helmet. These devices are all refinements of the crude diving helmets used back in ancient times by Alexander the Great, while recording plant and animal observations. These are some of the earliest underwater biological observations ever recorded. The earliest account of any type of diving apparatus is found in Aristotle and dates back to about 1000 B.C. . 1918. Sphyraena barracuda; Its morphology, habits, and history. Papers Tortugas Laboratory 12: 53-108. Carnegie Institution of Washington Publication Number 252. This article provides a general description of the great barracuda, Sphyraena barracuda made at the Tortugas Marine Laboratory, based on local collections and an examination of 12 large individuals using length/weight measurements, color and markings, jaws and teeth, internal organs, foods and feeding, and manners of breathing. Additional information is presented on their habits, how they may be caught, and parasites. An interesting historical side of the paper compiled from around the world includes verbatim quotes and descriptions of their great size, ferocity, fossil forms, nomenclature, habitats, and food poisoning in man. Accounts of their poisonous flesh in the West Indies date as far back as 1667. Largest sizes of West Indies individuals approach 8-10 feet in length, with some highly "dubious" reports of specimens reaching sizes of 18-20 feet in length. . 1913. Uterine gestation in the nurse shark, Ginglymostoma cirratum. Journal of the Elisha Mitchell Scientific Society 29: 8. Also, in Science, 1913, v.37, p.993. The breeding habits and embryology of this shark were studied at the Tortugas Laboratory in the summer of 1912. A brief account was published in the Year Book for 1912, p. 148- 150. 34 134. Halley, R. B. and R. P. Steinen. 1979. Groundwater observations on small carbonate islands of southern Florida. In Guide to sedimentation for the Dry Tortugas. Compiler R. B. Halley, p. 82-89. Tallahassee, Florida: South East Geological Society Publication. Observations are reported on the unusual hydrology of Loggerhead Key, a sandy key in the Dry Tortugas in comparison with observations on Cluett Key, a mud key which lies 200 km NE of Loggerhead in western Florida Bay. The ground water of Loggerhead and Cluett Keys differs significantly from the surrounding sea water, despite the relatively small size of the island. Climate alone does not determine the character of these ground waters; for example, Loggerhead Key is underlain by less saline ground water than Cluett Key despite the fact that it receives less rainfall. Ground water under such small islands such as these is formed from topography, sediment character, vegetation, and many more parameters that are themselves interrelated. They conspire to form ground water that not only differs from sea water, but also can react with the island sediments to change the character of the ground water. In this manner, island ground waters serve as geologic agents, hastening the alteration of marine carbonate sediments to limestone and dolomite. 135. Hanlon, R. T. and R. F. Hixon. 1986. Behavioral associations of coral reef fishes with the sea- anemone Condylactis gigantea in the Dry Tortugas, Florida USA. Bulletin of Marine Science 39, no. 1: 130-134. Over 30 small West Indian reef fishes dwell within the tentacular sphere of anemones, mainly to avoid predation. Most species swim carefully to avoid the stinging tentacles, but some species also have a physiological adaptation (skin mucus alteration) that allows them to be in full and vigorous contact with the tentacles in a manner similar to Indo-Pacific anemonefishes such as Amphiprion, Dascyllus and Premnas. The authors report herein six species of reef fishes that are facultative associates of the sea anemone Condylactis gigantea (Weinland) in the Dry Tortugas Islands. The fishes were not found associated with other anemones. One species, Labrisomus gobio, is a new record of a fish with both the behavioral and physiological adaptations to dwell unharmed among the stinging tentacles of Condylactis gigantea. 136. Hargitt, C. W. 1911. Cradactis variabilis: An apparently new Tortugan Actinian. Papers Tortugas Laboratory 3: 49-53. Carnegie Institution of Washington Publication Number 132. The author believes this species of actinian has never before been described, and names it variabilis. The specimens seem to have the capacity to move about more or less freely, and the frond-like organs situated about the margin of the oral disk and outside the outer cycloe of tentacles aid in such movement. The color is pale olivaceous-green to brownish; tentacles somewhat lighter; foliose organs darker, even brownish, with flake-white pads. The body is highly contractile, with a weak or diffused sphincter. The reproductive season seems to be in the spring and early summer. The habitat is chiefly in holes, crevices, or similar secluded places in the coral reefs or about the shoals where protection is available. 137. Harrington, B. A. and J. J. Dinsmore. 1975. Mortality of transient cattle egrets at Dry Tortugas, Florida. Bird-Banding 46, no. 1: 7-14. This article examines the idea presented by Browder (1973) that cattle egrets pass through the Dry Tortugas with seasonal regularity, and that large numbers die after landing. This study concludes that regular spring movement occurs with many egrets stopping at the island, and that many of the egrets that stopped apparently died from starvation, especially in late June and in early July. The mortality in 1968 was higher that in 1970. 138. Harris, J. E. 1937. The mechanical significance of the position and movements of the paired fins in the Teleosti. Papers Tortugas Laboratory 31: 171-89 (issued Oct. 1936). Carnegie Institution of Washington Publication Number 475. 35 In the course of the evolution of the modern teleostean fish, a series of fairly well-defined changes has taken place in the body form and in the shape and position of the fins. The present paper discusses the mechanical factors concerned in the evolution of the teleost type of fish. A comparison of this type with the dogfish suggests that the development of an air bladder has been the primary factor involved in the change in general body form. The reduction in specific gravity of the fish, consequent upon this primary change, has removed the need for a lift force on the body during free swimming. The asymmetrical (heterocercal) tail has therefore disappeared. For the same reason, the pectoral fins are no longer needed as elevating planes, and become free to move up toward the mid line of the body to act as brakes in stopping and turning movements. The forward motion of the pelvic fins is a mechanism for producing a balanced vertical force and a balanced pitching moment. These fins are normally used in conjunction with the pectorals. The independent movements of the pectoral fins are then discussed. All types of movement so far observed are variations on a fundamental form, in which the metrachronal oscillation of the fin rays generates an undulating fin surface. The observed variations in form of the fin beat can be produced by varying the phase difference between the beat of successive rays, and also by making the oscillation of the fin ray asymmetrical. The characteristics of the pectoral musculature associated with such variations are pointed out, and illustrated by reference to a number of fish types. 139. Hartman, C. G. 1931. The hypophysis of fishes. Carnegie Institution of Washington, Year Book 30: 381-82. Studies on the influence of the hypophysis on menstruation and various forms of uterine bleeding in sharks were carried out. 140. Hartmeyer, R. 1911. Polycistor (Eudistoma) mayeri nov. sp. from the Tortugas. Papers Tortugas 141. Laboratory 3: 89-93. Carnegie Institution of Washington Publication Number 132. A new species Polycitor (Eudistoma) mayeri , a new ascidian collected in 1907 at the Tortugas is described as the largest and most beautiful ascidian of the Tortugas. It was collected in the deeper water of the Southwest Channel near Loggerhead Key, on sandy bottoms, where it is abundant. The color is pale yellow, with a reddish or violet tint. From the western Atlantic only five species of this genus have been described, and all of these are mentioned by Van Name from the Bermudas, but all these species have four rows of stigmata in the branchial sac and are in many other respects quite different from this species. . 1908. Reisebilder aus Westinidien mit besonderer Berucksichtigung der korallenbildungen. Deutsch. Gessel. Fur Volkstumlich Natuirkunde or Same Title in Meereskunde Jahrg. 3, Heft 2, 40 Pp 3, no. 2: 1-40. (No abstract available) 142. Harvey, E. N.. 1911. Effect of different temperatures on the medusae, Cassiopea, with special reference to the rate of conduction of the nerve impulse. Papers Tortugas Laboratory 3: 27-39. Carnegie Institution of Washington Publication Number 132. During the summer of 1909 a study was made of the effects of water temperatures on the nerves and muscle tissue of Cassiopea. Temperatures in the moat at Fort Jefferson ranged from 27°C to approximately 32-33°C. Activity limits and thermal death points of nerve and muscle were measured. It was found that nerve conduction rates fall off in rate with rise of temperature to a definite maximum, similar to that for enzyme action and for other life processes. 36 143. 144. 145. . 1914. The relation between the rate of penetration of marine tissues by alkali and the change in functional activity induced by the alkali. Papers Tortugas Laboratory 6: 131-46. Carnegie Institution of Washington Publication Number 183. The present study, made at Tortugas in the summer of 1911, is a continuation of permeability investigations undertaken at Columbia University in 1910 to 1911. The author's aim has been twofold. First, to compare the permeability of the cells and tissues of salt-water organisms with those of fresh-water forms. Second, to determine the relation between the rate of penetration of the alkali and the appearance of structural or functional changes in the cell. The author thinks that the presence of a sufficient number of OH ions within the egg may aid in breaking down the granules and that this breaking down increases also the degree of swelling of the egg. Cytolysis in Holothuris appears to be largely of this type, since NaOH enters before the increase in volume begins. From this point of view both theories of cytolysis contain an element of truth. Swelling of marine eggs is due both to an increase in permeability of the surface and also to the breakdown of lipoid or protein granules within. The latter tends to increase the swelling pressure or the osmotic pressure of the egg, but is secondary to the increase in permeability of the surface. . 1921. Studies on bioluminescence XIII: Luminescence in the Coelenterates. Biological Bulletin 61: 280-287. (No abstract available). . 1923. Studies on bioluminescence. XV. Electroreproduction of oxyluciferin. Journal of General Physiology 5: 275-84. This work was on the light-producing reaction in the luminous crustacean, Cypridina. Oxyluciferin may be reduced to luciferin at cathodes when an electric current is passed through the solution, or at cathodes formed by metal couples in solution, or at cathodes of oxidation-reducation cells of the NaCl - Pt - Na)S type. It is also reduced at those metal surfaces (Al, Mn, Zn and Cd) which liberate nascent hydrogen from water, although no visible hydrogen gas separates from the surface. Molecular hydrogen does not reduce oxyluciferin even though very finely divided, but will reduce oxyluciferin in contact with palladium. Palladium has no reducing action except in the presence of hydrogen, and apparently acts as a catalyst by virtue of some power of converting molecular into atomic hydrogen. Conditions are described under which a continuous luminescence of luciferin can be obtained. This luminescence may be used as a test for atomic hydrogen. It is suggested that the steady luminescence of bacteria is due to continuous oxidation of luciferin to oxyluciferin and reduction of oxyluciferin to luciferin in different parts of the bacterial cell. 146. Hatai, S. 1916. Changes in the chemical composition of starving Cassiopea xamachana. Carnegie 147. Institution of Washington, Year Book 15: 206-7. Studies were conducted on chemical changes occurring in Cassiopea during starvation. The constancy of water content suggests that Cassiopea is largely a jelly-like mass, and remains so throughout its life. In contrast, mammalian body-water content varies by age. . 1917. On the composition of Cassiopea xamachana and the changes in it after starvation. Papers Tortugas Laboratory 11: 95-109. Carnegie Institution of Washington Publication Number 251. For this study eight freshly caught normal Cassiopea, having different body weights, were subjected to starvation by placing the animal in filtered sea water. The results were as follows: 1. In general the smaller Cassiopea loses relatively more weight than the larger. 2. The percentage of water found in the entire body is nearly the same in all sizes of Cassiopea. However, the values of water content in the starved appear to be slightly higher than those found in the normal Cassiopea. 3. The nitrogen content of the entire body is 148. 3H) higher in the smaller than in the larger Cassiopea. 4. The absolute amount of nitrogen found in the starved Cassiopea is considerably higher than in the normal having the same body weight. It was noted that although high when compared with the normal, equal in weight to the starved animal, it is very low for the initial body weight of the starved animal. This shows that the nitrogen also has been consumed during the period of starvation. 5. The nitrogen contents for the different parts of the body are simular in their relations to those found in the normal Cassiopea. 6. The loss in weight of the different parts is of such a character that their proportions in the starved remain similar to those in the normal Cassiopea. . 1917. On the composition of the medusa Cassiopea xamachama. Proceedings of the National Academy of Science 3: 22-24. In this study, an examination was made of three different parts of Cassiopea, mouth- organs, umbrella, and velar margin to determine whether starving specimens lose weight uniformly, or whether the loss is dissimilar in the three parts. Results indicated that the smaller Cassiopea loses relatively more weight than does the larger Cassiopea. The percentage of water is similar through the entire body, the nitrogen content is higher in the smaller than the larger individuals, and nitrogen is much higher in the starved Cassiopea than in the normal specimen with the same body weight. Results are compared with Mayer's experiments, which showed nitrogen loss to be constant during the entire period of starvation. Differences may be due to the size of animals used in his studies, as larger individuals show little variation in nitrogen loss, whereas small Cassiopeas show large variations in nitrogen loss due to body size. 149. Hayes, F. R. 1932. Nitrogen in echinoid ontogeny. Carnegie Institution of Washington, Year Book 150. 31: 284-85. The chemical embryology of the echinoid egg was investigated, as well as variations in two sources of energy available in the egg: protein and lipins. . 1934. Variation in size and in nitrogen requirements during early development of the sea- urchin, Echinomtera lacunter. Papers Tortugas Laboratory 28: 181-93 (issued Mar. 1933). Carnegie Institution of Washington Publication Number 435. After the penetration of a spermatozoon, the developing echinoderm egg receives nothing from the outside except water and salts, until the comparatively advanced larva begins to eat. The morphological phenomena of ontogeny can be brought about only by the expenditure of energy, which must come from materials already present in the egg at the time of fertilization. The problems of chemical embryology include (a) a determination of the amount of energy required to produce structural changes, and (b) an investigation of the chemical transformations taking place. The work here reported deals with a certain phase of the chemical embryology of a common tropical sea-urchin, Echinometra lacunter. Studies of the first 24 hours of development of the eggs of this form were carried on during the summer of 1932 at the Tortugas Marine Station of the Carnegie Institution. Eggs of the sea-urchin, Echinometra lacunter, were concentrated with a hand centrifuge and then diluted with 500 times their volume. Analyses of primary amino nitrogen groups and of total nitrogen were made, and the ratio of the former to the latter calculated. From 4 hours onward the ratio of primary amino groups to total nitrogen increases. This does not mean, however, a synthesis of the former at the expense of the latter, but rather that in the combustion which provides the developing embryo with energy, some source of nitrogenous fuel other than NH) groups is being used. There is a marked loss in the quantity of nitrogen per egg during the period of development succeeding the first four hours. One million eggs contain some 13 milligrams of nitrogen, of which about 28 per cent is in the form of NH) groups. 38 151. Heard, R. W. and D. G. Perlmutter. 1977. Description of Colomastix janiceae, new species. A commensal amphipod (Gammaridea Colomastigidae) from the Florida Keys, U.S.A. Proceedings of the Biological Society of Washington 90, no. 1: 30-42. During November of 1968 and 1973 and June of 1970 more than 100 specimens of an undescribed commensal amphipod belonging to the genus Colomastix Grube, 1861 were collected from loggerhead sponges, Spheciospongia vesparia (Lamarck), in the lower Florida Keys. Additional specimens of this new species, collected from Dry Tortugas, Florida were borrowed from the Division of Crustacea of the U.S. National Museum of Natural History for examination. 152. Helwig, E. R. 1933. Regeneration in Jotrochota birotulata (Porifera). Carnegie Institution of Washington, Year Book 32: 271-73. The development and formation of cells over time was examined, from cross-sections made from the branches of the sponge, Jotrochota birotulata. 153. Hendee, E. C. 1931. Formed components and fertilization in egg of the sea-urchin Lytechinus variegatus. Papers Tortugas Laboratory 27: 99-105. Carnegie Institution of Washington Publication Number 413. This investigation of the eggs of Lytechinus variegatus collected during the summer of 1925 at the Tortugas was undertaken to determine if any substance of the egg was involved in fertilization. Certain cytoplamic substances (macrosomes, hyaloplasm, chondriosomes, fat droplets, and extra-nuclear basophilic granules) were demonstrated both before and after fertilization. Lipid granules, present in the mature unfertilized egg, disappeared upon fertilization. 154. Hendrix, S. A. and R S. Braman. 1995. NOx variation in the southeastern Gulf of Mexico. Florida Scientist 58, no. 3: 292-97. An automated system capable of providing speciation and concentration information for several atmospheric NOx compounds was used to obtain diurnal and location variation data during a five-day research cruise in the southeastern Gulf of Mexico approximately one mile west of Fort Jefferson, Dry Tortugas between May 18 and May 22, 1987. Speciation of these nitrogen compounds was achieved by selective preconcentration onto a series of chemically coated glass hollow tubes. Analysis was performed by thermally desorbing the collected analytes into a chemiluminescence detector providing sub parts-per-billion level determination. 155. Hess, W. N. 1937. Reactions to light in Ptychodera bahamensis. Papers Tortugas Laboratory 31: 77-86 (issued Aug. 1936). Carnegie Institution of Washington Publication Number 475. Little attention has been given to the study of light reactions in any of the Enteropneusta, and nothing is known, apparently, concerning the distribution or even the existence of photoreceptors in this important group of animals. The purpose of this investigation was to continue work on reactions to light and the photoreceptors in animals, using at this time a more highly evolved species than the earth-worm on which the earlier work was done. During the study, Ptychodera bahemensis responded negatively to ordinary intensities of light. The movements of Ptychodera, when exposed to light were slow and deliberate and there was little evidence of trial and error movements. The entire surface of the body was sensitive to light, the most sensitive regions being on the proboscis and collar. Removal of different parts of the body involving the central nervous system caused little if any decrease in the percentage of negative responses to light. The reaction time of the proboscis was greatly increased when it was removed from the rest of the animal. This is taken to indicate that the central nervous system functions to speed up responses greatly, but is not essential for responses. Removal of the proboscis together with the basal peduncle makes 156. 39 it impossible for the animal to orient when stimulated by light. This would seem to suggest that the peduncle contains a coordinating center for certain bodily movements, or that the animal has been rendered incapable of orienting, due to removal of that portion of the body containing most of the notochord. . 1940. Regional photosensitivity and photoreceptors of Crangon armillatus and the spiny lobster, Panulirus argus. Papers Tortugas Laboratory 32: 153-61 (issued Sept. 1940). Carnegie Institution of Washington Publication Number 517. Crayfish from which the eyes have been removed are sensitive to light in the region of the sixth abdominal segment, but no responses occurred when other regions were illuminated. The discovery that freshly molted Crangon armillatus are sensitive to light in other regions of their bodies, in addition to the sixth abdominal segment, led to this investigation. Results of this study indicated that Crangon armillatus is usually sensitive to light in many regions of its body, irrespective of how much time has elapsed since the last molting period. Freshly molted spiny lobsters (Panulirus argus) are sensitive to light in many regions of their bodies. Old spiny lobsters, with hard exoskeletons, from which the eyes have been removed are usually not sensitive to light of the intensity used in these experiments. The margins of the uropods of freshly molted Crangon armillatus and spiny lobsters are not sensitive to light, but the basal two-thirds of these appendages are sensitive to light. Adult Crangon armillatus and recently molted spiny lobsters react when illuminated from above after the sixth abdominal ganglion has been shielded by black cardboard and also after the ventral nerve cord has been cut between the fifth and sixth abdominal segments. This shows that photosensitivity in these eyeless animals is not limited to the sixth abdominal ganglion. Newly hatched Crangon armillatus with normal eyes swim toward the light with their caudal ends foremost irrespective of the number of abdominal segments that have been removed. Crangon armillatus and spiny lobsters from which the eyes have been removed do not usually orient to light, but respond by random movements. When their bodies are heavily pigmented, or if they are in poor physical condition, they do not respond at all. However, if they do respond their responses are usually much slower than those of freshly molted animals in good physical condition. The sixth abdominal segment of these eyeless spiny lobsters and crayfishes is the most photosensitive region of their bodies. However, in Crangon armillatus and the American lobster Homarus americanus all the abdominal segments appear to be equally sensitive to light. On the basis of regional photosensitivity of the uropods, it seems probable that the cell bodies of the neurons which connect with the peripheral spines are sensitive to light and hence function as photoreceptors. 157. Hoffman, W., and Jr. and P. C. Patty W. B. Robertson. 1979. Short-eared owl on Bush Key, Dry Tortugas, Florida. Florida Field Naturalist 7, no. 2: 29-30. The short-eared owl (Asio flammeus) is an uncommon but regular winter visitor to Florida. This record represents the second summer record of Asio flammeus in Florida, and the first record for the Dry Tortugas. The authors suggested that the bird in question had been on Bush Key for some time, subsisting on the abundant tern chicks. 158. Holmes, C. W. 1984. Carbonate fans in the Florida Straits. Society of Economic Paleontologists and Mineralogists Annual Meeting (Abstracts) 1: 39. No abstract available. 159. Hooker, D. 1911. Certain reactions to color in the young loggerhead turtle. Papers Tortugas Laboratory 3: 69-76 ahd illustrations. Carnegie Institution of Washington Publication Number 132. During the summer of 1907 observations and a series of experiments were made on the habits and early life history of young loggerhead turtles, which identified reactions to color 40 and geotropism as the determining factors for the causes of young hatchlings to reach the water. Based on day/night experiments on Loggerhead Key, hatchlings did not orient towards the sun or the odor of the water, but exhibited positive phototropism by responding to large surfaces of light of low intensity. After entering the water, the animal swam out to sea apparently attracted by the darker blue of the deeper water. Young turtles displayed positive geotropism when all possible negative geotropic reactions had been exhausted. 160. Hopkins, D. L. Locomotion/physiology of marine amoebae. Carnegie Institution of Washington, 161. 162. 163. Year Book.: ; 1929,v.28,286-288: 1930, v.29,335-337. The chemical and physical factors in the locomotion of marine amoebae collected from the tidal pools at Tortugas and cultured in the laboratory were examined. Relationships between sea-water salt and locomotion were determined by concentration and dilution. Highest rates of locomotion were found in normal sea-water and could be a useful criterion in classification and determining physiological condition in amoebae. Jaap, W. C. 1985. An epidemic zooxanthellae expulsion during 1983 in the Lower Florida Keys coral reefs: hyperthermic etiology. Proceedings of the Fifth International Coral Reef Symposium , 143-48. Moorea, French Polynesia: Antenne Museum-Ephé. Extensive reef coral zooxanthellae expulsion occurred from Key Largo to Dry Tortugas, Florida, during September 1983. Coral bleaching was intensive between Pelican Shoal and Sand Key Reef off Key West. Coral discoloration extended to depths exceeding 14 m but was especially severe in shallow (1-2 m) spur and groove habitats. Approximately 75-95% of all Millepora complanata and Palythoa caribaeorum were bone white, but most colonies remained viable. Affected M. complanata (bladed fire coral) retained the ability to inflict pain from dactylzooid nematocysts. Some individuals (5 to 10%) had fine algal growth indicating death on all or parts of their skeletons. Although 15 species of cnidarians, principally Scleractinia, were affected, some species (Madracis mirabilis, Porites porites, Montastraea cavernosa, Dendrogyra cylindrus ) appeared to be immune. A quantitative sample at Eastern Sambo Reef on 6 October documented 11 species and 209 colonies; M. complanata comprised 32.5% of all colonies. Transmission eletromicrographs did not reveal epidemic pathogenic organisms in affected coral tissues. Warm, calm weather prior to the expulsion was conducive to elevated seawater temperature. A seawater thermograph deployed off Marquesas Key recorded temperatures of 32.3 degrees C. during the period. . 1980. Stony coral community structure at Long Key Reef, Ft. Jefferson National Monument, Dry Tortugas, Florida. (abs.). Florida Scientist 43 (Suppl. 1). Stony coral populations at Long Key Reef were studied during summers of 1975-76 under National Park Service sponsorship. Plotless line transects (13, 25 m L ) were sampled in depths of 0.5-21.3 m. Abundance, cover, and diversity were greatest in depths greater than 8 m. Of 34 species encountered, only 23 were censused quantitatively. Montastraea annularis contributed 20% of all colonies and 37% of cover. Species richness was highest (11) on transects in 7.6-12.5 m depths. Shannon-Weiner diversity values H' log SUB-2 computed by transects for individual colonies ranged from 1.0-3.0. Pielou's eveness (J') values ranged from 0.36-1.00. Community relationships based on Morisita index values detected an assemblage dominated by M. annularis in 8-13 m and a M. cavernosa community in 18-21 m depths. Temporal comparison using the Morisita index revealed strong community stability during 1975-76. Jaap, W. C. and J. Wheaton. 1992. Summary of preliminary results, long-term ecological coral reef studies, Ft. Jefferson National Monument, Dry Tortugas. Prepared for the National Park Service Workshop, 28-30 April 1992, Miami, Florida. 41 Coral reefs exist over time scales of thousands of years. Processes of change in the geological-time context occur slowly, e.g. sea level change correlated with glacial and interglacial periods. The etiology of change is often poorly understood. For example, in 1878 a perturbation identified as, "black water” decimated Acropora spp. at Dry Tortugas. Determining what black water was may never be known. Long-term ecological research seeks to uncover processes that occur slowly or in which effects lag years behind the causes. In the absence of long-term research, serious misjudgments can occur in attempts to manage the environment. The National Park Service was interested in developing a reef resource monitoring plan for Dry Tortugas reefs and collaborated with the Florida Marine Research Institution in a joint study of reef resources. The goals of these studies included testing methods, acquiring a data base on coral reef benthic and fish communities to better understand the etiology of change, and isolating natural from anthropogenic changes. Five study sites were selected in 1989. Repetitive sampling was executed as precisely as possible using several different methods. These methods included: transect sampling, video sampling, quadrat sampling, photographic sampling, recruitment sampling, and environmental sampling. Results indicated that eleven octocoral, 22 scleractinian, and one milleporan species were enumerated on transects, while quadrats indicated 29 octocoral, 26 scleractinian and 1 milleporan species over the study's three year duration. Octocorals were consistently most diverse at Pulaski Shoal (20-21 species). Only 42 of 212 plates recruited scleratinian corals. This yielded an average of 0.35 recruit per plate. A total of 187 milleporan corals recruited to 212 plates for an average of 0.88 recruit per plate (34.9/m?). The only octocoral recruit recorded was the gorgonacean Briareum asbestinum, whose common name is corky sea fingers. 164. . 1994. Summary of preliminary results, long-term ecological coral reef studies, Ft. Jefferson National Monument, Dry Tortugas. Bulletin of Marine Science 54, no. 3: 1-10. Narrative same as in reference no. 163. 165. Jaap, W.C., J. L. Wheaton and K. B. Donnelly. 1990. Materials and methods to establish multipurpose, sustained, ecological research stations on coral reefs at Dry Tortugas. Diving for Science... 1990. Proceedings of the American Academy of Underwater Science, Tenth Annual Science Diving Symposium, 193-203. American Academy of Underwater Sciences: Costa Mesa, California. Sustained research requires precise, repetitive data acquisition to accurately evaluate patterns of change in species abundance and community structure. Permanent reference markers are essential to resample stations over time. The methods described here use solid markers from which several sampling devices can be deployed. A hydraulic drill is used to core 18-in deep holes into rock. A square stainless steel stake is inset, aligned, and cemented into each hole. Quadrats, photogrammetric and video apparatus, and recruitment arrays are deployed on or in reference to the stakes. Transects are extended between stakes. The method is suitable for coral reef and other hard-bottom investigations. 166. Jaap, W. C., J. L. Wheaton , K. B. Donnelly, B. J. Kojis and J. E. McKenna Jr. 1994. A three year evaluation of community dynamics of corals at Ft. Jefferson National Monument, Dry Tortugas, Florida. Bulletin of Marine Science 54, no. 3: 1077. Narrative same as in reference no. 167. 167. Jaap, W.C., J. L. Wheaton, K. B. Donnelly, B. L. Kojis and J. E. McKenna Jr. 1993. A three-year evaluation of community dynamics of corals at Ft. Jefferson National monument, Dry Tortugas, Florida, USA. (abs.). Proceedings of the 7th International Coral Reef Symposium, page 164. Guam: University of Guam. A study to evaluate methods and begin a long-term ecological research program at Ft. Jefferson was initiated at five reef sites in May 1989. Benthos was mapped and 42 photographed within quadrats (5 x 2.56 m per site). Attached biota and substrates were measured along 20- to 25-m transects (3 per site). Recruitment arrays were constructed of PVC pipe, flat stock, and ceramic tiles (10.8 x 10.8 cm) and were secured to the reference stakes. A carriage-mounted video camera, suspended on cables between two "T" poles secured to the stakes, was pushed the length of a transect. Results implied relative stability of the reef communities over three years. Dominant biota as determined by abundance and cover remained similar. Classification analyses of station time-series also corroborated relative stability. Recruitment of Millepora, Octocorallia, and Scleratinia was variable; most recruits were found in cryptic refuge. The heterogeneity, high-relief, and multi- layered canopy of these coral reef habitats restricts the usefulness of medium and long distance (>1.5m) photography and video. We conclude that multiple sampling methods are superior to a single sampling procedure. 168. Jaap, W. C., W. G. Lyons, P. Dustan and J. C. Halas. 1989. Stony coral (Scleractinia and Milleporina) community structure at Bird Key Reef, Ft. Jefferson National Monument, Dry Tortugas, Florida. Florida Marine Research Publication 46: 1-31. Stony coral community structure at Bird Key Reef was investigated during 1975 using 30 continuous 25-m line transects in depths of 0.5 to 21.3 m. Thirty-two species, 872 colonies, and 198 cm of coral cover were sampled quantitatively. Most species, colonies, and live coral cover occurred seaward of 8-m depths on spur and groove substrate. Montastrea annularis, M. cavernosa, and Siderastrea siderea constituted more than 50% of all cover. Species diversity (Shannon index, log sub (2)) ranged from 1.0 for individual transects. Diversity and envenness values computed from cover data were generally lower than values computed from abundance data, reflecting M. annularis dominance. Numerical community classification (Czekanowski's quantitative coefficient) revealed three groups and an ecotone, each related to depth and substrate: 9 transects in 1 to 6 m depths dominated by Porites asteroides and Diploria clivosa; an ecotone of 6 transects in 5 to 6 m depths; 5 transects in 6 to 9 m depths dominated by S. siderea; and 10 transects in 8 to 21 m depths dominated by M. annularis. 169. Jacobs, M. H. 1914. Physiological studies on certain protozoan parasites of Diadema setosum. Papers Tortugas Laboratory 6: 147-57. Carnegie Institution of Washington Publication Number 183. It has been shown by the author and others that different species of protozoa have certain physiological characteristics, often almost as stiking as their morphological ones, and which are probably of considerable significance in the interpretation of their habits of life and their relation to their environment. It occurred to the author to test a series of forms which naturally live under essentially the same environmental conditions, and which may be assumed to have done so for many past generations, in order to see whether they show greater likenesses than a number of forms selected at random, or whether each has preserved its individuality in spite of the similarity of its environment. The general results of the experiments performed show surprising differences in the resistance of the parasites of Diadema to various unfavorable conditions. In some cases the most resistant form may live several hundred times as long as the least resistant one. Comparing all of the results obtained, it is therefore seen that the similar habit of life of the four forms in question has not brought about physiological similarity except in certain adaptive characters which are a sine qua non for continued existence in the same host (e.g. ability to resist the digestive Juices of the latter, etc.). In other respects they are just as different as almost any four free- living forms that might be selected and the evidence of these experiments shows that the physiological characters of an organism are not merely the result of its environment, but may be as fundamental and characteristic as its morphological ones. 43 170. Jefferson, J. P. J. Y. Porter and T. Moore. 1879. On the destruction of fish in the vicinity of the Tortugas during the months of September and October 1878. Proceedings of the U.S. National Museum, Smithsonian Institution Press 1: 244-46. The information in this report is relative to the die-off of large numbers of fish due to a black water event in the Gulf of Mexico during the months of September and October 1878. 171. Jennings, H. S. 1909. Behavior of sea-anemones. Journal of Experimental Zoology 2: 447-72. The study of the behavior of sea anemones (Stoichactis helianthus and Aiptasia spp.) was made at the Carnegie Research Laboratory, Dry Tortugas using specimens collected in the shallow waters near Fort Jefferson. Changes in behavior due to varying states of metabolism for S. helianthus were examined using red meat, crab hard parts and filter paper as food. After satiation, food is rejected through various reactions dependent upon internal processes. Descriptions of food ingestion are described. For Aiptasia spp., experiments suggested that when the animals were hungry, they took both red and filter paper; when satiated they took neither. Other topics of study included climatization to stimuli (light), and reactions modified as a result of past experiences of the organism. Results are compared to other lower groups of animals. 172. Jindrich, V. 1972. "Biogenic buildups and carbonate sedimentation, Dry Tortugas reef complex, Florida." Ph.D. Dissertation, Geology, State University of New York at Binghamton. The Dry Tortugas, a horseshoe-shaped complex of carbonate banks and coral reefs, is located at the southern terminus of the Florida limestone shelf. The complex rises to the surface waters from a drowned Pleistocene surface that forms a circular platform having a general depth of 17-21 m. Three basic biogenic buildups (facies) comprise the reef complex: 1) detrital lagoonal bank, 2) Montastrea reef bank and, 3) Acropora palmata reef. These facies lie adjacent to one another and are also present in vertical succession as individual growth stages of varying thickness and lateral extent. A zone of Acropora cervicornis is developed as a transition between the Montastrea and A. palmata growth stages. The present organic assemblages and topography bear evidence of dominantly lateral progradation and cumulative storm effects that are linked to the slow eustatic sea- level rise for the past several millennia. Long-continued storm degradation is manifested by 1) continuous removal of A. palmata and its replacement by storm-resistant coralline algae and Millepora sp. to produce truncated rocky surfaces, 2) abundant reef rubble, 3) erosion of spur-grooves, and 4) development of intertidal rubble reef flats. Sediments ranging from cobble-sized rubble to medium silt are composed of Halimeda, coral and mollusc grains; coralline algae and foraminifers are present in minor amounts. Variations in texture and constituent particle composition are interpreted to be mainly a result of mode of sediment transport and effect of grain shape. Broadly-defined grain size populations produced by three modes of transport have characteristic assemblages of constituent particles. The populations include a gravel-sized surface creep population, sand-sized saltation population, and very fine sand-to silt-sized suspension population. Strong mixing occurs between the gravel and sand population on the storm-degraded shoals, and between the sand and silt population on the lagoon bottom. Sand flanking the reefs and reef banks shows minimum mixing hence good degree of sorting. Incongruous mixtures of the in- place fraction and varying proportions of the transported populations constitute detrital lagoonal banks as a substrate stabilized by seagrass and coral growth. The gravel-sand and sand-silt mixtures are related to deposition under highly variable energy conditions. Variability in energy conditions does not cause strong population intermixing on beaches. For the same reason, beach sediments show a high degree of sorting in all size grades from cobbles to fine sand. 44 173. Jones, N. 1938. Investigations on ascidians. Carnegie Institution of Washington, Year Book 37: 84. The summer of 1938 was devoted to the study of the structure, development, budding, and colony formation of Ecteinascidia tortugensis, a new ascidian species. The small ascidian, one of the commonest during the season, occurred in large numbers on the under sides of rocks just below low-water mark on both Bush Key and Long Key. Data report provided by Plough and Jones, pp 97-98. 174. Jones, R. D. 1991. An improved fluorescence method for the determination of nanomolar concentrations of ammonium in natural waters. Limnology and Oceanography 36, no. 1: 814-19. An improved fluorescence method is described for measuring nanomolar concentrations of NH, in natural waters. This method is based on the conversion of NH, to NH; and subsequent diffusion of NH; across a microporous hydrophobic Teflon membrane into a flowing stream of 0-phthaldialdehyde reagent to produce a fluorescent adduct. The product is detected fluorometrically with a lower detection limit of better than 1.5 nM. Up to 30 determinations h-1 can be made. The method works well in freshwater or salt water. Field tests of the method in the Dry Tortugas and Gulf Stream gave NH, concentrations that ranged from 18.0 nM in Gulf Stream waters to 2,254.7 nM in interstitial waters from coralline reef sands. The method can be used to measure near real-time NH, concentrations in situations where it was previously difficult or impossible. 175. Jones, R. S. and M. J. Thompson. 1978. Comparison of Florida reef fish assemblages using a rapid visual technique. Bulletin of Marine Science 28, no. 1: 159-72. Species composition, species diversity, and relative abundance of 4 coral reef fish communities in John Pennekamp State Park, Key Largo, Florida, are compared with 4 communities at Fort Jefferson National Monument in the Dry Tortugas using the species- time, random-count technique. The technique is similar to species-area methods, but time replaces area. Fish communities at Pennekamp Park showed the highest overall number of species and scores (reflecting species abundance, and species diversity). Two artificial reefs (shipwrecks) included in the study both show closer relationships to adjacent reefs than to wreck-specific species. 176. Jordan, D.S. 1904. Notes on fishes collected in the Tortugas Archipelago (by Dr. Joseph C. Thompson). Bulletin of the United States Fish Commission for 1902 22: 539-44. An additional sixteen species of fish are described for the Tortugas based on collections made by J.C. Thompson while on the northward cruise of the steamer Chesapeake. 177. Jordan, D. S. and J. C. Thompson. 1905. The fish fauna of the Tortugas Archipelago. Bulletin of the United States Bureau of Fisheries for 1904 24: 229-56. The shallow water fishes of the Tortugas, as suggested by A.G. Mayer to David Starr Jordan, are unsurpassed in variety and abundance anywhere along the Atlantic coast of the United States, and based on the nearness of the Gulf Stream and the winds and currents, pelagic fish from all over the Gulf of Mexico and the West Indies may be drifted by the Tortugas. Collections made by Thompson while on duty as a medical doctor at the Garden Key Naval Station resulted in an annotated fish list containing 218 species known to occur at the Dry Tortugas at the time. 178. Jordan, H. E. 1908. The accessory chromosome in Aplopus mayeri. Anatomischer Anzeiger Bd 32: 284-95. The purpose of this paper is to trace the accessory chromosome in the phasmid, Aplopus mayeri from material collected from Loggerhead key, Florida. The accessory chromosome appears in the resting stage of the secondary spermatogonia as a chromatin nucleolus characteristically close to the nuclear wall. Both the primary and secondary spermatogonia 72), 180. 181. 182. 45 have a metaphase group of 35 chromosomes. The accessory chromosome can be traced as a specific structure from the resting stage of the last order of spermatogonia through all the various phases of synapsis and maturation, until it disintegrates in the head of the ripening spermatozoa. . 1917. Aortic cell clusters in vertebrate embryos. Proceedings of the National Academy of Science 3: 149-56. Aortic cell clusters are described among various animal groups (pig and chick) and compared to 12-day loggerhead turtle embryos and mongoose embryos. Various aspects of the hemogenic activity of embryonic endothelium are described consequent to the inherent capacity of endothelium to produce hemoblast, and not in connection with an associated toxic substance. . 1917. Atresia of the esophagus in the embryo of the loggerhead turtle, Caretta caretta: A normal developmental condition. Papers Tortugas Laboratory 11: 345-60. Carnegie Institution of Washington Publication Number 251. A series of 26 embryos of the loggerhead turtle were collected and used originally for a study of the history of the primordial germ-cells. It was noticed that the esophagus was solid for a greater or less extent, approximately from the point of origin of the respiratory anlage to its bifurcation into the bronchi, from the eleventh to the thirty-second day of incubation. Points of special significance in regard to this material are: (1) the relatively longer persistence of the occlusion than has yet been described for any other form; (2) the absence of contributory yolk in the stenosed area; (3) close relation to the point of origin of the respiratory anlage, which fact may disclose its possible functional significance . . 1917. Embryonic history of the germ-cells of the loggerhead turtle (Caretta caretta). Papers Tortugas Laboratory 11: 313-44. Carnegie Institution of Washington Publication Number 251. The wide discrepancies in the published accounts of the origin and early history of the germ-cells in vertebrates provided the stimulus for this investigation. Twenty-five embryos of the loggerhead turtle (Caretta caretta), ranging from the second day (5 somites, 2mm. length) to the thirty-second day of incubation, were employed in this investigation. Results indicate that the primordial germ-cells migrate during the second day from the yolk-sac endoderm, where they are widely scattered caudally, into the lateral border of the area pellucida on each side of the embryonic disk. The germ-cells migrate by amoeboid activity. The migration period is not sharply limited. A certain number of germ-cells migrate out of the regular germ-cell route and go astray. The total number of primordial germ-cells counted in a 12-day embryo is 352. Occasional cells may divide by mitosis, or undergo degeneration, at any stage of their history or at any point of the route. No germ- cells were found contributing to the formation of the Wolffian duct. The germ-cells do not differ from young somatic cells in the character of their mitochondrial content. No transition stages between coelomic epithelial cells and germ-cells appear up to the 32-day stage. The evidence derived from a study of the Caretta embryos is in complete harmony with the idea of a single uninterrupted line of sex-cells from primordial germ-cells to odgonia and spermogonia, and with the hypothesis of a vertebrate Keimbahn or continuous germinal path. . 1908. The germinal spot in echinoderm eggs. Papers Tortugas Laboratory 1: 1-12. Carnegie Institution of Washington Publication Number 102. This paper reports the results of further studies of the prematuration stages of echinoderm eggs of additional species of echinoderms, a star-fish (Echinaster crassispina), and a brittle-star (Ophiocoma pumila). In Echinaster crassispina the chromosomes are derived exclusively from the nucleolus. In Ophiocoma pumila the chromosomes arise exclusively 46 183. 184. 185. from the nuclear reticulum. In some species the chromosomes arise from a chromatin- nucleolus, in others from a chromatic reticulum, and in still others in part from one source and in part from the other. The eggs of different forms differ in that some have only a chromatin-nuleolus, without distinct plastin ground-substance, resting in an achromatic nuclear reticulum (Echinaster); others possess both chromatin-nucleolus and plasmosome as well as a chromatic nuclear reticulum (Ophiocoma); and still others possess a double nucleolus (chromatin nucleolus and plastin ground-substance), with the chromosome complex gathered in a mass in the achromatic reticulum (Asterias). The function of the germinal spot then appears, in part at least, to be that of a storehouse of material which is to contribute to the formation of the chromosomes . . 1917. The history of the primordial germ cells in the loggerhead turtle. Proceedings of the National Academy of Science 3: 271-75. This study attempts to trace germ cell history in the loggerhead turtle and compare it to observations for other vertebrates. Embryos were collected for study from specimens taken on Loggerhead Key, Dry Tortugas during the Summer of 1914. The germ cell history of Caretta is very similar to that first described for Chrysemys and to that described for dogfish. . 1917. The microscopic structure of striped muscle of Limulus. Papers Tortugas Laboratory 11: 273-90. Carnegie Institution of Washington Publication Number 251. The study of the skeletal muscles of Limulus was undertaken with two chief objects in view: to test a conclusion suggested by earlier studies on the intercalated disks of vertebrate cardiac muscle, namely, that these disks are properly interpreted as "irreversible contraction bands" and to seek additional evidence in further refutation of the recently revived hypothesis that striped muscle can be interpreted in terms of "muscle-cells" and intercellular myofibrillae. It was found that both the skeletal and the cardiac muscles of Limulus consist of trabeculae of finely granular sarcoplam. In cardiac muscle the main trabeculae and their branches form a loose-meshed syncytium. Neither type of muscle contains mesophragmata. Very rarely an intercalated disk of the simple-comb type appears in the cardiac muscle. Both types are very similar in respect of the presence and arrangement, in the same phase of contraciton, of Q and J disks, and the telophragmata. The evidence is unequivocal against an interpretation of structure in terms of "muscle- cells" and intercellular myobibrillae. The nuclei of the growing muscles multiply by amitotic division. In essential structure the cardiac and skeletal muscles of Limulus are closely similar, indicating a close functional similarity. The structure serves, moreover, as a splendid illustration of the "law of biogenesis," in that it is practically identical with a stage in the early histogenesis of striped muscle of teleosts . . 1908. The relation of the nucleolus to the chromosomes in the primary odcyte of Asterias forbesii. Papers Tortugas Laboratory 1: 37-72. Carnegie Institution of Washington Publication Number 102. The primary object of this investigation was was to contribute to the subject of the relation between nucleolus and chromosomes during maturation. In summary, synizesis occurs in the odcyte of the first order at the very beginning of the growth-period (size of nucleus 5 microns). The growth-period is passed through rapidly. During the latter half of the growth-period all the chromatin, with the exception of what is held by the chromosomes, becomes stored in the enlarging nucleolus. The nucleolus consists of a plastin ground- substance infiltrated and covered over with chromatin. The chromosomes do not arise out of the nucleolus. The number of chromosomes in the prophase of the first polar mitosis is 18. They vary somewhat in size (one is considerably larger than the rest), all have a characteristic dumb-bell shaped appearance, and some are clearly double (bivalent). The 47 two maturation divisions effect a double longitudinal fission of the original bilobed chromosomes. The reduced number of chromosomes is again 18. Observations on Hipponoé esculenta agree in essential points with those made on Asteria forbesii and support the conclusions regarding the origin of the chromosomes, the function of the nucleolus, and the reduction phenomena. 186. . 1908. The spermatogenesis of Aplopus mayeri. Papers Tortugas Laboratory 1: 13-36. Carnegie Institution of Washington Publication Number 102. The object of the present investigation is primarily to trace the history of the accessory chromosome through the various stages in the process of spermatogenesis in the phasmid Aplopus mayeri. The material upon which the investigation is based was obtained from the Loggerhead Key, Florida. Primary spermatogonia divide both mitotically and amitotically. In the latter instance cell-division is frequently not consummated and a bi- or multi-nuclear cell results. In the first order of the secondary spermatogonia the accessory chromosome appears in the resting-stage. During synapsis the accessory chromosome lengthens into a club-shaped structure attached by its lesser end to the presynaptic thread, undergoes partial longitudinal division, closes up again during the height of synapsis, and returns again to its previous characteristic form and location in the nucleus of the growing primary spermatocyte. The second maturation division is equational, effecting a longitudinal division of univalent chromosomes. The accessory also divides equationally in the cells containing this element. A dimorphism of spermatozoa results; the accessory chromosome possessed by one-half probably represents a sex-determinant. The history of the accessory chromosome gives evidence that it at least possesses a strict morphological and probably also a physiological individuality. 187. Kaas, P. 1972. Polyplacophora of the Caribbean region. P. Wagenaar and L. J. Van Der Steen Hummelinck, 1-162. Studies on the Fauna of Curacao and Other Caribbean Islands, ed. P. Wagenaar and L. J. Van Der Steen Hummelinck. The Hague: Martinus Nijhoff. This paper includes Tables of Distribution of Polyplacophora of the Caribbean . The author took into consideration the whole of the Florida coast as far north as Fernandina, E. Florida and the Keys, the Dry Tortugas, W. Florida, the Gulf of Mexico, and also the Bermudas. Thiele's description of his single 5.5 specimen of Ischnochiton hartmeyeri from Bird Key Reef is translated into English here by Kaas. 188. Kale, H. W. 1985. Florida birds - Dry Tortugas. Florida Naturalist 58, no. 2: 6. A sighting of a great black-backed gull is made at the Dry Tortugas, and a scarcity of land birds is reported. 189. Kellner, Carl. 1907. Embryology of the appendicularian, Oikopleura. Zoological Anzeiger, Bd. 31: May. The appendicularia of the Dry Tortugas specimens of Salpae were collected. Appendicularia of the genus Oikopleura and their "houses" were examined and found in surface waters. Their anatomy and histology are described. 190. Kille, F. R. 1936-1937. Regeneration in holothurians. Carnegie Institution of Washington, Year Book. Note: published as follows: 1936, v. 35, p. 85-86; 1937, v. 36, p. 93-94. Histological studies were conducted on sea-cucumbers of the genus Holothuria, to determine the manner in which the digestive system is reconstituted following autotomy by means of electrical stimuli. 191. Kopac, M. J. 1936. Electrical resistance of Valonia. I. Changes in the resistance with time in impaled coenocytes. Papers Tortugas Laboratory 29: 359-86 (issued Mar. 1936). 48 Carnegie Institution of Washington Publication Number 452. During the summers of 1933 and 1934 the author worked at the laboratory of the Carnegie Institution located on Loggerhead Key, Dry Tortugas, Florida. Several species of Valonia were found growing abundantly on the various coral reefs of the Dry Tortugas. A study of the electrical resistance of impaled Valonia coenocytes by using a technique more highly refined that that employed by previous investigators was initiated. Although only V. ventricosa was used in this study, the methods developed and used here may be extended to the study of other species of Valonia. Glass microcapillaries, with tips ranging from 0.025 to 0.1 mm. in diameter and filled with vacuolar sap, served as microsaltbridges leading from the vacuole to a calomel half-cell. A larger glass tube (the macrosaltbridge), filled with sea-water, was used as a saltbridge leading from the sea-water surrounding the coenocyte to another calomel half-cell. The Valonia coenocytes were impaled on the tip of the microsaltbridge with the aid of a micromanipulator. It was found that coenocytes with a high chloroplastid density have a constant Rp several times higher than coenocytes with a low chloroplastid density. It is postulated that only the inter-chloroplastidal protoplasm is capable of conducting a current. The average initial Rp in type A punctures was 60 to 65 per cent of the constant Rp. In some coenocytes a constant Rp was reached in a few minutes. This increase in Rp is caused largely be the redistribution of those chloroplastids around the microtip which were disturbed by the puncture. The average initial Rp in type B punctures was 2 to 3 per cent of the constant Rp. After the chloroplastids are redistributed in this hyaline zone, the disintegrated chloroplastids are extruded, and the tiny vauoles are eliminated, no further increase in Rp takes place . 192. Kunkel, B. W. 1934. The selective action of certain adverse environmental conditions on the hermit crab (Clibanarius tricolor Gibbes). Papers Tortugas Laboratory 28: 215-44 (issued Aug. 1933). Carnegie Institution of Washington Publication Number 435. The problem of selection is undoubtedly a very complex one. The characters which enable one organism rather than another to survive are difficult to ascertain; a favorable variation of one part may be accompanied by an unfavorable variation of another, so that selection may have no effect upon the first feature. The present study has to do with a phase of the selection problem which, on the whole, has received rather scant attention from investigators. The selective effect of certain adverse conditions on a population has been studied. The problem is that of determining how a given species may respond to a change in environment, of determining the morphological difference between those individuals which succumb to a certain change in the normal environment and those which are able to withstand the change. The material upon which the present study is based was collected and the experiments were made at the Tortugas Laboratory. The small hermit crab Clibanarius tricolor Gibbes was selected for the experiments. 193. LeCompte, M. 1937. Some observations on the coral reefs of the Tortugas. Carnegie Institution of Washington, Year Book 36: 96-97. Particular attention was paid to the distribution and adaptation of the corals on the reef west of Loggerhead Key. A baseline of about 2500 yards is verified, documenting large heads of Orbicella (Madrepora) annularis, extensive growths of Acropora, and areas of gorgonians and algae. Beach rock development, coral feeding habits, and the effects of boring animals on corals are discussed. 194. Leitch, James L. 1936. The water exchanges of living cells. II. The application of a photographic method to the determination of the non-solvent volume of the eggs of Echinometra lacunter. Papers Tortugas Laboratory 29: 349-58 (issued Mar. 1936). Carnegie Institution of Washington Publication Number 452. Photography has been applied to the study of living cells and tissues. Leitch raised the 49 question of the feasibility of a photographic method in the study of the osmotic behavior of cells. The present paper outlines such a method and discusses some of the factors involved in the study of the water exchanges of the eggs of the sea-urchin, Echinometra lacunter, using measurements of photographs of eggs at equilibrium in dilute sea-water solutions. It was shown that photography can be employed in the study of the water exchanges of living cells. The non-solvent volume of the eggs of Echinometra lucunter is 36 per cent when calculated after from 60 to 90 minutes’ exposure to experimental solutions. Longer exposures to the experimental solutions result in a higher non-solvent volume of 48 per cent which is associated with pronounced vacuole formation. The appearance of vacuoles after the attainment of the first equilibrium is discussed and three different explanations proposed. . 1937. The water exchanges of living cells. IV. Further studies on the water relations of the eggs of the sea-urchin, Echinometra lacunter. Papers Tortugas Laboratory 31: 53-70 (issued July, 1936). Carnegie Institution of Washington Publication Number 475. The application of a photographic method to the determination of the non-solvent volume of the eggs of the sea-urchin, Echinometra lacunter, and also the effect on the non-solvent volume determinations of the length of time of immersion of eggs in diluted sea-water solutions, was demonstrated in another publication (Leitch,1936). The present paper considers the utilization of this photographic method for the study of the swelling and shrinking of the eggs of the same sea-urchin, the effect on the water relations of these cells of the time between that of spawning and that of introducing the eggs into the experimental solutions, and an analysis of equations which have been developed and applied by several investigators to explain the kinetics of water exchanges of living cells. Results showed that an analysis of the equations developed to interpret the kinetics of water exchanges of cells the so-called permeability "constants" are not constant for the eggs of Echinometra lacunter but vary with the dilutions of sea-water used and also with different intervals of time in the same dilution. The permeability constants for swelling and shrinking do not coincide, being between 0.250 and 0.650 for the former process and between 0.180 and 0.580 for the latter. The introduction of the correction for the non-solvent volume into the equations does not produce a better agreement between the constants for the two processes. The use of the photographic method (Leitch, 1936) is further substantiated for the determination of non-solvent volumes and is extended to the study of the swelling and shrinking of eggs. Approximately two hours from the time of spawning the non-solvent volumes is greatly increased, from 30 to 53 per cent of the initial volume. There is a slight retarding effect on the water exchanges of the eggs brought about by standing. The time at which the effect of standing at room temperature appears in the values of the non-solvent volume and rate of penetration of water is correlated with a sharp decrease in the percentage of development, a slight increase in the volume of the eggs and a cytolysis-like phenomenon which finally ends with the complete disintegration of the eggs. The production of fertilization membranes as a criterion of non-injury of the egg cell is shown to be inadequate and the percentage of development of normal larvae is urged as a better test of normality . 195. 196. Lessios, H. A., D. R. Robertson and J. D. Cubit. 1984. Spread of Diadema mass mortality through the Caribbean. Science 226: 335-37. Populations of the ecologically important sea urchin Diadema antillarum suffered severe mass mortalities throughout the Caribbean. This mortality was first observed at Panama in January 1983; by January 1984 it had spread to the rest of the Caribbean and to Bermuda. The sequence of mortality events in most areas is consistent with the hypothesis that the causative agent was dispersed by major surface currents over large distances. However, some of the late die-offs in the southeastern Caribbean do not fit this pattern. Several lines 50 of indirect evidence suggest that the phenomenon is due to a water-borne pathogen. If so, this is the most extensive epidemic documented for a marine invertebrate. 197. Linton, E. 1908. Helminth fauna of the Dry Tortugas. I. Cestodes. Papers Tortugas Laboratory 1: 198. 199. 200. Lipman, 201. ———. 157-90. Carnegie Institution of Washington Publication Number 102. This report is based on data collected at the Marine Biological Laboratory, Tortugas, Florida, June 30 to July 18, 1906. A list of the hosts which were examined for parasites, and a summary of the results of that examination, together with a few food notes are presented. A few extracts from notes made at the time the material was collected are presented. Acanthocephala are presented. The species found in the frigate mackerel was Echinorphynchus pristis. Few nematodes were found in the nurse-shark. New species of parasites are described. . 1910. Helminth fauna of the Dry Tortugas. II. Trematodes. Papers Tortugas Laboratory 4: 11-98. Carnegie Institution of Washington Publication Number 133. The collection here described was made at the Marine Biological Laboratory, Tortugas, Florida, in the summers of 1906, 1907, and 1908. The fishes examined were from the shallow waters of the reef. The distribution of parasites together with food notes have already been published in the Year Book of the Carnegie Institution of Washington for the years above named. This paper includes a list of Tortugas trematodes and their hosts, key to the genera and species described, and descriptions of species, habitats, etc, including: trematodes from loggerhead turtles and from fish. . 1907. Note on the habits of Fierasfer affinis. American Naturalist 41, no. 481: 1-4. Observations of the Fierasfer affinis entering its host, tail first are made. C. B. 1929. The chemical composition of sea water. Papers Tortugas Laboratory 26: 249- Vile Carnegie Institution of Washington Publication Number 391. In his studies on marine bacteria and related subjects, the author realized the need for more accurate and complete analytical data on the inorganic components of sea-water and determined to obtain them.. Two samples were analyzed from the Atlantic area and they were both from the Gulf Stream, and taken near Loggerhead Key in the Tortugas. The data render it clear that a large part of the ions important to algae are removed from solution in sea-water by a rise in pH of that medium, which is well within the range of daily rise in pH of sea-water carrying an active algal flora under the proper conditions of light and temperature. 1924. A critical and experimental study of Drew's bacterial hypothesis on CaCO; precipitation in the sea. Papers Tortugas Laboratory 19: 179-91. Carnegie Institution of Washington Publication Number 340. Based on a series of experiments to explain the precipitation of CaCO; in sea water, it was found that there are several ways to explain CaCO3, where it occurs in seawater, without introducing Drew's hypothesis or any other bacterial hypothesis. These explanations of the phenomenon seem adequate to account for the qualitative and quantitative differences in CaCO; as found under different conditions in seawater. Changes in water and air temperatures, and marine plant activity, which Drew and others clearly appreciated and understood, yet have introduced a purely gratuitous bacterial hypothesis based on what appears to be sound experiments. 51 202. . 1929. Further studies on marine bacteria with special reference to the Drew hypothesis on CaCo; precipitation in the sea. Papers Tortugas Laboratory 26: 231-48. Carnegie Institution of Washington Publication Number 391. Bacterial populations in the open sea are very small. Mixed or pure cultures of organisms isolated from sea-water are incapable of precipitating CaCO; from sea-water to which no salts have been added, or even in the presence of added KNO3 Mixed or pure cultures of such organisms are incapable of precipitating CaCO; in a sea-water medium if KNO; and organic matter as the sugars or similar forms free from calcium are added to the medium. Upon the basis of evidence in this and in earlier papers the Drew hypothesis is shown to be untenable, and at the very least uproved. This strong probability is reemphasized in the purely physical-chemical nature of CaCO; precipitation on a large scale in nature. 203. Locker, S. D. A. C. Hine and E. A. Shinn. 1991. Sea level geostrophic current control on carbonate shelf-slope depositional sequences and erosional patterns, South Florida platform. AAPG Bulletin 75, no. 3: 623. High-resolution seismic reflection profiles across the shelf-slope margin between the Dry Tortugas and Key West, Florida, indicate that sea-level fluctuations and the eastward flowing Florida Current are major controls on late Quaternary sequence stratigraphy. The study area, a transition zone between the open south Florida shelf and the lower Florida Keys island/reef system, is typified by a shallow shelf with reef margin adjacent to a deeper lower-shelf/slope. The lower-shelf/slope is composed of stacked or prograding sequences that downlap and pinchout on the Pourtales Terrace. Strike oriented stratigraphic sections exhibit many sea-level controlled features such as lowstand erosion, transgressive unconformities, and highstand system tracs. Lowstand reefs, notches, or barriers are observed as deep as 150m below present sea level. Depositional styles change along-slope from west to east. The western portion of the study area is characterized by thick, low amplitude prograding sequences related to abundant supply of sediment through off-shelf transport during high sea-levels as well as along-slope reworking by the Florida current. Part of this section has been severely eroded by along-slope current producing localized cur and fill structures and widespread erosional unconformities. To the east, a thinner section of high-amplitude reflections is common seaward of the lower Florida Keys reef tract system. Again, along-slope current erosion and winnowing of sediment supplied by the adjacent margin is evident. This study provides new evidence of how a strong geostrophic boundary current along with fluctuating sea levels have interacted to control depositional sequences on a carbonate slope in the Florida/Bahamas platform complex. 204. Longley, W. H. 1917. Changeable coloration in Brachyura. Proceedings of the National Academy of Science 3: 609-11. Studies on changes in the color of brachyuran crabs (Ocypoda and Callinectes sp.) at the Marine Laboratory, and in the field on Loggerhead Key, Dry Tortugas, demonstrate adaptive coloration based on temperature variations and the color of the substratum upon which the specimen is resting. It is expected that in future studies the same general rules of adaptation for fishes will apply to crabs. 205. . 1918. Haunts and habits of tropical fishes. American Museum Journal 18: 79-88. Observations are recorded at the Dry Tortugas using underwater photography. Habits of the shallow water reef species were photographed in water less than 10 feet deep. Emphasis is placed on the biological significance of color in fishes with their surroundings. Fish color change may be evoked by offering them food by hand at different locations. The foods and feeding habits of reef fish are discussed in this report. 206. ————. 1918. Marine camoufleurs and their camouflage: the present and prospective significance of facts regarding coloration of tropical fishes. Smithsonian Report (1920): 475-85. a2 207. 208. 209. Fish are used as an example of an animal which uses color patterns, not as a struggle for existence as hypothesized by Darwin, but as a means of expressing its biological significance by displaying a natural system of camouflage. Some colors in fishes are not changeable, but seem to be correlated to definite habits. In the case of those that are changeable, there is conclusive evidence that they are displayed under specific conditions. For example, transverse bands are shown when the species is inactive. However, upon movement the bands are replaced by stripes. . 1916. Observations upon tropical fishes and inferences from their adaptive coloration. Proceedings of the National Academy of Science 2: 733-37. The conception that species have been multiplied by divergent evolution of related strains is based on many observations. If the Darwinian hypothesis is true, the character of organisms should be largely of an adaptive sort, but its adherents have been content to support this position by inputting utility to structure and habits. It has not been proved in fishes that some color combinations ward off enemies nor that pigmentation is functionally conspicuous. Many of the brightly colored fishes of the Tortugas have been studied to evaluate their coloration objectively. Most species exhibit countershading with darkest shading on the upper surface and lighter shading on the mid-ventral or lower line. Thirteen species of fish studied exhibit color changes based on their surroundings observed from boats or from the bottom using diving equipment and photography. Correlation of color with habitat has been documented. Some examples suggest that red fish are rarely seen during the day, gray fish with diurnal activity patterns are found near large coral heads, lighter blue fish are habitually found swimming well above the bottom in moderate depths, and those species largely found over grass beds are of green color or have a green color phase. As far as this class of animals is concerned, Longley postulates that there is no ground for the belief that bright color is correlated any way with armament or distastefulness. Problems of mimicry resemblance are unresolved, however the observations presented in this abstract undermine many speculative explanations of animal coloration in terms of natural selection and replace them with something which may not be dismissed from consideration. . 1936. Species studies and the species problem. American Naturalist 70: 97-109. (No abstract available). . 1917. Studies upon the biological significance of animal coloration. I: The colors and color changes of West Indian reef fishes. Journal of Experimental Zoology 23: 536-601. Studies were carried out at the Dry Tortugas to determine the biological significance of changes in color of reef fishes. It was found that fishes are countershaded; color changes, which are common even among the most gaudy, tend to assimilate them with their environment; and in general, their colors repeat those of their surroundings. Specially defended types are not unlike others in pigmentation, nor inferior to them in their ability to effect adaptive color adjustments. Finally, there is no evidence that brightly colored species enjoy greater immunity from attack than their fellows, for they constitute a large proportion of the food and may be readily identified in the stomach contents of predaceous forms. These statements, which rest upon a great body of verifiable observations, are consistent with the Darwinian hypothesis, but inconsistent with the assumption that animals of high color possess more than minimal conspicuousness under natural conditions. They impel one to reject the hypotheses of warning and immunity coloration, signal and recognition marks, and sexual selection, at least in so far as they may ever have been supposed to apply to these forms. Upon the contrary, they confirm Thayer's conclusions regarding the obliterative function of color and pattern, emphasize the common occurrence of adaptive characters among animals, and suggest that their evolution has been guided throughout by natural selection. 53 210. . 1917. Studies upon the biological significance of animal coloration. II: A revisional working hypothesis of mimicry. American Naturalist 51: 257-85. In this report, various hypotheses proposed by the author and other investigators relating changes in animal coloration in relation to habits are discussed. The author postulates that bright colors of tropical fishes are correlated with the animal's habits from work achieved at the Dry Tortugas. Other coloration hypotheses are provided dealing with butterflies and lizards, as well as warning coloration in bright and dull-colored insects. These ideas submitted by the author constitute working hypotheses to be tested by other biologists. 211. Longley, W. H. and S. F. Hildebrand. 1940. New genera and species of fishes from the Tortugas, Florida. Papers Tortugas Laboratory 32: 223-85 (issued Sept. 1940). Carnegie Institution of Washington Publication Number 517. Thirty new genera and species of fishes described in these pages resulted from studies carried on for many years, principally at Tortugas, Florida, by the late Dr. William H. Longley, whose untimely death occurred before he had fully completed a study of his collections and a manuscript embodying a complete account of his field observations. The present writer has made further studies of the Tortugas collections, and has endeavored to extract interesting facts from Dr. Longley's notes on those species not treated in his unfinished manuscript. 212. Longley, W. H. and S. F. Hildebrand. 1941. Systematic catalogue of the fishes of Tortugas, Florida; with observations on color, habits, and local distribution. Papers Tortugas Laboratory 34: 1-331. Carnegie Institution of Washington Publication Number 535. Observations on the fishes of the Tortugas Atoll were made by the senior author spanning a period of over 25 years. An inventory of 442 species is included, covering a wide range of habitats: bare sand, seagrass beds, coral reefs, channels between the keys, and deep waters over 600 feet in depth a few miles southward. Over 300 species were associated with coral reef habitat. This study represents the first fish survey conducted on the Florida Reef Tract. Field observations were made largely with the use of a diving helmet, which enabled the investigator to observe and photograph the fish in their native habitat, and to give information as to their behavior, feeding and habits, and especially adaptive coloration. Much of Longley's work is documented by the world's first underwater color photography. Following the death of Dr. Longley, Dr. Hildebrand undertook the editing and the completion of the manuscript. Material added by him bear his initials. 213. Lucké, B.. 1937-1938. Studies on the tumors of fishes (of the snapper family Lutjanidae) . Carnegie Institution of Washington, Year Book. Note: published as follows; 1937, v.36, p. 98-99; 1938, v.37, p. 92-94. Certain kinds of tumors found on thirty nine fish belonging to several species of snappers closely resemble human neoplasms arising from nerves. They arise in the subcutaneous tissue and appear as flattened oval masses. No tumors of this kind were found on other species of fish. Epithelial growths were found in thirty specimens of Halichores radiatus, from a total of six thousand fish observations. Multiple papillomas of the skin and the eye were reported in a green turtle caught off Cape Sable. 214. ————. 1942. Tumors of the nerve sheaths in fish of the snapper family (Lutjanidae). Archives of Pathology 34: 133-50. Fish of the snapper family Lutjanidae are commonly afflicted with tumors which resemble the nerve sheath tumors of man called variously neurinoma, neurolemmoma, schwannoma, or neurofibroma. Neoplasms of this kind have been observed in 76 fish of three species, the gray snapper (Lutjanus griseus), the dog snapper (L. jocu), and the schoolmaster (L. apodus). Most of the fish were collected from the Dry Tortugas. Many other fish families 54 were examined, however no tumors of the kind were found. The tumors generally were found along the course of the subcutaneous nerves, particularly of the head and dorsal regions, as solitary or multiple, relatively large firm white masses. Like human neoplasms, the tumors of fish are usually composed of two kinds of tissue: one compact and richly fibrocellular; the other loose reticulated and poorly cellular. The component cells and intercellular fibers of the tumors appear to be essentially the same, and arranged in similar patterns, in fish and man. Unlike human tumors, the fish tumors, though well circumscribed, are usually not encapsulated. Nerve sheath tumors appear to be more common in certain fish species than in man. The frequency of occurrence of these tumors, which can be maintained for long periods in marine aquariums, renders them favorable material for studies of neoplasms. 215. Lucké, B. and H. G. Schlumberger. 1949. Neoplasia in cold-blooded vertebrates. Physiological Reviews 29, no. 2: 91-126. This review complements an earlier review to source material, abstracts of all the reports in the literature dealing with tumors in fishes, reptiles, and amphibians. In regards to fishes of the Tortugas, a review is presented on the senior author's work on tumors of nerve tissue, as described in Lucke (1942, reference no. 214). 216. Lynts, G. W. 1968. Analysis of recent foraminiferal fauna from the Dry Tortugas, Florida. (abs.). Geological Society of America Special Paper 101: 128-29. This analysis of total foraminiferal population is based upon 16 samples collected in 1960 from the Dry Tortugas, Florida. Fourteen samples represent reoccupation of stations sampled by Cushman (1922) in his original description of the foraminiferal fauna. Q- modal factor-vector analysis indicates that the fauna is characterized by three assemblages (factors) which account for 89 per cent of the total information (sum of squares of all entries in data table). In general, these assemblages are characterized by a few dominant species. One of the assemblages, Assemblage III, is directly related (r= +0.911) to depth of water. The total population of the 14 samples representing reoccupation of Cushman's stations were compared with the total population indicated by Cushman (1922). Comparison was made using F-ratios and percentage of number of species occurring in both samples (Sc). F-ratios estimate degree of variation between samples, whereas Sc measures variation in species composition. F-ratios indicated that at seven of the 14 stations there were significant differences in total population between the 1919 and 1960 collections. Sc's ranged from 18.3 to 56.9 per cent and showed no relationship to degree of variation estimated by F-ratios. This variation in foraminiferal fauna between collections may represent either real changes in populations or apparent variations. If variations are real, they reflect changes in the ecosystem over the 41 years. If variations are apparent, they may represent either inability to resample microhabitats or vagaries in taxonomic discrimination. 217. Lyons, W. G. 1980. Polyplacophora of Dry Tortugas Florida with comments on /schnochiton hartmeyeri. Bulletin of the American Malacological Union, Inc. 46: 34-37. 450 specimens and 14 species of chitons were collected during 1978-79 from a station near Garden Key. Only Acanthochitona sp. and Stenoplax purpurascens were relatively common. 218. Mann, A. 1936. Diatoms in bottom deposits from the Bahamas and the Florida Keys. Papers From the Tortugas Laboratory 29: 121-28. Note: This is Appendix 1 to Calcareous shallow water marine deposits of Florida and the Bahamas by Eldon Marion Thorp . The twenty-four samples of calcareous sand collected by Doctor Vaughn in 1914 between Cape Florida and Key West and at Tortugas were examined at the time these samples were . . | | q 25) received. A list of stations at which diatoms were collected is given, including Tortugas, with a list of the diatoms found. Species of the genus Mastogloia are very abundant in these Florida samples, and the author has found them to be so in all collections from Florida Waters. In other parts of the world they are relatively much less abundant. 219. Manter, H. W. 1942. Gasterostomes (Trematoda) of Tortugas, Florida. Papers Tortugas Laboratory 33: 1-19 (issued June, 1940). Carnegie Institution of Washington Publication Number 524. A report of the Gasterostomatus trematodes collected in 1930, 1931, and 1932 is given here. Fifteen species are reported; nine are considered new. 220. . 1934. The genus Helicometra and related trematodes from Tortugas, Florida. Papers Tortugas Laboratory 28: 167-80 (issued Mar. 1933). Carnegie Institution of Washington Publication Number 435. Observations on the trematode genera Helicometra, Helicometrina and a new related form are based on material collected at the Carnegie Biological Laboratory at Tortugas, Florida. The genus Helicometra is represented at Tortugas by three species, H. execta, H. torta, and H. fasiata. The characteristics of each of these are described. A key is given to the species of the genus. H. execta is recorded from 6 additional hosts, making a total of 10. A mutilated specimen of H. torta showed this species has little or no power of regeneration. Helicometra fasciata from three hosts at 50 to 60 fathoms is reported for the first time from America. Metacercariae of Helicometrina nimia are described encysted in the muscles of the shrimps, Lysmata intermedia and Crangon formosum. Cercariae from Columbella mercatoria identified as Cercaria J of Miller were found to encyst readily in the muscles of Lysmata intermedia. Helicometrina parva, a new species is described . 221. ————. 1934. Some digenetic trematodes from deep-water fish of Tortugas, Florida. Papers Tortugas Laboratory 28: 257-345 (issued Jan. 16, 1934). Carnegie Institution of Washington Publication Number 435. The parasitic fauna of ocean depths is practically unknown. Extensive fish population occurs at all depths, very little study has been made on the helminths of these fish. During the summers of 1930, 1931, and 1932 collections were made of parasites from fish trawled from depths varying from 40 to 582 fathoms at Tortugas, Florida. Most of these hauls were made about 10 miles south of Loggerhead Key. Fish taken from these depths were commonly parasitized by helminths and especially by trematodes. It was found that the trematode fauna of the deep-water fish is practically as abundant and as varied as is the rich trematode fauna of the reef fish. A description is given of 49 species of trematodes collected from approximately 90 species of fish from depths of 40 to 582 fathoms. 721 individual fish were examined. Approximately 80 per cent of the host species were infected with trematodes, a percentage comparable with the degree of infection found in fish of shallow water. One new subfamily (of the family Heterophyidae), 11 new genera and 33 new species are described. Seven species of trematodes, from deep water only at Tortugas, are identical with forms well known from northern regions. Studies from shallow water at Tortugas show practically no similarity to northern forms. The deep-water trematode fauna is more like the surface fauna of Maine, Great Britain or Norway than like the shallow-water fauna at Tortugas, only a few miles away. This tendency to resemble surface trematodes of cold-water regions suggests that temperature is an important factor in the distribution of marine fish trematodes. This study emphasized the fact that a gradient of changing environment (such as depth) is reflected not only in the free-living population of a region but also in their parasites. 222. Marsh, G. 1940. The effect of light on the inherent E. M. F. of Valonia ventricosa. I. Intensity and time relations. Papers Tortugas Laboratory 32: 65-84 (issued Oct. 1939). 56 Carnegie Institution of Washington Publication Number 517. The interpretation of the electrical changes produced in green plants by light has been retarded by the confused nature of the published results. In order to interpret electrical changes in a tissue in terms of some underlying process it is essential to obtain precise information concerning the distribution of E.M.F. within the tissue and the conditions of summation of the potentials of the individual cells included in the electrical circuit. The present paper reports the effect of visible light at known intensities upon the inherent E.M.F. of the coenocytic alga Valonia ventricosa. Results indicated that when the intensity of incident light is altered, the inherent E.M.F. of an impaled Valonia cell undergoes a characteristic cycle of change with definite time relations, following which a steady level is reached. The steady level of E.M.F, plotted against the logarithm of the light intensity rises from the dark potential along a sigmoid curve to a maximum at about 250 foot-candles, then descends along a similar curve toward the dark potential. The decline in potential with light intensity beyond the maximum was reversible. No injury was detected at any intensity. The effect of intermittent light with equal light-dark periods was similar to that of continuous light of half the intensity, save in one experiment, wherein the effect was similar to that of continuous light of increased intensity. It is concluded that the effect of light on the E.M.F. is due primarily to the release of oxygen in photosynthesis. 223. ————. 1940. The effect of light on the inherent E. M. F. of Valonia ventricosa. II. The relative energy absorption spectrum. Papers Tortugas Laboratory 32: 99-120 (issued May 1940). Carnegie Institution of Washington Publication Number 517. : The interest in the relative effectiveness of different wave-length bands of visible light upon bioelectric potentials centers about the question of the nature of the agent in the living cell which absorbs the radiant energy, and its contribution to the electromotive mechanism. For the green plants three principal lines of evidence have been adduced to support the conclusion that chlorophyll is the photosensitive agent. Results indicated that the steady E.M.F. of impaled Valonia ventricosa illuminated with light of limited spectral composition was matched with white light. The ratio of the intensity of white to that of filtered light for an E.M.F. match was independent of the magnitude of the E.M.F. matched and of the absolute intensities. The relative energy absorption is shown to compare fairly well for the filter series with the relative absorption of chlorophyll mixtures over the same spectral range calculated from the determinations of the absorption coefficient published by Zscheile. It is concluded that chlorophyll is the photosensitive material absorbing the radiant energy responsible for the effect of light upon the protoplasmic E.M.F. in Valonia. The chlorophyll system is, therefore, an intimate part of the electromotive mechanism. 224. . 1937. Effect of temperature upon the inherent potential of Valonia. Papers Tortugas Laboratory 31: 1-16. Carnegie Institution of Washington Publication Number 475. The effect of temperature upon the potential is of prime importance in the determination of the nature of the underlying electrochemical process. The electromotive force of a system in thermodynamic equilibrium (including the diffusion potential) is proportional to the absolute temperature (Q 10 of 1.04 or less within the biological range of temperatures). The E.M.F. found across the protoplasmic layer of Valonia is not a thermodynamic one. It is produced by an oxidation-reduction system in flux equilibrium at phase boundaries within the cell. The E.M.F. is not primarily determined by the external medium. The influence of the salt content of the sea-water upon the inherent potential is fundamentally no different from the influence of the composition of the medium upon any other biological process, as respiration, irritability, contractility, etc., where specific electrolytes in different proportions condition, but do not cause, the process. S/ 225. Mast, S.O. 1911. Behavior of the loggerhead turtle in depositing its eggs. Papers Tortugas Laboratory 3: 63-67. Carnegie Institution of Washington Publication Number 132. The nesting behavior of a single loggerhead turtle is described. 226. Matthai, G. 1915. Preliminary report on the comparative morphology of the recent Madreporaria around Tortugas. Carnegie Institution of Washington, Year Book 14: 209. General observations were recorded on the common corals of the Tortugas. The only species that extruded larvae was Favia fragum. 227. Mayer, A. G.. 1908. The annual breeding swarm of the Atlantic Palolo. Papers Tortugas Laboratory 1: 105-12. Carnegie Institution of Washington Publication Number 102. The habits of the "Atlantic palolo" are quite similar to those of the palolo worm of Samoa and the Fiji Islands. The worms are, however, specifically different, the Atlantic palolo being Eunice fucata Ehlers, and the Pacific worm E. viridis Gray. The annual swarming of the Atlantic palolo has been observed only at Tortugas, Florida, although the worm is abundant in the Bahamas and other parts of the West Indies. 228. . 1911. The converse relation between ciliary and neuro-muscular movements. Papers Tortugas Laboratory 3: 1-25. Carnegie Institution of Washington Publication Number 132. A series of experiments on marine invertebrates were conducted beginning at the Tortugas Laboratory, and later at Woods Hole, Mass, and the New York Aquarium involving the effects of ions of blood salts, magnesium, sodium, calcium, ammonium, potassium, and hydrogen on neuro-muscular systems in relation to maintaining ciliary movements. In each case they are the exact apposite of their effects upon ciliary movements of invertebrates studied. Studies were carried out on invertebrate organisms abundant at Tortugas including annelid larvae, Limulus, veligers , actinian larvae, larvae of the Atlantic palolo worm, Eunice fucata, and ctenophores, Cassiopea. Preliminary reports of the research were published in the Biological Bull., Woods Hole, v. 17 (341-342); in the Proceedings of the Soc. for Experimental Biology and Medicine, 1909, No. 7, (19-20), and in the Carnegie Year Book for 1909, p. 152. 229. ————. 1914. The effects of temperature upon tropical marine animals. Papers Tortugas Laboratory 6: 1-24. Carnegie Institution of Washington Publication Number 183. Tropical marine animals commonly live within 5° C. of their temperature of maximum activity and within 10° to 15° C. of their upper death temperature. In marine tropical forms even a few degrees of heat or cold cause a marked depression in movement. In tropical Scyphomedusae this depression of movement appears to augment about as the square of the change in temperature from that of the optimum. Time is an important factor in these experiments, for animals can withstand a higher degree of heat if the temperature be raised quickly than if it be raised slowly. It appears that the reef corals at Tortugas, Florida, live in water which is commonly within 10° C. of their upper death-temperature, and if the ocean were heated to 38° C. (100.4° F.) only one species, Siderastraea radians, could survive. Next to Siderastraea radians the most resistant coral is S. siderea. It is associated in its habitat with Orbicella annularis one of the most sensitive of the reef corals, which is killed at 14.1° and 36.8° C. In general, however, the corals of the shallow-reef flats, such as Siderastraea radians, Porites furcata, and Maeandra areolata, are the most resistant both to heat and cold, while those of deep water, such as Madrepora palmata , Eusimilia knorri, and Oculina diffusa, are the least resistant. As a result, we are led to conclude that were the water cooled by an exceptionally prolonged norther to 13.9° C. for 9 hours, 58 230. 23s Siderastraea radians , S. siderea, and Maeandra areolata would survive without apparent injury while Porites furcata, P. clavaria, Maeandra clivosa, and Favia fragum would also survive, but with more or less injury. This temperature would be fatal to Orbicella annularis, Porites astraeoides, and Madrepora muricata (cervicornis). . 1922. Hydrogen-ion concentration and electrical conductivity of the surface water of the Atlantic and Pacific. Papers Tortugas Laboratory 18: 61-85. Carnegie Institution of Washington Publication Number 312. The hydrogen-ion concentration of sea-water was determined by placing 0.4 c.c. of 0.1 per cent of the red dye thymolsuphonemphthalein in 70 per cent alcohol, in a test-tube of resistance glass, 24 mm. in caliber, then adding sea-water so as to make up 30 c.c. of solution. A series of such tubes, ranging from 7.95 to 8.3 pH, was standardized by Professor J.F. McClendon and presented to the author who restandardized these tubes at intervals of two years by comparison with determinations of pH made by a Leeds and Northrup potentiometer. In order to avoid writing negative exponents, the symbol "pH" to indicate the negative logarithm of the hydrogen-ion concentration was devised. Despite its artificiality, one soon finds that the pH system gives a clearer idea of the alkalinity or acidity of a solution than does a direct expression of the hydrogen-ion concentration. In testing water, pH 7 would indicate practical neutrality; pH above 7, alkalinity; and below 7 acidity. The carbon-dioxide tension of the sea-water was calculated from the pH and the temperature by the method devised by McClendon, Gault, and Mulholland (1917, Carnegie Inst. Wash. Pub. No. 251, p. 36). McClendon found that the pH of sea-water falls 0.01 for 1° C. decline in temperature. The salinity of the sea is expressed in grams of total salts per 1,000 grams of sea-water, and was determined by the well-known method of using a standard AgNO; solution with K,CrO, as an indicator, and testing against a sample of standard sea-water obtained from Professor Martin Knudsen. Upon being taken from the sea, the water was tested for temperature and pH, and a sample was preserved for determination of salinity. In connection with these tests of hydrogen-ion concentration, the electrical conductivity of the sea-water off Tutuila, Samoa and Tortugas, Florida was determined by Kohlrausch's method. At Tortugas, Florida, the conductivity of sea-water having 20.06 grams of chlorine in 1,000 grams of water, corresponding to a salinity of 36.24, was determined by the same apparatus, and with a portion of the same KCI solution used in Samoa. In lagoons such as that of Tortugas, Florida, and in closed shallow areas, McClendon found there was a diurnal variation in the pH, the water becoming more alkaline by day and relatively acid during the night. This was attributed to the effect of photosynthesis by plant life, which is active in daylight but ceases during the night. Over shallow regions, where the water may become impounded in tide-pools at low tide, the effect of photosynthesis is often very marked, the pH changing greatly while the temperature may change but little. The rise in pH was due to the loss of Co) resulting from photosynthesis. . 1914. The law governing the loss of weight in starving Cassiopea. Papers Tortugas Laboratory 6: 55-82. Carnegie Institution of Washington Publication Number 183. The medusae were always starved in the purest sea-water which was either dipped from the ocean in glass or canvas buckets or pumped into glass reservoir tanks through hard-rubber pipes by means of a hard-rubber pump. The medusae were starved side by side in one and the same glass aquarium, but when this was impossible the aquaria were of similar size and form and were placed side by side, so as to be subjected to similar environmental changes. The decline in weight of two normal medusae of Cassiopea xamachana starved each in one liter of sea-water, changed once in 24 hours, and kept in the diffuse daylight of the laboratory at Tortugas, Florida, from June 8 to 20, 1913. One medusa, A, was starved in sea-water which had been passed through two glass funnels each holding two sheets of 232% 233% 234. 235. 236. ZBI 59 Chardin filter paper. The other medusa, B, was starved in sea-water, which, in addition to having been filtered through the Chardin Filters, was also filtered through a bacteria-proof porcelain filter. It appears that all food had been removed from the water by Chardin filters and the medusa in the bacteria-free sea-water starved more slowly than the one in the sea-water which had not been passed through the porcelain filter. . 1908. Marine laboratories, and our Atlantic coast. The American Naturalist 42: 533-36. In this general article by Dr. Mayer concerning the importance of marine laboratories along our Atlantic Coast, the Tortugas is mentioned as having a certain remoteness from the busy world and consequent freedom from interruption peculiarly favorable to the conduct of research. . 1915. The nature of nerve-conduction in Cassiopea. Proceedings of the National Academy of Science 1: 270-274. Narrative same as in reference no. 234. . 1917. Nerve-conduction in Cassiopea xamachana. Papers Tortugas Laboratory 11: 1-20. Carnegie Institution of Washington Publication Number 251. By means of Professor E.G. Conklin and the authorities at Princeton the author used the facilities at the Biological Laboratory in Guyot Hall, where the kymograph records taken at Tortugas were studied and the results tabulated. The object of this research was to obtain an accurate quantitative determination of the rate of nerve-conduction in natural and in diluted sea-water at constant temperature, and also to estimate the effects of various artificial sea-water solutions containing all or some of the sodium, magnesium, calcium, and potassium cations of sea-water. The effects of temperature upon nerve-conduction are also of great importance. These studies were carried out in June and July 1916, upon Cassiopea xamachana, a thizostomous scyphomedusa which is abundant in the salt-water moat surrounding Fort Jefferson at Tortugas, Florida. In summary, nerve-conduction is due to a chemical reaction involving the cations of sodium, calcium, and potassium. Magnesium is non-essential. Observations do not support the "local action" theory of Lillie (1916). . 1918. Nerve-conduction in diluted and concentrated sea-water. Papers Tortugas Laboratory 12: 179-83. Carnegie Institution of Washington Publication Number 252. Ring-shaped strips of subumbrella tissue of the scyphomedusa Cassiopea xamachana were deprived of marginal sense-organs and placed in concentrated sea-water in order to determine the effect of concentration of electrolytes upon their rate of nerve-conduction. Experiments made in 1917 upon Cassiopea found that the rate has increased while the electrical conductivity has diminished. The injurious effects of concentrated sea-water upon regeneration and growth have been studied by Loeb, and by Goldfarb (1914), and there is a general resemblance between their curves and those in this paper for the rate of nerve-conduction, excepting that for regeneration somewhat dilute sea-water seems to be more favorable than normal sea-water, whereas in nerve-conduction the highest rate is obtained in slightly concentrated sea-water . . 1908. A plan for increasing the efficiency of marine expeditions, marine laboratories and our Atlantic coast. American Naturalist 42: 533. Narrative same as in reference no. 232. . 1914. The relation between the degree of concentration of electrolytes of sea water and the rate of nerve conduction in Cassiopea. Papers Tortugas Laboratory 6: 25-54. Carnegie Institution of Washington Publication Number 183. 60 238. ese). 240. 241. If sea-water be diluted with distilled water, or with a 0.9 molecular solution of dextrose, thus preserving its normal osmotic pressure but reducing the concentration of the cations of sodium, magnesium, calcium, and potassium, the rate of nerve-conduction increases as dilution proceeds, becoming most rapid in 90 per cent sea-water + 10 per cent distilled water or dextrose. The sodium cation is an active stimulant for nerve-conduction. Experiments with the magnesium cation show that it is not a stimulant for nerve- conduction. In very slight excess the potassium cation produces a permanently stimulating effect, as does sodium, but in denser concentration it produces momentary stimulation of the rate of nerve-conduction followed by depression. In all essential respects the effects of potassium are similar in kind, but more marked in degree, to those of sodium. —. 1908. Rhythmical pulsation in Scyphomedusae.II. Papers Tortugas Laboratory 1: 113-31. Carnegie Institution of Washington Publication Number 102. The following paper presents the results of a continuation of studies, the first report of which appeared in publication No. 47 of the Carnegie Institution of Washington, 1906. The present paper aims to correct certain errors in the previous report, and to announce some new results. Conclusions presented suggested that sea-water is a balanced fluid neither inhibiting nor stimulating pulsation in Cassiopea xamachana. The stimulus which causes pulsation is due to the constant formation of sodium oxalate in the terminal entodermal cells of the marginal sense organs. This sodium oxalate precipitates calcium, as calcium oxalate, thus setting free sodium chloride and sulphate which act as nervous stimulants. Pulsation is thus caused by the constant maintenance at the nervous centers in the sense-organs of a slight excess of sodium over and above that found in the surrounding sea-water. In Cassiopea the pulsation-stimulus is conducted by the diffuse nervous network of the subumbrella, and is independent of the muscles which may or may not respond to its presence by contraction. Strong primary nervous and muscular excitement followed by exhaustion and sustained muscular tetanus is produced in Lepas or in Cassiopea by a solution containing the amounts and proportions of NaCl+KCl+CaCl? found in sea-water. —. 1900. Some medusae from the Tortugas, Florida. Bulletin of the Museum of Comparative Zoology at Harvard College 37, no. 2: 1-82. This extensive article on the medusae of the Dry Tortugas includes an alphabetical list of species described, table showing the wide geographical range on some Tortugas Medusae, and comparisons of the Tortugas Fauna with that of Southern New England, the Tropical Atlantic, Fiji Islands and Tropical Pacific. . 1902. The Tortugas, Florida as a station for research in biology. Science (Washington, DIE) Mg 190-192. The advantages of establishing a tropical research marine biological laboratory in the Dry Tortugas over other Caribbean sites are discussed. The nearness of the Florida Current, an extension of the Gulf Stream, to the Tortugas is a distinct advantage over other sites because of its richness in pelagics, especially small juveniles and larvae during the summer. Other sites are richer in coral, such as Jamaica, but they are further from the Gulf Stream and are impacted by land runoff. . 1918. Toxic effects due to high temperature. Papers Tortugas Laboratory 12: 173-78. Carnegie Institution of Washington Publication Number 252. The experiments cited below appear to indicate that death from high temperature may be due to the accumulation of acid in the tissues. Reef corals from Tortugas, Florida, were kept at a constant temperature in warm ocean-water for 60 minutes in a thermostat, in the dark the temperature remaining constant within about 0.1° C. throughout the hour. In this manner the temperature was found that is just sufficient to kill the coral. The results are as follows: Acropora muricata 34.7, Orbicella annularis 35.6, Porites astraeoides 35.8, 61 Porites clavaria 36.4, Maeandra areolata 36.8, Porites furcata 36.85, Favia fragum 37.05, Siderastrea radians 38.2. It is apparent that those corals which live in cool, relatively agitated water, free from silt, are those that can not withstand high temperatures, whereas those which live in the hot, silt-laden shallows near shore are, generally speaking, forms which can resist high temperature. Favia fragum is however an exception. It seems possible that death from high temperature may be due to the accumulation of acid (possibly H,COs;) in the tissues, the rate of formation of this acid being related to the rate of metabolism of the tissues. Thus animals of the same class having a high rate of metabolism, as measured by oxygen consumption, are more sensitive to heat and to CO) than those having a low rate of metabolism. 242. . 1922. The tracking instinct in a Tortugas ant. Papers Tortugas Laboratory 18: 101-7. Carnegie Institution of Washington Publication Number 312. Monomorium destructor Jerdon, a tropicopolitan ant of East Indian origin, was identified in Florida . It is a small, reddish-brown ant, a great pest in the wooden buildings of the Tortugas laboratory. These pests have killed rats confined in cages within 24 hours. The experiments described were made on the flat wooden floor of the laboratory. To attract the ants, a number of recently killed houseflies were impaled upon a pin. The pin with its lure of flies was then thrust into the floor in front of a foraging ant, which would often pass within 0.25 inch of the lure without perceiving the flies; but if its course were such that it came appreciably nearer than 0.25 inch, the ant suddenly turned toward the flies, and without apparent excitement appeared to "inspect" them, spending a half minute or more crawling over them and stroking them with its antennae. This "finder ant" soon leaves the flies without carrying off any piece of them, but instead of moving off in the erratic and tortuous path it was pursuing before it found the flies, it now goes in a fairly straight path toward some crevice in the floor, out of which there soon pours an excited swarm of its nest-mates, who proceed toward the flies in a fairly straight path. When an ant returns to the nest it pursues a fairly straight path which is more or less right in direction, but when the ant has gone the correct distance, it begins to wander in more or less tortuous courses until it finds the nest. 243. McClendon, J. F. 1917. Diurnal changes in the sea at Tortugas, Florida. Proceedings of the National Academy of Science 3: 692. The only diurnal change noted in the Gulf Stream was a change in temperature of about 1 degree C and the resulting change in oxygen tension. However, marked differences were found in temperature, pH, CO >, and O; concentration in shallow water where light could reach the bottom. The temperature, O, concentration and O, tension were lowest and the CO), concentration and CO, tension highest at5 A.M. The temperature, O2 concentration and O, tension were highest and CO, concentration and CO) tension were lowest at 3 P.M. during July at Dry Tortugas. The magnitude and exact time of minima and maxima varied from day to day, and varied a great deal with station location. Studies on the effects of these changes on organisms were made. The limiting factor for plants was nitrogen, while the limiting factor for animals was food and oxygen. 244. ———. 1917. Effect of oxygen tension on the metabolism of Cassiopea. Proceedings of the National Academy of Science 3: 715-16. Experiments on the effect of oxygen tension on the metabolism of Cassiopea were carried out at the Tortugas Laboratory by using the umbrella of Cassiopea to maintain a layer of cells in seawater at 30 degrees C. The metabolism varied with oxygen concentration. This may be true for all animals, however there is a distinction between the metabolism of vertebrate cells and Cassiopea. Vertebrate cells give out lactic acid when asphyxiated whereas Cassiopea may remain without oxygen for seven hours without giving out CO, or any other acid. Although details of the experiments will be published elsewhere, it was 62 245. 246. 247. 248. concluded that changes in the threshold of stimulation of the respiratory and basomotor centers may affect metabolism in man and animals. . 1917. The effect of stretching on the rate of conduction in the neuro-muscular network in Cassiopea. Proceedings of the National Academy of Science 3: 703. The experiments on Cassiopea collected at the Dry Tortugas, tend to support the conclusions reached by Carlson, that stretching the nerve does not change the rate of the nerve impulse, and that the conducting substance itself, can be stretched and relaxed. . 1917. The equilibrium of Tortugas sea water with calcite and aragonite. Proceedings of the National Academy of Science 3: 612-19. This report provides information on the continuing controversy on the solubility of Calcium chloride in sea water. The precipitation of CaCO; at Tortugas was studied by T.W. Vaughan, R.B. Doyle, and G.H. Drew. Drew observed that denitrifying bacteria, Pseudomonas calcis obtained from sea water was capable of changing calcium nitrate to calcium carbonate in culture media, and supposes that a similar process occurs in seawater. This study attempts to determine the nitrates or nitrites. If the pH is maintained (by plants) at 8.2 at the Tortugas, the introduction of calcite crystals would result in a lowering of the calcium content of Tortugas seawater by about 4.5%. . 1914. Experiments on the permeability of cells. Papers Tortugas Laboratory 6: 123-30. Carnegie Institution of Washington Publication Number 183. One of the most important steps in the analysis of life was the discovery of oxygen. Ever since that time it has been known that animals absorb free oxygen and give it out in a combined form. In this experiment three methods of procedure were followed: (1) the use of cell masses as partitions (on eggs of Lytechinus); (2) the use of quantities of eggs suspended in a liquid medium (on eggs of Fundulus); (3) experiments on individual eggs (of Arbacia). The permeability of the egg to ions and perhaps some other substances increases on fertilization. The unfertilized egg is perhaps in a dormant condition and the increase in permeability probably allows a rapid interchange with the surrounding medium necessary for activity (development). Whereas this supposed significance of permeability has not been proven, the sea-urchin;s egg is not an exception. The relation of permeability to oxidation can hardly be determined until more is known about the mechanism of animal oxidations. These seem to depend on structure since complete oxidations cease when structure is completely destroyed. Reference is made only to oxidations resulting in the formation of CO:. Oxidizing enzymes such as tyrosinase, which are independent of structure, do not completely oxidize the substances acted on. . 1911. On adaptations in structure and habits of some marine animals of Tortugas, Florida. Papers Tortugas Laboratory 3: 55-62. Carnegie Institution of Washington Publication Number 132. This article discusses the habits of some marine animals of the Tortugas. Many of these animals were thigmotactic and remained in glass tubes rather than in the open. They learned to find the tubes when removed from them. Such was the case with five species of the Alpheidae, one of the Pontoniidae, Typion tortugae Rathbun, and Gonodactylus aertedii. All the anemones were thigmotactic on their bases. These same animals were heliotropic. The crustaceans were negatively heliotropic and the anemones kept their bases from the light, while Cradactis variabilis Hargitt hid all but the tips of the fronds and tentacles from the light. In removing its base from the light, Stoichactis helianthus, which lives on coral heads, makes snail-like movements similar to Metridium, while Cradactis, which lives in holes in decayed coral heads, crawls on its tentacles. 63 249. . 1918. On changes in the sea and their relation to organisms. Papers Tortugas Laboratory 12: 213-58. Carnegie Institution of Washington Publication Number 252. The sea and air form the circulating media for the living organisms of the world. The local composition of the sea is distinctly affected by living organisms. The local changes in the composition of the sea are the subject of the present paper. These changes are due chiefly to organisms, but partly to meteorological causes. The water evaporated is returned with addition of fixed nitrogen from electric discharges or falls on the land and is returned with various salts, chiefly CaCO3, and with fixed nitrogen and other products of organisms. Various seaweeds absorb CO) thus leaving an excess of CaCO, which has a very low solubility and is constantly being precipitated in certain warm seas, and is precipitated within the bodies of organisms in the surface waters of all seas. In working out the relation of H-ion concentration (pH) to the solubility of CaCO; in sea-water, it was found that all sea-water is supersaturated with CaCO3, and will lose some of it if shaken with calcite or aragonite crystals. The pH is influenced by plant and animal life and arises at Tortugas to 8.35 during the day over well-lighted bottoms rich in vegetation, and falls to 8.18 during the night. It may be said, therefore, that conditions in shallow water over eelgrass or other seaweed or corals (with symbiotic algae) favor the precipitation of CaCO3. The question arises whether the occasional high pH of Tortugas sea-water is sufficient to explain the precipitation of CaCO3. The author's experiment showed that if the pH of sea-water should be maintained (by the action of plants) at 8.2 while it was agitated with calcite crystals, the loss of CaCO; would be about 0.001 N, or 0.0005 M, or 0.1 gram per liter. This would cause a deposit of 10 kg. per square meter of bottom in water 100 meters deep. This would cause a lowering of the total calcium content of Tortugas sea-water by about 4.5 per cent. The actual precipitation of CaCO; was most noticeable in the Marquesas lagoon. At 4 p.m., July 30, the pH was 8.46 and there was a precipitate of CaCO; coming down in the water and encrusting the eel-grass. 250. ————. 1917. The standardization of a new calorimetric method for the determination of the hydrogen-ion concentration, CO, tension and CO, and O; content of sea water, of animal heat, and of CO) of the air, with a summary of similar data on bicarbonate solutions in general. Journal of Biological Chemistry 30: 265-88. Experiments were conducted on pH, CO, tension, CO, and oxygen content of Dry Tortugas seawater and seawater from other oceanic areas using a Leeds & Northrup potentimeter and a 0.1 KCL calomel electrode. It was found that neither the salinity nor the alkaline reserve in seawater of the tropical or temperate oceans change sufficiently to noticeably change the relation of pH to CO; tension, although the alkaline reserve does change sufficiently to affect the total CO) greatly. 251. McClendon, J. F., C. C. Gault and S. Mulholiand. 1917. The hydrogen-ion concentration, CO2 tension, and CO2 content of sea water. Papers Tortugas Laboratory 11: 21-69. Carnegie Institution of Washington Publication Number 251. Narrative same as in reference no. 250. 252. Meeder, J. F. 1979. Corals and coral reefs of the Dry Tortugas, Florida. in Guide to sedimentation for the Dry Tortugas. R.B. Halley (Compiler), 46-47. S.E. Geological Society Pub. This paper presents a description of the corals from two localities in the Dry Tortugas, on a back reef environment of a fringing reef near Garden Key, and the second, a series of patch reefs off Loggerhead Key. The general setting, ecology, distribution, and types of corals are discussed for each locality. Forty-one of the forty-two species of corals reported at the Tortugas are covered in a field key and later described in this paper. 64 253. Miller, H. M. Jr. 1926-1929. Behavior of trematode larvae. Carnegie Institution of Washington, Year Book. Note: Published as follows: 1926, v.25, p. 243-244: 1929, v. 28, p. 295. Anatomical/morphological descriptions of six larval tremeatodes infesting the mollusk, Cerithium litteratum, taken from Bird Key Reef Porites beds, were provided. Percentages of occurrence of the 6 cercariae were given. The behavior of the members of this morphological group are described in detail, including aspects of their life history. 254. Miller, R. A. and H. B. Smith. 1931. Observations on the formation of the egg of Echinometra lacunter. Papers Tortugas Laboratory 27: 47-52. Carnegie Institution of Washington Publication Number 413. The study of the ovaries of Echinometra was undertaken in the hope that by the use of some of the newer cytological methods, it might be possible to extend our knowledge of the processes of odgenesis, particularly of those concerned in the formation of yolk in the chinoid egg. This paper presents the observations that have been made, and the conclusions that have been drawn. It was found that in Echinometra lancunter, undifferentiated cells along the wall of the ovary are uniform in appearance, although they are destined to develop into two entirely different kinds of cells. As development proceeds, some of the cells become odgonia, while others enlarge and disintegrate to form deutoplasmic bodies. These are more coarsely granular than was the cytoplasm from which they were formed. The granules exhibit different affinities for stains. Fully formed nutritive spheres are of two kinds, granular and non-granular. The former are composed entirely of cytoplasmic, or of both cytoplastic and nuclear material. The nutritive spheres group themselves around the o6gonia in a follicular arrangement. Eventually the nutritive spheres enter the egg and disappear as such, forming the yolk content of the cytoplasm, which becomes homogeneous and evenly granular. The deutoplasmic granules are smaller and more diffuse in the mature eggs than in the odcytes. The nutritive spheres have been shown to be composed of phospholipins suspended in a homogeneous medium. It is probable that they are largely lecithin in content. Their origin is not known. 255. Millspaugh, C. F. 1907. Flora of the sand keys of Florida. Columbian Museum 118, no. (Bot. Ser. 2): 191-245. A list of species and details of vegetation, as well as elevational descriptions and size dimensions of sand keys westward of Key West, including the Tortugan group, were compiled by the author during the winter through spring of 1904. 256. Mitchell-Tapping, H. J. 1981. Particle breakdown of recent carbonate sediment in coral reefs. Florida Scientist 44, no. 1: 21-29. Skeletal particles of the major components of the carbonate sediment of the reef shoal environment were examined using the scanning electron microscope. This examination revealed no set pattern of skeletal breakdown according to microarchitectural structure, as postulated by the Sorby principle, but that such a breakdown depends on mineralogical composition, wall thickness, grain size and pattern, density, and the amount of cementation and bloerosion. To investigate general particle-size abundances and deficiencies in carbonate sediment, samples were taken from the reef crest, back-reef-rubble and open- sand ecozones of the reef shoal environments of sites from the Bahamas, Dry Tortugas, Lower Florida Keys, Grand Cayman Island and the U.S. Virgin Islands. Size analyses of these samples showed that the sediment is moderately sell-sorted, coarsely-skewed and leptokurtic. Although particle-size abundances (or modes) exist in each individual site, there is no particular particle-size abundance that is common to all the sites. It is inferred that the particle abundances (or modes) for each site are a product of the sorting potential of the wave energy and that this sorting potential is the major control of the breakdown of 65 sand-sized skeletal particles rather than the microarchitectural structure as proposed by the Sorby principle. 257. Moritz, C. E. 1936. Embryology of the sea-hare, Aplysia protea and of Crangon armillatus. Carnegie Institution of Washington, Year Book 35: 90. Observations from aquaria are recorded on the early development of Aplysia, from embryo to 5 days beyond hatching. Adults were collected from the moat on Garden Key and Bird Key Reef. 258. Multer, H. G. 1975. Field guide to some carbonate rock environments; Florida Keys and Western Bahamas. Fairleigh Dickinson University 40: 175 pp. This report presents the most recent compilation on Holocence sediments in the western Bahamas and in the Florida Keys including the Dry Tortugas. Selected geologic literature pertinent to local environments is noted within the text, with full bibliographic citations following each subject area. Carbonate sand beach and beach rock environments from the Florida Keys, Bimini, and Loggerhead Key are compared. In summary, Holocene sediments of these areas today present a vast array of textures and constituent particles characteristic of environments which have been subjected to fluctuating sea levels and storm action. Such data may be used to interpret ancient environments. 259. . 1971. Holocene cementation of skeletal grains into beachrock, Dry Tortugas, Florida. Carbonate Cement. O. P. Bircker. Baltimore, Maryland: Johns Hopkins Press. A discussion is presented on the origin of beach rock at Loggerhead Key, Dry Tortugas. Present evidence suggests that cementation is due to alternate wet and dry salt water spray conditions with skeletal grains providing nuclei for precipitation from a supersaturated calcium carbonate solution. The limited ground water conditions and lack of grain solution for providing aragonite cement are two evidences in favor of the above cited evaporation origin for the cement in this rock. Ginsburg (1953) reached similar conclusions for beach rock in the same area. 260. Murphy, L. E. 1993. Dry Tortugas National Park, Submerged Cultural Resources Assessment, L. E. Murphy. Submerged Cultural Resources Unit, Southwest Region, National Park Service, Santa Fe, New Mexico. This volume describes and assesses the known and potential archeological resources in Fort Jefferson National Monument, later redesignated Dry Tortugas National Park. The emphasis is on submerged cultural sites, particularly shipwrecks. The importance of linking the natural resources with submerged cultural resources is provided by identifying the biological influences on the cultural resources. The Dry Tortugas and South Florida geological development and environmental succession is summarized with focus on the postglacial development of the Florida-Reef Tract, depositional environments, coral reef and sand key development as well as Late-glacial and Postglacial succession of environments. An overview on the physical oceanography of the eastern Gulf of Mexico concentrating on the Dry Tortugas with emphasis on currents and climate that affect shipping vessel casualties and site preservation is provided. Recommendations for future research and resources management are given. 261. Nance, J. M., E. F. Klima and F. J. Patella. 1986. Review of the Tortugas pink shrimp fishery from May 1984 to December 1985, Galveston, Texas, NOAA/NMES, Southeast Fisheries Center, Galveston, Texas. NOAA Tech. Memo.177. Commercial pink shrimp fishing data from the Tortugas (Dry Tortugas Islands, Florida) fishery were reviewed for biological year 1984 (May 1984-April 1985) and the first 8 months of biological year 1985 (May 1985-December 1985). Pink shrimp landings were Just over 11.0 million pounds in biological year 1984 with 17,000 days of fishing 66 expended. This computed to a CPUE value of 643 pounds per day. Pink shrimp landings for biological year 1985 are estimated to be around 9 million pounds with 15,000 days of fishing expended. The predicted CPUE value for 1985 should be around 600 pounds per day. Biological year 1984 experienced two extended periods of pink shrimp recruitment into the Tortugas fishing grounds. 262. O'Neill, C. W. 1976. Sedimentology of East Key, Dry Tortugas (abs.). Florida Scientist 39 (Suppl. 263. 1), Fortieth Annual Meeting of the Florida Academy of Sciences at Eckerd College, St. Petersburg, Florida March 18,19,20, 1975: 10. East Key of the Dry Tortugas rests on a large crescent shaped bank and oscillates about a stable core in response to seasonal variations. A simple strand/dune plant community is largely responsible for short term stabilization of this central core. Historical studies covering 200 years show that East Key varies in its bank position on long term basis and has in the past been of much greater areal extent. In addition, historical records show the Dry Tortugas group has decreased in extent from 11 keys to the present seven. This effect is thought to be due to a combination of eustaic sea level change and storm degradation. . 1976. "Sedimentology of East Key, Dry Tortugas, Florida." University of South Florida. Ph.D. Dissertation. The Dry Tortugas platform is a complex of reefs, banks, and shoals which lie 65 nautical miles west of Key West, Florida. This reef platform most closely resembles the resorbed reef of Maxwell's (1968) classification. A review of historical records covering a 463-year period indicates that the Dry Tortugas island group is undergoing progressive degradation and has been reduced from 11 rocky islets in 1513 to 7 at present. This reduction is thought to be due to the combined effect of episodic events and the Holocene transgression. East Key maintains a position on the windward segment of the Tortugas group. Short-term changes to the key are basically in response to regular seasonal variation and cause the island to oscillate to the north and south about a plant-stabilized core. Long- term changes reflect the effect of episodic events and entail inundation, major shifts inposition, and changes in the orientation of the key. Mean grain size of East Key sediments is near 0.900 or coarse sand. The sediments are nearly without exception moderately well sorted with an average value of 0.570. The mean skewness value is -0.22 (coarsely skewed). Texture was compared to morphology. The most striking correlation occurred on the foreshore where there is a distinct tendency for sediments to become coarser moving seaward from the berm. The average percentage composition of East Key sediments, by constituent, was found to be 52 percent Halimeda, 35 percent coral, 6 percent Mollusca, 3 percent coralline algae, 1 percent Foraminifera, 1 percent echinoid fragments and 2 percent miscellaneous plus unknown. The most significant correlation between composition and morphology occurred on the foreshore, where percent Halimeda generally increased seaward from the berm. This general increase in percent Halimeda correlates with a tendency for sediments to become coarser moving seaward from the berm. 264. Ogden, J.C., W.C. Jaap, J. W. Porter, N. P. Smith, A. M. Szmant and D. Forcucci. 1993. SEAKEYS: A large scale interdisciplinary study of the Florida Keys Reef Tract (abs.). Proceedings of the Seventh International Coral Reef Symposium, v. 2. Mangilao, Guam: University of Guam Marine Laboratory. The SEAKEYS program has established a research framework which encompasses the large geographic and long time scales of natural marine processes and ecosystem variation. The core of the program is a series of instrumented, satellite-linked monitoring stations which span the 220 mile coral reef tract. Mesoscale physical oceanographic studies are concentrated in the major channels potentially linking Florida Bay and the population centers of the Keys with the reef tract. Simultaneously, nutrient studies are probing the possibility of sewage and agrichemical contamination, complicated by natural sources of 67 nutrients. A series of long-term photomosaic stations have tracked coral community dynamics for more than 5 years. The design of the SEAKEYS program may provide an example for long-term research on coral reefs elsewhere. 265. Osburn, R. C. 1914. The Bryozoa of the Tortugas Islands, Florida. Papers Tortugas Laboratory 5: 181-222. Carnegie Institution of Washington Publication Number 182. In the summer of 1908 the writer had the privilege of spending the month of June at the Carnegie Institution Laboratory of Marine Biology on Loggerhead Key of the Tortugas Islands. The entire period was devoted to a close search for the bryozoa inhabiting the shallow waters about the reefs, on the piles of the old government dock on Garden key, in the moat of old Fort Jefferson on the same key, and in dredging the shallow waters about the islands down to 22 fathoms. Comparatively little work has been done on the bryozoa of the Florida and West Indian regions. By comparing with lists from other regions where the bryozoa have been carefully worked, it will be seen that the bryozoan fauna of the Tortugas and of the Florida-West Indian regions 1s fairly rich in species and fairly representative of tropical and semi-tropical regions. 266. Payne, F. 1937. Early development of Ptychodera bahamensis. Papers Tortugas Laboratory 31: 71-76 (issued July, 1936). Carnegie Institution of Washington Publication Number 475. The author records some observations made on Ptychodera bahamensis during the summer of 1933 while working at the Carnegie Laboratory at Tortugas, Florida. The animals were found most abundantly on the coral reef near Fort Jefferson in water which at low tide was from one to two feet deep. The larvae described by Weldon, Morgan, and Stiansy and assigned to Ptychodera bahmensis do not agree with the author's own observations of the larvae of Ptychodera bahamensis which have been followed from the fertilized egg. It seems conclusive that more than one species is involved or that errors have been made on the part of someone. Even though the author has not followed development through to metamorphosis, it seems clear that the tornariae, described by Weldon and Morgan are different from the tornaria of Ptychodera bahamensis, assuming his identification is correct. 267. . 1933-1938. Embryology and cytology of the balanoglossid, Ptychodera bahamensis... Carnegie Institution of Washington, Year Book. Note: published as follows: 1933, v. 32, p. 277-78; 1938, v. 37, p. 84. Embryological work was conducted on the supposed protochordate, Ptychodera bahamensis, starting with the fertilized egg. Development was followed as far as the tornaria. 268. Pearse, A. S. 1934. Animals in brackish water ponds and pools at Dry Tortugas. Papers Dry Tortugas Laboratory 28: 125-42 (issued Dec. 1932). Carnegie Institution of Washington Publication Number 435. During the summer of 1931 two ponds and three pools were studied on Long and Garden Keys, Dry Tortugas. Because of the remoteness of the Dry Tortugas, the animals which live in these isolated habitats are of particular interest. Results were as follows: Five brackish water pools and ponds were studied on Long and Garden Keys. These were variable in salinity and temperature, and limited in extent. They showed various stages of evolution from sea to fresh water and conditions of life in them were more or less severe. The animals in the ponds and pools were resistant to environmental variations. They lived in salinities between 0.6 and 6.4 per cent and endured temperatures above 42° C. Some of the small bodies of brackish water on Dry Tortugas contained curious mixtures of marine and fresh-water animals. Callianassas, marine snails, mullets, and needle-fishes lived with 68 269. 270. PHIM Piles dragon-fly nymphs, water boatmen, surface bugs, aquatic beetles and midges. Lists of the animals collected and observed in each pond or pool are given. Each pond or pool contained certain characteristic animals, which had become dominant in that particular habitat. As the small pools trended toward fresh water, insects were increasingly dominant in them. Insect populations show great pressure and spread into all available habitats. When small pools are cut off from the ocean and are gradually transformed into fresh-water habitats, insects are the pioneers and soon become dominant. . 1934. Freezing points of bloods of certain littoral and estuarine animals. Papers Tortugas Laboratory 28: 93-102 (issued Dec. 1932). Carnegie Institution of Washington Publication Number 435. It is well known that the bloods and body fluids of marine invertebrates and elasmobranchs have about the same osmotic pressure as the ocean water in which they live, though the salt content of such fluids is usually a little less than that of the surrounding medium. Those of teleost fishes (marine, fresh-water and land) and of fresh-water and land animals generally have osmotic pressures which are much below those of sea-water. Littoral crustaceans and fishes are of particular interest because they represent various stages of adjustment to life on land. At Tortugas, conditions are particularly favorable for the study of such adjustments for the littoral group. The observations cited in this paper appear to justify the conclusion that crabs and fishes which take up terrestrial life or air-breathing have bloods of lower osmotic pressures than comparable marine or fresh-water animals. The attainment of land life by marine animals is apparently associated with a reduction in salinity and stabilization of the contents of the blood. . 1934. Inhabitants of certain sponges at Dry Tortugas. Papers Tortugas Laboratory 28: 117- 24 (issued Dec. 1932). Carnegie Institution of Washington Publication Number 435. Many sponges are veritable living hotels. Their canals are densely populated with a variety of animals and some species have not been found elsewhere. During the summer of 1931 the writer studied the animals which occurred in five species of sponges at Dry Tortugas. The number of animals which live in big sponges is enormous. Though each sponge appears to be occupied to its full capacity, the number of animals per cubic centimeter of sponge is apparently influenced by depth and the size of the sponge itself. In loggerheads there are relatively more guests in smaller sponges and in deep water. Among the species of small sponges there are striking differences in the number of animals present and these may be due to inherent qualities. For example Stematumenia foetida (Schmidt) contained very few animals, whereas the slightly smaller Spongia officinalis L. was crowded. . 1918. Notes on certain amphipods from the Gulf of Mexico, with descriptions of new genera and new species. Proceedings of the U.S. National Museum 43, no. 1936: 369-79. This is a report on a portion of the amphipods from the Gulf of Mexico in the collection of the United States National Museum. The collections are from several sources and extend over a long period of years. The greater part of them had not yet been examined. Those described were taken chiefly by the steamers Fish Hawk and Albatross of the United States Bureau of Fisheries. . 1929. Observations on certain littoral and terrestrial animals at Tortugas, Florida, with special reference to migrations from marine to terrestrial habitats. Papers Tortugas Laboratory 26: 205-23. Carnegie Institution of Washington Publication Number 391. In the past, various types of animals have migrated from the ocean into fresh-water streams or lakes, and from there gained a foothold on land. On the shores of all oceans, animals may be found which are partially adjusted to life in fresh water or on land. The Dry 69 Tortugas contain no fresh water, and therefore offer an excellent opportunity to study littoral animals which have no immediate contact with fresh-water habitats. In summary, at Tortugas certain reef, beach and land animals were studied with reference to migrations from sea to land. Hermit crabs which have become more or less adjusted to life on land show a progressive reduction in the number of gills. Crabs which have migrated landward show a progressive lessening of gill-volume. Beach animals which show any landward trend usually live longer when kept in air than when kept in fresh water. Animals which have attained some degree of ability to live on land have often also acquired a greater degree of ability to resist the extraction of constituents of body fluids into fresh water. Animals which migrate from the sea and become established on land do not do so on account of one "lure" or one "danger." Each habitat has certain advantages and certain disadvantages. A continually changing animal must continually make adjustments to a continually changing environment, and when it migrates to a new habitat, must make many compromises between new advantages and dangers, old necessities and new requirements, and old habits and new abilities. NB). . 1934. Observations on the parasites and commensals found associated with crustaceans and fishes at the Dry Tortugas, Florida. Papers Tortugas Laboratory 28: 103-15 (issued Dec. 1932). Carnegie Institution of Washington Publication Number 435. During the summer of 1931 the writer had opportunity to conduct post mortems on various crustaceans at Dry Tortugas from June 3 to August 22. Dr. Waldo Schmitt furnished and identified many of the specimens. The following parasitic isopods were taken from fishes: 2 Cymothoa oesturm (L.) from the gill cavity of Caranx ruger (Bloch), July 26; 1 Rocinela signata Schioedte and Meinert from the gills of Promicrops itaiara Lichtenstein in July and 1 from the gills of Lutianus analis (Cuvier et Valiencennes), June 27; 8 Excorallana tricornis (Hansen) from the nose of Promicrops itaiara Lichtenstein, July, and 31 from the gill cavity of Epinephelus moro (C. et V.). The occurrence of the parasites and commensals associated with crustaceans depends upon a variety of factors - host specificity; habitat; habits, structure and physiology of hosts and parasites, etc. In general the greatest number of species of parasites occurred in or near the littoral zone. However, great numbers of parasites per host were encountered among some land crustaceans. 274. ————. 1929. Two new mites from the gills of land crabs. Papers Tortugas Laboratory 26: 225-30. Carnegie Institution of Washington Publication Number 391. During July and August 1928, mites were found at Dry Tortugas on the gills of the land hermit crab, Cenobita diogenes (Latreille), and on the Nassau crab, Gecarcinus lateralis (Freminville). These crustaceans visit the ocean only once each year when they hatch out their young. No mites were found on the gills of the ghost crab, Ocypoda albicans (Bosc), which often visits the ocean and bathes its gills. Mixed or pure cultures of bacterial populations are incapable of precipitating CaCO; in a sea-water medium if KNO3 and organic matter as the sugars or similar forms free from calcium are added to the medium. A number of different forms of bacteria in the sea possess the power of precipitating CaCO; in appropriate media containing a large excess of soluble salts, but only under such conditions. Among such organisms there is great variability to perform the task in question, depending on the composition of the medium. 275. Perkins, H. F. 1908. Notes on the medusae of the Western Atlantic. Papers Tortugas Laboratory 1: 133-56. Carnegie Institution of Washington Publication Number 102. The Marine Biological Laboratory in the Dry Tortugas is well situated for the study of many of the lower marine animals, their behavior, and the conditions of life, particularly 70 the coelenterates. One quite unique feature occurs in the Tortugas in the presence of the old fortification and surrounding moat of Fort Jefferson. The moat affords remarkably favorable conditions for the growth and multiplication of the lower forms of plants and animals sheltered by the sea-wall, its shallow water warmed by the sun and kept from stagnation by the agitation and partial change of the tides. The writer has for several years been interested in the causes of migration and segregation of Medusae and has had the privilege of examining specimens of this genus from Jamaica, and has studied the characteristics of the specimens found in the Bahama Islands and at the Tortugas. There was less of peculiarity in all the surroundings, the temperature of the water, storm influence, and food supply being normal for the shores of coral islands. The only points of difference to be noted in the medusae are with reference to size and color-pattern. The main features of the two species, Cassiopea xamachana and Polyclonia frondosa are presented. 276. Petrovic, C. A. and J. King Jr. 1973. Bird records from the Dry Tortugas. Florida Field Naturalist 277. 278. Phillips, 279. 1, no. 1: 5-8. During a visit March 25 to April 4, 1967 to the Tortugas, a total of 70 species of mostly land birds were recorded. Detailed observations on twenty specimens rarely seen at the Tortugas are presented. . 1972. Common elder and king rail from the Dry Tortugas Florida. Auk 89, no. 3: 660. The authors watched the early spring migration at the Dry Tortugas Islands, which lie in the Gulf of Mexico about 70 miles west of Key West, Florida, and recorded 70 species, the majority land birds. The two records reported here represent significant additions to the species known distribution . A. H. 1917. Analytical search for metals in Tortugas marine organisms. Papers Tortugas Laboratory 11: 89-93. Carnegie Institution of Washington Publication Number 251. This study concerns the problem stated in Year Book no.14, page 193 of the Carnegie Institution of Washington. A large number of specimens were collected to be analyzed for metals. The metals determined were iron, manganese, zinc, copper, and lead. For the determination of zinc, copper, and lead, when the dried material was sufficient, 20 grams were used as a sample; when it was not possible to use 20 grams, the results are all calculated to 20 grams.. . 1922. Analytical search for metals in Tortugas marine organisms. Papers Tortugas Laboratory 18: 95-99. Carnegie Institution of Washington Publication Number 312. Included in the material collected at the Tortugas and analyzed for metals (some of the results of which were reported in the annual report of the Carnegie Institution for 1917) was a brown spotted holothurian, Stickopus mobii, which was analyzed by the methods there indicated. The element vanadium was found in the holothurian material and heretofore has never been reported from seawater. Vanadium has been reported from freshwater and in the blood of an acidian from the Bay of Naples. This vanadium content of the blood does not seem to be a characteristic of all acidians, as two other species from the Tortugas yielded no vanadium, neither did two other species of holothurians yield vanadium. Two species, a chordata and a echinoderm contained vanadium, indicating that other forms may use vanadium as an oxygen carrier in their vascular systems. The source of vanadium in sedimentary rock and coals has always been somewhat of a puzzle. It is possible that such forms as Stickopus mobii may concentrate vanadium and that in depth, could easily be fixed and held as a constituent of the sedimentary rocks thus formed. 71 Another possibility of the fixation of vanadium under the above conditions is the presence of hydrogen sulphide which is constantly liberated from muds of mangrove lagoons. 280. . 1918. A possible source of vanadium in sedimentary rocks. American Journal of Science 46: 473-74. Narrative same as in reference no. 279. 281. Pichard, S. L. and J. R. Paul. 1991. Detection of gene expression in genetically engineered microorganisms and natural phytoplankton populations in the marine environment by messenger RNA analysis. Applied Environmental Microbiology 57, no. 6: 1721-27. A simple method that combines guanidinium isothiocyanate RNA extraction and probing with antisense and sense RNA probes is described for analysis of microbial gene expression in planktonic population. Probing of RNA sample extracts with sense-strand RNA probes was used as a control for nonspecific hybridization or contamination of mRNA with target DNA. This method enabled detection of expression of a plasmid- encoded neomycin phosphotransferase gene (nptII) in as few as 10 super(4) Vibrio cells per ml in 100 ml of seawater. We have used this method to detect expression of the ribulose-1, 5-bisphosphate carboxylas large-subunit gene (rbcI) in Synechococcus cultures and natural phytoplankton populations in the Dry Tortugas, Florida. During a 36-h diel study, rbcL expression of the indigenous phytoplankton was greatest in the day, least at night(1100, 0300, and 0100 h, and variable at dawn or dusk (0700 and 1900 h). These results are the first report of gene expression in natural populations by mRNA isolation and probing. 282. Pitts, R. F. 1936. Clearance values of sucrose and creatinin in the kidneys of the red grouper, Epinephelus striatus. Carnegie Institution of Washington, Year Book 35: 90-91. Studies were made on the excretion of urinary creatine nitrogen from the red grouper, Epinephelus striatus. 283. Plan Development Team, Reef Fish Management Plan South Atlantic Fishery Management Council. 1990. The potential of marine fishery reserves for reef fish management in the U.S. southern Atlantic, Coastal Resources Division. Marine fishery reserves (MFRs), areas with no consumptive usage, are recommended as a viable option for management of reef fisheries in the U.S. southern Atlantic region. MFRs are designed to protect reef fish stocks and habitat from all consumptive exploitation within specified geographical areas for the primary purpose of ensuring the persistence of reef fish stocks and fisheries. Fishery reserves are intended to protect older and larger fishes. This will benefit reef fisheries by protecting critical spawning stock biomass, intra-specific genetic diversity, population age structure, recruitment supply, and ecosystem balance while maintaining reef fish fisheries. The MFR concept is easily understandable by the general public and possibly more easily accepted than some other management strategies. Fishery reserves provide some insurance against management and recruitment failures, simplify enforcement, and have equitable impact among fishery users. Data collection needs solely for management are reduced and management occurs without complete information and understanding about every species and interaction. Use of fishery reserves will establish U.S. leadership in producing model strategies for cooperative international reef resource management in the Caribbean. Large resident fishes that wander out of reserved can help maintain certain trophy fisheries. MFR sites with natural species equilibrium will allow measurement of age, growth, and natural mortality for fisheries purposes and will provide a basis for other educational, economic, and scientific benefits. Because there is no fishing within MFRs, impacts of hook and release mortality are eliminated and the temptation for incidental poaching is reduced. A mixed management strategy is recommended where 20% of the shelf is MFR, while the remaining 80% is 72 managed for optimal yield by any of several traditional options. Coordinated fishery reserve efforts in state waters would enhance the benefits of MFRs. Obstacles to fishery reserves include automatic resistance to new approaches in U.S. marine fisheries, opposition by some local special interests near proposed reserves, and uncertainty concerning the size, location, and number of reserves necessary to ensure persistence of the reef fish fisheries. The incentive for deliberate poaching may be increased within the reserves; thus, at-sea surveillance and enforcement may be necessary. New artificial reefs may be needed to replace those lost by inclusion within fishery reserves. Other fishery management plans should be coordinated to control trolling and other fishing activities within reserves that may impact reef fishes. The short-term impacts on total harvest caused by placing fishing habitat into fishing reserves should be compensated for by long-term fishery benefits. The Dry Tortugas is listed as a potential marine fishery reserve site. 284. Plantier, T. L. 1988. "A comparison of reproductive success in early and late breeding sooty terns Sterna fuscata in the Dry Tortugas." MS.Thesis, Florida Atlantic University, Boca Raton. Evidence indicates that earlier-nesting birds are often older, choose preferred nest sites, and have greater reproductive success than those nesting later. The sooty terns at Bush Key appear to follow a similar pattern. The first birds arrive at the west end of the breeding grounds three weeks earlier than birds at the east end and behaviorally appear to be older and more experienced. The west birds settled in the more desirable habitats (the west end was cooler than the east end) and laid larger eggs, hatched larger chicks, enjoyed greater hatchability, fed their chicks at a lower frequency when they were young, and had greater reproductive success than birds in the east. This was accomplished through a combination of choosing physically and thermally more favorable habitat, which was more centrally located, being more persistent incubators and brooders, and, by nesting earlier, having larger, less-easily eaten chicks by the time avian predators arrived on the island. 285. Plough, H. H. and N. Jones. 1940. Ecteinascidia tortugensis, species Nova; with a review of the perophoridae (Ascidiacea) of the Tortugas. Papers Tortugas Laboratory 32: 47-60 (issued Oct. 1939). Carnegie Institution of Washington Publication Number 517. During the season of 1936 at the Tortugas Laboratory the senior investigator undertook a study of the regeneration of pieces cut from the growing stolons of several species of the family Perophoridae (Ascidiacea Phlebobranchia), which grow in profusion at many places in the Tortugas area. A new member of the family Perophoridae is described and named Ecteinascidia tortugensis from its type locality, the Dry Tortugas Key, Florida. It is shorter than other Ecteinascidia, lies on the ventral side attached along the test, has the siphons on the dorsal side widely separated and opening in opposite directions, and possesses a marked secondary loop in the intestine. This species reaches sexual maturity early in July at the Tortugas, about two weeks later than E. conklini. A brief account of the development is given. The structure and growth habits of E. tortugensis indicate that it is intermediate between E. turbinata and Perophora. They suggest a relationship of the Perophoridae with the Ascidiidae. 286. Porter, J. W. 1977. Pseudorca strandings. Oceans 10, no. 4: 8-16. This article provides information and observations surrounding the stranding of thirty false killer whales (Pseudorca crassidens) on the Dry Tortugas Islands near Florida in July 1976. The herd appeared to be protecting an injured male, as evidenced by aspects of social behavior and agnostic behavior directed at sharks and the author. Among other suggestions, the author postulates that the injured male was unable to feed due to parasitic infestation of the ears and consequent impairment of echolocation, which caused the whale to beach in order to avoid drowning in its weakened state. Other strandings are discussed in light of the information obtained. 13 287. Porter, J. W., J. F. Battey and G. J. Smith. 1982. Perturbation and change in coral reef communities. Proceedings of the National Academy of Science 79, no. 5: 1678-81. Ninety-six percent of surveyed shallow-water Dry Tortugas reef corals died during the severe winter of 1976-77. Data from skeletal stains indicate that death occurred during the mid-January intrusion of 14 degree C water onto the reef. In deeper water, community parameters such as percent cover, species number, and relative abundance showed no significant change. However, an analysis of competitive interactions at the growing edges of adjacent colonies reveals a 70% reduction in space competition during this environmental disturbance. These results can explain high variability in the growth rate of Floridian reefs and demonstrate the importance of obtaining long-term spatial information to interpret successional dynamics of complex communities. 288. Porter, J.W., O. W. Meier, L. Chiang and T. Richardson. 1993. Quantification of coral reef change (Part 2): the establishment and computer analysis of permanent photostations in the Florida SEAKEYS survey (abs). Proceedings of the Seventh International Coral Reef Symposium, v.1. Mangilao, Guam: University of Guam Marine Laboratory. Photostations in five of six locations in the Florida Keys reveal a decline of monitored coral reef resources during the 1980's when up to 40% of the coral died in some protected areas. Reductions in the number of species and extraordinary shifts in the pattern of species abundance occurred in addition to loss of live coral cover. While normally associated with catastrophic physical disturbances, this coral mortality occurred during a period without major hurricanes in Florida. Relocatable photostations reveal a multiplicity of causes for this decline. These include: (1) mortality due to "white band" and "black band" disease, (2) direct and delayed mortality from "coral bleaching," caused by abnormally elevated sea temperatures, (3) some mechanical damage, and (4) an increase in cover by algae. The establishment and sequential analysis of remote sensing data acquired from permanent photo-stations will be described in detail, as well as limits to the interpretability of these photogrammetric data. 289. Potthoff, T. and W. J. Richards. 1970. Juvenile bluefin tuna, Thunnus thynnus (Linnaeus), and other scombrids taken by terns in the Dry Tortugas, Florida. Bulletin of Marine Science 20, no. 2: 389-413. The identification and seasonal distribution of juvenile scombrids in the waters near the Dry Tortugas, Florida, are described. Specimens were collected (1960 through 1967) from regurgitated food of terns. Fishes identified were Thunnus thynnus, Thunnus atlanticus, Euthynnus alletteratus, Auxis spp., and Katsuwonus pelamis; sizes ranged from 24-146 mm. standard length. For the first time, juvenile bluefin tunas are reported in the Dry Tortugas region; their presence may indicate that spawning of the species takes place in the area. Identification methods are discussed, with special emphasis on features of the axial skeleton and the number of gillrakers over the ceratobranchial bone of the first gill arch. A method is presented for estimating the standard length of damaged specimens on the basis of the length of the vertebral column. 290. Powers, P. B. A. 1933. Ciliates infesting the tortugas echinoids. Carnegie Institution of Washington, Year Book 32: 278-79. Based on studies conducted on the ciliates of the Tortugas sea-urchins, Echinoida, it was found that when sea-urchins were infested, they made excellent reservoirs for certain species of ciliates in which to conduct detailed studies of their internal morphology, cytoplasmic inclusions and neuromotor apparatus. 291. ——. 1936. Studies on the ciliates of sea-urchins: a general survey of the infestations occurring in Tortugas Echinoids. Papers Tortugas Laboratory 29: 293-326 (issued Dec. 1935). Carnegie Institution of Washington Publication Number 452. 74 292. Pratt, H. 293. 294. With the accumulation of data concerning the ciliate infestations of the alimentary tract of echinoids, it became of increasing interest to have a complete record of the ciliates infesting sea-urchins about Tortugas. The writer spent the summer of 1933 at the Tortugas Laboratory. During this time twelve well-defined species of ciliates were found distributed among seven species of sea-urchins. The present paper gives a complete account of the general morphology of these ciliates, as well as a description of the various associations encountered. Twelve species of ciliates are described which infest the alimentary tract of seven species of sea-urchins from the region about the Dry Tortugas. Eight of the twelve are new. Only five of these twelve species show any marked host specificity. The remaining seven species all show a definite host preference. All of these ciliates are associated with their host when the latter are found occurring near the tide line. In sea- urchins taken below a depth of ten fathoms, only four species of ciliates are found : Cryptochilidium bermudense, Anophrys elongat, Cohnilembus coeci and form M. The nature of this infestation is one of endocommensalism, there being no present evidence to indicate pathogenic tendencies for any of its members. Data concerning the geographical distribution of the ciliates infesting sea-urchins from the localities of Beaufort, North Carolina, Bermuda and Tortugas have been summarized. It is suggested that the center of this infestation will be found in the sea-urchins from the region of the Lesser Antilles and that this infestation has been carried northward along with its host, through the agency of the Gulf Stream. S. 1910. Monocotyle floridana, a new monogenetic trematode. Papers Tortugas Laboratory 4: 1-9. Carnegie Institution of Washington Publication Number 133. The genus Monocotyle was established by Taschenber, in 1878, for a worm which he had found on the gills of the eagle-ray (Myliobatis aquila) at Naples, and which he named Monocotyle myliobatis. The only other known species is Monocotyle ijimae, which was discovered in Japan in the mouth of Trygon pastinacea, and described by Goto in 1894. The worm herein described makes the third member of the genus and was taken from the gills of the whip-ray (Myliobatis freminvillei) in the Gulf of Mexico and studied at the Marine Biological Laboratory at Dry Tortugas, Florida. It differs in certain features from the two other species of the genus, but in the general shape and size of the body, the form and structure of the suckers, down to the smallest details, and the general arrangement of the genital organs it shows a close relationship to them, especially to M. ijimae. . 1910. Parallel transport in tropical trematodes. Science 31: 471-72. The digenetic trematodes, as well as other internal parasites, have probably in their phyletic history followed somewhat different rules of descent from those of other animals. The fact that they live inside of other animals and have a very complex life history must affect their phyletic development, in that migrations are very much limited, and their structure is very uniform in the parasites themselves. It is probable that where there are apparently related species of digenetic trematodes living in widely separated localities, they possess the same or similar structural features. This does not necessarily indicate that there is a close genetic relationship between them. These facts are well illustrated by the several species of digenetic trematodes belonging to the genus Helicometra, which were found in certain fishes of the Tortugas, Florida, and also occur in the Meditteranean Sea. The species of this peculiar genus are thus taken as an indication, not that they necessarily bear a close genetic relationship to one another, but that similar or identical environmental conditions exist for them in these places, so that they have come to possess in the course of time a structure so similar that they are included in one and the same genus. . 1916. The trematode genus Stephanochasmus Looss in the Gulf of Mexico. Parasitology 8, no. 3: 229-38. 75 Two species of the genus Stephanochasmus were found in fishes examined for parasites at the Dry Tortugas: S. casus Linton and S. sentus Linton. The anatomy of these worms have | several interesting and unique features. S. casus Linton is described in this article. 295. ————. 1913. The trematode parasites of the loggerhead turtle. Science 37: 264-65. The studies of trematode parasites of the loggerhead turtle in the Mediterranean and the Gulf of Mexico, Dry Tortugas are briefly discussed and compared. Nineteen species occurred in the turtles of the Gulf of Mexico. Eight of these species also occur in the Mediterranean Sea. The most numerous trematode occurring in the Dry Tortugas is Cymatocarpus undulatus. A more detailed discussion is found in the Archives de Parasitologie article by Pratt. 296. ———. 1912. Trematodes of the Gulf of Mexico. Verhandlungen des VIII Internationalen Zoologen- Kongresses, 780-781. Jena, G. Fischer. This is a discussion of the article written by Pratt in no. 133 Carnegie Institution of Washington, listing the trematodes collected at the Dry Tortugas and the hosts they live in. 297. ————. 1916. Trematodes of the loggerhead turtle (Caretta caretta) of the Gulf of Mexico. Archives De Parasitologie: 411-27. Five species of trematodes are discussed in this paper. Reference is made to the studies of Linton (1910), as well as studies made in the Mediterranean Sea. The five trematode species were found in turtles captured on Loggerhead Key in the Dry Tortugas: Wilderia elliptica, Pachypsolus tertius, and Rhyditodes secundus, Pelsiochorus cymbiformis and Cymatocarpus undulatus. 298. Raim, A. W., W. Cochran and R. D. Applegate. 1989. Activities of a migrant merlin during an island stopover. Journal of Raptor Research 23, no. 2: 49-52. Activities of a radio-tagged merlin (Falco columbarius) which was trapped and identified as an adult female by George Allex and Daniel D. Berger, were observed from 10-16 April 1977 on Loggerhead Key. 299. Reighard, J. 1908. An experimental field-study of warning coloration in coral-reef fishes. Papers Tortugas Laboratory 2: 257-325. Carnegie Institution of Washington Publication Number 103. This paper embodies a search for the biological significance of the conspicuousness which it attempts to show characterizes many of the coral-reef fish of the Tortugas region. After showing that this conspicuousness is not a secondary sexual character and that it serves neither for protective nor aggressive resemblance, its value as a warning character is subjected to experimental test. Experimental evidence is presented to show that the gray snapper, the commonest predaceous fish, discriminates certain colors, forms associations with rapidity, and retains these for a considerable time (memory). If any of the coral-reef fishes possess a combination of consicuousness with such unpleasant attributes as render then unpalatable, the gray snapper should have learned to avoid them at sight and their conspicuousness would then have a warning significance. It is shown that when atherina, an inconspicuous fish which serves normally as the food of the gray snapper, is given an artificial warning color and at the same time rendered unpalatable, it is after a brief experience, no longer taken as food by the gray snapper. Artificially colored atherinas thus come to have a warning significance for the gray snapper and are avoided, even when not unpalatable, although normal atherinas are still readily eaten. The conclusion is thus reached that the existence of a warning coloration or of warning conspicuousness in coral fishes is easily possible. The conclusion is reached that the conspicuousness of coral-reef fishes, since it is not a secondary sexual character and has no necessary meaning for protection, aggression, or as warning, is without biological significance. The coral-reef 76 fishes have no need of aggressive inconspicuousness because their food consists of invertebrates, chiefly fixed. They have no need of protective inconspicuousness because the reefs and their agility afford them abundant protection. Selection has therefore not acted on their colors or other conspicuous characters, but these have developed in the absence of selection and through internal forces. They are the result of race tendency unchecked by selection . 300. . 1907. The photography of aquatic animals in their natural environment. Bulletin of the United States Bureau of Fisheries 27: 41-68. This paper describes the photography of aquatic organisms in their native environment and under normal conditions by carrying the camera into the field. Photos and diagrams are provided for cameras and apparatuses that remain above the surface of the water and cameras that are submerged. 301. Reynolds, J. E. III and J. C. Ferguson. 1984. Implications of the presence of manatees, Trichechus manatus near the Dry Tortugas Islands, Florida USA. Florida Scientist 47, no. 3: 187-89. | Two West Indian manatees (Trichechus manatus) were observed 61 km northeast of the Dry Tortugas Islands, a location not normally considered to be part of the species range. ) When spotted, the animals were swimming in a soutwesterly direction, away from Florida. Observations such as this, of manatees far from freshwater, raise the question of whether manatees require regular access to freshwater for osmoregulation, as suggested in the literature. 302. Reynolds, J. E. III and J. C. Steinmetz. 1983. Dry Tortugas: products of time. Sea Frontiers 29, no. 2: 66-75. This article discusses the general formation of the islands and their history. 303. Richards, O. W. 1934-1936. Growth studies in the ascidian, Phallusia nigra, and hermit crab, Caenobita clypeatus. Carnegie Institution of Washington, Year Book Note: published as follows 1934, v. 33, p. 261; 1936, v. 35, p. 92. The early growth and development of the ascidian Phallusia nigra was recorded over time using motion and still photography. Claw size ratios to body size of the hermit crab, Coenobita clypeatus, were examined by correlation methods. 304. Ricklefs, R. E. and S. C. White. 1981. Growth and energetics of chicks of sooty tern, Sterna fuscata and common tern, Sterna hirundo. Auk 98, no. 2: 361-78. The energy budgets of chicks of the common tern (Sterna hirundo) were measured on Great Gull Island, New York. Also measured were the sooty tern (S. fuscata) on the Dry Tortugas, Florida The respiratory energy requirement was determined by measuring oxygen consumption in a closed system. The growth energy requirement was calculated from the lipid and protein contents of a series of chicks spanning the range between hatching and fledging. Energy budgets calculated for the two species differed in several ways. (1) Maintenance metabolism was lower in the sooty tern owing to its warm environment. (2) Sooty terns allocated more of their energy intake to lipid accumulation from an earlier age. (3) In the sooty tern, the allocation of energy to growth initially was high, but its absolute amount decreased steadily throughout the growth period. In the common tern, both growth and maintenance energy allocations increased rapidly during the first half of the development period. (4) In sooty tern chicks energy metabolism approached its maximum rate (135 kJ/day) by the end of the first third of the development period, after which it leveled off. In the common tern, energy metabolism increased from about one-quarter of its maximum during the first five days after hatching to its maximum of 200kJ/day during the third week of the postnatal development period. Although these observations support the hypothesis that slow growth in pelagic seabirds is selected to Vi reduce the energy requirement of the chick, the energy budgets also suggest that a doubling of the growth rate by the sooty tern would increase the maximum energy requirement of the chick by only 20% and the total feeding requirement of the adult by only 5%. Moreover, the levels of water in muscles suggest that the sooty tern develops mature function earlier than does the common tern, which in itself might be sufficient to account for the slower growth of the first species. 305. Ricklefs, R. E. and S. C. White-Schuler. 1978. Growth rate of the brown noddy on the Dry Tortugas Florida USA. Bird-Banding 49, no. 4: 301-12. Growth rates within seabird species can vary with locality, season, and year. In this study noddy tern checks captured on Bush Key, Dry Tortugas, June, 1972, were weighed and measured. Growth increments were used to calculate a composite wing length growth curve to estimate the ages of chicks. A logistic curve was fitted to describe the relationship between weight and age. Growth constants of the fitted curve (growth rate K = 0.153, asymptote A=160 grams, and age at inflection ti=14.0 days) were similar to values reported for the brown noddy on Kure Island and Manana Island, Hawaii. Also reported are outer primary and rectix lengths and body temperatures of nestlings and adults. 306. Riley, G. A. 1938. Study of the plankton in tropical waters. Carnegie Institution of Washington Year Book 37: 98. The small quantity of plankton in tropical waters as contrasted to higher latitudes is investigated, and when compared to a similar survey underway in Long Island, N.Y., the indication is that chlorophyll and plant pigments are one-twenty-fifth the amount found in New York. 307. Riska, D. E. 1986. An analysis of vocal communication in the adult brown noddy, Anous stolidus. Auk 103, no. 2: 359-69. The author analyzed vocal signals of marked adult Brown Noddies (Anous stolidus) throughout their nesting season in the Dry Tortugas, Florida from 1979 to 1982. The basic unit of the adult repertoire is a wide-band click, less than 4 msec duration, ranging in frequency from 200 to 3,300 Hx. He identified nine temporal arrangements of these clicks, which form the notes of the calls. These calls differ little in frequency range, but they differ in the mean frequency of the most intense sound energy band, in note duration, in the number of clicks per note, and in internote interval. These calls are used in different contexts, which sometimes overlap. Frequency, note duration, and length varied among individuals for some calls. No tonal elements characteristic of calls of brown noddy nestlings remain in the adult repertoire. 308. ———. 1986. "Communication behavior of the brown noddy (Anous stolidus) and sooty tern (Sterna fuscata), Dry Tortugas, Florida (vocalizations, laridae, signals, colonial, breeding)." Ph.D. Dissertations, University of California at Los Angeles. The basic unit of the adult repertoire is described as a wide-band click, less than 4 msec duration, in the frequency range 200 to 3300 Hz. Nine calls differ in temporal arrangements of clicks, mean frequency of the most intense sound energy band, note duration, number of clicks per note, and inter-note interval. Frequency, not duration, and inter-note interval do not differ between sexes. The nestlings of the brown noddy produce three structurally different vocalizations within one day after hatching. Postures of chicks and contexts in which these signals are used differ. The repertoire is composed of frequency-modulated tonal elements and broad-band bursts of sound with little resemblance to the adult repertoire. Juvenile bush-nesting noddies begin flying when 40- 48 days old, after which they are still fed at their nests. Adult noddies accept a substituted nestling differing from their own in size, color and plumage stage, up to at least 20 days post-hatching. The adult Sooty Terns produce eight structurally different vocalizations, 78 and nestlings produce three, in the frequency range 300-7000 Hz. Postures differ for each call, but contexts in which these are used overlap. The range of frequencies in which young birds call extends higher than that of adults, but the frequency-modulated tonal elements characteristic of nestling vocalizations remain complex in the adults. 309. Rivas, L. R. 1951. Preliminary review of the western North Atlantic fishes of the family 310. Roberts, 311. Roberts, Scombridae. Bulletin of Marine Science of the Gulf and Caribbean 1, no. 3: 213-30. This paper brings up to date the taxonomy of the western North Atlantic mackerels and tunas. In addition to a key to the genera and species, a complete synonymy, a diagnosis and pertinent comments are given under each species. H. H., L. J. Rouse Jr., N. D. Walker and J. H. Hudson. 1982. Cold water stress in Florida Bay and northern Bahamas, a product of winter cold air outbreaks. Journal of Sedimentary Petrology 52, no. 1: 145-55. During January 1977 three consecutive cold fronts crossed south Florida and the northern Bahamas which depressed shallow-water temperatures below the lethal limit for most reef corals. Digital thermal infrared data acquired by the NOAA-5 meteorological satellite, in situ water temperatures, and meteorological data were used to study the thermal evolution of Florida Bay and Bahama Bank waters. The third and most important frontal system depressed Florida Bay water below 16 degrees C, a thermal stress threshold for most reef corals, for 8 days. Coral mortality at Dry Tortugas was up to 91 percent during the 1977 event. Coral and fish kills were also reported from other parts of the Florida Reef Tract and northern Bahamas. Study results show that cold-water stress conditions can exist over vast shallow-water areas and have residence times of several days. H., H. Lawrence , J. Rouse Jr. and N. D. Walker. 1983. Evolution of cold water stress conditions in high-latitude reef systems: Florida Reef Tract and the Bahama Banks. Caribbean Journal of Science 19, no. (1-2): 55-60. Thermal depression of shallow bank and bay waters accompanying the passage of severe cold fronts can stress high latitude coral reef systems, such as those of the Florida Reef Tract and northern Bahama Banks. Laboratory and field experiments suggest that sustained temperatures below 16 degrees C are detrimental to most reef-building corals. Time-series satellite imagery provides a data base for assessing the thermal variability of waters interfacing with reef systems. Digital thermal infrared data acquired by the NOAA- 5 meteorological satellite were used to study thermal evolution of Florida Bay and Bahama Bank waters during a succession of three cold-air outbreaks (January 1977). These studies indicate that the temperature of subtropical bank and bay waters is subject to depression below 16 degrees C accompanying the outbreak of unusually cold air. This superchilled water can have a residence time of days. The cooling process creates water masses that are out of density equilibrium with warmer ocean water. Offshelf movement of the cold, dense water occurs at particular sites, as shown by time-series satellite data. The absence of coral reefs opposite tidal passes in the Florida Keys is attributed to this process, which has probably limited development of the entire reef tract. 312. Robertson, W. B. Jr. 1978. Species of special concern sooty tern. Birds 2, no. Edited by H. Kale: 89-90. A description, range, and habitat of the sooty tern are given along with its life history and ecology at the Dry Tortugas. Its classification is based not on its abundance, but it is because the Dry Tortugas colony is a major Florida wildlife resource. Aside from the Tortugas no other location in Florida is suitable for sooty tern nesting. This colony affords a means of monitoring the general health of offshore Gulf waters of southern Florida. gS S113), . 1964. The terns of the Dry Tortugas. Bulletin of the State Museum , Biological Science 8, no. 1: 1-95. New information from unpublished sources and from published records hitherto overlooked permit a re-evaluation of the history of the Dry Tortugas and of the terns that inhabit them. The geography and ecology of the 11 keys that have variously comprised the group since it was first mapped in the 1770's are described and their major changes traced. The recorded occurrences of the seven species of terns reported nesting on the Keys are analyzed in detail. The sooty tern colony has fluctuated from a low of about 5,000 adults in 1903 to a reported peak of 190,000 in 1950; for the past four years it has remained steady at about 100,000. The brown noddy population, which reached a peak of 35,000 in 1919, was reduced by rats to about 400 adults in 1938; it is in the neighborhood of 2,000 today. A colony of 150 to 450 roseate terns has nested in most years from 1917 to the present. About 500 least terns nested regularly trom 1918 to 1932, then unaccountably dwindled to a few pairs by 1937 and shortly afterward disappeared. Royal and sandwich terns nested abundantly in the mid-19th century, and a colony of royals may have existed as late as 1890. Both species are believed to have been extirpated from the Tortugas by egging. No verifiable evidence exists for the nesting of the common tern, which has been reported several times. The black noddy, first reported for the continental United States at Dry Tortugas in 1960, has been found there each summer since. 314. . 1969. Transatlantic migration of juvenile sooty terns. Nature 222: 632-34. From 1959 to 1968, 70,000 adult and 130,000 juvenile sooty terns (Sterna fuscata) were banded at Bush Key, Dry Tortugas, Florida. By December 1968, 29 juveniles were recovered in West Africa. It appears that the primary biological function of the transatlantic migration is to avoid intraspecific competition and this adaptive value becomes evident when the migration of juveniles is seen in the context of the rigidly structured sooty tern population. It may be evidence for a successful evolutionary mechanism. 315. Robertson, W. B. Jr. and B. Given. 1980. Ruddy quail dove Geotrygon montana again at Dry Tortugas Florida USA. Florida Field Naturalist 8, no. 1: 23-24. About noon on December 15, 1977, a cold day with severe northwesterly squalls, Given found and photographed a large, reddish dove on the second tier of Fort Jefferson, Dry Tortugas, Florida. This record is the fifth report of the species from Florida and the second from Dry Tortugas. 316. Robertson, W. B. Jr. and C. R. Mason. 1965. Additional bird records from the Dry Tortugas. Florida Naturalist 38: 131-38. Sprunt (1962-63) summarized what was known about the occurrence of birds at the Dry Tortugas through the Summer of 1962. In this paper the authors report on recent bird records up to April 1965. Comments relate to 12 species new to the list or those known from either one or two records. Sprunt listed 227 species of birds for the Tortugas, the authors add 12 to bring the total to 239 species. 317. Robertson, W. B. Jr. and L. C. Below. 1975. A red-headed woodpecker at Dry Tortugas. Florida Field Naturalist 2, no. 1: 20. On May 5, 1973 Mr. and Mrs. G.H. Perbix of Cincinnati and Mrs. Below, members of the tern-banding party then at Dry Tortugas, visited Loggerhead Key and at once noticed an adult red-headed woodpecker (Melanerpes erythrocephalus) in the large Australian Pines (Casuarina equisetifolia) near the dock. We find only one other report of the red-headed Woodpecker at Dry Tortugas. Howell (1932:308) wrote that the species was unknown in the Florida Keys "...except for a single occurrence on the Tortugas - a bird seen there on a number of days early in June." The red-headed woodpecker is not known to occur outside 80 the United States but the present record inevitably raises the question: was the bird migrating across the Gulf or was it merely a vagrant? 318. Robertson, W. B. Jr. D. R. Paulson and C. R. Mason. 1961. A tern new to the United States. Auk 78: 423-25. This note provides a description of the black noddy, Anous tenuirostris collected at Dry Tortugas. This is the first of this species collected in the United States. Two specimens were taken from Bush Key during July 1960. The bird occurs nearly world-wide in the warmer seas, but is absent from most of the Atlantic Ocean north of the equator and most of the Caribbean Sea. 319. Robinson, A. H. 1976. Marine, island and coastal parks in the United States National Park system: A review and progress report in 1975. International Conference on Marine Parks and Reserves., pp. 226-27. Gland, Switzerland: IUCN. This paper provides a basic introduction to critical marine habitats and the planning and management of marine parks and reserves, including interpretation and environmental education in marine parks. Progress in the creation of marine parks and reserves is reviewed, and a special report on marine park systems in the Pacific region is included. Fort Jefferson National Monument is discussed as the first underwater preserve established in the United States. 320. Schaeffer, A A. 1925. Experiments on the influence of temperature and dilute and concentrated sea-water on ameboid movement. Bulletin of the Ecological Society of America: 11. The reactions of various species of amoebas to different concentrations of sea-water have been used during the past several years at Tortugas as important aids in the identification and fixation of species. The rate of movement of several species was studied in various concentrations of sea-water indicating that the optimal concentration of sea-water is below the norm in every case when measured by the rate of cell-coordinated movement. 321. ————. 1926. Taxonomy of the amebas: with descriptions of thirty-nine new marine and freshwater species. Papers Tortugas Laboratory 24: 1-116. Carnegie Institution of Washington Publication Number 345. The purpose of this report is to set forth a description of 39 new species and 11 new genera of amebas (Amoebaea), and to propose a preliminary system of classification of the amebas, based on their general morphology. General observations on structure, physiology, distribution, and methods of investigation are provided. The changes of form which amebas undergo is a fundamental morphological characteristic of amebas, and forms the basis of a natural classification. For the purpose of quickly recognizing a species other characteristics are more valuable, such as the nucleus, vacuoles, crystals, resistance to dilutions and concentrations of sea-water, etc. A brief discussion of these characteristics with reference to specific descriptions is given along with colored drawings and photographic text-figures. 322. Schmitt, W. L. 1924-1932. Systematic-ecologic studies of the decapod crustacea. Carnegie Institution of Washington, Yearbook. Note: published as follows: 1924, v.23, p. 200-201; 1925, v.24, p. 230-231; 1930, v.29, p. 34371931, v:30) pe 3892 193825v. 37, p. 279: Very striking color characteristics/variations affecting chela, and often the appendages, are noted among snapping shrimp, Synalpheus, and giant isopod crustaceans. Bathymetric distribution of decapods are investigated. 323. Schnell, G. D. 1974. Flight speeds and wing beat frequencies of the magnificent frigate bird. Auk 91, no. 3: 564-70. 81 Wingbeat frequencies and flight speeds of magnificent frigatebirds were recorded with a Doppler radar in the Dry Tortugas, Florida. The flapping rate averaged 2.84 beats per second (SD 0.14) and was not significantly correlated with flight speed, providing further evidence that the birds’ wingbeat frequency is essentially constant within species. The flapping rate is somewhat higher than predicted from the theory of mechanical oscillators when the distance from the end of the wing to the first articulated joint is used as an estimate for the average effective wing length. Flight speeds of birds in a flat calm averaged 22.55 mph. The highest average ground speed of 30.17 mph was obtained from frigatebirds flying in a 6 to 8 mph wind, and the lowest of 16.00 mph for birds flying into the 65 mph wind. Airspeeds were greater for frigatebirds flying into the wind than for those moving across or with the wind. 324. Schreiber, R. W., W.B. Robertson Jr. and T. Bellow. 1976. Nesting of brown pelicans, Pelecanus occidentalis, on the Dry Tortugas, Florida. Florida Field Naturalist 3, no. 2: 47-48. On June 14, 1974 Bush Key, Dry Tortugas Ted Bellow and C. Winegarner found 5 brown pelican nests about 12 feet above ground in the white mangroves (Laguncularia racemosa) along the north shore. Nineteenth-century records of pelicans breeding on the Dry Tortugas are ambiguous. ...on the Tortugas (1860) it thus appears that a few pairs did breed on the Tortugas in the mid-1900's, but by late in the century none did so. Our record is the first reported nesting of this species in the 20th century on the ornithologically well-known islands (Robertson and Mason, 1965). Three of the nests found in 1974 contained two eggs each, one nest was empty, and the fifth was not checked. 325. Schroeder, P. B. and J. H. Davis. 1971. Ecology vegetation and topography of the Dry Tortugas updated to 1970. Quarterly Journal of the Florida Academy of Science (Supp! 1): 12-13. The half-dozen islets of the Dry Tortugas have been ecologically studied periodically since the turn of the century. In November and February a year ago, a field party from the University of Miami made a topographical and vegeational study of several of these keys. The pertinent information gathered at that time is now available and provides continuity with the studies of Millapaugh (1907), Bowman (1918) and Davis (1942). The keys studied have changed from barren coral and sand to substantial islets largely covered with vegetation. The configuration of one of these has been completely altered. All the keys have been changed considerably in shape. Vegetational communities have shown similar changes and maturity. Mangrove areas (red and white) have become established and enlarged. Australian pines and other exotics, introduced to Loggerhead Key, have spread over much of the island and now are found on Bush Key. 326. Scott, W. E. D. 1890. On birds observed at the Dry Tortugas, Florida, during parts of March and April, 1890. The Auk: A Quarterly Journal of Ornithology 7, no. 4: 301-14. The list of birds observed at the Tortugas includes eighty species, fifty seven of which were land birds. The author states that no land birds breed on any of the keys group, and that the stay of any land bird is of very short duration. 327. Seaman, W. Jr. and D. Y. Aska. 1974. Research and information needs of the Florida spiny lobster fishery. State University System of Florida Sea Grant Program, Miami FL. 64 pgs. In response to a number of fishermen in South Florida, the State University System of Florida Sea Grant Program became involved in research on the spiny lobster, Panulirus argus. When additional research needs were expressed, Florida Sea Grant decided to become better informed on the subject, and evaluate its potential for service to the persons dependent on this fishery resource. A meeting of persons and organizations involved in the biology and/or utilization of the spiny lobster fishery in Florida was called to identify broadly the problems and information needs of persons dependent on the spiny lobster 82 : resource, to assess existing sources of information and their possible applications, and to identify priorities and actions needed to resolve user problems. 328. Shinn, E. A., J. H. Hudson, R. B. Halley and B. Lidz. 1977. Topographic control and accumulation rate of some Holocene coral reefs: South Florida and Dry Tortugas. Proceedings, Third International Coral Reef Symposium, RSMAS, Univ. of Miami, Coral Gables FL. p.1-7. Core drilling and examination of underwater excavations on 6 reef sites in south Florida and Dry Tortugas revealed that underlying topography is the major factor controlling reef morphology. Carbon-14 dating of coral recovered from cores enables calculation of accumulation rates. Accumulation rates were found to range from 0.38 m/1000 years in thin Holocene reefs to as much as 4.85 m/1000 years in thicker buildups. Cementation and alteration of corals were found to be more pronounced in areas of low buildup rates than in areas of rapid accumulation rates. Acropora palmata, generally considered the major reef builder in Florida, was found to be absent in most reefs drilled. At Dry Tortugas, the more than 13-meter thick Holocene reef did not contain A. palmata. The principal reef builders in this outer reef are the same as those which built the Pleistocene Key Largo formation, long considered to be a fossilized path reef complex. 329. Shinn, E. A. 1984? Geologic history, sediment, and geomorphic variations within the Florida Reef Tract. Advances in reef sciences, abstracts and schedule of presentations: a joint meeting of the Atlantic Reef Committee and the International Society for Reef Studies , 113-14. Miami, Florida: University of Miami. A combination of core drilling, high resolution seismic profiling, and constituent particle analysis reveal these major aspects of Holocene reef development and sediment distribution within the Florida reef tract: (1) reef distribution and shape are controlled by underlying Pleistocene limestone topography; (2) accumulation of sand and rubble occurs in forereef and backreef areas; and (3) composition of sediment and coral distribution are controlled by the reef tract trend relative to prevailing wind and exposure to Gulf of Mexico water. 330. Shinn, E. A., B. H. Lidz, R. B. Halley, J. H. Hudson and J. L. Kindinger. 1989. Reefs of Florida and the Dry Tortugas: Miami to Key West, Florida, July 2-7, 1989 . 28th International Geological Congress. Field Trip Guidebook (American Geophysical Union), no. T176. Washington, D. C.: American Geophysical Union. This field guide concentrates on explaining the distribution of Holocene coral reefs, the relationship between topography and Holocene sea-level rise, and the compositional and thickness variation of sediments produced in and adjacent to the reefs. A discussion and speculation of the future of the reefs under a stable sea, and a lowered sea-level is included. Also attached is a key to the Stony Corals of the Florida Keys and Dry Tortugas, a species list, illustrations of geologic cross-sections, aerial and underwater photographs of reefs and coral. 331. Shoemaker, C. R. 1934. Two new genera and six new species of Amphipoda from Tortugas. Papers Tortugas Laboratory 28: 245-56 (issued Nov. 1933). Carnegie Institution of Washington Publication Number 435. New genera and species of Amphipoda are described from specimens collected at the Tortugas, including Socarnes concavus, Gitanopsis tortugae, Heterophilas seclusus, Pontogeneia longleyi, Ampithoe divursia, Leucothoides pottsi (new species); and Heterophlias, Leucothoides (new genera). 332. Silberman, J. D., S. K. Sarver and P. J. Walsh. 1994. Mitochondrial DNA variation and population structure in the spiny lobster Panulirus argus. Marine Biology 120: 601-8. Adult spiny lobsters (Panulirus argus) were collected from nine locations including the 83 Tortugas, throughout the tropical and subtropical northwest Atlantic Ocean and examined for mitochondrial DNA (mtDNA) variation. 187 different mtDNA haplotypes were observed among the 259 lobsters sampled. Haplotype diversity was calculated to be 0.986 and mean nucleotide sequence-diversity was estimated to be 1.44%; both of these values are among the highest reported values for a marine species. Analysis of molecular variance (AMOVA) and phenetic clustering both failed to reveal any evidence of genetic structure within and among populations of P.argus The present data are consistent with high levels of gene flow among populations of P.argus resulting from an extended planktonic larval stage and strong prevailing ocean currents. 333. Smayda, T.J., Y. Shimizu, C. R. Tomas and D. G. Baden. 1993. The influence of phosphorus source on the growth and cellular toxin content of the benthic dinoflagellate Prorocentrum lima. Toxic Phytoplankton Blooms In The Sea., 565-70. The relationship between toxin content and nutritional status of the toxic marine phytoplankton species Prorocentrum lima was examined in a clonal culture isolated from the Dry Tortugas, Florida, grown with inorganic phosphate and glycerol phosphate enriched media. Growth, alkaline phosphatase activity and okadaic acid content were measured. Phosphate enriched cultures exhibited rapid growth rates(0.75 div/d), moderate terminal densities of 134,779 cells/ml and low alkaline phosphatase activity (<14 fg/cell/min). Cells grown with glycerol phosphate had lower growth rates, between 0.16 and 0.45 div/d, but higher maximal densities, >200,000 cells/ml, and had alkaline phosphatase activity an order of magnitude greater than those grown in inorganic phosphate. When comparing toxin levels at 20 and 30 days, cells grown on the organic phosphate enrichments had consistently higher per cell values (11.2 and 14.2 pg/cell, respectively) than those with inorganic phosphate (7.5 and 8.9 pg/cell), respectively). Phosphorus source effected growth, maximal densities, and okadaic acid content of P. lima. ; 334. Smith, H. G. 1937. Contribution to the anatomy and physiology of Cassiopea frondosa. Papers Tortugas Laboratory 31: 17-52 (issued July 1936). Carnegie Institution of Washington Publication Number 475. This research was undertaken to extend our previous scanty knowledge on the physiology of feeding and digestion in the Scyphozoa. Cassiopea was selected as the experimental material for two reasons, it is a member of the Rhizostomeae in which the mode of feeding is particularly interesting owing to the sub-division of the mouth, while certain species, including the one studied, possess zooxanthellae. Other aspects of the structure and physiology of species of this genus have been extensively studied, notably at the Tortugas Laboratory, by Mayer and others. It was originally intended to work on C. xamachana, which was very abundant at one time in the moat at Fort Jefferson. Recent changes in the conditions in the moat, the result of silting up, have caused the complete disappearance of the species although from this locality. In the absence of this species, C. frondosa was investigated and this although less hardy than C. xamachana, proved satisfactory material. Experiments have been made on the effect of starvation in light and in darkness on the medusae. In light, specimens were kept alive for 15 days, and in darkness for 7 days. Numerous algae were ejected by way of the gastric filaments and plaited membranes at the base of the filaments and the medusae became brown in color. They also shrank considerably in size. The effect of the zooxanthellae on phosphorus excretion has been studied, the amount of phosphorus in the sea-water surrounding one specimen being reduced to zero within 24 hours. Finally, feeding, digestion and symbiosis in C. frondosa have been discussed. It has been suggested that the variation in pH in the coelenteron affects the activity of the jellyfish. The association with zooxanthellae is probably similar in nature to that which occurs in the Madreporaria. 84 335. South Florida Area : Synthesis of available biological, geological, chemical, socioeconomic and 336. Spence, cultural resource information. 1990. OCS Study, MMS 90-0019. U.S. Department of the Interior, Minerals Management Service, Atlantic OCS Regional Office. This study summarizes the available biological, geological, chemical, and socioeconomic information in south Florida in relation to the potential effects of offshore gas and oil exploration and development. The synthesis will help Federal and state policy makers make informed decisions about future lease offerings and environmental restrictions on offshore oil and gas operations. The Dry Tortugas is included as part of the South Florida Reef Tract. In summary it would be very difficult to protect the mangroves, reefs, seagrass beds and their associated assemblages from large oil slicks. Severe weather would make it impossible. The Dry Tortugas experienced an oil spill from the beaching of Brother George in 1964. Birds were killed . Some coral may have been killed around the Tortugas from the 3,100 barrel spill, but it did not affect other areas further to the east of the site. If a large oil spill did occur here it would take 100+ years for the oldest coral heads to regrow and achieve the same level of pre-spill structural complexity. The effects of an oil spill on other flora and fauna of the Florida Reef Tract can only be guessed. J., and O. W. Richards. 1940. Native cellulose in the ascidian Phallusia nigra. Papers Tortugas Laboratory 32: 163-67 (issued Sept. 1940). Carnegie Institution of Washington Publication Number 517. Many organic compounds of high molecular weight are readily identifiable from the characteristics of the X-ray diffraction diagrams. Cellulose and its derivatives have been extensively examined by X-ray diffraction methods in the search for a complete solution of the structure and crystallite arrangement of the cellulose molecule. From the analytical standpoint, X-ray diffraction diagrams not only confirm the initial chemical identification of cellulose by Schmidt (1845), but they also show the presence of crystallites and their orientation. The Phallusia nigra was collected in the moat of Fort Jefferson and the tunic was removed on return to the Tortugas Laboratory. The result, namely, the recognition of native cellulose and the preferred orientation of the crystallites in Phallusia nigra, is naturally anticipated from previous observations on other ascidian tests. This method provides a useful analytical "tool" for use in zoological investigation. 337. Sprunt, A. Jr. 1963. Birds of the Dry Tortugas. Florida Naturalist: 22-26, 52-53. 338. ——. 339, ———_. 340. ———. 341. ———. This is a continuation of the listing from the 1962 series on listings of birds of the Dry Tortugas. 1962. Birds of the Dry Tortugas 1857-1961. Florida Naturalist 35: 35-40, 82-85, 129-32. A brief discussion of the history of bird studies of the Dry Tortugas is given. Special attention is paid to the migratory birds passing through the Dry Tortugas in hope of shedding light on trans-Gulf migration. 1947. Blizzard of birds: the Tortugas terns. National Geographic Magazine February: 213-30. This article gives a history of the Tortugas terns up to 1947. Boobies and noddies are included also. 1950. Bridled tern, Sterna a. Melanoptera, taken at Dry Tortugas. Auk 67, no. 4: 514. This article provides an account of the first Sterna melanoptera recorded at the Dry Tortugas, and the fifth specimen recorded in Florida. 1950. A list of birds of the Dry Tortugas Keys, 1857-1949. Florida Naturalist 23: 49-60, 73-78, 105-11. 342. 343. 85 A listing of the birds of the Dry Tortugas is given. Land birds-pigeons through vireos- including warblers through sparrows and water birds. . 1951. Some observations on the fall migration at Dry Tortugas. Auk 68: 218-26. The author arrived at the Tortugas following a hurricane August 26-27, which seemed to have no effect on the Tortugas, or the birds there. He found the birds to be in good physical condition, with no signs of exhaustion. Birds were tame and could be approached. A listing of the birds sighted is given. . 1948. The tern colonies of the Dry Tortugas keys. Auk 65: 1-19. The first post-war (1945-46) status report on tern populations inhabiting the Keys of the Dry Tortugas is presented in this paper. A brief history on population counts dating back to 1832 by Audubon is given, as well as a description, mostly vegetative, on Keys utilized by terns for nesting activities. Tern springtime arrival and summer departure are discussed, along with numbers of eggs produced, nesting locations and tern behavior. Based on the square-yard unit system, it was determined that the population count for Sooties was 97,200, while a count of 550 was found for the noddies by numbers of nests. The tern populations have suffered virtually no damage during the occupation of the islands by naval forces. Aside from weather, predation by natural enemies includes sand-crabs and man-o'-war-birds. The tern colonies appeared safe, but certain topographical changes, such as the recent increase in vegetation may be problematical. 344. Stevenson, J. O. 1938. The tern colonies of Dry Tortugas. Bird-Lore 40, no. 5: 305-9. This article describes briefly the history of the tern colonies of the Dry Tortugas. The author visited the Tortugas on May 24, 1937, two years after a hurricane swept through the islands destroying Bird Key, the historic breeding grounds for thousands of sooty and noddy terns. 345. Steward, F. C. 1940. The growth of Valonia ventricosa J. Agardh and Valonia ocellata Howe in culture, with a note on the sap composition of Valonia ocellata Howe at Tortugas. Papers Tortugas Laboratory 32: 85-98 (issued Oct. 1939). Carnegie Institution of Washington Publication Number 517. So much physiological work has been done using species of Valonia that their mode of development has special interest. Living material of V. ventricosa and V. ocellata was collected at the Dry Tortugas, Florida. These species were chosen because of the difference in their morphology. Valonia ventricosa J. Agardh and V. ocellata Howe have been kept alive for over two years from their original collection. Vesicles of considerable size (V. ventricosa) and with all the characteristics of the plant in nature have been grown attached to a suitable substratum. The development of the vesicle and rhizoids from aplanospores is illustrated by a series of photographs. V. ventricosa also produces filaments which penetrate the substratum and from which close clusters of vesicles arise as they do in the normal habitat. The appearance of the aplanospore and growing vesicle between crossed Nicols is described and its bearing on the structure of the wall indicated. Valonia ocellata produces pear-shaped vesicles, cylindrical rhizoidal processes (which it is shown may become long and branched), and apparently proliferated masses composed of small cellular segments. The growth and development of all these structures from aplanospores, or the product of "segregative division" have been observed and are recorded by photographs. 346. Steward, F. C. and J. C. Martin. 1937. The distribution and physiology of Valonia at the Dry Tortugas, with special reference to the problem of salt accumulation in plants. Papers Tortugas Laboratory 31: 87-110 (issued Oct. 1936). Carnegie Institution of Washington Publication Number 475. 86 This paper presents the results of a survey, made during the summers of 1933 and 1934 of the physiological behavior of the two species of Valonia which are most abundant at Tortugas, Florida. One may well ask what justification there can be for yet another paper on Valonia. Whatever the legitimate claims which may be made for such attention, they are somewhat counterbalanced by the inaccessibility of Valonia, which has prevented that examination by a variety of investigators which is the best safeguard against overemphasis. Valonia macrophysa occurs at Tortugas only in the moat of old Fort Jefferson. This organism demands complete protection from the effects of swell and surf. The growth obtained on a horizontal ledge is luxuriant; that on an inclined or vertical surface sparse and irregular. In the protected locations it demands, V. macrophysa is exposed to and withstands, a wide range of light conditions and diurnal fluctuations in the composition of the external medium. Valonia ventricosa is abundantly obtained on Bird Key Reef. The distribution of V. ventricosa is complementary to that of V. macrophysa, and the solution of the problem whether the species are distinct, raised thereby , must await adequate transplant experiments. The range of sap composition which V. ventricosa and V. macrophysa exhibit at Tortugas in sea water is described. Differences occur in the composition of the sap of V. macrophysa grown in different parts of the moat of Fort Jefferson. The principal causal factor appears to be the light condition which it obtains during growth. In general the conditions which produce the most abundant growth of V. macrophysa \ikewise produce the greatest concentration of potassium and lowest concentration of sodium. 347. Stockard, C. R. 1908. Habits, reactions, and mating instincts of the "Walking Stick," Aplopus 348. 349. mayeri. Papers Tortugas Laboratory 2: 43-59. Carnegie Institution of Washington Publication Number 103. This investigation of a protectively adapted insect is important to show definitely whether the actions of such an animal are coordinated with its protective structure. It is concluded that the habits of Aplopus mayeri on its food-plant Suriana maritima are as truly protectively adapted as is its singular stick-like appearance. . 1911. The influence of regenerating tissue on the animal body. Papers Tortugas Laboratory 3, no. 41-48. Carnegie Institution of Washington Publication Number 132. It is stated that when the adult animal body begins to regenerate new tissue in order to replace a lost part, or when abnormal secondary growths arise, the condition of growth- equilibrium is disturbed and such a disturbance is followed by changes which affect the usual physiological condition of the body. The question as to whether the changes following normal regenerative growth are in any way similar to those effects resulting from malignant or abnormal secondary growths arises. . 1908. Studies of tissue growth. I. An experimental study of the rate of regeneration in Cassiopea xamanchana (Bigelow). Papers Tortugas Laboratory 2: 61-102. Carnegie Institution of Washington Publication Number 103. The author responds to the studies of Zeleny (1903 and 1905) in which he suggested that the greater the degree of injury within limits, the more rapid the rate of regeneration. Zeleny suggested that the animal with the greater number of appendages removed might exercise the regenerating ones more than the animal with less: activity should increase the rate of regeneration in animals. The author tests the influence of rest and activity on regenerating tissues of medusa and finds no increase in the regeneration rate from activity. Rate of regeneration was also tested against food consumption, distance of cuts from the margin of the medusa disks, cuts from different parts of variously shaped surfaces, removal of oral epithelium of different sizes and at different distances, and the influences of changed chemical conditions on regeneration. 87 350. Stoddart, D. R. and F. R. Fosberg. 1981. Topographic and floristic change, Dry Tortugas, Florida, 1904-1977. Atoll Research Bulletin 253: 1-56. Topographic and floristic surveys of the Dry Tortugas Keys in 1904, 1915, and 1937 have been used in discussions of the changing relationships between area and floristic diversity on small islands over time, and of the processes of colonization and extinction. It is shown that earlier topographic surveys are in general too unreliable to be so used. A list of Dry Tortugas plants, including all published records was as well as new collections made in 1962 and 1977, is presented, together with maps of the keys made in 1977. The total flora of about 130 species includes at least 35 native species, including 5 species of sea-grasses and 4 species of mangroves. Introduced species are largely confined to the two largest islands, and the floras of the smaller keys are dominated by a small number of native species. 351. Stone, R. G. 1931-1932. Effect of irradiation by radium upon regeneration in marine annelids. Carnegie Institution of Washington, Yearbook. Note: published as follows: 1931, v. 30, p. 395; 1932, v. 31, p. 279. The effect of combined beta and gamma radiations upon regeneration in polychaetes is studied. Histological material is being used to determine the source of new tissue in regenerated segments and to discover what tissues are affected by radiation. 352) . 1934. Radium radiation effects on regeneration in Euratella chamberlin. Papers Tortugas Laboratory 28: 157-66 (issued Jan. 1933). Carnegie Institution of Washington Publication Number 435. Regeneration in the polychaete annelids has been investigated in some instances but the histological changes are not so well known as in the oligochaetes. The influence of X-rays and radium upon regeneration in various animals has been demonstrated, but the polycheates have seldom been used in these investigations. It has been found that the effects of radiation are often limited to specific tissues; by reason of their greater susceptibility they may be injured or destroyed by the exposure. During the summers of 1931 and 1932 the author was able to study the effects of radiation upon polychaete regeneration at the Tortugas Laboratory. In the sabellid Euratella chamberlin, posterior regeneration of abdominal segments is rapid and complete. Regeneration is inhibited by sufficient exposure to the beta and gamma rays of radium. Similar exposure to gamma rays alone has no effect upon the amount of regeneration. No structural changes were observed in the radiated worms to account for this change. It is suggested that ionization induced by the beta rays is responsible for the failure of regeneration . 393), . 1936. Regeneration in the cirratulid Cirrineris. Papers Tortugas Laboratory 29: 1-12 (issued Nov. 1935). Carnegie Institution of Washington Publication Number 452. There has been considerable study of polychaete regeneration , but the observations are not as extensive as those among the oligochaetes. This investigation of Cirrineris was undertaken to determine the extent of segment replacement and the source of the new tissues. Material was secured at the Tortugas Laboratory. In summary a head region and six to seven segments posterior to it are regenerated when more than this number are removed. Posterior regeneration is rapid and complete; the approximate number of segments removed is replaced. Wound closure is effected in the same manner at both ends of Cirrineris. The edges of the everted intestine unite with the epidermis to close the body cavity. New nervous tissue arises by proliferation and inward migration of cells from the adjacent epidermis. The old nerve cells do not participate in regeneration. Material for regeneration of the intestinal lining arises by proliferation within the gut epithelium. Mesodermal structures regenerate from old mesodermal tissues. Replacement material is supplied by (a) nuclei and cytoplasm from muscles cells and connective-tissue elements 88 after degeneration of their differentiated cytoplasm; (b) peritoneal cells which furnish most of the new material. 354. Stoneburner, D. L. and C. S. Harrison. 1981. Heavy metal residues in sooty tern, Sterna fuscata tissues from the Gulf of Mexico and North Central Pacific Ocean. Science of the Total Environment 19, no. 1: 51-58. The comparison of mean cadmium, mercury and selenium concentrations in the eggs, feathers and body tissues of breeding sooty tern (Sterna fuscata) from the Dry Tortugas, Florida, and Lisianski Island, Hawaii, supports the hypothesis that a physiological mechanism exists which functions in the detoxification of heavy metals. The data collected from two geographically isolated populations of this pelagic bird indicate that the mechanism responds in a uniform manner to widely different environmental levels of heavy metals. The data and observations suggest that the mechanism evolved in response to natural fluxes of heavy metal concentrations in the marine ecosystem, not in response to | recent injections of heavy metal laden industrial wastes. 355. Stoneburner, D. L., P. C. Patty and William B. Robertson Jr. 1980. Evidence of heavy metal accumulations in sooty terns. Science of the Total Environment 14, no. 2: 147-52. Sooty terns from the population that nests at Bush Key, Dry Tortugas, Florida, had substantial burdens of Cd, Hg and Se. Analysis of selected tissues, feces and eggs by neutron activation techniques showed highest levels of Hg in eggs, feathers, and blood; of Cd in kidney and bone; and of Se in kidney, liver, and feathers. The concentrations of Cd, Hg, and Se in the eggs suggests that the heavy metals are being transmitted to succeeding generations. The significance of the concentrations, their effect on the reproductive success of the population, and the question of whether or not the metals transmitted to eggs represent "bio-magnification" merit further work. 356. Strom, R. N.,R.S. Bramen , W. C. Jaap, P. Donan, K. B. Donnelly and D. F. Martin. 1992. Analysis of selected trace metals and pesticides offshore of the Florida Keys. Florida Scientist 55, no. 1: 1-13. Trace metal and pesticide contents of sediments and producer and consumer organisms were analyzed from samples taken from eighteen stations off the Florida Keys from Biscayne National Park to the Dry Tortugas. Samples were analyzed for total mercury, tin (inorganic and organic), arsenic (inorganic and methylated), lead, copper, cadmium, and halogenated pesticides. Pesticide concentrations were below detection limits. In general, concentrations of trace metals increased from sediments to producers to consumers at each station. Though the concentrations tended to be low, some deviations were ascribed to human inputs. Fewer significant correlations were observed than expected, possible because of the dependence of the uptake mechanism upon the ability of the system (sediment, producer, consumer) to remove trace metals from particular materials. Sponges have this ability and may represent a useful means of monitoring the quality of the environment on a sustained basis. The results are generally consistent with a relatively clean environment with some localized anthropogenic effects. 357. Stromsten, F. A. 1911. A contribution to the anatomy and development of posterior lymph hearts of turtles . Papers Tortugas Laboratory 3: 79-87 Carnegie Institution of Washington Publication Number 132. This article concludes that the development of the posterior lymph hearts of turtles is initiated by he vacuolation of the postiliac mesenchymal tissue during the middle and latter part of the second week of development of the loggerhead turtle. The spongy tissue thus formed is then invaded by capillaries from the first two or three dorsolateral branches of the caudal portion of the postcardinal veins. The final stage in the development of the posterior lymph hearts is reached by the dilation and confluence of these veno-lymphatic 358. B59) 360. 361. 362. 363. 364. 89 sinuses, from before backward, forming a pair of sac-like organs, each with a single central cavity. ——. 1910. The development of the posterior lymph hearts of the Loggerhead turtle Thalassochelys caretta. Proceedings of the Iowa Academy of Science 17: 227-28. Observations made on the lymphatic systems of turtles indicates their origin is more or less independent of the venous system. Later investigations confirm this view, suggesting that posterior lymph hearts of the loggerhead turtle are developed from embryonal cappillaries, which have been captured and modified by the mesenchymal spaces of the post-iliac regions of the body. Tandy, G. 1931. The superficial structure of coral reefs; plant succession upon prepared substrata. Carnegie Institution of Washington, Year Book 30, 32: 395; 26S. Plant and animal successions were examined on concrete cubes planted in the water at three sites: Fort Jefferson moat, an iron wreck east of Loggerhead Key, and northwest of Loggerhead Key. Tartar, V. 1938-1939. Regeneration in the starfish Linckia and in the protozoan Condylostoma. Carnegie Institution of Washington, Year Book. Note: published as follows: 1938, v. 37, p. 99-102; 1939, v. 38, p. 230-31. Regeneration experiments were conducted on starfish with and without arms and isolated arms. Under normal conditions polarity of arms is not altered by isolation. Tube feet cell differentiation was examined in relation to color changes. In the ciliate, Condylostoma, the normal form and typical arrangement of cytoplasmic differentiations may easily be altered. Tashiro, S. 1914-1915. Further studies on CO) in sea water and CO) production in tropical marine animals. Carnegie Institution of Washington, Year Book. Note: published as follows: 1914, v. 13, p. 170; 1915, v. 14, p. 217-19. Studies were conducted on the presence of "free CO 2 in sea water." A rapid method to estimate amounts of CO, produced in sea water by marine animals was devised. Taylor, J. B. 1981. Premetamorphic veligers of Fort Jefferson Dry Tortugas, Gulf of Mexico, and Beaufort Inlet, North Carolina. Bulletin of the American Malacology Union, Inc. 50: 29- 30. (No abstract available). Taylor, W. R. 1928. The marine algae of Florida, with special reference to the Dry Tortugas. Papers Tortugas Laboratory 25: 1-219. Carnegie Institution of Washington Publication Number 379. The study of marine vegetation of the Dry Tortugas was originally undertaken to provide a simple check-list of algae of the islands for use of persons visiting the Carnegie Laboratory there, with a description of the more important ecological features and records of the locations where plants of experimental importance might be found. When it was discovered that information about Florida algae in general was scanty, the study extended to a thorough study of all available Florida material. Records of the occurrence of marine algae on the east coast of Florida and the Florida Keys were collected. This is the first time, since Harvey, Farlow and Melvill, that an attempt was made to list completely the Florida algae. . 1925. The marine flora of the Dry Tortugas. Revue Algologique 2: 113-35. The marine algae of the Dry Tortugas are listed, and a description of the distribution is given of the important types throughout the area. 90 365. Teas, H. J. and P. B. Schroeder. 1971. Vegetation analysis in the Dry Tortugas by remote sensing. Quarterly Journal of the Florida Academy of Science 34, no. (Suppl 1): 13. Detailed ground truth observations were carried out on the four large islands in the Dry Tortugas using aerial photography and 12S image enhancement equipment. Several vegetation associations (strand-beach, strand-dune, strand-scrub) are distinguished and a number of plant species identified (Rhizophora, Laguncularia, Bursera, Conocarpus, Casuarina, Cocos, and Phoenix). 366. Tennent, D. H. 1911. Echinoderm hybridization. Papers Tortugas Laboratory 3: 117-51. 367. 368. Carnegie Institution of Washington Publication Number 132. The Toxopneustes female x Hipponoé male and the reciprocal cross Hipponoé female x Toxopneustes male were easily made after allowing the eggs to stand in sea-water for some hours before fertilization. In the embryos of both crosses made in ordinary sea-water, which was alkaline, the Hipponoé influence showed a tendency to predominate. It is suggested that the variations in the alkalinity of the sea-water, which have been brought about artificially, may correspond to normal seasonal changes. The results of this and of other investigations show species tendencies toward different grades of temperature and of alkalinity. The explanation of the preponderance of one character over another in echinoderm hybrids seems to lie in the reaction of the species toward a complex of factors. . 1920. Evidence on the nature of nuclear activity. Proceedings of the National Academy of Science 6: 217-21. The author describes the results of the Arbacia eggs and other materials examined. Basophilic bodies found are not in the nature of chromidia, but are the result of indirect nuclear activity. . 1942. The photodynamic action of dyes on the eggs of the sea urchin, Lytechinus variegatus. Papers Tortugas Laboratory 35: 1-153. Carnegie Institution of Washington Publication Number 539. The work recorded in this paper was begun as a study of the experimental modification and control of cell division in the egg of the sea urchin, Lytechinus variegatus. Early in the investigation the photodynamic effects of the dye neutral red were found to be striking and it was decided to undertake a study of the effects produced by other dyes. Transmission of visible light by some of the filters was so low that the intensity of the light transmitted was not sufficient to produce photodynamic effect. With dyes that produced a photodynamic effect, irradiation of a solution of the dye resulted in the formation of a photocompound. This photocompound was the active agent in the production of the photodynamic effect. The threshold for violent surface reaction (blister cytolysis) of Lytechinus eggs in 1:150,000 solution of neutral red in sea water lay at about 2500 foot-candles. From this point to about 4300 foot-candles violent surface reaction usually stood at about 2 per cent. Between 7000 and 7500 foot-candles it increased to 20-25 per cent, and between 800 and 9500 foot- candles it increased to 75-90 per cent. At intensities from 300 to 10,000 foot-candles there was a regular increase in the violence of the surface reaction and complete inhibition of the cleavage processes. Irradiation in some of the solutions of dye at temperatures above 32° C. resulted in injury from which the eggs did not recover. In blister cytolysis the formation of blisters starts at a single point. Adjacent blisters come into contact with one another until the entire surface is covered. The content of these blisters is liquid, is clear on the living egg, and in the fixed egg seems to be the same as the cytoplasm of the egg with all formed components removed. It is conceivable that these components could be filtered out, but there is no evidence of the accumulation of granules at the point where the cytoplasm might have been extruded . 9] 369. Tennent, D. H. and V. H. Keiller. 1911. The anatomy of Pentaceros reticulatus. Papers Tortugas Laboratory 3: 113-16. Carnegie Institution of Washington Publication Number 132. This account is a description of the anatomy of Pentaceros reticulatus. Figures are used to illustrate the organs which are described. Those which seem of greatest interest are the intestinal caeca. These were found in some instances to be greatly distended, stimulation causing their contraction. In this behavior we support the idea of the analogy of the intestinal caeca of the starfish to the respiratory trees of the holothurian, an idea which has been based upon the similarity of position of these organs. 370. Tennent, D.H., C. V. Taylor and D. M. Whitaker. 1929. An investigation on the organization in a sea-urchin egg. Papers Tortugas Laboratory 26: 1-104. Carnegie Institution of Washington Publication Number 391. In this report the eggs of the sea urchin, Lytechinus, were studied from samples taken at the Tortugas. The differentiation of ectoderm-forming substance over the entire surface of the egg begins before fertilization by the exclusion of the endoderm-forming material from the superficial layers of the egg. The number and relative distribution of micromeres is independent of the plane of section and of the size of the fragment. There is no localization of micromere-forming material. 371. Tennent, D. H., M.S. Gardiner and D. E. Smith. 1931. A cytological and biochemical study of the ovaries of the sea-urchin Echinometra lacunter. Papers Tortugas Laboratory 27: 1-46. Carnegie Institution of Washington Publication Number 413. The investigations upon which this paper is based constitute a new method of attack on the problem of the functional significance of chondriosomes, Golgi bodies and other "inclusions" in protoplasm. In 1926 a definite research program for histochemical and biochemical study of the eggs and ovaries of the sea-urchin Echinometra lacunter was begun. Summarizing: analytical figures for the percentage of lipids and of glycogen are given. The amount of the latter, 12.42 per cent, and 12.72 per cent of the dried extracted tissue is high. In addition, the presence of cerebrosides and sphingomyelin are indicated. The lipid composition of this tissue seems to be complex. The unsaturation of several preparations used in the study of staining reactions was determined, to find out if there was any correlation between unsaturation and staining with osmic acid (see Section I). On the whole the lipids are probably more unsaturated than similar preparations from mammalian tissues. 372. Thiele, J.. 1916. Molluskenfauna Westindiens. Zoologische Jahrbucher Supplement II: 109-32. A listing of the mollusks of the West Indies is given and a preliminary catalogue of the shell-bearing marine mollusks and brachiopods of the southeastern coast of the United States. This article is in German. 373. Thompson, M. J. and T. W. Schmidt. 1977. Validation of the species/time random count technique sampling fish assemblages at Dry Tortugas. Proceedings of the Third International Coral Reef Symposium, No.1:283-288. Miami, Florida: RSMAS, University of Miami. Ichthyofauna at four coral reef sites in Fort Jefferson National Monument, Dry Tortugas, are compared during summers 1975 and 1976. Samples were taken using the species/time random count technique, a newly developed visual censusing method based upon the rate at which species are encountered by a free swimming observer. Data were collected by different observers during the two years’ sampling. Within nine fish families dominating the Tortugas ichthyofauna, the rank of five did not vary at all between 1975 and 1976 samplings. Among the four families exhibiting changes in abundance, only the Serranidae showed a variation greater than 10.0%. The marked variation of 25.8% within this family is attributed to identification problems within the genus Hypoplectrus. Overall numbers of 92 species and relative species abundances within each sampled coral reef area showed minimal variation between years. The species rank correlation coefficient (Spearman's r s) between two years of observations was 0.92. High correlation between results from two different observation teams shows the species/time random count technique to be a highly reliable method of comparing coral reef fish assemblages. 374. Thorp, E.M., A. Mann, T. W. Vaughan and F. J. Haight. 1936. Calcareous shallow-water marine deposits of Florida and the Bahamas. With appendices: 1.Mann, A. Diatoms in bottom deposits from the Bahamas and the Florida Keys; 2. Vaughn, Thomas Wayland. Current measurements along the Florida Coral Reef Tract with notes on current observations, Florida Keys, June, October, November, 1914. See separate entries for Appendices. Papers Tortugas Laboratory 29: 37-143 (issued Dec. 1935). Carnegie Institution of Washington Publication Number 452. Determinations of the quantities of material derived from organic and inorganic sources have yielded the following results: Coralline algae, collectively, are shown to be the organic group that makes the largest contribution of organically secreted calcium carbonate. Next in order of magnitude are the mollusks, followed in descending order by foraminifera, madreporarian corals, alcyonarian spicules, worm tubes, crustacean fragments, and Bryozoa. The principal non-calcareous mineral is quartz. Quantitative counts of alcyonarian spicules show that they are relatively minor components of the sediments, being exceeded by madreporarian fragments in a ratio of about 2.5 to 1. Terrigenous minerals are remarkable scarce. A very small amount of volcanic glass and a few species of heavy minerals occur well distributed over the region. The sources of all the volcanic glass and some of the heavy minerals are thought to be distant and that they are wind blown. Coal and ashes brought from outside sources by human agencies have been introduced into the sediments of Tortugas lagoon, and, in smaller quantities, in a few other places. 375. Tomas, C. R. and D. G. Baden. 1993. The influence of phosphorus source on the growth and cellular toxin content of the benthic dinoflagellate Prorocentrum lima. Fifth International Conference on Toxic Marine Phytoplankton, 565-70. St. Petersburg, Florida: Florida Marine Research Institution. The relationship between toxin content and nutritional status of the toxic marine phytoplankton Prorocentrum lima was examined in a culture from the Dry Tortugas, grown with inorganic phosphate and glycerol phosphate. Phosphorus source affected growth, maximal densities, and okadaic acid content of Prorocentrum lima. 376. Torrey, H. B.. 1927-1928. Effect of thyroxin on division rates of various cells. Carnegie Institution of Washington, Year Book. Note: published as follows: 1927, v. 26, p. 228-229; 1928, v. 27 , p. 287. Thyroxin depressed cell division and differentiation in eggs of sea-urchin (Echinometra lacunter), ascidians (Phallusia nigra), and hydroids (Pennaria tiarella), collected at Tortugas. 377. Treadwell, A. L. 1911. Eunicidae of Tortugas. Bulletin of the American Museum of Natural History 30: 1-12. Systematic accounts of six species of polychaetous annelids are provided from specimens collected in the dead coral rock around Fort Jefferson during 1908. Some species are redescribed because of their earlier incomplete descriptions. Brief notes on their abundance and distribution are included. 378. Treadwell, A. L. 1921. Leodicidae of the West Indian region. Papers Tortugas Laboratory 15: 1- LS 93 Carnegie Institution of Washington Publication Number 293. A systematic study based on specimens of the family Leodicidae is presented. Collections were made at the Dry Tortugas and Key West region of Florida, and in Bermuda, Porto Rico, Montego Bay, Jamaica, and Tobago. Collecting was done along shore or in comparatively shallow water. The Leodicidae are a well-defined family in which the most constant structures are internal rather than external. There is always a well-developed jaw apparatus, composed of bilaterally arranged series of chitinous plates developed in a pharyngeal pouch, and capable of protrusion for feeding purposes through the mouth. The structure of these jaws was used by Ehlers as a basis for classification, though the external organs are a more convenient means of recognition. Se) . 1917. Polychaetous annelids from Florida, Porto Rico, Bermuda, and the Bahamas. Papers Tortugas Laboratory 11: 255-68. Carnegie Institution of Washington Publication Number 251. This paper is a preliminary description of some new species belonging to the Polychaetous annelids, as well as new species of other families which have been collected incidentally in this work, including a new sabellid belonging to the collection of the American Museum of Natural History. 380. Ubelacker, J. M. 1982. Review of some little-known species of syllids (Annelida: Polychaeta) described from the Gulf of Mexico and Caribbean by Hermann Augener in 1924. Proceedings of the Biological Society of Washington 95, no. 3: 583-93. The types of six little-known syllid species described by Augener in 1924 from the Dry Tortugas, Florida, and from St. Thomas and St. Croix in the West Indies, were reexamined. Haplosyllides floridana is a sexual form herein synonymized with it. Eusyllis antillensis and Syllis (Typosyllis) tigrinoides are synonyms; the latter name is retained. Syllis (Typosyllis) fuscosuturata has previously been synonymized with Branchiosyllis exilis corallicoloides and remains a valid species. 381. Vaughan, T. W. 1914. The building of the Marquesas and Tortugas Atolls and a sketch of the geologic history of the Florida Reef tract. Papers Tortugas Laboratory 5: 55-67. Carnegie Institution of Washington Publication Number 182. The study of the geology and the geologic processes of the Florida reef tract, and especially of the Tortugas and the Marquesas, has been continued since 1910. It is now possible to outline the salient geologic episodes in the history of the entire Florida Reef tract and to Institution comparisons with other coral-reef areas. 382. ———.. 1910. A contribution to the geologic history of the Florida Plateau. Papers Tortugas Laboratory 4: 99-185. Carnegie Institution of Washington Publication Number 133. This paper is the outgrowth of the author's association with two organizations, the United States Geological Survey and the Carnegie Institution of Washington. The author has visited all the principal keys between Miami and Key West, to collect and study bottom samples, particularly the deposits accumulating behind the keys, and to examine several important living coral reefs around the Tortugas. The scope of the paper was enlarged to trace the geologic history of the Floridian Plateau from Oligocene to Recent time. This is to be regarded as only a sketch of the geologic development of the Floridian Plateau, as many problems need solution and many phases of its history need further investigation. Perhaps its principal value may be in directing attention to some of the unsolved problems. It is necessary to know more accurately the amount of water discharged by the streams and the quantities of solids borne by them to the sea. The chemical processes of precipitation have not been sufficiently studied. There is also great need for more extensive studies of the marine bottom deposits within the 100-fathom curve. The deep wells recently put down on 94 383. 384. S85: Key Vaca, Big Pine Key, and Key West have given valuable data, but deep wells are also needed on the Marquesas, and the Tortugas, in order to discover what underlies the surface formations. It is hoped this paper may serve as a convenient summary of the present knowledge of the geologic history of this interesting region, perhaps present an interpretation somewhat different from those preceding, and be a stimulus to further investigation. . 1915. Coral-reefs and reef corals of the southeastern United States; their geologic history and significance. Bulletin of the Geological Society of America 26: 58-60. The geologic history of the extensive coral reefs of the southeastern United States and near-by West Indian Islands was outlined, and the bearing they have on the theory of coral reef formation was indicated. The author stated his conclusions regarding the Florida coral reefs as follows: (1) Corals have played a subordinate part, usually a negligible part in the building of the Floridean plateau; (2) every conspicuous development of coral reefs or reef corals took place during subsidence; (3) in every instance the coral reefs or reef corals have developed on platform basements which owe their origin to geologic agencies other than those dependent on the presence of corals. The conclusions in this report are summarized as follows: Critical investigations of corals as constructional geologic agents are bringing increasing proof that they are not as important as was believed. All known modern offshore reefs which have been investigated grow on platforms which have been submerged in Recent geologic times. No evidence has been presented to show that any barrier reef began to form as a fringing reef and was converted into a barrier by subsidence. There were platforms in early Teritiary time on the site of many of the present-day platforms, and evidence has yet been adduced to prove long-continued, uninterrupted subsidence in any coral-reef area. The width of a submerged platform bordering a land area is indicative of the stage attained by planation movement. The importance of coral reef studies to geology suggests they are only a conspicuous incident in time. . 1936. Current measurements along the Florida coral reef tract. Papers Tortugas Laboratory 29: 129-41. Carnegie Institution of Washington Publication Number 451. Note: This is Appendix 2 to Calcareous shallow water marine deposits of Florida and the Bahamas by Eldon Marion Thorp. During June and July 1914, while studying the phenomena associated with the Florida Coral-Reef Tract the author initiated a series of current measurements by using Ekman current meters. South of the Tortugas a non-tidal current toward the west is clearly indicated. The data here presented are inadequate for positive conclusions regarding the Counter Current. . 1915. The geologic significance of the growth rate of the Floridian and Bahamian shoal- water corals. Journal of the Washington Academy of Science 5, no. 17: 591-600. The object of this investigation has been to aid in understanding the amount of work stony corals may do as constructional geologic agents, and especially in the formation of coral reefs. This subject needs to be studied from at least five different view points , e.g.: (1) the quantity of material contributed by corals and that contributed by other agents must be estimated and the respective proportions determined; (2) in coral reef areas the ratio of the area covered by corals to that not covered by corals should be estimated; (3) the relations of coral reefs, continuity and discontinuity must be determined; (4) marine bottom deposits must be analyzed according to the source of the material, and the percentage of the calcium carbonate contributed by the differing agents estimated; (5) the rate of growth of corals needs to be known. There is no single formula for the growth rate of corals, as it varies by species and ecologic conditions. Observations/experiments on the growth rates of Tortugas corals are as follows: (1) Colonies obtained from the planule whose history was known, 386. 387. 388. 95 and were planted off the moat wall and on the NW side of Loggerhead Key; (2) Colonies cemented to tiles and planted at the same sites as above; Colonies naturally attached at sites described above. The reef species of greatest concern and importance is Orbicella (Montastrea) annularis followed by in importance, Maeandra strigosa, M. labyrinthiformes, and Siderastrea siderea. The upward growth is critical of the massive heads Orbicella (Montastrea) annularis which form the strong framework of the reef and averages | foot in 43.54 years or 7 mm /year and which might form a reef 150 feet thick in between 6500 and 7600 years. A table on the average annual growth rates of corals from the Florida region is provided. . 1914. The platforms of barrier coral reefs. Bulletin of the American Geographical Society 46: 426-29. The author states that there are three kinds of coral reefs: fringing or shore reefs which occur along the strand line, barrier reefs which occur at varied distances off shore and have lagoons from one to as much as forty fathoms depth between them and the strand line, and atolls, which are ring-like and enclose lagoons. As the relations of barrier reefs and atolls to the platforms on which they stand constitute the essential part of the theory of development of Recent reefs, the discussion of coral reef theory has been waged over the interpretation of these relations. The object of this paper is to point out the relations of barrier coral reefs to the last dominant change in position of the strand line and to indicate the organisms forming Recent barrier reefs have played in building the reef platforms. . 1914. Preliminary remarks on the geology of the Bahamas, with special reference to the origin of the Bahamian and Floridian Oolites. Papers Tortugas Laboratory 5: 47-54. Carnegie Institution of Washington Publication Number 182. The author presents a preliminary summary of the information compiled (to 1914), on the origin of calcium carbonate sediments in south Florida, and the Bahamas, using various hypotheses developed by the leading geologists of the time. These geologists included Alexander and Louis Agassiz, Sanford, and Drew who worked in the Dry Tortugas and believed the precipitation of calcium carbonate was due to the effects of denitrifying bacteria. . 1916. The results of investigations of the ecology of the Floridian and Bahamian shoalwater corals. Proceedings of the National Academy of Sciences 2: 95-100. This paper presents a summary of the knowledge on the ecology of shallow water corals in the Florida-Bahamian region, with a detailed description of new information on food preferences of corals, and salinity and water temperature tolerances, based on studies conducted at the Dry Tortugas and the upper Florida Keys area. Mayer, at the Tortugas Laboratory, found that temperatures of 13.9 °C would exterminate the principal Floridean corals; similar results were found for corals around Australia. Light experiments at Fort Jefferson suggested vigorous coral growth in well lit wharf areas, and little growth in piling areas of perpetual shading. Tests conducted at the Marine Laboratory suggested that corals could survive at salinities of 27-38 ppt, but not as low as 19 ppt. Other conditions necessary for vigorous coral growth are maximum water depths of 45 meters and rocky or firm bottoms, without silty deposits. The growth rate of corals was determined by planting planulae in the laboratory, by measuring colonies which had been cemented to disks, and fixed on heads of stakes driven into the sea bottom. Measurements of colonies naturally attached were also made. Plantings at the Tortugas were made at Loggerhead Key, and around Fort Jefferson. The more massive the coral, the slower the growth; while the more ramose (Acropora palmata) and the more porous the skeleton, the more rapid the growth. The growth rate of the principal reef builders (massive corals) in the Florida region, Orbicella (Montastrea) annularis is from 5-7 mm per year and would form a reef of 150 96 389. 390. 391. feet in thickness from 7620 to 6531 years. A. palmata may build a similar thickness in 1800 years. . 1914. Sketch of the geologic history of the Florida coral reef tract and comparisons with other coral reef areas. Journal of the Washington Academy of Sciences 4: 26-34. The author presents the two hypotheses for the formation of atolls: one attributes atolls to the submarine solution of the interior of a mass of limestone; the other accounts for them by constructional agencies. The author believes the solution theory is disproved by a chemical examination of sea-water from a Tortugas lagoon. He believes the Marquesas and the Tortugas are constructional phenomena and owe their configuration to the prevailing winds and currents. . 1918. The temperature of the Florida Coral-Reef Tract. Papers Tortugas Laboratory 9: 319-39. Carnegie Institution of Washington Publication Number 213. The temperature data presented were assembled primarily for their bearing on the effect temperature exerts on the bathymetric and geographic distribution of coral reefs. Temperature is also one of the most important factors in determining the geographic distribution of sea-level and near sea-level reefs. Vaughan, T. W., M. A. Goldman, J. A. Cushman, M. A. Howe and others. 1918. Some shoal-water bottom samples from Murray Island, Australia, and comparisons of them with samples from Florida and the Bahamas. Papers Tortugas Laboratory 9: 235-97. Carnegie Institution of Washington Publication Number 213. The present paper is a preliminary contribution to the study of the marine bottom deposits in three coral-reef areas: Murray Island, Australia; the Bahamas, and southern Florida. Mechanical analyses have been made of all samples except those obtained in 1915, and the results of the chemical analyses of a selected set are presented. An attempt has been made to outline a method of studying calcium carbonate bottom deposits, in the hope that progress may be made toward an adequate classification of such sediments. The Tortugas Lagoon samples are coarser than those in Marquesas Lagoon, and those from the latter locality are coarser than the Bahama sample from South Bight and the west side of Andros Island. Some terrigenous material, mostly quartz sand, is washed into Biscayne Bay, Florida, and into the sounds south of it, but otherwise practically none reaches the key and reef region. The Florida area is therefore a perfect example of limestones forming in shoal water near a land area which is not crossed by large streams. The Fe,0; content of the Florida samples seems somewhat higher, up to about 0.37 per cent, than that of the Bahama samples. Reconsideration of the evidence bearing upon the precipitation of CaCo3 in tropical and subtropical waters and the possibility of its re-solution by ocean-water leads to the conclusion that precipitation is resulting from both organic and inorganic agencies, and that no appreciable re-solution is taking place in the water. 392. Visscher, J. P. 1930-1931. Distribution of barnacles with special reference to behavior of larvae. Carnegie Institution of Washington, Year Book Note: published as follows: 1930, v. 29, p. 346; 1931, v. 30, p. 397. A study of the barnacles was made, more than twenty species being found. Several species appear to be new to science. Behavior of larvae appeared to vary depending on habitat, as certain barnacles were found on crabs, others above the tide on pilings, still others only on coral and on the spiny lobster, Panulirus argus. 393. Vukovich, F. M. 1988. On the formation of elongated cold perturbations off the Dry Tortugas. Journal of Physical Oceanography 18, no. 7: 1051-59. The life cycle of a cold perturbation on the boundary of the Loop Current in the Gulf of : 97 Mexico was studied over the period of 18 March to 22 May 1984, approximately a 60-day period. The study focused on the behavior of the surface and subsurface area of the cold perturbation as it moved along the boundary of the Loop Current. The area of the perturbation was defined by an alongflow-scale length, which is the scale length parallel to the unperturbed flow of the Loop Current, and the crossflow-scale length, which is the scale length perpendicular to the unperturbed flow of the Loop Current. 394. Vukovich, F. M. and G. A. Maul. 1985. Cyclonic eddies in the eastern Gulf of Mexico. Journal of Physical Oceanography 15, no. 1: 105-17. Cold-domed cyclonic eddies juxtaposed to the cylconic shear side of the Gulf Loop Current are observed in simultaneously obtained hydrographic, current meter mooring, and satellite data as cold perturbations on the northern extreme of the current and grow either into a cold tongue or a quasi-stable meander off the Dry Tortugas, Florida. Areal shipboard surveys show closed isopleths of temperature and salinity, and surface geostrophic current speeds relative to 1000 db are in excess of 100 cms super(-1). The diameter of the cold domes varied from 80 to 120 km. 395. Wallace, W. S. 1908. A collection of hydroids made at the Tortugas during 1908. Carnegie Institution of Washington, Year Book 7: 136-37. At least fifty species of hydroids were collected. A tentative list of those identified is provided. 396. Wartman, W. 1929. Studies on Echinometra. Carnegie Institution of Washington, Year Book 28: Die (No data report provided). 397. Watson, J. B. 1908. The behavior of noddy and sooty terns. Papers Tortugas Laboratory 2: 187- Daas Carnegie Institution of Washington Publication Number 103. The work presented in this report is preliminary in nature. Following a general description of the two species, a geographical situation and present history of the tern colony at the Tortugas is given. All observations were recorded during the nesting season on Bird Key, a small coral island covered in part by bay cedar, mixed with cactus in the central western parts of the island, with little vegetation elsewhere. Observations on their foods and feeding habits indicated that the birds usually feed in groups, never swim nor dive, but skim along the surface picking up small fish being attacked by larger fish. Feeding distance from the Key was estimated between 4 and 10 knots. Mating has been suggested prior to arriving at the Tortugas, although some indications of sexual activity occurred for the noddies, but not the sooties. Noddies nest in vegetation, while sooties build nests in sand. Usually one, sometimes two eggs are laid, with a period of incubation for the noddy from 32 to 35 days. The parents alternately feed the young at intervals from 1 to 4 hours; their general conduct does not greatly change at the arrival time of the young. However, two days after the arrival, the parents are more ferocious; both species return to normalcy as the chicks gain strength. The birds become exhausted caring for their young, and collect upon the beach for "sunning". Egg coloration tests indicated that neither species recognized its own egg. As for the noddy's nest environment, it could be disturbed without affecting the bird, as long as the egg position was not changed. Tests conducted using Porter's learning maze indicated that noddy's were slower than sooties, because of their longer standing time. Further maze tests using darkness and maze rotation were inconclusive. Other in captivity tests showed that the sooty is highly excitable and nervous, whereas the noddy is stolid and indifferent. 98 398. Watson, J. B. and K. S. Lashley. 1915. Homing and related activities of birds. Papers Tortugas Laboratory 7: 1-104. Carnegie Institution of Washington Publication Number 211. The present series of studies on the behavior of birds is a direct outgrowth of an investigation made on the noddy and sooty terns nesting on Bird Key, Tortugas, Florida. The homing "instinct" is the central topic in all the papers. In the 1907 investigation the fact appeared that terns possess a homing sense, behaving exactly as do homing pigeons when sent away from their nests and young. The 1907 investigation already referred to is concerned largely with instincts in terns-those of feeding, nesting, brooding, etc. In general the problems of proximate orientation are relatively simple and straightforward. On the island of Bird Key the terns make their adjustment to the nest, mate, young, etc., on the basis largely of visual habits. There is no evidence of any remarkable or unusual sensitivity, nor of the functioning of any hypothetical sense-organ. The present paper seems to call for a separation between proximate orientation and distant orientation. Mathematical considerations show that at such distances the goal can not possibly (directly) visually stimulate the bird, even granting absolute visual acuity and complete absence of haze, etc. This work has shown further, in the terns at least, that there is no special Spiirsinn-special tactual or olfactory mechanism situated in the nasal cavity which may function in homing. The task of explaining distant orientation is an experimental one, which must yield positive results as soon as proper methods are at hand. Two lines of investigation offer hopeful results: the rearing of homing pigeons in a cote, or the rearing of the birds in a wire-covered yard attached to a cote. We could tether individual birds to the top of the cote by cords which would permit a view only of the neighborhood immediately surrounding the cote. With these experiments upon homing, work upon the sensory equipment of the homing pigeon should be carried on. It is just possible that these animals possess on certain parts of the body, tactual and thermal mechanisms which may assist them in reacting to slight differences in pressure, temperature, and humidity of air columns. The experiments and conclusions on homing proper can be found on pages 59 and 60. These results, which not settling the question of the sensory mechanism by means of which the birds return to the nests, do remove all doubts about the fact that the noddy and sooty terns can return from distances up to 1,000 miles in the absence of all landmarks. The problem of homing has thus become defined, and experimental work of a definite kind is needed for its solution . 399. Wells, R. C. 1922. Carbon-dioxide content of sea water at Tortugas. Papers Tortugas Laboratory 18: 87-93. Carnegie Institution of Washington Publication Number 312. It is generally considered that the carbon-dioxide content of sea-water may be increased by accessions from the air, by animal life, by the decay of organic matter in the sediments on the bottom or elsewhere, by the solution of carbonate rocks, by the contributions of rivers, and by gas vents beneath the sea. Sea-water may lose carbon dioxide to the air, to plants, and in the formation of carbonate rocks and the carbonaceous parts of organisms. Mere evaporation and precipitation also alter the carbon-dioxide concentration somewhat if other conditions remain unchanged. The writer made determinations on sea-water from Tortugas, Florida, in June 1919 taken directly from the sea at various points about Loggerhead Key, which reveal unmistakable diurnal variations. The water has sufficient contact with plants and sea-weeds to show the effect of photosynthesis on its CO, content. There is a loss of CO, by day and a gain by night. Plant life appears to be the chief agency in causing a daily variation in the CO, content. Determinations of CO, should probably be made soon after the time the samples are collected, on account of the possibility of the decay of organic matter, such as algae, in preserved samples. The average "excess base" found at Tortugas corresponds to a normality of 0.00239. This titration includes everything that consumes acid; it represents chiefly bicarbonate, about 0.00183, some 99 carbonate, about 0.00041, and other substances that contribute to the alkalinity, about 0.00015. The methods used in arriving at these figures were provided along with a record of determinations made at Tortugas and the relation between the carbon-dioxide content of the water and time of day. 400. Westinga, E. and P. C. Hoetjes. 1981. The intrasponge fauna of Spheciospongia vesparia (Porifera, Demospongiae) at Curacao and Bonaire. Marine Biology 62, no. (2-3): 139-50. The infauna of 35 individuals of Spheciospongia vesparia (Lamarck, 1814) of different volumes and from different sites and depths have been inventoried and compared. The number of sponge-inhabiting taxa is logarithmically related to sponge volume. Biomass and total number of the animals contained in the sponge are directly proportional to sponge volume. Numerical and taxonomic composition of infaunas from different sampling sites is fairly constant. Biomass and total number of sponge inhabiting animals is not significantly different for any of the four sampling sites. Several taxa, however, are more abundant in sponges from one or more localities. The ratio of total biomass to total number of intrasponge fauna is found to be significantly smaller for sponges collected in deep water than in shallow water. Differences from and similarities with Pearse's results (1932,1950) on the infauna of the same sponge species at Dry Tortugas and Bimini are discussed. The relation of the number of contained taxa and the volume of a sponge is compared with the relation of island size and number of taxa present according to MacArthur and Wilson's island theory (MacArthur, 1972). Finally the erratic occurrence of some taxa as opposed to the highly regular occurrence of some other taxa is discussed. It is concluded that the composition of the sponge-infauna in specimens larger than 11 is highly constant and that the sponge-inhabiting fauna constitutes an ecological community. 401. Westrum, B. L. and P. A. Meyers. 1978. Organic carbon content of seawater from over three Caribbean reefs. Bulletin of Marine Science 28, no. 1: 153-58. Seawater samples from transects crossing three Caribbean coral reefs, including the Dry Tortugas, showed variations in concentrations of organic carbon. Total organic carbon increased substantially over two fringing reef crests. Most of this increase occurred in the particulate fraction at the seaward edge of the crest but in the dissolved fraction at the landward edge. Back reef levels of total organic carbon were lower than those seaward of the reef. These observations support the hypothesis that organic carbon can be physically removed from the benthos at the turbulent reef crest and be subsequently utilized in backreef areas. The reef-flat formation studied in January 1975 in the Dry Tortugas was situated off the western shore of Loggerhead Key. This study indicates that organic matter contributed at the crest is available as a resource to only a limited portion of the backreef community - that part located directly behind the crest. The observed decrease in TOC levels implies quick biological utilization or loss through physical processes. Thus, despite continual input, no net accumulation of organic matter occurs in the backreef area, and this region can be described as being relatively depleted in organic carbon. If large coral formations are present, as at the Dry Tortugas location, they can contribute organic matter to the surrounding seawater. 402. Wheaton, J. 1980. Ecology of gorgonians (Octocorallia: Gorgonacea) at Dry Tortugas, Florida . Florida Scientist, 43 (suppl. 1), 20. This study reports the species composition and distribution of the gorgonian fauna of Long Key Reef, Dry Tortugas during the summers of 1975-1976. 23 species were recorded. Additional samples increased the number of species to 35. Most shallow reef gorgonians were Plexaura. 403. Wheaton, J.L., W.C. Jaap, B. L. Kojis, G. P. Schmahl, D. L. Ballantine and J. E. McKenna Jr. 1993. Transplanting organisms on a damaged reef at Pulaski Shoal, Ft. Jefferson National 100 Monument, Dry Tortugas, Florida, USA: An experiment to enhance recruitment. (abs.). Proceedings of the Seventh International Coral Reef Symposium, p. 639. Mangilao, Guam: University of Guam. Grounding of the 475-ft. freighter, Mavro Vetranic, at Pulaski Shoal Reef, Dry Tortugas, on 30 October 1989, damaged 3,465 m’ of reef surface. After one year, minimal recruitment of macrobenthos, principally the alga Dictyota, had occurred. An experiment was designed to test effects of adding relief and transplanting sponges, octocorals, and scleractinian corals on recruitment of biota to the damaged area. In Sept. 1991, one control and two experimental sites, each 9-m’, were selected, marked, mapped, and photographed. Large reef rocks were placed in one experimental plot to provide relief and refuge. More than 185 organisms (73 species of algae, Porifera, and Cnidaria) were transplanted and cemented into five of the nine square-meter subunits in the other experimental plot. Transplanting was labor intensive, requiring 64 man-hours to collect, move, and cement organisms. We then rephotographed and mapped the plots. Sites will be monitored to determine if recruitment of macrobenthic organisms is enhanced . 404. Whitaker, D. 1926. Organization of echinoderm egg, and a measurable potential difference between the cell interior and outside medium. Carnegie Institution of Washington, Year Book 25: 248-55. Egg development, investigated in the sea-urchin, Lytechinus, suggested that the differentiation of ectoderm begins before fertilization by the exclusion of the endoderm- forming substances from the superficial layers of the egg. Micromere-forming substances do not differentiate before fertilization. 405. White, S.C., W. B. Robertson Jr. and R. E. Ricklefs. 1976. The effect of Hurricane Agnes on growth and survival of tern chicks in Florida. Bird-Banding 47, no. 1: 54-71. In June, 1972 Ricklefs and White were studying the energetics of nestling growth in sooty terns (Sterna fuscata) at the Dry Tortugas, when Hurricane Agnes passed west of the area. High winds, heavy rain, rough seas, and low temperatures prevailed for more than a week. Robertson worked in the colony from 28 June, about a week after the storm subsided, to 6 July. It is reported here the effects of Hurricane Agnes on the growth and survival of young sooty terns and brown noddies (Anous stolidus ). 406. Wichterman, R. 1942. Cytological studies on the structure and division of three new ciliates from the littoral earthworm of Tortugas. Papers Tortugas Laboratory 33: 83-103 . Carnegie Institution of Washington Publication Number 524. During the summer of 1939 this study of Protozoa inhabiting the intestine of the littoral earthworm Pontodrellus bermudensis Beddard, was begun. The study revealed three previously undescribed ciliates: Hysterocinita pontodrila, n. s.p.; Anoplophyra macroneucleata, n. sp.; and Maupasella leptas, n. sp. This paper describes the ciliates and gives an account of fission in each species. Of the 230 worms examined, 64% were infected with the ciliates. Generally a worm was parasitized with two different species. Observations on the length of life of the ciliates in seawater were recorded. Encystment was not encountered. The presence of stages in the life history of acephaline gregarines and nematodes was noted. 407. . 1942. A new ciliate from a coral of Tortugas and its symbiotic zooxanthellae. Papers Tortugas Laboratory 33: 105-11. Carnegie Institution of Washington Publication Number 524. A new ciliate was found on the coral Eunicia crassa E. and H. and is described as Paraeuplotes tortugenesis, n. gen. and n. sp., and is placed in the family Paraeuplotidae, n. fam. The coral it was found on is a member of the Alcyonaria fauna, and is commonly found in the Caribbean, and is abundant on the reefs of the Tortugas. The morphology of 101 the ciliate is discussed, as well as the presence of zoozanthellae. The question 1s posed "what is the nature of the symbiosis between the protozoan and the zooanthellae it contains?". 408. Williams, O. L. 1932. Studies on the nematodes of Tortugas fishes. Carnegie Institution of Washington, Year Book 31: 291-92. Observations of more than 800 fishes representing about 175 species during the summer of 1932 demonstrate that the incidence of infestation with nematodes is lower in fishes of the Tortugas than in the cooler, shallow waters found farther north. 409. Willier, B. H. 1936. A study of the early embryology of the Loggerhead sea turtle and of sharks. Carnegie Institution of Washington, Year Book 35: 92. The embryological development of the Loggerhead turtle (Caretta caretta) was examined from the time of egg laying to within a few days of hatching. Significant observations are presented. 410. Wilson, C. B. 1936. Parasitic copepods from the Dry Tortugas. Papers Tortugas Laboratory 29: 327-47 (issued Dec. 1935). Carnegie Institution of Washington Publication Number 452. Two collections of parasitic copepods contained in the present paper were made at the Marine Laboratory of the Carnegie Institution in the Dry Tortugas, involving the handling of a large number of the local fishes. In addition to the specific objects of investigation it was soon noted that the fish were more or less infested with parasitic copepods and isopods. Upon identification, seven of the species are new to science, and two others have been made the types of new genera. The other species have been obtained before either in the waters around the Dry Tortugas, the Bahamas, the Bermudas or the West Indies. 411. Winegarner, C. E., W. B. Robertson and W. Hoffman. 1984. Anolis sageri sageri (brown anole) USA: Florida: Monroe Co: Dry Tortugas, Garden Key. Herpetological Review 15, no. 3: 77-78. Three males and one female specimen were taken on a large pile of bricks and rubble just east of the moat surrounding Ft. Jefferson, April 8-10 1983. Population currently seems limited to this small portion of the island, so introduction may have been very recent. A construction barge moored adjacent to the collection site from October 1981 to June 1982 possibly was a source of colonizing individuals. However the regular arrival of Park Service boats and private vessels are other possibilities. 412. Wolfe, C. A. 1989. "Growth of the Brown Noddy (Anous stolidus) in the Dry Tortugas (Florida)." Master of Science, Florida Atlantic University, Boca Raton. The author discusses the slow growth rate of the brown noddy nestlings in the Dry Tortugas as to what would be predicted based on adult body size and mode of development. This prolonged growth pattern is typical of tropical pelagic seabirds. An intraspecific comparison of growth rates among several populations of brown noddies , indicates that growth of body mass of the Tortugas noddies is significantly faster, the development period shorter, and the asymptotic size smaller than in Pacific populations. However, there were no differences among the populations in the rates of wing or culmen growth. The Bush Key nestlings appear to receive a higher quality diet that contains proportionally more fish, while Pacific nestlings receive substantial amounts of squid. The Pacific nestlings also seem to be subjected to a thermally more stressful microclimate, which may necessitate the allocation of proportionally more of their total energy to thermoregulation and less to growth. 102 413. 414. 415. Woolfenden, G. E. and W. B. Robertson Jr. 1991. A banded red knot seen at the Dry Tortugas. Florida Field Naturalist 19, no. 4: 106-7. The red knot (Calidris canutus) is a locally abundant and winter visitor on both coasts of Florida, however it is rare at the Dry Tortugas, with only three sightings prior to a sighting made by the authors during May-June 1988. It was suggested that all sightings at the Tortugas represented birds of a knot population that winters along the Atlantic coast of Patagonia in South America. . 1975. Least terns nest at the Dry Tortugas. Florida Field Naturalist 2, no. 1: 19-20. On July 1, 1973, as members of the tern-banding party landed on Middle Key, Dry Tortugas, they saw 4 adult-plumaged Least terns and one fledged juvenile in the company of 8 adult-plumaged Roseate Terns, Sterna dougallii. Search of the island, a barren sand bank with only a small area above high tide, revealed 2 Least Tern nests, one with 2 eggs and the other with one egg, and 4 Roseate nests, each with 2 eggs. It is of interest that Least Terns have again attempted to nest at Dry Tortugas after an absence of almost 25 years. Woolfenden, G. E., S.C. White, R. L. Mumme and W. B. Robertson Jr. 1976. Aggression among starving cattle egrets. Bird-Banding 47, no. 1: 48-53. Cattle egrets (Bubulcus ibis) flying over the Gulf of Mexico often land at the Dry Tortugas. Food suitable for cattle egrets is scarce locally and many egrets die at the Dry Tortugas, presumably from starvation. In June 1975 an infestation of a sea grape tree by caterpillars of the moth Sarasota Plumigerella Hulst. provided a natural, albeit limited, food supply at which we observed cattle egret behavior. From several observers, we were able to compare aggression, feeding frequency, plumage condition and death weights of the starving birds. Of special interest was the opportunity to test relationships between aggressiveness and feeding frequency under the unusual circumstances of starving birds competing for a concentrated, but limited food supply. 416. Woolfenden, G. E and W. B. Robertson Jr. 1975. First nesting of the house sparrow at Dry Tortugas. Florida Field Naturalist 3: 23-24. This paper describes the first occurrence of the House sparrow at the Dry Tortugas. During mid-June 1974, nest building and copulation was observed in a coconut palm east of the moat bridge on Garden Key. Four eggs were found later, but no further inspections were made. House sparrows that reach the Tortugas are considered true migrants or birds from the West Indies that accompanied north-bound migrants of other species. 417. Yamanouchi, S. 1929-1935. Life histories and cytology of marine algae. Carnegie Institution of Washington, Year Book . Note: published as follows: 1929, v. 28, p. 297; 1930, v. 29, p. 346; 1931, v. 30, p. 371; 1932, v. 31, p. 259; 1933, v. 32, p. 265; 1934, v. 33, p. 263; 1935, v. 34, p. 75. Local populations of marine algae, Phaeophycae and Chlorophyceae were studied with emphasis on Caulerpa. Reproductive phases of many specimens were collected for later morphological and cytological study. 418. Yonge, C. M. 1936. Studies on the biology of Tortugas corals. I. Observations on Maenadra areolata Linn. Papers Tortugas Laboratory 29: 185-98 (issued Dec. 1935). Carnegie Institution of Washington Publication Number 452. Maenadra areolata is one of the commonest corals of the Caribbean and Florida reefs. It is a highly specialized species adapted for life in a restricted environment. It thrives best on the flats behind the reefs. It has no firm basal attachment, it can not resist the impact of the waves of rough seas. At the Tortugas it is very common in sheltered areas on the inner side of the circle of reefs where wide stretches of sand occur. The best collecting ground is the lee of Bird Key Reef. Feeding is entirely by means of the tentacles, there is no reversal 103 of ciliary currents. Not only is sediment removed very rapidly from the surface, but colonies can completely uncover themselves within twelve hours after being buried in the sad. Unlike Fungia which uncovers itself by the exclusive action of cilia, M. areolata first distends the tissues with water. Distension for cleansing is essentially different from expansion for feeding. Planulation, so far as can be determined at present, has a lunar rhythm, culminating about the time of new moon. After an initial stage when upward and outward growth are about equal, outward growth predominates, an oval or rounded colony with a small basal attachment being finally produced. The stage at which detachment occurs must vary with environmental conditions. Colonies may be formed from a single planula or from the fusion of several. M. areolata is a species highly adapted for life on sand occupying in the Atlantic, the same habitat occupied by the Fungiidae in the Indo- Pacific. Adaptability in reef-building corals is discussed. 419. . 1936. Studies on the biology of Tortugas corals. IJ. Variation in the genus Siderastrea. Papers Tortugas Laboratory 29: 199-208 (issued Dec. 1935). Carnegie Institution of Washington Publication Number 452. This paper presents data on stony corals on the Tortugas reefs. The genus Siderastrea is represented at the Tortugas by the two species, S. siderea and S. radians. S. siderea forms larger rounded colonies which were not observed above the level of low-water springs. S. radians is essentially a shore-living species possessing the physiological adaptations characteristic of all shore-living animals. S. radians is capable of great modification both in the form of the skeleton as a whole and also in the size and shape of the corallites and in the number, slope and thickness of the septa. This species has been enabled, as a result, to occupy a variety of habitats, the extremes being represented by the surf region on the beach rock on the one hand, and by the still, sediment-laden water in the moat at Fort Jefferson on the other. The relation between form and environment in corals is discussed and the general conclusion reached that the great success of the Madreporaria is probably due to the presence of species highly specialized for a particular environment and also of others which can be modified for existence in a variety of different environments . 420. . 1937. Studies on the biology of Tortugas Corals. III. The effect of mucus on oxygen consumption. Papers Tortugas Laboratory 31: 207-14 (issued Oct. 1937). Carnegie Institution of Washington Publication Number 475. Experiments are described which indicate that a large proportion of the apparent utilization of oxygen by corals is actually due to oxidation of mucus secreted by them during the course of the experiment. The amount of mucus varies greatly in different genera and may also be increased at certain times, e.g. during planulation. In view of this source of error it is impossible to accept their face value figures which claim to represent either the absolute or the comparative rates of respiration in different corals, or general conclusions which are based on these figures. Oxidation of mucus may be expected to affect the apparent rate of respiration in all aquatic animals which normally secrete mucus. 421. Yonge, C. M. and H. M. Nicholas. 1940. Structure and function of the gut and symbiosis with zooxanthellae in Tridachia crispata (Oerst.) Bgh. Papers Tortugas Laboratory 32: 287- 301 (issued Sept. 1940). Carnegie Institution of Washington Publication Number 517. During the visit of the senior author to the Tortugas Laboratory in the season of 1934, a number of specimens of a very interesting and beautiful species of opisthobranch mollusk were collected a low tide on the surface of the reefs. Examination revealed the invariable presence of brown unicellular algae, or zooxanthellae, within their tissues. Tridachia crispata is an elysoid opisthobranch with the body extremely flattened dorsoventrally and extended into undulating body folds laterally and terminally. It occurs under stones on the reefs of the Torugas group and elsewhere in the West Indies. The feeding and digestive 104 systems are described. These have the typical elysoid structure with modifications, notably in the digestive diverticula, correlated with the excessive flattening of the body. Like the other members of the Elysiidae, T. crispata is a highly specialized herbivore. Zooxanthellae are habitually present in a restricted zone a short distance from the margin of the body fold. They occur freely within the connective tissue and increase by division. There is no evidence that the animal normally consumes them, but reasons are given for the suggestion that they may be of value to the animal by removing waste products of metabolism produced within the body fold . 422. Zeleny, C. 1907. The effect of degree of injury, successive injury, and functional activity upon regeneration in the scyphomedusan Cassiopea xamachana. Journal of Experimental Zoology 5: 265-74. This study is part of a series of experiments at the Dry Tortugas Marine Lab on the internal factors controlling regeneration in Cassiopea and other forms, including the degree of injury and successive removal of a part and rhythmical pulsations of the disk. The removal of 6 to 8 arms constitutes the most favorable degree of injury for the regeneration of each arm. When comparing the rate of regeneration of disks, where the disk was made to pulsate rhythmically with cases without pulsation, there is no advantage in favor of the pulsating ones, but rather a retardation. Other tests of successive injury upon regeneration were made on chelae of the gulf-weed crab, Portunus sayi, which reveal that the second regeneration is greater than the first. However, when the age factor is removed the two are exactly alike. 423. . 1908. Some internal factors concerned with the regeneration of the chelae of the gulf-weed crab (Portunus sayi). Papers Tortugas Laboratory 2: 103-38. Carnegie Institution of Washington Publication Number 103. The primary object of the experiments described was twofold: the quantitative determination (1) of the effect of successive removal of an organ upon its power to regenerate and (2) of the character of the changes, if any, produced in the uninjured parts of the animal by such removals. It was found that (1) individuals of Portunus sayi with a cephalo-thoracic length between 3-9 and 14.5 mm. show but a slight correlation between the length of the molting period and the size or age of the animal. (2) The amount of regeneration of the right chela between the same limits of size is likewise but slightly correlated with the length of the molting period, but is very closely correlated with the size of the animal. (3) The specific amount of regeneration of the right chela increases slightly with increase in size or age of the animal.(4) The specific length of the left chela in uninjured individuals increases slightly with increase in size or age of the animal.(5) The proportion between the amount of regeneration of a chela and the length of the chela in uninjured individuals of the same size is constant, uninfluenced by the size of the animal. (6) In single individuals the third regeneration is greater than the second and the second is greater than the first. (7) When the correction for change in the power of regeneration with size or age is made, it is found that successive removal neither retards nor accelerates the regeneration of the right chela. (8) The right chela is slightly larger that the left in a great majority of the individuals. (9) The removal and regeneration of the right chela produces no change in the growth of the uninjured left chela. 424. Zheng, W. and E. S. Van Fleet. 1988. Petroleum hydrocarbon contamination in the Dry Tortugas USA. Marine Pollution Bulletin 19, no. 3: 134-36. The present study extends a previous work westward to the point where Florida Keys island chain intersects the Gulf Loop Current. Since the Dry Tortugas are located in this unusual area, they provide an ideal location for examining the fate of petroleum discharged into the eastern Gulf of Mexico. Beach tar samples were collected along 1 m wide transects at 18 stations according to the procedures established by CARIPOL (1980). The 105 distribution of Dry Tortugas beach tar ranged from 0.6 g m super (-2) to 22.1 g m super (- 2) dry weight with an average of 9.2 plus or minus 7.8 g m super (-2). There appear to be no strong correlations between Dry Tortugas beach tar concentrations and either predominant wind direction or major Gulf Loop Current circulation patterns. 106 Agassiz, A., 1, 2 Andres, B., A., 3 Applegate, R. D., 298 Aska, D. Y., 327 Austin, O. L. Jr., 4 Awbrey, F. T., 23 Baden, D. G., 333, 375 Bailey, E., 5 Bailey, P. L., 6 Baker, B., 7 Ball, S.C., 8 Ballantine, D. L. 9, 403 Barnes, G. W., 72 Bartsch, P., 10, 11, 12, 13, 14 Battey, J. F., 287 Bellow, T. H., 15, 16, 324 Below, L. C., 315 Bennett, F. M., 17 Berrill, N. J., 18 Blinks, L. R., 19 Bohnsack, J. A., 20 Bortone, S. A., 21 Boschma, H., 22 Bowles, A. E., 23 Bowman, H. H. M., 24 Boyden, A., 25 Bradbury, R. C., 26 Braman, R. S., 154 Bramen, R. S., 356 Breder, C. M. Jr., 27, 28 Brinley, F. J., 29 Brooks, H. K., 30 Brooks, W. K., 31, 32 Brown, D.E.S., 33 Brown, W.Y,. 34 Bullington, W. E., 35 Burkenroad, M., 36 Caira, J. N., 37 Calder, D. R., 38 Carrier, J. C., 39, 40 Cary, L.R., 41- 48 Cate, C.N., 49 Ceurvels, A. R,. 105 Chambers, E. L., 50 Chiang, L., 288 Child, C. A., 51 Clapp, R. B., 52 Author Index Clark, H. L., 53 Clark, L. B., 54 Cochran, W. W., 298 Cole, L. J., 55 Collie, M. R., 56 Colman, J., 57 Conger, P.S., 58 Conklin, E. G., 59, 60 Coonfield, B. R., 61, 62 Coutiére, H., 63 Cowles, R. P., 64, 65 Criales, M.M., 66 Cubit, J. D., 196 Cushman, J. A., 67, 391 Cutright, P. E., 68 Dall, W. H., 69 Darby, H. H., 70-72 Davis, G. E., 73- 82 Davis, J.H., 325 Davis, J. H. Jr., 83, 84 Davis, R. A. Jr., 85 de Laubenfels, M. W., 86- 88 de Renyi, G. S., 89 Deflaun, M. F., 90 Dinsmore, J. J., 91, 92, 137 Dodrill J. W., 81, 82 Dole, R. B., 93 Domeier, M. L., 94 Donaldson, H. H., 95 Donan, P., 356 Donnelly, K. B., 166-168, 356 Doyle, M.M., 98 Doyle, W. L., 96-98 Drew, G. H., 99 Dustan, P., 100, 101,163 Edmondson, C..H., 102 Erseus, C., 103 Farfante, I. P., 104 Feinstein, A. A., 105 Feltham C. B., 111 Fenimore Johnson, E. R., 72 Ferguson, John C., 301 Field, R. M., 106, 107 Fisk, E. J., 108 Forcucci, D., 264 ECO Fosberg, F. R., 350 Gardiner, M. S., 369 Gault, C.C., 251 Gee, H., 110, 111 Gersh, I.. 112 Gilmore, R. G., 113 Ginsburg, R.N., 114 Given, B., 316 Goldfarb, A. J., 115-122 Goldman, M. A., 391 Goodrich, H. B., 123 Gordon, M., 124 Goy, J. W., 125 Grave, C., 126-128 Gudger, E. W., 129-133 Haight, F. J., 374 Halas, J.C., 101 Halley, R.B., 134, 328, 330 Hanlon, R. T., 135 Hargitt, C. W., 136 Harper, D. E., 20 Harrington, B. A., 137 Harris, J.E., 28, 138 Harrison, C.S., 354 Hartman, C.G., 139 Hartmeyer, R., 140, 141 Harvey, E.N., 142-145 Hatai, S., 146-148 Hayes, F.R., 149, 150 Heard, R. W., 151 Helwig, E.R., 152 Hendee, E. C., 153 Hendrix, S.A. 154 Hess, W.N., 54, 155, 156 Hildebrand, S. F., 211, 212 Hine, A. C., 203 Hixon, R. F., 135 Hoetjes, P.C., 400 Hoffman, W., 157, 411 Holmes, C. W., 158 Hooker, D., 159 Hopkins, D. L., 160 Howe, M.A., 391 Hudson, J. H., 310, 328, 330 Hutton, R.F., 105 Jaap, W.C., 101, 161-168, 264, 356, 403 Jacobs, M.H., 169 107 Jefferson, J. P., 170 Jehl, J. R., 23 Jennings, H.S., 171 Jindrich, V., 172 Jones, N., 173, 285 Jones, R. D., 174 Jones, R.S., 113,175 Jordan, D. S., 176, 177 Jordan, H. E., 178-186 Kaas, P., 187 Kale, H. W., 188 Keiller, V.H., 369 Kellner C., 32, 189 Kille, F. R., 190 Kilma, E. F., 261 Kindinger, J. L., 330 King, J. Jr., 276, 277 Kojis, B. J., 167, 168, 403 Kopac, M. J., 191 Kunkel, B. W., 192 Lashley, K.S., 398 Le Compte, M., 193 Lee, T.N., 66 Leitch, J.L., 194, 195 Lessios, H. A., 196 Lidz, B. H., 328, 330 Linton, E., 197-199 Lipman, C. B., 200- 202 Locker, S. D., 203 Longley, W. H., 204-212 Lucké, B., 213-215 Lynts, G. W., 216 Lyons, W.G., 163, 217 Mann, A., 218, 374 Manter, H. W., 219-221 Marsh, G., 222- 224 Martin, D. F., 356 Martin, J.C., 346 Martin, L. K., 40 Mason, C.R., 316, 318 Mast, S. O., 225 Matthai, G., 226 Maul, George A., 394 Mayer, A. G., 227-242 McClellan, D.B., 20 McClendon, J. F., 243-251 McKenna, J. E. Jr., 167, 168, 403 108 Meeden Jars) 252 Meier, O. W., 288 Meyers, P.A., 401 Miller, H. M. Jr., 253 Miller, R. A., 254 Milligan, M.R., 103 Millspaugh, C. F., 255 Mitchell-Tapping, H. J., 256 Moore, T., 170 Moritz, C. E., 257 Mulholland, S., 251 Multer, H. G., 258, 259 Mumme, R. L., 415 Murphy, L. E., 260 Nance, James M., 261 Nicholas, H.M., 421 -Nicoll, P. A., 128 O'Neill, C. W., 85, 262, 263 Ogden, J.C., 264 Osburn, R. C. , 265 Patella, F.J., 261 EV Iiy, 12 (C IS, S55) Paul, J.R., 281 Paulson, D.R., 318 Payne, F., 266, 267 Pearse, A. S., 268-274 Perkins, H. F., 275 Perlmutter, D.G., 151 Petrovic, C. A., 276, 277 Phillips, A. H., 278-280 Pichard, S. L., 281 Pitts, R. F., 282 Plan Development Team, Reef Fish Management Plan, South Atlantic Fishery Management Council., 283 Plantier, T. L., 284 Plough, H. H., 285 Porter, J. W., 264, 286-288 Porter, J. Y., 170 Potthoff, T., 289 Powers, P. B. A., 290, 291 Pratt, H. L. Jr., 39, 40 Pratt, H. S., 292- 297 Pritchard, M.H., 37 Raim, A., 298 Rebenack P., 21 Reighard, J., 299, 300 Reynolds, J. E. II, 301, 302 Richards, O. W., 303, 336 Richards, W. J., 289 Richardson, T., 288 Ricklefs, R. E., 304, 305, 405 Riley, G. A., 306 Riska, D. E., 307, 308 Rivas, L. R., 309 Roberts, H H., 310, 311 Robertson, D.R., 196 Robertson, W. B. Jr., 4, 5, 34, 52, 92, 157, 312- 318, 324, 355, 405, 411, 413-416 Robinson, A. H., 319 Rouse, L. J. Jr., 310, 311 Sarver, S. K., 332 Schaeffer, A. A., 320, 321 Schlumberger, H.G., 215 Schmahl, G. P., 403 Schmidt, T. W., 373 Schmitt, W. L., 322 Schnell, G. D., 323 Schreiber, R. W., 324 Schroeder, P. B,. 325, 365 Scott, W. E. D., 326 Seaman, W., Jr., 327 Shimizu, Y., 333 Shinn, E. A., 203, 328-330 Shoemaker, C. R., 331 Siegel, D. M.., 21 Silberman, J. D., 332 Smayda, T. J., 333 Smith, G. J., 287 Smith, H. B., 254 Smith, H. G., 334 Smith, N. P., 264 Snoek E., 105 South Florida Area Study 335 Spence, J., 336 Sprunt, A. Jr., 337-343 Steinen, R. P., 134 Steinmetz, J. C., 302 Stevenson, J. O., 344 Steward, F. C., 345, 346 Stockard, C. R., 347-349 Stoddart, D. R., 350 Stone, R. G., 351-353 109 Stoneburner, D. L., 354, 355 Yamanouchi, S., 417 Strom, R. N., 356 Yonge, C. M., 418-421 Stromsten, F. A., 357, 358 Szmant, A. M., 264 Zeleny, C., 422, 423 Zheng, W., 424 Tandy, G., 359 Tartar, V., 360 Tashiro, S., 361 Taylor, C. V., 371 Taylor, Jane B., 362 Taylor, W. R., 363, 364 Teas, H. J., 365 Tennent, D. H., 366-371 Thiele, J., 372 Thompson, J.C., 177 Thompson, M. J., 175, 373 Thorp, E. M., 374 Tomas, C. R., 333, 375 Torrey, H. B., 376 Treadwell, A. L., 377-379 Ubelacker, J. M., 380 Van Fleet, E.S., 424 Vaughan, T. W., 374, 381-391 Visscher, J. P., 392 Vukovich, F. M., 393, 394 Walker, N. D., 310, 311 Wallace, W.S., 395 Walsh, P.J., 332 Wartman, W., 396 Watson, J. B., 397, 398 Wells, R.rC., 399 Westinga, E., 400 Westrum, B.L., 401 Wheaton, J., 164-168, 402, 403 Whitaker, D. M., 371, 404 White, S.C. 305, 405, 415 White-Schuler, S. C., 304 Wichterman, R., 406, 407 Williams, O. L., 408 Willier, B. H., 409 Wilson, C. B., 410 Winegarner, C. E., 15, 411 Wolfe, C. A., 412 Woolfenden, G.E., 4,5, 413-416 110 Subject Index Accipiter striatus, 3 Acropora, 241 Acropora cervicornus, 30, 73, 74, 193 Acropora palmata, 74, 172, 328, 388 Actinian, 60, 136 Alcyomeum, 48 Alcyonaria, 48, 407 Alpheidae, 63, 70, 248 Alpheus, 63, 70 Amphipods, 151, 271, 331 Anchor damage, 73 Animal succession, 57 Anisonema vitrea, 102 Annelids, 119; 120, 228, 351, 352, 353, 377, 378, 379, 380 Anous stolidus, 34, 307, 308, 405, 412 Ants, 55, 242 Aplopus, 178, 186, 347 Aplysia protea, 257 Ascidians, 18, 126, 127, 128, 140, 228, 285, 303, 336, 376 Asio flammeus, 157 Astroea longispina, 43 Astropecten duplicatus, 65 Atlantic palolo (Worm), 227, 228 Atmospheric compounds, 154, 249 Audouinella ophioglossa, 9 Audubon, 10 Avicennia, 83 Banded red knot, 413 Barnacles, 392 Bathydrilus oligochaeta, 103 Beach rock, 106, 107, 114, 259 Beach tar, 424 Belonidae, 27 Bioluminescence, 144, 145 Bird history, 338, 339, 343, 344, 397 Birds, 4, 5, 10, 17, 108, 137, 157, 316, 326, 335, 337, 338, 339, 341, 342, 344, 413 Black noddy, 313, 318 Black phoebe, 108 Black water event, 163, 170 Blood relationships, 25 Bluefin tuna, 289 Botanical ecology, 24 Brackish-water ponds, 268 Breeding, 11, 13, 14, 284, 308 Briareum, 48, 163 Brown anole, 411 Brown noddy, 34, 305, 307, 308, 313, 397, 398, 405, 412 Brown pelican, 15, 324 Bryozoans, 265 Bubulcus ibis, 415 Calcium carbonate, 99, 106, 107, 201, 202, 246, 249, 259, 274, 385, 391 Calidris canutus, 413 Cardinal, 16 Cardinal cardinalis, 16 Cassiopea, 44, 45, 47, 142, 228, 237, 244, 245, 334 Cassiopea xamachana, 116, 146, 147, 148, 231, 234, 235, 238, 275, 334, 349, 422 Cattle egrets, 137, 415 Caulerpa, 417 Cerions, 11, 12, 13, 14 Chart, 109 Ciliates, 35, 290, 291, 360, 406, 407 Clibanarius, 192 Cnideria, 22, 41, 42, 46, 48, 59, 98, 135, 146, 147, 148, 161, 162, 168, 171, 172, 193, 228, 231, 234, 235, 237, 241, 244, 245, 334, 349, 395, 402, 403, 418, 419, 422 Coelenterates (Cnideria), 144, 275 Color patterns, 61, 62, 123, 130, 204, 205, 206, 207, 209, 210, 299 Common tern, 304 Condylactis gigantea, 135 Conocarpus, 83 Coral bleaching, 161, 288 Coral growth, 335, 385, 388 Coral mortality, 287, 288, 310, 311 Coral polyps, 22 Coral reef, 1, 2,57, 73, 161, 162,163; 16s-steve 168, 252, 264, 287, 288, 310, 311, 359, 381, 382, 383, 384, 385, 386, 387, 390, 403 Coral reef sampling, 165, 167, 288, 359, 403 Coral reef structure, 74, 79, 100, 168, 172, 335, 385, 387, 389, 391 Corals, 98, 226, 229, 241, 248, 252, 287, 383, 402, 403, 418, 420 Crabs, 33, 71, 192, 204, 257, 274, 303, 422, 423 Crangon armillatus, 62, 70, 71, 89, 156, 257 Crustaceans, 66, 71, 75, 78, 81, 89, 104, 125, 145, 151, 156, 192, 204, 248, 261, 274, 303, 322, 331, 410, 422, 423 Cultural resources, 260 Currents, 384, 389, 393, 394, 424 Cytology, 33, 43, 50, 59, 87, 96, 97, 98, 117, SAT I2 NAS 49 ISON IS2 153578; 179, 180, 181, 182, 183, 184, 185, 186, 191, 194, 195, 222, 223, 224, 244, 247, 254, 257, 266, 267, 320, 321, 336, 345, 346, 349, 353, 360, 366, 367, 368, 370, 371, 376, 404, 406, 409, 417 Diplectrum vittatum, 21 Diadema, 196 Diatoms, 58, 218 Diplectrum formosum, 21 Diporia clivosa, 168 Diving helmet, 131 DNA,90, 332 Dove, 315 Echinaster, 65, 182 Echinoderms, 53, 118, 121, 122, 149, 150, 153, 182, 185, 190, 194, 195, 196, 247, 254, 280, 290, 291, 360, 366, 368, 369, 370, 371, 376, 404 Echinometra lacunter, 53, 150, 194, 195, 254, Sil, SHG) Ecology, 83, 84 Effect of light on organisms, 28, 65, 72, 155, 156, 159, 222, 223, 334, 346, 368 Effect of photosynthesis, 230, 246 Effects of radiation, 351, 352 Effects of temperature on organisms, 224, 229, 234, 241, 244, 287, 310, 311, 388, 390, 393, 394 Empidonomus varius, 26 Epinephelus, 282 Epinephelus morio, 273 Evolution, 138 Exococetidae, 27 Fish, 21, 27, 28, 29, 33, 36, 39; 61, 68, 94, 112, MB IS 24 29M IG OMS2lS3 135385 139, 170, 176, 177, 205, 206, 207, 209, 210, DEO NSS ANA S22 1295 282,283; 289, 299, 309, 373, 408, 410 Fish communities, 175, 212, 283, 373 Fish kill, 310 Fish sampling techniques, 175, 205 Fish tumors, 213, 214, 215 Flight speed (Birds), 323 Florida Keys National Marine Sanctuary, 7, 20 Food habits, 21, 92, 171, 205, 397, 421 Foraminiferae, 67, 216 Frigate birds, 323 Gastropods, 89 LU Geologic history, 381, 382, 383 Geology, 30, 83, 106, 107, 114, 134, 172, 203, 256, 258, 259, 260, 262, 263, 302, 328, 329, 374, 381, 382, 383, 385, 391 Ginglymostoma cirratum, 39, 112, 129, 133 Gorgonacae, 41, 42, 46, 402 Gorgonia, 48, 402 Great black-backed gull, 188 Growth, 303, 304, 305, 346, 348, 405, 412 Habits, 64, 132, 133, 248, 272, 273, 275, 347, 398, 418, 419, 421 Haemulidae, 36 Halichores, 213 Halichores bivittatus, 124 Halimeda, 263 Halimeda hummii, 9 Hatching success, 23 Hemirhamphidae, 27 Histology, 190 Holothuria, 190, 279, 280, 369 Homing instinct, 10, 398 House sparrow, 416 Hydroids, 38, 116, 376, 395 Hydrology, 134 Hypoglossum rhizophorum, 9 Hypoplectrus, 94, 373 Insects, 8, 347 Invertebrates, 6, 8, 11, 12, 13, 14, 18, 22, 25, 32, 35, 37, 38, 41, 42, 43, 44, 45, 46, 47, 48, 49, 50, 51, 53, 54, 55, 58, 59, 60, 62, 63, 64, 65, 66, 67, 69, 70, 71, 73, 74, 75, 76, 77, 78, 80, 81, 82, 86, 87, 88, 89, 98, 100, 101, 102, 103, 104, 117, 118, 119, 120, 121, 122, 125, 126, 127, 128, 135, 136, 140, 142, 143, 144, 145, 146, 147, 148, 149, 151, 152,.153, 155, 156, 161, 162, 169, 171, 172, 178, 182, 183, 185, 186, 187, 189, 190, 192, 193, 195, 196, 197, Ie, ID, Bil, PIO, QAO, PA, P27, Q29), Z3i- 235, 238, 239, 242, 244, 245, 246, 247, 248, 253, 254, 261, 265, 266, 267, 270, 271, 272, 275, 279, 280, 285, 290, 291, 295, 294, 295, 296, 297, 303, 322, 331, 334, 347, 351, 352, 353, 360, 366, 367, 368, 369, 376, 377, 378, 379, 380, 392, 395, 400, 402, 408, 418, 419, 421, 422 Iotrochota, 87, 152 Ischnochiton, 187, 217 Land-birds, 3, 16, 26, 276, 277, 315, 317, 326, 341, 415, 416 112 Larval shrimp, 66 Launcularia, 83 Least terns, 414 Leodice fucata, 54 Limulus, 184, 228 Linckia, 360 Lipogramma anabantoides, 113 Littoral, 269, 272 Loggerhead turtle, 159, 179, 180, 181, 183, 198, 295, 297, 357, 358, 409 Long Term Ecological Research (LTER) 163, 167, 225, 264 Longevity, 34 Lutjanus griseus, 124, 214 Lytechinus variegatus, 50, 153, 247, 368, 370, 404 Maeandra aerolata, 22, 229, 241, 418 Maps, 1, 74 Manatees, 301 Mangrove, 83, 325, 365 Marine algae, 9, 19, 96, 97, 191, 222, 223, 224, 345, 346, 363, 364, 417 Marine amoebe, 160, 320, 321 Marine bacteria, 99, 110, 111, 202 Marine fishery reserves (MFR), 283 Marine Laboratory, 232, 236, 240 Marine parks, 79, 319 Masked booby, 52 Mastogloia, 218 Mating, 39 Medusae, 44, 45, 228, 231, 234, 239, 275, 349 Merlin, 298 Metals in organisms, 278, 279, 280, 354, 355, 356 Migration, 17 Millepora complanata, 161, 172 Mites, 274 Mollusks, 11, 12, 13, 14, 49, 69, 89, 187, 217, DIRE) eed, 92, C19) | Monocotyle, 292 Monospotus indicus, 9 Montastrea annularis, 162, 168, 172, 241, 385 Montastrea cavernosa, 168 Morphodynamics, 85, 192 Mosquitoes, 8 Myliobatis, 292 Nematodes, 408 Neoplastic growths, 124 Nesting, 15, 324, 343, 397, 414, 416 Nurse shark, 39, 40 112, 129, 133, 197 Ocypoda, 64 Oikopleura, 189 Oil spill, 335 Ophicoma, 182 Orbicella (Madrepora), 193, 229, 241, 388 Osmotic pressure, 269 Owl, 157 Panulirus argus, 75, 77, 78, 89, 156, 327, 332 Parasitic copepods, 410 Parasitic isopods, 273, 410 Parasitic worms, 37, 197, 198, 199, 219, 220, 221, 253, 292, 293, 294, 295, 296, 297 Pedibothrium, 37 Penaeus duorarum, 66, 261 Pesticides, 356 PH, 230, 250, 251, 334 Physiology, 47, 97, 111, 135, 160, 169, 334, 345, 346, 357, 358 Phytoplankton, 333, 375 Plankton, 306 Plectognath, 28 Plexaura, 46 Pollution, 335, 424 Polycistor, 140 Polyplacophora, 217 Pomacentrus, 29 Pomacentrus leucostictus, 61 Porites, 229, 241 Porites asteroides, 168 Portunus sayi, 422, 423 Promicrops (Epinephelus) itajara, 273 Protozoan, 102, 160, 169, 406, 407 Pseudocyphoma, 49 Ptychodera, 43, 89, 155, 266, 267 Pycnogonida, 51 Rails, 277 Rat, 95 Recreational headboat fishery, 20 Red-headed woodpecker, 317 Red tide, 105 Reef fishes, 20 Reef formation, 2, 41, 42 Reefs, 30 Regeneration, 6, 44, 45, 47, 115, 116, 120, 152, 190, 235, 285, 348, 349, 351, 352, 353, 360, 422, 423 Replenishment reserves, 7, 20, 283 Rhizophora, 83 Richardina spinicincta, 125 Roseate terns, 313, 414 Sabellids, 6, 352, 379 Sabines's gull, 56 Salinity, 93, 116 Salpa floridian, 31 Scleractinia, 161 Scyphomedusa (jellyfish), 59, 234 Sea-birds, 4, 15, 23, 34, 52, 56, 91, 92, 188, 284, 298, 304, 305, 307, 308, 312, 313, 318, 323, 324, 340, 341, 354, 355, 397, 398, 405, 412, 414 Sea-cucumber, 190 Sea-level change, 203, 258, 262, 263 Sea turtles, 159 Sea-urchin, 50, 117, 121, 122, 150, 153, 194, 195, 196, 247, 254, 290, 291, 366, 368, 370, 371, 404 Sea-water composition, 72, 90, 93, 115, 174, 200, 201, 202, 230, 235, 237, 238, 243, 246, 249, 250, 251, 281, 320, 361, 389, 399, 401 SEAKEYS, 264 Sedimentation, 85, 172, 256, 258, 259, 262, 263, 329, 374 Sharks, 39, 139 Sharp-shinned hawks, 3 Sicyonia penaeoidea, 104 Siderastera, 168, 229, 241, 385, 419 Snapping shrimps, 63, 322 Sooty terns, 4, 5, 23, 91, 92, 284, 304, 312, 313, 314, 354, 355, 397, 398, 405 Speculata advena, 49 Spheciospongia vesparia, 151, 400 Sphyraena barracuda, 132 Spiny lobster, 75, 76, 77, 78, 80, 81, 82, ISO, S27, Sw Sponges, 86, 87, 88, 101, 151, 152, 270, 400, 403 Sport harvest, 75, 76, 78, 79, 81, 82 Starfish, 65, 360, 369 Starvation, 231, 334, 415 Stephanochasmus, 294 Sterna fuscata, 4, 5, 91, 92, 284, 304, 314, 354, 405 Sterna hirundo, 304 Sterna melanoptera, 340 Sting ray, 68 Storms, 17, 405 Sula dactylata, 52 Syllids, 380 Synentognathi, 27 113 Topography, 1, 84, 325, 328, 329, 343, 350 Transatlantic migration, 314 Transplanting organisms, 403 Trematodes, 220, 221, 253, 292, 293, 294, 295, 296, 297 Trichechus, 301 Tunicata, 31, 32 Valonia, 19, 96, 97, 191, 222, 223, 224, 345, 346 Vanadium, 279, 280 Variegated flycatcher, 26 Vegetation, 24, 84, 255, 325, 350, 365 Vessel groundings, 335, 403 Vocal signals, 307, 308 Worms, 6, 54, 103, 119, 120, 155, 197, 198, DY, By ALVZ, Psa, 22), P23)5 P2O, PM, SIS, 353, 377, 378, 379, 380, 408 Zooxanthellae, 98, 334, 407, 421 *# U.S. GOVERNMENT PRINTING OFFICE: 1998- 434-814 ATOLL RESEARCH BULLETIN NOS. 443-449 NO. NO. NO. NO. NO. NO. NO. 443. 444. 445. 446. 447. 448. 449. THE EVOLUTION OF A HOLOCENE FRINGING REEF AND ISLAND: REEFAL ENVIRONMENTAL SEQUENCE AND SEA LEVEL CHANGE IN TONAKI ISLAND, THE CENTRAL RYUKYUS BY H. KAN, N. HORI, T. KAWANA, T. KAIGARA, AND K. ICHIKAWA CHECKLIST OF THE SHOREFISHES OF OUVEA ATOLL, NEW CALEDONIA BY MICHEL KULBICKI AND JEFFREY T. WILLIAMS ON THE ORIGIN OF DRIFT MATERIALS IN THE MARSHALL ISLANDS BY D.H.R. SPENNEMANN DISTRIBUTION OF RAT SPECIES (RATTUS SPP.) ON THE ATOLLS OF THE MARSHALL ISLANDS: PAST AND PRESENT DISPERSAL BY D.H.R. SPENNEMANN A POSSIBLE LINK BETWEEN CORAL DISEASES AND A CORALLIVOROUS SNAIL (DRUPELLA CORNUS) OUTBREAK IN THE RED SEA BY ARNFRIED ANTONIUS AND BERNHARD RIEGL MARINE ALGAE FROM OCEANIC ATOLLS IN THE SOUTHWESTERN CARIBBEAN (ALBUQUERQUE CAYS, COURTOWN CAYS, SERRANA BANK, AND RONCADOR BANK) _BY GUILLERMO DIAZ-PULIDO AND GERMAN BULA-MEYER SCIENTIFIC STUDIES ON DRY TORTUGAS NATIONAL PARK: AN ANNOTATED BIBLIOGRAPHY BY T.W. SCHMIDT AND L. 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