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Age) OT je December 31, 1972 ATOLL RESEARCH BULLETIN Bacterial Counts in Surface Open Waters of Eniwetok Atoll, Marshall Islands by Louis H. DiSalvo Marine Studies on the North Coast of Jamaica edited by Gerald J. Bakus Fish Diversity on a Coral Reef in the Virgin Islands by Michael J. Risk Recolonization of a Population of Supra- tidal Fishes at Eniwetok Atoll, Marshall Islands by William A. Bussing Some Marine Benthic Algae from Truk and Kuop, Caroline Islands by Roy T. Tsuda Additional Records of Marine Benthic Algae from Yap, Western Caroline Islands by Roy T. Tsuda and Mary S. Belk 157. 158. 159. 160. 161. 162. Carbonate Lagoon and Beach Sediments of Tarawa Atoll, Gilbert Islands by Jon N. Weber and Peter M. J. Wood- head Birds Seen at Sea and on an Island in the Cargados Carajos Shoals by R. Pocklington, P. R. Willis and M. Palmieri Geomorphology and Vegetation of Iles Glorieuses by R. Battistini and G. Cremers Reef Islands of Rarotonga by D. R. Stoddart with list of vascular ~ plants by F. R. Fosberg South Indian Sand Cays by D. R. Stoddart and F. R. Fosberg Island News and Comment am Issued by THE SMITHSONIAN INSTITUTION Washington, D.C., U.S.A. 151. 152. 153. 154. 155. 156. ATOLL RESEARCH BULLETIN Bacterial Counts in Surface Open Waters 157. of Eniwetok Atoll, Marshall Islands by Louis H. DiSalvo Marine Studies on the North Coast of Jamaica 158. edited by Gerald J. Bakus Fish Diversity on a Coral Reef. in the Virgin Islands by Michael J. Risk 159. Recolonization of a Population of Supra- tidal Fishes at Eniwetok Atoll, Marshall Islands by William A. Bussing 160. Some Marine Benthic Algae from Truk and Kuop, Caroline Islands by Roy T. Tsuda : , , 161. Additional Records of Marine Benthic Algae from Yap, Western Caroline Islands 162 by Roy T. Tsuda and Mary S. Belk ‘i Issued by Carbonate Lagoon and Beach Sediments of Tarawa Atoli, Gilbert Islands by Jon N. Weber and Peter M. J. Wood- head Birds Seen at Sea and on an Island in the Cargados Carajos Shoals by R. Pocklington, P. R. Willis and M. Palmieri Geomorphology and Vegetation of Iles Glorieuses by R. Battistini and G. Cremers Reef Islands of Rarotonga by D. R. Stoddart with list of vascular plants by F. R. Fosberg South Indian Sand Cays by D. R. Stoddart and F. R. Fosberg Island News and Comment THE SMITHSONIAN INSTITUTION Washington D.C., U.S.A. December 31, 1972 ACKNOWLEDGMENT The Atoll Research Bulletin is issued by the Smithsonian Institution as a part of its Tropical Biology Program. It is co- sponsored by the Museum of Natural History, the Office of Environ- mental Sciences, and the Smithsonian Press. The Press supports and handles production and distribution. The editing is done by the Tropical Biology staff, Botany Department, Museum of Natural History. Final typing of this issue was contributed by the U. S. Air Force (AF/LGFC, Hq USAF). The Bulletin was founded and the first 117 numbers issued by the Pacific Science Board, National Academy of Sciences, with financial support from the Office of Naval Research. Its pages were largely devoted to reports resulting from the Pacific Science Board's Coral Atoll Program. The sole responsibility for all statements made by authors of papers in the Atoll Research Bulletin rests with them, and statements made in the Bulletin do not necessarily represent the views of the Smithsonian nor those of the editors of the Bulletin. Editors F. R. Fosberg M.-H. Sachet Smithsonian Institution Washington, D. C. 20560 De Re Stoddart Department of Geography University of Cambridge Downing Place Cambridge, England ATOLL RESEARCH BULLETIN No. 151 BACTERIAL COUNTS IN SURFACE OPEN WATERS OF ENIWETOK ATOLL, MARSHALL ISLANDS by Louis H. DiSalvo Issued by THE SMITHSONIAN INSTITUTION Washineton.) D. C., U. S. A. December 31, 1972 » 2 ‘ i He i i { . {i Ay s t eer i a a WA, eh hi Ae yao mn ale , vf Q © DAY. ‘ lies) 4 Y en n ave d ae! fa) ‘igs ney BoA ie r < ; , : 7 i iH ¥ P 53 > Ae : fi S x 5 : ore ri M ean aes = ai lay eit * = i ey H r a ‘ vt 7 6} z : a4 eeu = Se SRE Ber We. 4, a ¢ ely 2 eee Be) f) ~ va i ie f fq = % L : 4 ra 4 i 2 *., ; a e a ‘i 2 v j ‘ = al i — rice } 2 bet oli i i As) ae > + i & ar - , F e i y " ya ia ‘+ 4 ; , ; hy uw ° A 7 , i i r ? rE - ‘ 7 t 1 1 i ; : S : i oo oo Se and a ( ' - i a8 7 ; yas eae uh f ten - \¥ ‘ os é i s ° , Gi oe ‘ f i : = R fie at F : Pye 0 x i i Oy i i _ - a . = M i‘ - =e j ns a Wiss ° = 1 y eae \ i e a ; ; i bate Joa \ ; 5 , ‘ a 4 - > > (i Tbe on BACTERIAL COUNTS IN SURFACE OPEN WATERS OF ENIWETOK ATOLL, MARSHALL ISLANDS by Louis H. DiSalvo’* INTRODUCTION AND METHODS During my investigations of coral reef microbiology at Eniwetok Atoll, Marshall Islands in 1968, large numbers of bacteria were found inhabiting coral reef internal sediments (10‘-10 colonies/g; DiSalvo, 1969). With the high level of metazoan activity on reefs it was expected that sediment processing and disturbance might cause the release of bacteria, contributing to the organic particulate enrichment of lagoon waters (Marshall, 1965; Johannes, 1967). To my knowledge, no bacteriological data were available which compared oceanic counts to lagoonal counts near an atoll, and therefore the following measurements were made both in reference to the work at hand and as a general contribution to atoll ecology. Open water sampling for aerobic heterotrophic: bacteria was carried out on six bi-weekly dates in June and July 1968. A total of 24 samples of surface seawater were obtained at oceanic and lagoonal stations. In addition, a total of 14 samples were obtained downstream from the Japtan reef at the surface and at 2 meters depth. Sample locations are shown in Figure 1. Surface (top 5 cm) water samples were obtained in sterile 200 ml prescription bottles from a boat. Skin diving was employed to obtain the 2m samples. Samples were kept on ice for return to the laboratory, and plated for bacterial counts within five hours of collec- tion. The plating medium was ZoBell 2216e (Oppenheimer and ZoBell, 1952) solidified with 1.5% (w/v) agar. Duplicate pour plates were made using 0.1 ml raw seawater inocula and 10 ml medium which had been cooled to 40-45°C. Plates were incubated at ambient tempera- ture (30°C) for 48 hours and then counted for total colonies. RESULTS AND DISCUSSION Bacterial counts for numbered samples located by Figure 1 are listed in Table 1. These counts are within order of magnitude limits established by other investigators using similar methods on waters near non-atoll reefs (Table 3). Table 2 lists counts for two depths immedi- ately behind the Japtan Reef. Summarized results in Table 3 suggest there was no significant difference between oceanic and back-reef counts. Conclusions from these data are tentative and preliminary. Comparisons with plate counts of other workers in Table 3 have been made for convenience, although they may be invalidated by slight differences in methodology and materials. The limitations of the plate count technique in obtaining ‘‘true counts’? are well known (ZoBell, 1946; Jones and Jannasch, 1959) and present results can only be considered as a measure of relative microbial activity. The counts are —— ee *Present address: Naval Biomedical Laboratory, Naval Supply Center, Oakland, Calif. 94625. (Manuscript received July 1970--Eds.) 2 within higher ranges expected for clear tropical pelagic waters, and it may therefore be hypo- thesized that the waters surrounding the atoll are enriched in some way, perhaps by local up- welling or organic enrichment from the reefs. The small numbers of sample platings preclude rigorous statistical interpretations of the data. More seriously, I was unable to sample the same water mass flowing from ocean to lagoon over the Japtan Reef for the before-after measurements. Both these factors were the result of logistical problems, helping to explain why there are essentially no microbiological data available for atoll waters. Since ithas previously been suggested that the Japtan Reef was in a steady state with regard to the import and export of matter and energy (Odum and Odum, 1965), it is not unreasonable to deduce from the available data that this reef is in equilibrium with regard to input and output of heterotrophic bacteria. Future studies should attempt to confirm present results under various seasonal and tidal regimes, and extend the measurements to representative depths within and outside the atoll. Such measurements might be useful in studies of outer slope upwelling and local enrichment due to the coral reefs. ACKNOWLEDGEMENT The author wishes to thank the Eniwetok Marine Biological Laboratory and the University of Hawaii for support and facilities used to accomplish this research. LITERATURE CITED DiSalvo, L.H. 1969. Regenerative functions and microbial ecology of coral reefs. Ph.D. Thesis. Univ. of N. Carolina, Chapel Hill. 289 p. Gee, A.H. 1932. Lime deposition and the bacteria. I. Estimate of bacterial activity in the Florida Keys. Pap. Tortugas Lab., Carnegie Inst., Washington, D.C. 28: 67-82. Gundersen, K.R. and D.B. Stroupe. 1967. Bacterial pollution of Kaneohe Bay, Oahu. Univ. Hawaii Water Resources Res. Ctr., Tech. Rpt. 12. 24 p. Johannes, R.E. 1967. Ecology of organic aggregates in the vicinity of a coral reef. Limnol. Oceanogr. 12: 89-195. Jannasch, H. and G.E. Jones. 1959. Bacterial population inseawater as determined by several methods of enum=2ration. Limnol. Oceanogr. 4: 128-139. Marshall, N. 1965. Detritus over the reef and its potential contribution to adjacent waters of Eniwetok Atoll. Ecology. 46: 343-344. ~-e------- 1968. Observations on organic aggregates in the vicinity of coral reefs. Mar. Biol. 2: 50-53. Odum, H.T. and E.P. Odum. 1955. Trophic structure and productivity of a windward coral reef community at Eniwetok Atoll. Ecol. Monogr. 23: 291-320. Oppenheimer, C.H. and C.E. ZoBell. 1952. The growth and viability of sixty-three species of marine bacteria as influenced by hydrostatic pressure. J. Mar. Res. 11: 10-18. 3 Sisler, F.D. 1962. Microbiology and biochemistry of sediments and overlying water, p. 64-69, in, P.E. Cloud, Jr., Environment of calcium carbonate deposition west of Andros L., Bahamas. Geol. Surv. Prof. Pap. 350. U.S. Govt. Printing Off., Washington, D.C. 138 p. ZoBell, C.E. 1946. Marine microbiology. Chronica Botanica Pub., Waltham, Mass. 240 p. Table 1. BACTERIAL COUNTS FOR ENIWETOK ATOLL OPEN WATERS Sam ple* Sample Colonies/m1** Colonies/ml *For sample location, see Fig. 1. **Colonies/ 0.1 ml plated x 10. Table 2. BACTERIAL COLONY COUNTS FOR JAPTAN [. BACK- REEF WATERS* Date Surface samples Samples from 2m depth** 1968 Sample # Colonies/ml Sample # Colonies/ml 7/15 1 2 lua 3 4 5 7/19 6 Mean values * Water depth approximately 4 meters. ** Obtained by diving. Table 3. SUMMARY OF COLONY COUNTS FOR ENIWETOK ATOLL OPEN WATERS Stations Number of samples Ocean (nos. 1-5, 7-11 Lagoon (all Japtan*, nos. 20-24) Previous data, near reefs Gee (1932)' Sisler (1962)? Gundersen and Stroupe (1987)° Sieburth (unpub.)* 10 19 Colonies/m1 272 = 100 222 (6) 100,150, 700** < 10-103 90,60,1360,12,42** 1352;0** 215,235,250, 700** *See table 2. **Individual results. *Open water near Florida reef tract (Gulf Stream, Bird Key Harbor, Marquesas lagoon). * Water over Great Bahama Bank west of Andros I. * (a) immediately outside northern Kaneohe Bay, Hawaii. (b) Kauai I., Hawaii, 1.5-2.5 miles offshore. * unpolluted lagoon stations, Majuro Atoll, Marshall Is., Feb-Apr. 1970. Unpub. data of J.MMcN.Sieburth, U.Rhode I. Narragansett Marine Laboratory, Kingston. Japtan back-reef stations “x Japtan |. <— We LAGOON NORTH PACIFIC OCEAN Figure 1. Southeast quadrant of Eniwetok Atoll, Marshall Is. Gres Oxine 165°15'W), to show sampling stations. Approximate current pattern is designated by small arrows. Contour interval 10 meters. sara ng pa ATOLL RESEARCH BULLETIN No. 152 MARINE STUDIES ON THE NORTH COAST OF JAMAICA edited by Gerald J. Bakus Issued by THE SMITHSONIAN INSTITUTION Washington, D. C.., U.S. A. December 31,. 1972 — tin ae ai 1 & i! an , a a aed { E Oth aera pe: : soe at Diets w es i’ — 5 ion lek Eb ‘ _ We G 25s Wks = 2 ee ss eral Homiwnlt Cork am f . 7 - teat = : ‘ ; ees TE = a, ne tn ; ii i J \ ] : “Ae H ‘ : eee L ; 20 eC OTR : } : 7 i i, ne —T = i _" ‘0 , : wy i208 7 ; 1 y ' : ' ‘i foe : : ( , ¢ ae tom ; a ” - 7 : | ; v ‘ ' = . iat : : r : ; 4 ' - ¢ >» i ie MARINE STUDIES ON THE NORTH COAST OF JAMAICA Edited by Gerald J. Bakus’ INTRODUCTION During July and August of 1970 a group of 12 students and several instructors participated in research and teaching in the second Organization for Tropical Studies (OTS) course in Tropical Marine Biology. The primary site was the Discovery Bay Marine Laboratory and immediate environs on the north coast of Jamaica. The laboratory provided facilities for maintaining live marine organisms, under the direction of Mr. Norman Copland, Manager. Discovery Bay consists of a variety of marine communities, particularly sandy bottoms, Thalassia beds, and coral patch reefs. Among the most striking communities is an exceptional sponge bed located at a depth of 50-75 feet (15-23m) just off Colombus Park, with certain specimens approaching the size of a washtub. The bay is contained by a reef crest of zoanthids, scleractinian corals, sea urchins, and numerous associated organisms. Beyong the reef crest, scleractinian corals slope gently seaward before the island shelf plunges into the depths. Underwater visibility on the outside of the reef crest often approaches 150 feet (46m). Sclerac- tinian corals are magnificently developed along this narrow shelf. Sclerosponges occur as cavernicolous cryptofauna in waters 8 to 92 meters deep (Hartman and Goreau, 1970). Goreau (1959) presented a detailed report on the ecology of coral reefs on the north coast of Jamaica. The low incidence of exposed soft-bodied invertebrates and the low standing crops of exposed fleshy algae on the reef slope are in part interpreted as the evolutionary product of tropical predator-prey and grazer-plant interactions (Bakus, 1969). The abstracts presented below summarize studies conducted during the OTS program. A complete report of our research activities is on file in the OTS North American Office. LITERATURE CITED Bakus, G.J. 1969. Energetics and feeding in shallow marine waters. Internat. Revue Gen. Exper. Zool. 4: 275-369. Goreau, T.F. 1959. The ecology of Jamaican coral reefs. Ecology 40(1): 67-90. Hartman, W.D. and T.F. Goreau. 1970. Jamaican coralline sponges: their morphology, ecology and fossil relatives. Symp. Zool. Soc. London No. 25: 205-243. *Allan Hancock Foundation, University of Southern California, Los Angeles, California (Manuscript received Nov. 1970--Eds.) ABSTRACTS Distributional patterns of rocky intertidal gastropods in Discovery Bay, Jamaica By Lenora H. Atsatt, Dept. of Environmental and Population Biology, Univ. of California, Irvine Distributional patterns of rocky intertidal gastropods in Discovery Bay, Jamaica, were examined in an attempt to develop a quantitative method of describing habitat differentiation. The relationship of common species of gastropods of the families Neritidae and Littorinidae to a series of physical variables, including texture, moisture content, slope angle, exposure, and zonation was analyzed using chi-square tests. Each species was found to differ in habitat preference from the other species for at least one of these variables. Tectarius muvicatus, a littorine, was distinct from all other species in habit preference, while the three Nevita spp. were less clearly segregated. Littorina ziczac and Echininus nodulosus, two littorines which were relatively unsegregated with regard to the physical variables, were found to be positively associated. Activity patterns in Tectavius muricatus were examined in an attempt torelate behavioral response to distributional patterns. Results indicated that moisture level is the most critical factor involved for the initiation and termination of activity, and that combinations of other en- vironmental stimuli are responsible for direction of motion. Modelexperiments onthe releasing mechanism of cleaning behavior in the shrimp Periclimenes pedersoni By Patrick L. Colin, School of Marine andAtmospheric Sciences, Univ. of Miami, Miami, Florida Periclimenes pedersoni exhibits cleaning behavior towards anesthetized fish, paper fish models, paper fish shapes, and paper rectangles. When satiated with food the shrimp did not exhibit any cleaning behavior towards any of the models. Food deprivation for periods greater than 24 hours lowered the selectivity of the cleaning response to models. A white paper model with inked features produced cleaning responses most often; more often than an anesthetized cardinal fish, Apogon maculatus. Contrast between ground color and markings was most important in releasing cleaning behavior. Movement of the model inhibited release of cleaning, and olfactory clues were not important. Some observations on the metabolism and growth of Thalassia testudinum Konig and its epibiota By David D. Dow, Dept. of Zoology, Univ. of Georgia, Athens On the basis of bottom area, Thalassia plants were found to have a metabolic rate of 3.47 g 09/m2 per day. On the basis of leaf area the Thalassia blades with epibiota had a metabolic rate of 2.27¢ 09/m2 per day, while the epibiota alone accounted for 0.18 ¢ 09/m? per day. Exclosure experiments suggested that grazing by reef fishes and invertebrates does signifi- cantly affect the standing crop of Thalassia but not the blade length distribution. 3 The distribution and abundance ofanemones and theiy commensal shrimp in Discovery Bay, Jamaica By Catherine P. Engel, Dept. of Biological Sciences, Univ. of California, Santa Barbara A two-week study of the distribution and abundance of the anemones and their associated shrimps was done in Discovery Bay, Jamaica. Transect surveys were made in mixed Thalassia (10 ft. depth), a shallow inshore shelf (1-2 ft. depth), the reef crest (1-2 ft. depth), and the back reef (2-3 ft. depth), with twenty-four 2m2 samples in each area. The anemones that were surveyed included Bartholomea annulata, Stoichactis helianthis, Condylactis gigantea, Heterac- tis lucida, Lebrunia coralligens, anda burrowing anemone (unidentified). The anemone species were most evenly ranked in the mixed Thalassia and the mean number of anemones per 2m? was greatest in the reef crest sample. No shrimgs were associated with Heteractis, Lebrunia, or the burrowinganemone. Peviclimenes yucatanicus and Periclimenes pedersoni were found on Bartholomea and Condylactis. Periclimenes Sp. was found only on Stotchactis. Alpheus armatus was associated with Bartholomea. Thor sp. was found with Stotchactis and Bar- tholomea. Thor amboinensis was associated with Bartholomea, Condylactis, and Stoichactis. Heteromysis actiniae was found with Bartholomea. The greatest number of shrimp species present was in the mixed Thalassia area. The distribution and abundance of both the anemones and their shrimps were variable from one area to another. This was primarily attributed to substrate and water movement. Patterns of distribution of Foraminifera on Thalassia testudinum By Malcolm G. Erskian, Institute of Ecology & Bodega Marine Laboratory, Univ. of California, Davis In Discovery Bay, Jamaica, the spatial distribution and density of epiphytic Foraminifera was studied. Two species of polythalamous Foraminifera, Planorbulina sp. (Family Planor- bulinidae) and Sorites sp. (Family Soritidae) were found epiphytically on turtle grass, Thalassia testudinum. Samples were collected by removing all of the Thalassia in an area of 0.1 square meter. Three replicate 0.1 square meter samples were taken at a depth of 3 meters. From each 0.1m“ plots, ten whole Thalassia plants were chosen and each blade divided into quadrats. The number of each species of Foraminifera was recorded. A co- efficient of dispersion (r) was calculated by the variance: mean ratio method. Planorbulina sp. and Sorites sp. were found to have an aggregated spatial distribution. Variance: mean ratios were found to be significantly greater than unity, except one station. Aggregation seems more pronounced in Planorbulina sp., r ranging from 1.4 to 2.4, than Sorvites sp., withr ranging from 0.9 to 1.4. There seems to be a niche partitioning between species. Planor- bulina sp. is found along the margins of the Thalassia blade ina higher proportion than to- ward the median, whereas Sorvites sp. is found in a higher proportion in the median. The epiphytic niche increases the area of available substrate when compared to the sediment surface. Thalassia was found to increase the available substrate to Foraminifera by a factor of 6, producing a very high population density when compared to a unit area of sediment surface. Planorbulina sp. averaged over 60,000 individuals per square meter sediment surface and Sorites sp. over 12,000. 4 A preliminary study on the toxicity of the sponge Haliclona rubens (Pallas) de Laubenfels By Gerardo Green-Macial, Dept. of Biological Sciences, Univ. of Southern California, Los Angeles A high degree of toxicity was found in the sponge Haliclona rubens. The toxin was ex- tracted by homogenizing 4g of live sponge in each of the following solvents: hexane, benzene, ether, chloroform, acetone, ethanol, methanol, and distilled water. After centrifugation and evaporation of the supernatent, the extracts were tested against squirrelfishes (Holocentrus vufus). The behavior of the fish was observed and timed. Control experiments were run each time. A coefficient of toxicity (of ) was calculatedaccording to the following formula: “= xe x 100 where: x = weight of the toxin, y= weight of the dead fish, t = time in which death Of the fish occurs. The effectiveness of the toxin and its solubility are inversely proportional to the value. In the case of H, rubens, the most effective solvent (lowest < value) isacetone (0.28 g of acetone-toxin extract in 2 liters of sea water, killeda 53.2 ¢ fish in 5.5 min.). Aspects of the biology of the bluehead wrasse Thalassoma bifasciatum By John B. Heiser, Dept. of Anatomy, Cornell University, Ithaca, New York Because of its shallow water diurnal habits and non-retiring nature, the labrid genus Thalassoma iS an appropriate organism for investigation of the morphological and behavioral adaptations that have contributed to the success of the suborder Labroidei in tropical marine shore waters. Pectoral muscular modifications, as wellas extensive development of the cephalic lateralis system, correlate with observed behavior patterns. The behavioral contexts of the varied color patterns were tentatively identified and found to be rather consistent. The depth distribution was examined and thought to correlate withthe environmental diversity of coralline areas, especially below 10 meters. The species is not uniformly distributed on all reef zones nor is there a uniform decline with depth, though a large percentage of the population is in water less than 30 meters. The species is an opportunistic micro-carnivore, including its cleaning behavior. Despite this latter behavior, it does not enjoy complete immunity from predation. Aggregate spawning by yellow phase fishes was repeatedly observed and analyzed into successive components. Spawning by bluehead phases was not observed but it is felt that such polymorphism may be an important key to the success of the Labroidei, many of whose members Show similar patterns. Observations on the associations and feeding of six species of prosobranch gastropods on Anthozoans By Alan C. Miller, Dept. of Biology, Univ. of Oregon, Eugene The associations and predation of prosobranch gastropods on anthozoans was studied in Discovery Bay, Jamaica. Coralliophila abbreviata was associated with 12 scleractinian coral species, Ricordea florida (Actiniaria), and Zoanthus sociatus (Zoanthidea). Coralliophila caribea was associated with 6 stony corals, 3 gorgonians, Zoanthus florida, andthe actiniarians Rhodactis sanctithomae and Ricordea florida. Two methods of feeding were observed for Coralliophila: the proboscis is inserted through the epidermis of corals into the gastroderm‘s where it is moved around; the proboscis is extended over the colony and inserted into the oral opening of an individual polyp. It could not be discerned what the Covalliophila were digesting, but zooxanthellae were found in the digestive tract. 4) Cyphoma gibbosum is associated with erect gorgonians. The many undigested zooxan- thellae in its fecal pellets suggest that it mainly digests animal tissues. Heliacus cylindricus and H, infundibuliformis (one individual each) were found among Zoanthus florida polyps. Feeding was not observed. One Calliostoma javanicum (Trochidae) was found, and it consumed part of an Agaricia agaricites colony (scleractinian coral) in the laboratory. Its fecal pellets contained many undigested zooxanthellae and undischarged nematocysts which suggests that it might be digest- ing the animal tissue. This has been observed before at the Discovery Bay Marine Laboratory and is of interest since trochids are considered to be herbivores. Relationships between type of locomotion, size, and speed in larger gastropod molluscs By Susanne E. Miller, Dept. of Zoology, Univ. of Washington, Seattle The speed and methods of locomotion in a wide size range of large Caribbean gastropod species were studied in order to determine the effects of absolute size on different types of locomotion. Cittarium pica and Fasciolaria tulipa move by retrograde alternate ditaxic waves of muscular contraction of the sole of the foot, Stvombus gigas by leaping movements, and Cassis species apparently by ciliary action. Leaping is the fastest method of locomotion and ciliary movement is slowest in these large gastropods. Absolute speed on suitable horizontal substrates seems to increase with size over the size range of the species tested, and absolute speed on vertical or inclined substrates also increases but at a lower rate. These effects of weight on speed are most apparent in the species with ciliary locomotion and in Fasciolaria tulipa, in which adhesion at higher speeds is poor. Speed is inversely proportional to size in Cittarium pica but directly proportional to sizein Strombus gigas. The limitations ofthe types of locomotion with absolute size are discussed with respect to different types of habitats. Preliminary study of six Jamaican blue crabs, genus Callinectes (Decapods: Portunidae) By Elliott A. Norse, Dept. of Biological Sciences, Univ. of Southern California, Los Angeles Results of a summer study of Jamaican blue crabs are presented. Ecological separation between the six Jamaican species is discussed. Collecting localities for Callinectes sapidus, C. bocourti and C, exasperatus indicate that these prefer mud bottoms to sand bottoms; C. danae is found on both bottom types, while C. marginatus and C. ornatus are members of sandy bottom communities. Callinectes bocourti and C. sapidus males are found in fresh water rivers; C. danae, C. exasperatusand C,. sapidus females live in estuaries and brackish bays while C. marginatus and C. ornatus are recorded from marine situations. Further separation based on feeding is proposed, with C. exasperatus and C. marginatus being able to exploit hard-bodied food organisms, while other species are not anatomically equipped to do so. An albinistic population of Callinectes ornatus is described in conjunction with im- plications on the evolutionary role of geographical isolation in speciation. Cannibalism is an important population control in Callinectes species. Laboratory studies of interspecific competition found males of C. exasperatus occupying the dominant position in shelter hierarchy while males of C, bocourti are more active in asserting feeding dom‘nance in a two-species experiment. Callinectes species are found to be the largest and most active carnivores resident in estuarine and brackish environments on Jamaica’s northern shore. Using a ‘“‘Number of Features of Difference’’ table, a tentative phylogeny of six Jama‘can anda seventh species of swimming crabs from the Pacific (Panama) is proposed. Callinectes marginatus and C. exasperatus are found to compose a closely related ‘‘species group’? while C. ovnatus, C, danae and C. arcuatus represent a second species group. 489-409 O - 73 - 2 6 On the biology of Ophioblennius atlanticus By Janet Osburn, Dept. of Zoology, Univ. of Washington, Seattle The blenniid Ophioblennius atlanticus is a common shallow water shorefish inhabiting regions of primarily dead coral and cobble rock. Like many other members of its family, this species maintains a stable home range, generally about two mSters in diameter. In- dividual fish forage over this more or less defined area in short bursts of feeding activity, biting detritus and fine algal layers from the substratum. The territory typically includes a station such as a particular rock or crack to which the fish returns between forays. Initial tagging experiments showed that when individuals were transplanted several meters from their hom? range, there was a tendency for them to return from such transfers. A study of predation on tropical holothurians at Discovery Bay, Jamaica By James D. Parrish, Graduate School of Oceanography, Univ. of Rhode Island, Kingston Evidence is presented for predation on littoral holothurians by large gastropods in the lagoon behind a fringing reef at Discovery Bay, Jamaica. An observation of consumption of an Actinopyga agassizi by the triton Charonia variegata in the field was followed by observa- tions in the laboratory of attacks by 4 specimens of C. variegata on 5 specimens of A. agassizi and by 5 specimens of C. variegata on 2 specimens of Ludwigothuria mexicana. In all, 4 A. agassizi and 2 L. mexicana were killed and eaten. There is strong evidence that Isostichopus badionotus is also attacked in the laboratory and in the field. The full sequence of attack and defensive behavior was observed in the laboratory. Actinopyga agassizi appears to release a noxious substance locally from the body surface under rather specific form: of irritation, including attack by C. variegata. This defense appears to be rather effective under some circumstances. Advanced hunger and periods of exposure in close proximity to the holothurian appear to give C. vaviegata sufficient resistance to the ‘‘noxin’’ to permit success- ful predation. Observations and repeated census in the field study area suggest that, based on numbers of holothurians and predaceous snails and normal range of movem=ents, this pre- datory interaction may be of ecological importance. Larval salinity acclimatization in the tropical shore crab, Sesarma ricordi By Jon D. Standing, Dept. of Zoology, Univ. of California, Berkeley Zoea of the euryhaline shore crab, Sesarma vicordt, from Jamaica, were collected in environments with high (36.8 %o) and low (4.2 %o) salinities and were subjected to ten acutely administered salinity conditions from 0 to 175% sea water for three days. One hundred per- cent of the high acclimatized zoea survived 100% S.W.; survival percentages decreased to forty-five percent at 25% S.W. and fifty-five percent at 150% S.W. By contrast, one hundred percent of the low acclimatized zoea survived the range from 10 to 100% S.W., and seventy and forty percent of the animals survived 5% and 150% S.W., respectively. More animals survive a graded salinity stress than an acute stress of the sam2 magnitude and over the same time period. Hyperosmotic stresses in the supralittoral pool habitat of the larvae are gradual but often extremely high. Hypo-osmotic stresses after rains may be extremely low because of the inability of larvae to swim into well developed freshwater lenses (rain water tends to float as a layer on top of the salt water of the pools). ATOLL RESEARCH BULLETIN No. 153 FISH DIVERSITY ON A CORAL REEF IN THE VIRGIN ISLANDS by Michael J. Risk Issued by THE SMITHSONIAN INSTITUTION WasiaenptyomuDir (C. se User A Weeienbery sis, 1972 ‘) Sasue (oe Laag we bye A in mi Die whe a 7 if hye : ; i: 3 e ne : t ; a A ; : i a , : ; 2 i i 7 ‘ i y hi : ' 7 Laat e X ; A, " . Arik et ant od “ waits dh \ PA Me it ha re ; , ‘ cm bev) Veen | h 7 & 5 y i ni i : . ' 7 = ‘ “a F Z ae A 4 a 4 ‘3 y nee tk , ? . : af, a , t Pa AL et | 4 ; 5 1 4 i= a me 4 g : - Bia cag var } i to hat 4 i v ‘ ‘ os : { ‘ Fi - a 7 t ‘ ii U i ue i = i 4 } As A , 3 i Lah ef = a $8 ; | F 1 i ; 2 Z — r at wl - a { e I : = : : i ak cat a HST ak it oe j « os , és t a : ¥ eee * = ¢ \» 2 { 1 : | ¥ : i eek pra Oh ing’ = i f ‘ ’ : n ‘ y 4 z ; = a, = ; ‘ F : F F “ ie at : oa i - : - FISH DIVERSITY ON A CORAL REEF IN THE VIRGIN ISLANDS by Michael J. Risk* INTRODUCTION Terrestrial ecologists have accumulated data relating bird species diversity to foliage height diversity (MacArthur and MacArthur, 1961; MacArthur, 1964; Recher, 1969); complexity of foliage pattern seems to be a better predictor of bird species diversity than is the tree species diversity. There is some indication that a similar relationship exists on coral reefs, between fish species diversity and substrate complexity. Talbot (1965, p. 453), in comparing fish collections from areas with either few or many species of corals present, states “‘It is probable that the greater variety of coral species...allows for a more varied fish population than the single species,’’ and, ‘‘It is probable that the more complex coral population in the mixed stand provides more ecological niches than are available in the single species stand.’’ Hiatt and Strasburg (1960, p. 118) found more species and individuals on ‘‘ramose’’ than on ‘““slomerate’’? coral heads: ‘‘Because glomerate coral heads are generally devoid of interstices in which smzll organisms can hide, such heads are visited only by fish species intent on browsing or grazing coral polyps.’’ Ina study on Conus, Kohn (1967, p. 257) found ‘‘Type III habitats are topographically the most complex...and these support the most diverse assem- blages of Conus.’’ The author attempted to measure the relationship between fish species diversity and the physical makeup of the coral substrate, following the work done on bird species diversity. PROCEDURE (a) Study area The area selected was a portion of a low-lying, shallow patch reef located in the eastern part of Greater Lameshur Bay, St. John, Virgin Islands. Substrate types within the area were live coral (Montastrea annularis, Millepora alcicornis, Porites furcata, and Agaricia agarvicites), dead coral, sponges (mostly /rcinza spp.) and carbonate sand. (b) Sample locations Two transects were laid out in the form of a ‘‘T’’, with the upright perpendicular to the shore, and the crosspiece further offshore and parallel to the shoreline. The ‘‘upright’? transect began about 50 meters from shore, in 4 meters »5f water, and extended 16 meters seaward, to a depth of 4.5 meters. The ‘‘crosspiece’’ transect was also 16 meters long, and 4.5 meters deep throughout its length. ‘Department of Biological Sciences, University of Southern California, Los Angeles, California 90007 (Manuscript received Feb. 1971--Eds.) (c) Determination of fish species diversity, Teles Only territorial fishes were considered in this study. Several days were spent diving and obServing in the area, in order to determine which species were territorial. When doubt existed about a particular fish, it was frightened into its home territory several times over the course of 20r3 days. Only fishes consistently returning to the same loca- tion were counted; appropriate members were three species of damelfish, a goby, a squirrelfish, and occasional filefishes, tangs, juvenile grunts and snappers. All caves, crevices, and holes were investigated, often using a light. Numbers of individuals of the selected species were then determined in each of the sixteen quadrats. Each area was counted at least twice a day for a period of three days. To reduce operator bias, previous values recorded on underwater slates were left on shore, and the quadrats counted beginning at different ends of the two transects (see Appendix A). Census values were in good agreement, and ranged from 4 to 7 species and from 7 to 21 individuals per square meter. Fish species diversity values were calculated using Shannon’s formula for H' (Shannon, 1948). Determination of substrate topographic complexity, ‘‘T’’ Actual surface area of a substrate as compared to its horizontal projection area was assumed to be a measure of spatial heterogeneity. The technique used was a modifi- cation of geological ‘‘point-counting,’’ wherein volumes of constituents are assumed proportional to their representation along line transects. In the present study, actual surface area was assumed to be a function of actual linear dimensions. The author used a fine-link chain, which could be draped and applied over a length of one meter. This was done eight times for each m2ter-square quadrat; in some cases four meters of chain were required to equal one ‘‘straight’’ meter. Actual surface area was assum2d proportional to the squares of these estimates of actual linear dimensions; therefore, the value of T for any one quadrat is the sum of the eight estim3ztes, squared. These values are record2d in Append:x B. Determination of substrate biological diversity ‘‘B’’ The substrate was aroditrarily divided into seven constituents: four species of living coral, dead coral, sponges, and carbonate sand. Different species of living coral were almost always separated by a band of dead coral; sponges grew only on this dead coral. Each constituent type was assumedto have areal coverage proportional to its intersect length along line transects. Again, in each of the sixteen quadrats, intersect length were determined along eight one-meter line transects. The result for each quadrat was therefore a total length of eight meters, divided among up to seven constituents. B was then calculated from these data, using Shannon’s formula for H'; values for each quadrat are recorded in Appendix C. Data processing Multiple regression analysis was performed on the data at the University of Southern California Computer Sciences Laboratory. Underwater data gathering The author is familiar with data-gathering by SCUBA, having conducted benthic surveys and studies in a variety of non-tropical habitats. The amount of time and energy required to measure only sixteen sample areas was staggering. 3 For this study, the author spent 80 hours underwater; including the preliminary study, more than 800 fishes were identified, and their territories noted; intersect values for substrate types were determined (to the nearest millimeter) along 144 meters of reef, and 1,028 meters of chain were stretched over the reef surface. To measure each meter- Square quadrat took at least five hours. RESULTS Significant relationships between the three main variables (Hg, T and B) were tested for. In addition, many subdivisions of the main variables, such as sand cover, total coral cover (live or dead), and total fishes, were investigated. Significant correlation exists only between fish species diversity, Hs, and substrate topographic complexity, T. Results of analysis of the three main variables are given in Table 1. Table 1. CORRELATION COEFFICIENT MATRIX Hy 1.000 *0.624 0.004 T 1.000 -0.161 B a i 1.000 *p < 0.05 Regression Equation: Hg = 1.06+ 0.004T+ 0.108B Multiple correlation coefficient 0.633. fish species diversity (‘‘nits?’) substrate topographic complexity substrate biological diversity He T B The regression coefficient for T is highly significant (p <0.01); that for B is not significant. Increase in the multiple correlation coefficient on addition of the B effects to those of T is only 0.009. DISC USSION Although the number of samples is quite small, there exists a striking positive correlation between fish species diversity and degree of substrate topographic complexity. This result is in accordance with predictions from terrestrial bird studies. It is interesting that there is no significant correlation between fish species diversity and the biological nature of the substrate; the choice of a territory would seem to be controlled by predation pressure, rather than by feeding preferences. There is also no significant positive correlation between total numbers of fishes and either substrate topographic complexity or substrate biological diversity; the relationship does not appear to be a simple one of ‘‘more holes, more fish.’? There apparently is occurring some selective partitioning of the habitat by individual fish species. ACKNOWLEDGMENTS Field work was carried out during the summer of 1969, when the author was a Teaching Assistant for the Organization for Tropical Studies course in Tropical Marine Biology. This was an invaluable opportunity to study and work on coral reefs. Drs. Gerald J. Bakus and Peter W. Frank have been generous of their time in reading and criticizing the manuscript. Any remaining errors or omissions are the author’s. LITERATURE CITED Hiatt, Robert W. and Donald W. Strasburg, 1960. Ecological relationships of the fish fauna on coral reefs of the Marshall Islands. Ecol. Monographs 30: 65-127. Kohn, Alan J. 1967. Environmental complexity and species diversity in the gastropod genus Conus on Indo-West Pacific reef platforms. Amer. Nat. 101: 251-260. MacArthur, R. and J.W. MacArthur. 1961. On bird species diversity. Ecology 42: 594-598. MacArthur, R. 1964. Environmental patterns affecting bird species diversity. Amer. Nat. 98: 387-397. Pianka, E.R. 1966. Latitudinal gradients in species diversity: a review of concepts. Amer. Nat. 100: 33-46. Randall, John E. 1963. An analysis of the fish populations of artificial and natural reefs in the Virgin Islands. Carib. J. Sci. 3: 31-47. Recher, Harry F. 1969. Bird species diversity and habitat diversity in Australia and North Africa. Amer. Nat. 103: 75-80. Shannon, F.W. 1948. The commonness, and rarity, of species. Ecology 29: 254-283. Talbot, F.H. 1965. A description of the coral structure of Tutia Reef (Tanganyika Territory, East Africa) and its fish fauna. Proc. Zool. Soc. London 145: 431-470. APPENDIX A POPULATIONS OF TERRITORIAL FISHES IN SAMPLE AREA ejjand smijaajdodAy Wayon} snyjuedeuoN sua|niaod sminyjuedy snjn) smjuad0j0H sminios uojnwaey (Ant) -ds snuefyny 1uosSdwoy} sidayouyeu5 SMNSAJYD UOpoYyJedsosa iA) Suoljiue|d snijuadewodny snosny Smjuadewodny 1 (Perpendicular to shore) Transect No. Total No. of Quadrat No. 4 3 2 4 3 1 (shallow) 8 (deep) Transect No. 2 (Parallel to shore — —— — — ——— —————————————————————————————————————— Total No. of fishes Quadrat No. oH Sc 8 (south) APPENDIX B VALUES OF T, SUBSTRATE TOPOGRAPHIC COMPLEXITY (m*) Transect No. 1 Transect No. 2 Quadrat No. T Quadrat No. ne 1 (shallow) TS) 1 (north) 36.5 2 95.7 2 73.6 3 28.2 3 Zilles 4 97.2 4 44,2 i) 85.8 i) 35.4 6 97.1 6 84.1 7 103.1 U Beets 8 (deep) Sie 8 (south) 60.0 APPENDIX C VALUE OF B, SUBSTRATE BIOLOGICAL DIVERSITY (‘‘nits’’) Transect No. 1 Transect No. 2 Quadrat No. B Quadrat No. B 1 (shallow) 1.34 1 (north) 1521 2 123 2 1.40 3 i ilY/ 3 1.06 4 He 4 1.34 5 1.34 5 1.43 6 0.98 6 1223 7 ibe lal 7 1.68 8 (deep) 117 8 (south) 1.08 ATOLL RESEARCH BULLETIN No. 154 RECOLONIZATION OF A POPULATION OF SUPRATIDAL FISHES AT ENIWETOK ATOLL, MARSHALL ISLANDS by William A. Bussing Issued by THE SMITHSONIAN INSTITUTION Washungtoni D.) Co5 Ui (S.A December 31, 1972 re ec A : p j : i : - * u% ‘ ie Rez, i . Viale, ay ita Wy ata. See ean re: alt ‘aap anal a ~ . _ - asttine 10 sr P teh Eh | | mena AARSERM, —~ - y= cee ae fe t ae 7 ig Wa ? = a ; : ‘ at ego) spin on : at 5 P R : 7 . : 7 ‘ . f ; is GF a: mn "3 a - if a a - : 2 , iss _ bi i : i ; I : . = : si RECOLONIZATION OF A POPULATION OF SUPRATIDAL FISHES AT ENIWETOK ATOLL, MARSHALL ISLANDS by William A. Bussing’ INTRODUCTION The field work and most of the identifications of fishes for this study were carried out during July 1965 at the Eniwetok Marine Biological Laboratory of the University of Hawaii while I was assisting Dr. Gerald J. Bakus ina study of the grazing effects of reef fishes. A small supratidal pool was noticed near a large quarry which we visited frequently during the fish-grazing study. Rotenone was applied to the pool early in July and yielded a large number of fishes and an especially large biomass of the moray eel Gymnothorax picta. It was decided at that time to repoison the pool at the end of July and compare the species composition, biomass and ecology of the fishes present in each collection in order to ascertain the rate and manner of recolonization of the fish population and something about its trophic structure. The phenomenon of plant succession has been frequently discussed and is rather well known. The accompaying animal succession has been less well studied, but seems to closely follow the same pattern as plant succession, e.g. certain pioneer species appear initially and are replaced by a sequential series of populations until a relatively stable climax community is established. Numerous studies have been done on succession of protozoa in culture jars (Bick, 1967a; 1967b), on the establishment of marine invertebrate communities on newly con- structed or specially prepared substrates (Reish, 1964, Poore, 1968) and on seral succession of fishes in ponds and marshes (Shelford, 1911a; 1911b). All these animal succession studies really involve plant succession as well. In both cases the flora and fauna are concurrently undergoing seral changes. The present study is unique in that only the fish population (and certain invertebrate species) were removed from the community. The rather sparse algal community was not apparently affected directly by the ichthyocide. DESCRIPTION OF HABITAT A general description of the Marshall Islands and their climate has been given by Hiatt and Strasburg (1960). The tidepool under consideration was located on the windward reef side of Eniwetok Island (11°21'N; 162°20'E) one of the chain of islands forming Eniwetok Atoll. The pool, although facing the open sea, was protected from excessive wave action by a shallow reef flat which extended about 200 meters offshore. With the aid of Coast and Geodetic Survey tide tables, the surface of the tidepool was estimated to lie about .95m above mean sea level. Daily high tides progressed from 1.1m on July 7 to 1.6m on July 29. Thus, the pool in question was flooded daily throughout the study period. The measured range of temperature variation in the pool during the study period was from 28° to 32°C. The lowest reading was taken in the morning on a rainy overcast day and probably ‘Departamento de Biologia, Universidad de Costa Rica, San José, Costa Rica. (Manuscript received March 1971--Eds.) 2 approximates the true low temperature for the pool during July. The highest temperature was recorded at midday on July 7, although this was surely surpassed on other days when measure- ments were not taken. The water in the pool was generally clear, but turned slightly brown in color after several hours of insolation. The study pool consisted of a 20m by 40m depression in old limestone pavement. The depth was a rather uniform 0.2m, but at one point reached 0.4m. The sides and much of the bottom consisted of coral rock and rubble; part of the bottom was covered by sand. Little life except a few fish were evident at first glance, but a considerable number of fishes of different species were collected by poisoning. Other conspicuous larger organisms revealed by turning over rocks and by the poison were xanthid crabs, hermit crabs (Diozgenidae) and octopods. The flora consisted principally of green and blue-green filamentous algae. MATERIALS AND METHODS All fishes were collected with dip nets and by hand after poisoning with Pro-Noxfish. Two collections were made: on July 7 and 22 days later on July 29. The behavior of fishes was observed intermittently before July 7 and during the period between the two collections. The majority of the fishes were identified at the Eniwetok Marine Biological Laboratory by means of the literature and by comparison with Laboratory reference specimens identified by Dr. John Randall. Specimens of each species were counted, the ranges in standard length taken and the moist formalin weight recorded. Later, the average weight was computed for each species in both collections (Table 1). The trophic level of each species of fish was determined using Hiatt and Strasburg (1960) where applicable. Direct examination of gut contents was made in other cases and on 15 soecimens of Gymnothorax picta. The percentages by weight of herbivorous, omnivorous and carnivorous fishes were calcu- lated for each collection (Table 2). Percentages for the first collection were determined twice: once including all fishes and the other excluding the moray eel Gymnothorax picta. This was done in order to better compare the trophic structure of each collection, because G. picta accounted for such a large percentage (44.8%) of the total biomz.ss of the first collection and was absent from the second collection. TROPHIC STRUCTURE OF TH FISH POPULATION Little has been said about the community organization of tidepools. I have been unable to find any reports on the trophic structure of the fishes of a tidepool community. Carnivorous or partially carnivorous fishes make up the greatest biomass in the first collection even if G, picta is excluded from the calculation. These top-heavy pyramids are, of course, based on fishes alone and do not represent the complete trophic structure of the pool. Likewise, such a predominance of secondary consumers is not surprising in a zone which receives daily a great accumulation of organic matter from the highly productive neritic zone. When biomass distribution of the first collection (excluding G, picta) is compared with the trophic structure of the second collection it is seen that the percentages of fishes repre- senting different trophic levels is roughly comparable (Table 2). Thus, the original trophic structure of the fish population was re-established within three weeks or sooner after the initial poisoning. RECOLONIZATION OF THE FISH POPULATION Thirty-three species belonging to 16 families were present in the two collections. Only five species, a snapper (Lutjanus gibbus), two tangs (Acanthurus elongatus and A, nigricans) and two puffers (Avothron meleagris and A. nigropunctatus) were present in the second collection, but not in the first. The species diversity was strikingly reduced in the second collection, although each family taken in Collection No. 1 was usually represented in the second collection by some specimens of the commonest species of that family. The total fish biomass was also greatly reduced even though the pool was inhabited by fishes the day after the poisoning, and free access to the pool was possible at least once a day (high tides of 1.1 to 1.6m) during the three weeks between poisonings. The average size of specimens in Collection No. 1 was 5.26gm (3.02 gm excluding G. picta) compared to 0.362gm in Collection No. 2. At 1300 hours on the day following the first poisoning, several small specimens of Abudefduf glaucus, Istiblennius edentulus and J. lineatus were seen behaving normally. The pool apparently had been washed clean of poison and these fishes had gained entry to the pool during the high of 1.1m at 0105 hours the night before. Two small morays (Gymnothorax bikiniensis) were seen lying on their backs in the middle of the pool at this time. Both were nearly dead and had probably entered the pool at night before it had been completely rinsed of poison. Juveniles of the commonest species showed the greatest ability to repopulate the pool. Small specimens of Abudefduf glaucus and Istiblennius edentulus accounted for the majority, by weight andnumber, of the fishes taken in the second collection. Adults of the above mentioned species are territorial to a varying degree andperhaps for this reason adults from surrounding waters did not readily reach the barren pool. This, however, does not explain the lack of the species which do not exhibit territorial behavior. In addition to the fishes, the only macroscopic organisms which appeared to be affected by rotenone were octopods and shrimps. Xanthid crabs and hermit crabs were not killed, and appeared to occur in equal numbers after the poisoning. The death of shrimps and possibly many smaller invertebrates couldexplain the decrease innumbers of omnivores and carnivores. Filamentous algae, the principal producer in the pool, seemed unaffected, but supported a much smaller herbivore population after the poisoning; acanthurids and mugilids decreased greatly in numbers and weight. Young of the snapper Lutjanus gibbus appear to have replaced the four species of Epine- phelus which were taken in the first collection. Thess species seem to have similar niches when young. They are carnivores and lurk around and under ledges searching for prey. The puffer Canthigaster solandvi was collected in equal numbers (10) in both collections, but the day before the second collection an estimated 200-309 specim0O= a a 5 ee \ 5 & Fonan | 2 ° 10" aN = ea Mesegon| 32-35 35' 40' 4s Map of Truk and Kuop showing collecting sites (map taken from Gressitt, 1954). SOME MARINE BENTHIC ALGAE FROM TRUK AND KUOP, CAROLINE ISLANDS’ by Roy T. Tsuda’ INTRODUCTION Truk (Fig. 1) consists of a lagoon, with about 40 volcanic islands and several reef patches, encircled by a barrier reef. The lagoon is from 45 to 65 kilometers in diameter and ranges in depth to 75 meters. Fringing reefs prevail along the coast of the larger lagoon islands. Wave-washed benches, which are present along the windward coast of non-lagoon volcanic islands, are absent. The barrier reef with its numerous passages is about 225 km in circum- ference and has approximately 50 small coral islets. Most of the barrier reef is awash during low tides. Kuop Atolllies only three km south of the southernmost terminus of the Truk barrier reef and for the purpose of algal distribution is merely a southern extension of Truk. Past records of marine benthic algae from Truk can be found in seven publications, while marine algae from Kuop Atoll have not been reported to date. The first paper to list marine algae from Truk was published by Okamura (1916) who cites 27 species. In a systematic com- pilation of 484 species of marine algae from the Pacific Ocean, Schmidt (1928) includes all the species reported by Okamura and four additional species from Truk. During the summer of 1960, Mr. Ernani Menez, presently with the Smithsonian Oceano- graphic Sorting Center, made a rather extensive collection of marine benthic algae from several islands in the Caroline Islands while a msmber of the Miami University-Collegiate Rebel Expedition to Micronesia. The collections were stored in the Department of Botany of the University of Hawaii until 1964 when Dr. Gavino Trono, Jr., presently with the University of the Philippines, undertook the task of identifying the specimens as part of his Ph.D. dissertation under the direction of Dr. Maxwell S. Doty. In the same year, Dr. George J. Hollenberg of the University of Redlands arrived at the University of Hawaii to resume his studies on the red algal genera Polysiphonia and Herposiphonia, and incorporated those specimens that he had examined from the Caroline Islands into his papers (Hollenberg, 1968a, 1968b, 1968c). These publications include four species of Polysiphonia and four species of Herposiphonia from Truk. The results of Dr. Trono’s study on the marine benthic algae from the Caroline Islands were published in two parts. Part I (Trono, 1968) lists 42 species of blue-green and green algae from Truk, while Part II (Trono, 1969) includes 48 species of brown and red algae from Truk and also incorporates the eight species reported by Dr. Hollenberg. Altogether, in the above mentioned papers, 102 species of marine benthic algae have been reported from Truk. Recently, opportunities to make additional collections of marine algae on Truk and Kuop have arisen in conjunction with the ‘‘crown-of-thorns’”’? starfish, Acanthaster planci, studies in Micronesia. Mr. Richard H. Randall of the University of Guam took the initiative to make a ‘Contribution No. 12, Micronesian Institute for the Natural Sciences, University of Guam. * The Marine Laboratory, University of Guam, Agana, Guam 96910. (Manuscript received Nov. 1970--Eds.) 2 moderate collection of marine algae from Truk and Kuop during the Westinghouse Acqanthaster survey in the summer of 1969. The following summer, the author was able to make additional collections of algae when he participated as member of the University of Guam’s Acanthaster monitoring team. The purpose of this paper is to provide a checklist of the species collected by Mr. Randall and the author. Seven additional specimens collected by Mr. Asterio Takesy, formerly a student at the University of Guam, are also incorporated in this paper. ACKNOWLEDGEMENT The author is grateful to Mr. Wyman X. Zachary, Director of Resources and Development, and to Mr. Peter T. Wilson, Fisheries Management Biologist, both of the Trust Territory of the Pacific Islands, for their financial and logistic support during the author’s stay on Truk. The author acknowledges the help of Dr. Robert S. Jones, Director of the University of Guam’s Marine Laboratory, Mr. M. Rodney Struck, staff member of the University of Guam’s Marine Laboratory, and Mr. Milton McDonald, Starfish Control Specialist for Trust Territory, who served as diving companions and at times contributed specimens to the overall collections. STATION DESCRIPTIONS The following is a listing of the 35 algal collecting stations which can be located on Fig. 1. The stations for Truk and Kuop are numbered consecutively but are separated merely for con- venience. Information for each station includes 1) the specific island from which the collection was made, 2)a brief description of the habitat, i.e., zone and depth, 3) name of collector, 4) date of collection and 5) the collection number(s) assigned to the algae at each station. Truk (7°28'N, 151952'E) STATION 1: Moen I., Mwaan Village, reef flat, 1 m deep, legit A. Takesy, May, 1969 (RT 2856-2861). 2: Osakura I., lagoon slope, 7m deep on dead coral rubble, lezit R.H. Randall, July 6, 1969 (RHR 359). 3: Moen I., east end, lagoon slope, 3-7 m deep on dead coral rubble, legit Randall, July 7, 1969 (RHR 369). 4: Fanuet I., reef margin, nearly awash, legit Randall, July 10, 1969 (RHR 371). 5: Fanuet I., lagoon slope, 3m deep attached to dead Acropora, legit Randall, July 10, 1969 (RHR 372). 6: Herit I., reef flat and lagoon slope, .3-3 m deep, legit Randall, July 18, 1969 (RHR 368). 7: Moen I., Goverament Dock outside of Fisheries Office, .5 m deep on cement wall, legit R.T. Tsuda, June 13, 1970 (RT 3367-3368). 8: Yanagi I., west lagoon slope, 10-17 m deep on dead Acropora and coral rubble, legit Tsuda, June 13, 1970 (RT 3370-3387). 9: Dublon I., north reef flat, .3 m deep on dead coral and sand, lezit Tsuda, June 13, 1970 (RT 3333.-3492). STATION 10: 11: 12: Leg 14: 5s 16: 17: 18: 19: 20: Qe 22: 23: 24: 745)8 26: 27: 28: 29: 3 Dublon I., north lagoon slope, 3-7 m deep in live Acropora bed, legit Tsuda, June 13, 1970 (RT 3405-3408). Dublon I., north lagoon floor, 17m deep on coral rubble and sand, legit R.S. Jones, June 13, 1970 (RT 3413-3415). Reef patch, .5km SW of Dublon I., 2m deep, legit Tsuda, June 14, 1970 (RT 3416-3417). Sunken Japanese Ship, ca. 4km SE of Etten I., 20-30m deep, legit Tsuda, June 14, 1970 (RT 3419-3434). Pis I., seaward side, intertidal zone, legit Tsuda, June 15, 1970 (RT 3437-3441). Pis I., west of channel, seaward reef margin (exposed) and seaward reef front (1m deep), legit Tsuda, June 16, 1970 (RT 3442-3449). Pis I., east of channel, seaward slope, 15-35m deep, legit Tsuda, June 16, 1970 (RT 3451-3461). North Pass, channel slope, 10-30m deep, legit Tsuda, June 17, 1970 (RT 3462-3463). Pis I., west of channel, seaward slope, 45m deep on dead coral, legit Tsuda, June 17, 1970 (RT 3465-3466). Pis I., boat harbor, 1m deep near shore, legit Tsuda, June 17, 1970 (RT 3467). Liball I., reef flat, .3-1m deep, legit Tsuda, June 17, 1970 (RT 3469-3487). Moen I., Wiicap, mangrove area, reef flat and reef front., .3-2m deep, legit Tsuda, June 18, 1970 (RT 3488-3505). Salat I., east coast, lagoon slope, 5m deep, legit Tsuda, June 19, 1970 (RT 3508-3514). Drift, north of Moen I., legit Tsuda, June 19, 1970 (RT 3515). Etten I., reef flat, .5-1m deep, legit Tsuda, June 20, 1970 (RT 3518-3519). Sunken Japanese Destroyer, east coast of Fefan I., 10-30m deep, legit Tsuda, June 20, 1970 (RT 3520-3535). Sunken Japanese Tanker, northwest coast of Dublon I., 10-30m deep, legit Tsuda, June 20, 1970 (RT 3536-3546). Scheiben I., lagoon slope, 7m deep, legit Tsuda, June 23, 1970 (RT 3558-3561). Kuop (6°59'N, 151959'E) Lagoon reef, 1-2 m deep on sandand coral, legit Randall, July 12, 1969 (RHR 360). Southwest side of atoll, seaward reef front, 7m deep, legit Randall, July 13, 1969 (RHR 364). 4 STATION 30: Southwest side of atoll, seaward reef margin, awash, legit Randall, July 13, 1969 (RHR 365a). 31: Southwest side of atoll, seaward reeffront, 7m <\sep, legit Randall, July 13, 1969 (RHR 365b-d). 32: Northwest side of atoll, seaward submarine cliff, 16-40 m deep, legit Randall, July 13, 1969 (RHR 366). 33: Northwest side of atoll, seaward submarine cliff, 3-16m deep, legit Randall, July 13, 1969 (RHR 367). 34: Northwest side of atoll, seaward reef flat and margin, exposed to .3m deep, legit Tsuda, June 21, 1970 (RT 3547-3550). 35: Northwest side of atoll, seaward submarine cliff, 10-30m (teep, legit Tsuda, June 21, 1970 (RT 3552-3557). SPECIES LISTING The following list includes specimens collected by A. Takesy (May 1969), R.H. Randall (July 1969) and the author (June 1970). All specimens are catalogued in the University of Guam Herbarium. Species preceded by an asterisk represent new records from Truk. DIVISION CYANOPHYTA (blue-green algae) Order Oscillatoriales *Calothrix confervicola (Roth) Ag. TRUK: Sta. 15 (RT 3443). *Hormothamnion solutum Bornet & Flahault TRUK: Sta. 21 (RT 3488). Microcoleus lyngbyaceus (Kutz.) Crouan TRUK: Sta. 6 (RHR 368e), Sta. 8 (RT 3370, RT 3382), Sta. 9 (RT 3399), Sta. 13 (RT 3430), Sta. 22 (RT 3510), Sta. 25 (RT 3523). The specimens vary from 13 to 40 in diameter. Schizothrix calcicola Gomont TRUK: Sta. 8) (RT 3371): The specimen appearsasa gelatinous ball with individual filaments 1 to 1.5 a in diameter. *Schizothrix mexicana Gomont TRUK: Sta. 4 (RHR 371a), Sta. 8 (RT 3383), Sta. 15 (RT 3445), Sta. 22 (RT 3511). *Spirulina subsalsa Gomont TRUK: Sta. 9 (RT 3399, intermixed with Microcoleus lyngbyaceus). DIVISION CHLOROPHYTA (green algae) Order Ulotrichales *Enteromorpha clathvata (Roth) Ag. TRUK: Sta 14 (RT 3441). The specimen consists of an immature branched thallus about mn ‘igh. Order Cladophorales Rhizoclonium samoense_ Setchell TRUK: Sta. 14 (RT 3439). Order Siphonales Bryopsis pennata Lamx. TRUK: Sta. 20 (RT 3470), Sta. 25 (RT 3528). *Caulerpa ambigua Okamura TRUK: Sta. 13 (RT 3433b). Caulerpa brachypus Harvey TRUK: Sta. 8 (RT 3372), Sta. 13 (RT 3426), Sta. 25 (RT 3529), Sta. 26 (RT 3541). The latter three specimens collected from sunken ships possess thin delicate elongated blades. Caulerpa cupressoides (West) C. Ag. TRUK: Sta. 9 (RT 3396). *Caulerpa filicoides Yamada TRUK: Sta. 13 (RT 3425), Sta. 26 (RT 3546). This species was one of the most dominant algae found on the sunken ships. Caulerpa lentillifera J. Ag. TRUK: Sta. 11 (RT 3413), Sta. 13 (RT 3429), Sta. 26 (RT 3544). Caulerpa racemosa (Forskal) J. Ag. TRUK: Sta. 6 (RHR 368b), Sta. 8 (RT 3373), Sta. 9 (RT 3392), Sta. 18 (RT 3465), Sta. 20 (RT 3473), Sta. 21 (RT 3492), Sta. 25 (RT 3525, RT 3533). KUOP: Sta. 35 (RT 3555). Caulerpa serrulata (Forskal) J. Ag. TRUK: Sta. 19 (RT 3467), Sta. 22 (RT 3513). KUOP: Sta. 35 (RT 3554). Caulerpa sertularioides (Gm:21.) Howe TRUK: Sta. 9 (RT 3390), Sta. 21 (RT 3495), Sta. 26 (RT 3540). Caulerpa taxifolia (Vahl) C. Ag. TRUK: Sta. 20 (RT 3481). Caulerpa urvilliana Montagne TRUK: Sta. 20 (RT 3476). KUOP: Sta. 28 (RHR 360b). *Caulerpa verticillata J. Ag. TRUK: Sta. 21 (RT 3496). Chlorodesmis fastigiata (C. Ag.) Ducker TRUK: Sta. 16 (RT 3460). KUOP: Sta. 34 (RT 3549). Chlorodesmis hildebrandtii A, & E.S. Gepp TRUK: Sta. 16 (RT 3491, epiphytic on Halimeda). 6 Codium edule Silva TRUK: Sta. 15 (RT 3446). This specimen consists of only a fragment about 3cm long. *Halimeda copiosa Goreau & Graham TRUK: Sta. 6 (RHR 368c), Sta. 10 (RT 3406), Sta. 12 (RT 3416), Sta. 13 (RT 3422, RT 3427b), Sta. 16 (RT 3459), Sta. 25 (RT 3532), Sta. 26 (RT 3545). KUOP: Sta 31 (RHR 365b), Sta. 32 (RHR 366a), Sta. 35 (RT 3553). Halimeda cylindvacea Decaisne TRUK: Sta 21 (RT 3505), Sta. 22 (RT 3508), Sta 24 (RT 3519), Sta 27 (RT 3560). Halimeda discoidea Decaisne TRUK: Sta. 8 (RT 3374), Sta. 9 (RT 3391), Sta. 13 (RT 3427), Sta. 16 (RT 3458), Sta. 20 (RT 3480), Sta. 25 (RT 3521), Sta. 26 (RT 3537). KUOP: Sta. 29 (RHR 364c) *Halimeda gigas Taylor TRUK: Sta. 8 (RT 3375), Sta. 11 (RT 3415), Sta. 13 (RT 3421), Sta. 21 (RT 3493), Sta. 26 (RT 3543). *Halimeda gracilis Harvey TRUK: Sta. 15 (RT 3447), Sta. 17 (RT 3462). KUOP: Sta. 29 (RHR 364a). Halimeda incrassata (Ellis) Lamx. TRUK: Sta. 7 (RT 3667), Sta. 8 (RT 3377), Sta. 10 (RT 3405), Sta. 16 (RT 3456), Sta. 21 (RT 3501), Sta. 26 (RT 3536), Sta. 27 (RT 3559). KUOP: Sta 29 (RHR 364b), Sta. 31 (RHR 365c). Halimeda macroloba Decaisne TRUK: Sta. 9 (RT 3388), Sta. 21 (RT 3494). Halimeda macrophysa Askenasy TRUK: Sta. 10 (RT 3408), Sta. 16 (RT 3457), Sta. 25 (RT 3529), Sta. 27 (RT 3559). *Halimeda micronesica Yamara TRIK: Sta. 20 ‘RT 3479). KUOP: Sta. 29 (RHR 364d). Halimeda opuntia (L.) Lamy. TRUK: Sta. 4 (RHR 371b), Sta. 8 (RT 3384), Sta. 9 (RT 3397), Sta. 20 (RT 3478). *Halimeda simulans Howe TRUK: Sta. 9 (RT 3389), Sta. 21 (RT 3497). *Halimeda stuposa Taylor TRUK: Sta. 2) (RT 3482). KUOP: Sta. 228 (RHR 36)a). Rhipilia orientalis A. & E.S. Gapp TRUK: Sza. 8 (RT 333), Sta. 25 (RT 3523), Sta. 26 (RT 3542). *Rhipilia tomentosa Kutz. TRUK: Sta. 16 (RT 3455). Specimen heretofore known only from the Atlantic Ocean repressnts the first record from the Pacific region. Tydemania expeditionis Weber van Bosse TRUK: Sta. 8 (RT 3386), Sta. 13 (RT 3420, RT 3423), Sta. 25 (RT 3524). Both the flabellate andi the glomerulous forms are represented in the collections. The flabellate form was dominant on the sunken Japanese ships. Udotea argentea Zanardini TRUK: Sta. 8 (RT 3379), Sta. 25 (RT 3522). KUOP: Sta. 23 (RH 360d), Sta. 32 (RHR 366d). *Udotea geppii Yamada TRUK: Sta. 13 (RT 3419), Sta. 25 (RT 3530). KUOP: Sta. 28 (RHR 360e), Sta. 35 (RT 3552). *Udotea palmetta Decaisne TRUK: Sta. 20 (RT 3475). KUOP: Sta. 28 (RHR 360f). Order Siphonocladales Boergesenia forbestt (Harv.) Feldmann TRUK: Sta. 22 (RT 3509). Cladophoropsis membranacea (Ag.) Boerg. TRUK: Sta. 22 (RT 3512). *Dictyosphaeria cavernosa (Forskal) Boerg. TRUK: Sta. 13 (RT 3424), Sta. 20 (RT 3484). KUOP: Sta. 34 (RT 3550). Microdictyon okamuyrai Setchell TRUK: Sta. 2 (RAR 359c), Sta. 20 (RT 3477). KUOP: Sta. 28 (RHR 360c), Sta. 33 (RAR 367b), Sta 35 (RT 3556). Valonia ventricosa J. Ag. TRUK: Sta. 8 (RT 3381). Order Dasycladales *Acetabularia clavata Yamada TRUK: Sta. 15 (RT 3444p). The collection is represented by a single specimen, 1 mn high. Neomeris vanbosseae Howe TRUK: Sta. 14 (RT 3440), Sta. 20 (RT 3472), Sta. 21 (RT 3503). DIVISION PHAEOPHYTA (Brown algae) Order Dictyotales Dictyopteris repens (Okamura) Boerg. TRUK: Sta. 13 (RT 3431), Sta. 15 (RT 3444a), Sta. 16 (RT 3451). *Dictyota apiculata J. Ag. TRUK: Sta. 16 (RT 3452), Sta. 21 (RT 3500b). 8 *Dictyota bartayresii Lams. TRUK: Sta. 5 (RHR 372b), Sta. 8 (RT 3385), Sta. 11 (RT 3414), Sta. 13 (RT 3434), Sta. 24 (RT 3518), Sta. 25 (RT 3539), Sta. 27 (RT 3561). *Dictyota patens J. Ag. TRUK: Sta. 4 (RHR 371c), Sta. 21 (RT 3500a). Lobophora variegata (Lamx.) Womersley TRUK: Sta. 5 (RHR 372c), Sta. 8 (RT 3387), Sta 13 (RT 3428), Sta. 15 (RT 3449), Sta. 16 (RT 3454), Sta. 18 (RT 3466), Sta. 20 (RT 3471, RT 3483). Padina boryana Thivy TRUK: Sta. 7 (RT 3368), Sta. 9 (RT 3398), Sta. 21 (RT 3502). *Padina sp. TRUK: Sta. 8 (RT 3378), Sta. 13 (RT 3432), Sta. 20 (RT 3487, sterile), Sta. 25 (RT 3531). *Zonarvia hawaiiensis Doty & Newhouse TRUK: Sta. 8 (RT 3376). The specimen represents the first collection made outside of Hawaii where it was original- ly described as a new species. Order Dictyosiphonales Hydroclathrus clathratus (Bory) Howe TRUK: Sta. 9 (RT 3400), Sta. 21 (RT 3499). Order Fucales *Sargassum duplicatum J. Ag. TRUK: Sta. 23 (RT 3515). *Sargassum sp. TRUK: Sta. 6 (RHR 368c). This specimen resembles S, duplicatum except for a cylindrical stem. Turbinaria ornata (Turn) J. Ag. TRUK: Sta. 9 (RT 3393), Sta. 14 (RT 3438), Sta. 15 (RT 3448). KUOP: Sta. 30 (RHR 365a). DIVISION RHODOPHYTA (red algae) Order Nemalionales Actinotrichia fragilis (Forskal) Boerg. TRUK: Sta. 2 (RHR 359f), Sta. 21 (RT 3489). *Galaxaura fasciculata Kjellman TRUK: Sta. 12 (RT 3417), Sta. 25 (RT 3535). *Galaxaura fastigiata Decaisne TRUK Sta. 25: (RT 3527). *Galaxaura oblongata (Ellis & Solander) Lamx. TRUK: Sta. 2 (RHR 359e). Order Cryptonemiales Amphiroa foliacea Lamx. TRUK: Sta. 2 (RHR 359d), Sta. 3 (RHR 369). Fosliella farinosa (Lamx.) Howe KUOP: Sta. 32 (RHR 366g, epiphytic on Lobophora variegata). *Halymenia sp. TRUK: Sta. 2 (RHR 359a). *Jania capillacea Harvey TRUK: Sta. 15 (RT 3443, epiphytic on Turbinaria ornata). *Metagoniolithon sp. TRUK: Sta. 2 (RHR 359b), Sta. 21 (RT 3504), Sta. 25 (RT 3534). Further studies are being made on this species which represents the first collection of this genus outside of Australia. *Peyssonelia rubra (Grev.) J. Ag. TRUK: Sta. 2 (RHR 359a), Sta. 16 (RT 3461). *Porolithon craspedium (Foslie) Foslie KUOP. Sta. 31 (RHR 365d). Order Gigartinales Cevatodictyon spongiosum Zanardini TRUK: Sta. 14 (RT 3437). Gelidiopsis intricata (Ag.) Vickers TRUK: Sta. 9 (RT 3401). Hypnea espevi Bory KUOP: Sta. 32 (RHR 366e). Order Rhodymeniales Champia compressa Harv. TRUK: Sta. 13 (RT 3433a, epiphytic on Halimeda). Order Ceramiales *Acanthophora spicifera (Vahl) Boerg. KUOP: Sta. 32 (RHR 366c), Sta. 34 (RT 3548a). *Crouania minutissima Yamada TRUK: Sta. 20 (RT 3469). *Dasyopsis pilosa Weber van Bosse KUOP: Sta. 32 (RHR 366f). Herposiphonia delicatula Hollenberg KUOP: Sta. 34 (RT 3547). *Hypoglossum attenuatum Gardner TRUK: Sta. 26 (RT 3538). 10 Laurencia majuscula (Harv.) Lucas TRUK: Sta. 20 (RT 3485). *Laurencia parvipapillata Tseng KUOP: Sta. 34 (RT 3548b). This species is represented by a single fragment, 1 cm long. *Polysiphonia scopulorum Harvey TRUK: Sta. 20 (RT 3474). KUOP: Sta. 28 (RHR 360g). Tolypiocladia glomerulata (Ag.) Schmitz & Hauptfleisch TRUK: Sta. 9 (RT 3395, epiphytic on Halimeda discoidea). *Vanvoorstia sp. TRUK: Sta. 16 (RT 3453), Sta. 17 (RT 3463). Preliminary examination seems to indicate that the specimens may represent a new species. DISC USSION Eighty-five species of marine benthic algae are reported inthis paper. Forty-one species or 48 percent of the listed species represent new records for the Truk area. These species consist of 4 Cyanophyta, 15 Chlorophyta, 7 Phaeophyta and 15 Rhodophyta. With the addition of these species, a total of 143 species are presently known from Truk. The author feels that this total represents approxim<-tely 75 to 80 percent of the algae actually present on Truk. It is 2.f interest to note that of the 41 species reported as new records, 13 of them were found exclusively in water deeper than 7 meters. The continual use of SCUBA will no doubt increase the number of species known from Pacific islands. One additional species is present in the collection which is not included in the present listing. This green alga seemsto representa new genus and will be reported on at a later date. LITERATURE CITED Gressitt, J.L. 1954. Insects of Micronesia, Introduction. Vol. I. Honolulu, Bernice P. Bishop Museum, 1-257. mM Hollenbers. G.f. 1968a. An account of the species of Polysiphonia of the central and western tropical Pacific Ocean. I. Oligostphonia Pac. Sci. 22(1): 56-98, 43 figs. Seo Seuoooe. 1968b. An account of the species of Polysiphonia in the central and western tropical Pacific Ocean. II Polysiphonia. Pac. Sci. 22(2): 198-207, 3 figs. ---------- 1968c. An account of the species of the red alga Herposiphonia occurring in the central and western tropical Pacific Ocean. Pac. Sci. 22(4): 536-559, 25 figs. Okamura, K. 1916. List of marine algae collected in Caroline and Mariana Islands, 1915. Bot. Mag. (Tokyo) 30:1-14, 9 figs., 1 pl. Schmidt, V.O.C. 1928. Verzeichnis der Meeresalgen von Neu-Guinea and dem westlichen Oceanien. Hedwigia 68:19-86. Trono, G.C. Jr. 1958. The marine benthic algae of the -Caroline Islands, I. Introduction, Chlorophyta, and Cyanophyta. Micronesica 4(2): 1-206, 19 pls. See ete 1969. The marine benthic algae of the Caroline Islands, Il. Phaeophyta and Rhodophyta. Micronesica 5(1):25-119, 11 pls. ATOLL RESEARCH BULLETIN No. 156 | ADDITIONAL RECORDS OF MARINE BENTHIC ALGAE FROM YAP, WESTERN CAROLINE ISLANDS by Roy T. Tsuda and Mary S. Belk Issued by THE SMITHSONIAN INSTITUTION Wesioniaacons We Cos We SouNs December 31, 1972 489-409 O- 73 - 4 ADDITIONAL RECORDS OF MARINE BENTHIC ALGAE FROM YAP, WESTERN CAROLINE ISLANDS* by Roy T. Tsuda’ and Mary S. Belk* INTRODUCTION Yap (9925'N. Lat., 138°02'E. Long.) consists of four closely grouped islands separated by narrow Shallow channels. Encircling these islands isa barrier reef with seven major channels that provide water circulation to the lagoonfrom the outside. The air temperature ranges from 25°C in January to 28°C in September, and the rainfall ranges from 1.14 inches (2.9cm) in January to 36.86 inches (93.6cm) in December. Gressitt (1954) provides details on the terrestrial topography. Past records of marine benthic algae from Yap can be found in nine papers (Reinbold, 1901; Okamura, 1904, 1916; Schmidt, 1928; Tokida, 1939; Hollenberg, 1968a, 1968b; Trono, 1968, 1969) where a total of 83 species have been listed. It is of interest to note that none of these authors have personally made any algal collections on Yap. The purpose of this paper istopresenta listing of the unreported species of marine benthic algae from Yap that the senior author collected while on an Acanthaster planci (crown-of- thorns starfish) survey trip during November 24 to December 1, 1970. Twenty-three species are listed in this paper, thereby increasing the total number of species known from this island group by 28 percent. A total of 106 species (8 Cyanophyta, 41 Chlorophyta, 15 Phaeophyta, and 42 Rhodophyta) are now known from Yap. It should be pointed out that these collections are reported here strictly for phytogeographic purpose. It is only through the continual col- lecting and reporting of additional species that a flora of a given area may one day be said to be ‘‘well-known’’ floristically. All specimens are cataloged inthe University of Guam Herbarium. ACKNOWLEDGE MIINT The senior author is grateful to Mr. Peter T. Wilson, Chief of Marine Resources, Trust Territory of the Pacific Istands, for providing the opportua‘ty for him to visit Yap. The senior author acknowledges the help of Dr. Daniel P. Cheney, Dr. James A. Marsh and Mr. M. Rodney Struck of the University of Guam’s Marine Laboratory, who served as diving companions and at times contributed specimens to the overall collection. The authors thank Dr. Yuzuru Saito, Hokkaido University, for examining the two species of Laurencia. ‘Contribution No. 15, The Marine Laboratory, University of Guam. * The Marine Laboratory, University of Guam, Agana, Guam 96910. (Manuscript received April 1971--Eds.) STATION DESCRiPTIONS STATION 1: Seaward terrace outside barrier reef, 3m deep, 4km SW of Tomil Harbor entrance, XI-24-70. 2: Inner lagoon reef, Enhalus bed, 1m deep, Nimpol, XI-25-70. 3: Inner lagoon reef, Enhalus bed, 1-2m deep, Pelak, XI-25-70. 4: Outer lagoon reef, 1-2m deep, Pelak, XI-25-70. 5: Outer lagoon reef, dead Acropora bed, 2m deep, Tomil Harbor, XI-27-70. 6: Outer lagoon reef, 2m deep, SW of Tomil Channel, XI-28-70. 7: Inner lagoon reef, 1-2m deep, Garin I., XI-28-70. 8: Inner lagoon reef, Enhalus bed, 1-2m deep, between Map I. and Ruming I., XI-29-70. 9: Lagoon channel, deep hole, 10-13m deep, off USCG Beach, 2.5km S of Gofenu Entrance, XI-29-70. 10: Outer lagoon reef, 1-2m deep, Tomil Harbor, XI-30-70. 11: Intertidal zone at shoreline, NE of Pelak Entrance, XII-1-70. SPECIES LISTING DIVISION CYANOPHYTA (blue-green algae) Order Oscillatoriales Calothrix confervicola (Roth) Ag. Sta. 10 (RT 4048, mixed with Microcoleus lyngbyaceus). Calothrix pilosa Bornet & Flahault Sta. 2 (RT 3969, mixed with Microcoleus lyngbyaceus and sand). Hormothamnion solutum Bornet & Flahault Sta. 2 (RT 3961b, epiphytic on Avvainvillea erecta), Sta. 10 (RT 4048, mixed with Microcoleus lyngbyaceus) Schizothrix mexicana Gomont Sta. 7 (RT 4003, tufts on coral), Sta. 9 (RT 4025, loose filaments on coral). Spirulina subsalsa Gomont Sta. 10 (RT 4048, mixed with Microcoleus lyngbyaceus). DIVISION CHLOROPHYTA (green algae) Order Ulotrichales Enteromorpha kylinii Bliding Sta. 5 (RT 3989) Order Cladophorales Chaetomorpha crassa (Ag.) Kutz. Sta. 3 (RT 3978) Order Siphonales Avrainvillea lacevata Harvey Sta. 10 (RT 4042) Bryopsis pennata Lamx. Sta. 1 (RT 3955b, epiphytic on Laurencia sp.) Caulerpa elongata Weber van Bosse Sta. 9 (RT 4021), Sta. 10 (RT 4041). Halimeda gigas Taylor Sta. 5 (RT 3984), Sta. 7 (RT 4005c), Sta. 8 (RT 4018c), Sta. 9 (RT 4022b) Halimeda simulans Howe Sta. 2 (RT 3968), Sta. 7 (RT 4005b). Order Siphonocladales Dictyosphaeria versluysii Weber van Bosse Sta. 2 (RT 3970) Order Dasycladales Neomeris vanbosseae Howe Sta. 8 (RT 4014), Sta. 11 (RT 4049) DIVISION PHAEOPHYTA (brown algae) Order Ectocarpales Ectocarpus indicus Sonder Sta. 7 (RT 4006c, epiphytic on Padina minor). Order Dictyotales Dictyota apiculata J. Ag. Sta. 8 (RT 4013p). Dictyota cervicornis Kutz. Sta. 2 (RT 3959), Sta. 7 (RT 4008), Sta. 8 (RT 4013a). Padina minor Yamada Sta. 8 RT 4017, Sta. 11 - RT 4050. DIVISION RHODOFPHYTA (red algae) Order Cryptonemiales Metagoniolithon sp. Sta. 2 (RT 3967), Sta. 6 (RT 3996). These specimens represent the second collection of this genus from Micronesia and are presently being examined by Dr. H. William Johansen of Clark University, Massachusetts. Order Gelidiales Gelidium pulchellum (Turn.) Kutz. Sta. 10 (RT 4039). Order Ceramiales Laurencia majuscula (Harvey) Lucas Sta. 9 (RT 4026, identified by Dr. Y. Saito of Hokkaido University). Laurencia sp. Sta. 1 (RT 3955a), Sta. 3 (RT 3979), Sta 4 (RT 3982). According to Dr. Saito, these specimens are similar externally to L. papillosa but lack the characteristic palisade cortical cells. This species will be described as ‘‘new’’ in a future paper after additional fertile specimens are examined. Polysiphonia scopulorum Harvey Sta. 9 (RT 4029). DISCUSSION The most conspicuous marine plant in the predominantly sandy inner lagoon reef is Enhalus acoroides, which forms a band of about 50 meters or more around the entire coast of the four islands. Various algal species, e.g., Dictyota cervicornis, Halimeda opuntia, Hali- meda macroloba and Caulerpa racemosa, are found interspersed among the Enhalus. Although the navigable lagoon channel which surrounds the inner reef possesses a sandy substratum with almost no algae, several larger depressions, 13 meters deep, are found east of Gagil and west of Rumung Island. The dominant algae along the coral rim of these depres- sions are Caulerpa elongata, Halimeda gigas, Tydemania expeditionis and Rhipilia orientalis. The outer lagoon reef consists of patches of live corals interspersed with sand pockets and coral rubble. Microcoleus lyngbyaceus and Caulerpaurvilliana are the most conspicuous algae inhabiting the sand pockets, while Polysiphonia scopulorum and Gelidiopsis intricata cover the branches of dead Acropora. Valonia ventricosa and Actinotrichia fragilis are abundant among the coral rubble in this zone. However, algae are almost non-existent in the live coral areas. Likewise, the reef margin, seaward reef front and the seaward submarine terrace, possess very few algae except for crustose corallines. The marine flora on the reefs of Yap is very limited in terms of standing crop. This situation is not at all unusual since other islands which possess live corals reveal this same condition. Although a direct correlation seems to exist between the greater diversity of fishes and a live reef, a negative correlation is present between a smaller algal number and a live reef situation. An obvious reason for a greater algal flora on dead reefs is the larger settling surface available to the algal spores and zygotes. LITERATURE CITED Gressitt, J.L. 1954. Insects of Micronesia I, Introduction. B.P. Bishop Museum, Honolulu. viii, 257 pp. Hollenberg, G.J. 1968a. An account of the species of Polysiphonia of the central and western tropical Pacific Ocean. I. Oligosiphonia. Pac. Sci. 22(1): 56-98, 43 figs. Sodsoodoaens, 1968b. An account of the species of the red alga Herposiphonia occurring in the central and western tropical Pacific Ocean. Pac. Sci. 22(4): 536-559, 25 figs. Okamura, K. 1904. List of marine algae collected in Caroline Island and Australia. Bot. Mag. Tokyo 18: 77-96. —SoSeoesoag 1916. List of marine algae collected in Caroline and Mariana Islands, 1915. Bot. Mag. Tokyo 30: 1-14, 9 figs., 1 pl. Reinbold, T. 1901. Meeresalgen von den Karolinen (meist von Yap) welche Prof. Dr. Volkens gesam melt hat. Hedwigia 40: 350-351. Schmidt, V.O.C. 1928. Verzeichnis der Meeresalgen von Neu-Guinea und dem westlichen Oceanien. Hedwigia 68: 19-86. Tokida, J. 1939. A list of marine algae of Micronesia. Kagaku Nanyo 2(1): 16-26. Trono, G.C. Jr. 1968. The marine benthic algae of the Caroline Islands, I. Introduction, Chlorophyta, and Cyanophyta. Micronesica 4(2): 1-206, 19 pls. So qoouonoss 1969. The marine benthic algae of the Caroline Islands, Il. Phaeophyta and Rhodophyta. Micronesica 5(1): 25-119, 11 pls. nnavend | in} i oY ail) Sy er a a) tad ek FRR: ine Int ast ila ctalaaradl ~ ani eaet: yee aesoyuey se f * - PA : | 4 =i . eS 6 has nag’ ch a te a Wit ee So hs ‘pa] om ia epi 2 mt ety va See | 7 fe i qe ty ees Nay ha a , at ME Dod alle? oe ches sto io one S eeul ; es = ; > td gee 2 si ¢ a, o 19 « oer a7 i 4 A : ; [ j \ i a PTA pity tude no hats dive yith ce gp : - woe | a htt oat rf - 4 ij Be ee OEMs ooh il deretrckes Ate dene ei Och e ones ed hi tail 4 be mtd + pgs tain tultwientY of) So omnler DIRS BH A a ute swwi “gl: . hh ‘Ove eu cle Rt ik 3 (q+ Ratan many peace 2 baie es ill re} a oT whakiel eileaiy et) ii Sip. cia Ge. ay, f Fi * 1% « id any ATOLL RESEARCH BULLETIN No. 157 CARBONATE LAGOON AND BEACH SEDIMENTS OF TARAWA ATOLL, GILBERT ISLANDS by Jon N. Weber and Peter M.J. Woodhead Issued by THE SMITHSONIAN INSTITUTION Washington. DiniCers Wi oi At December 31, 1972 173 E TARAWA 20 Y Ba Oo 5 | 10 a KILOMETERS Bairiki Figure 1: Locations of sampling stations in Tarawa Lagoon. Inset at lower right shows positions of fringing reef flat sediment samples on Bikenibeu Island. BL = lagoonal reef flat; BO = ocean reef flat; BL and BO sample numbers begin at shore ( 1) and end at reef edge ( 6). CARBONATE LAGOON AND BEACH SEDIMENTS OF TARAWA ATOLL, GILBERT ISLANDS by Jon N. Weber’ and Peter M..J. Woodh2ad SUMMARY A large, shallow lagoon, almost isolated from the open ocean, distinguishes Tarawa from other atolls where carbonate sedimentation has been investigated. Passes through the atoll reef are lacking except along the western periphery. Size distribution statistics (median grain size, coefficients of sorting and skewness) were obtainedfor 57 sediment samples from Tarawa Lagoon and 12 samples from the fringing reefflats. Samples were subdivided into ten separate grain size fractions and the mineralogy of each fraction was determined. Component analysis of the larger grains was undertaken to identify the different sources of carbonate particles. Lime mids are accumulating in shallow water (6-10 m) over a large area of the southeastern lagoon. Evidence provided by mineralogy, carbon isotopic composition, and electron micro- scopy indicates that the fines consist of a mixture of algae-secreted needles and particles derived from the physical and biological breakdown of skeletal detritus. The abundance of carbonate mids in shallow water is attributed to the absence of current winnowing. INTRODUCTION Tarawa possesses several features which distinguish it from other atolls where carbonate sedimentation has been studied. These features especially concern the depth of the lagoon and the relatively poor connection between it and the open ocean. The water in much of Tarawa Lagoon is murky, a fact which is readily evident from the air. At the extreme southeastern corner of the lagoon, underwater visibility is less than 2 m, and associated with the high degree of turbidity are fine-grained carbonate muds accumulating in shallow water. The lagoon at Tarawa differs significantly from those of Bikini, Eniwetok, and other atolls where detailed sediment studies were conducted by Emery etal. (1954). Tarawa Lagoon is shallow, averaging about 12 to 15 m. with a maximum depth of 25 m. Rongelap, Eniwetok, and Bikini Atolls, however, have lagoons averaging about 50 m in depth (maximum 70 m). Furthermore, Tarawa Lagoon is encircled by coral reefs at sea level except along the western margin where the atoll reef top is submerged to about 9 m. There is only one deep (20-25 m) pass into Tarawa, whereas Rongelap Lagoon is connected to the sea by nine passes with a maximum depth of 66 m, and Bikini has eight passes or channels with a maximum depth of 60 m.° These variations in lagoon physiography might exert considerable influence on the characteristics of sedimentary material accumulating on the atoll. The dominant sediment contributors in most coral reef environments appear to be Halimeda, corals, molluscs, foraminifera, and the coralline algae. In unusual circumstances, other types of sediment may form, for example, non-skeletal carbonates in the lagoon of Hogsty Reef (Milliman, 1967) or the pelagic foraminifera, benthonic foraminifera, and mollusc sands on the Seychelles Bank (Lewis and Taylor, 1966). Considerable variations in the abundance of carbonate grains from different sources are observed from one reef to another and often within a single reef complex. Halimeda and foraminifera, for example, are important in the Great Barrier Reef (Maxwell, 1968) and the Florida reef tract (Ginsburg, 1956) provinces. Over large areas of the floors of Eniwetok and Bikini Lagoons, Halimeda is a major sediment ——__ - = =--3— *Pennsylvania State University, University Park, Pennsylvania 16802. ?Memorial University, St. John’s, Newfoundland, Canada. (Manuscript received June 1971--Eds.) 2 component, often comprising between 75 and 100% of the total (Emery et al., 1954). In Fanning Lagoon, however, foraminifera and Halimeda are not substantial contributors (Roy, 1970). Halimeda fragments are also relatively scarce in-the sediments of Kure and Midway Atolls (Gross etal. 1969). At Pearl and Hermes Reef, carbonate of foraminiferal derivation is considerably less abundant than grains from algae, corals, and molluscs (Thorp, 1936). Thus the striking differences in the proportions of various carbonate contributors reported for Pacific islands and atolls is an additional incentive for investigating the sedimentary deposits of Tarawa. PHYSICAL SETTING Tarawa Atoll, part of the Gilberts Islands group, is located in the central Pacific Ocean just north of the equator (1930'N, 173°E). The atoll is triangular in shape, with approximate dimensions of 25 km E-W and 30 km N-S. Although the dominant surface ocean current is from the east, the wind regime changes from NE Trades in Jan.-Feb. to SE Trades in July- Aug. The coral reef is continuous and rises to the surface on the windward side of the atoll, supporting numerous islands and islets no more than a few feet above sea level. These islands are surrounded by reef flats as mich as 300 to 800 m wide on both lagoonal and sea- ward shores, and they are separated from each other by narrow channels which are generally em<:rgent at low tide. Along the western margin of the atoll, the reef is submerged to depths of about 8 to 10 m. The mzin entrance to the lazoon is a breach in the leeward, subm 0.5mm grains in sample TA-7. 4 The percentage of coral debris is highest in the west near the atoll reef margin. Coral fragments decline markedly in abundance towards the east, amounting to essentially zero per- cent at station TA-7. There is a slight increase in the quantity of coral produced grains in the lee of the seaward (eastern) atoll reef, for example at TA-9. Skeletal detritus attributed to mollusca is also a significant sediment comjzonent, and as shown in Fig. 6, this material is most abundant in the center of the lagoon. Compared with other carbonate sediments of coral reef environments, echinoderm (debris appears to be unusually common. The percentage of this component is low in the western lagoon but it steadily increases eastwards, reaching up to 20% of the > 0.5mm grains in the region where the fine carbonate muds are accumulating (e.g. TA-6, 7, 8, 9). Skeletal ossicles of the echinoid Echinometra, ophiuroids, and crinoids constitute the greater portion of recog- nizable echinodern-derived grains from samples near the western atoll reef area, whereas elsewhere in the lagoon, heart urchin debris predominates. It is noteworthy that foraminifera contribute very little sand size carbonate throughout the lagoon except in the beach areas and along the western atoll reef zone. The 0.4 to 0.8mm size fraction of beach sands is almost exclusivey composed of the foram Calcarina. In sample TA-28, the benthonic foraminifera constitute 25% of the 0.84 to 2.00mm and 45% of the 0.42 to 0.84mm size fractions. Most of the smaller tests are Amphistegina, whereas Heterostegina is the major contributor of the larger shells. Pelagic foraminifera were very seldom observed, even in samples from the western atoll reef margin. Because of their fresh appearance, alcyonarian spicules are conspicuous in sediment samples from the western lagoon. They tendto concentrate in the 0.42 to 0.84mm size fraction but did not exceed 5% of any sample. Although coralline algae amounted to 9% in one sample, this component along with bryozoa, crustacean, andanneliddebris, etc. are minor constituents of the > 0.5mm grains in the lagoonal sediments of Tarawa Atoll. Major sources of aragonite are the green algae, especially Halimzda, the scleractinian corals, pelecypods, and gastropods. Smaller quantities of this mineral might be derived from hydrozoans, the octocoral Heliopora, some bryozoans, scaphopods, cephalopods, and some annelids. Producers of high-magnesium calcite include echinoderms, the coralline and red algae, benthonic foraminifera, alcyonarians, some bryozoans, calcareous sponges, decapods, and some annelids. Among the low-magnesium calcite contributors are a few benthonic fora- minifera, and some pelecypods and gastropods. The foram Amphistegina secretes low- magnesium calcite (Fig. 7) which accounts for the unusually high percentage of LMC in sample TA-28. COMPARISON OF SEDIMENTS ON REEF FLATS FACING OCEAN AND LAGOON Reef flats around the islands extend both seawardand into the lagoon, usually for appreciable distances (Fig. 1). As a substantial proportion of the total sediment associated with the atoll is derived from these reefs, samples were collected from reef flat environments on each side of Bikenibeu Island for comparison with sediments deposited within the lagoon proper. Physical conditions as well as the relative abundance of carbonate contributors are significantly different on the two types of reef flat. That facing the ocean, for example, is subjected to constant, high-energy wave action. Coralline algae, regular echinoids, and corals such as #’octllopova and Favites are conspicuous on the exposed seaward reef flats, whereas irregular echinoids, Porites, and calcareous green algae are more abundant on the flats adjacent to the lagoon. The species composition of the molluscan fauna of the two environments is also different. As expected, the seaward reef surfaces retain little sediment with the exception of the beach and the small localized deposits in isolateddepressions. These sediments are relatively coarse grained throughout (av. median diameter 1.13; range of median diameter 0.53 to 1.80mm). 4) In contrast, the lagoonal reef flats are almost completely covered with carbonate sediment having an average median grain size of 0.48mm (range of median 0.32 to 0.67). In this en- vironment, median grain size decreases slightly from the middle of the flat to the edge of the lagoon. Samples from both reef flats are generally weil sorted as indicated by the low values of the Trask sorting coefficient (av. 1.8 for the lagoon flat and 2.1 for the ocean flat). A‘ Bikenibeu, the degree of sorting increases from mid flat to lagoon edge but decreases from mid flat to ocean edge, probably reflecting the fact that the only significant sediment accumula- tions in the exposed, high-energy wave zone on the seaward flat are found in comparatively protected holes and depressions. Pernaps the most conspicuous difference between sediments of the lagoon and ocean sides of the island is the effect of abrasion caused by wave action. Carbonate grains on the seaward flats have a much higher degree of roundness and polish. The vitreous luster enhances the color of the component grains and imparts a distinctive appearance to the sediment. Aragonite is the most abundant mineral in all reef flat samples (Fig. 8) and in general, the lagoonal reef flat sediments contain a higher percentage than those on the seaward flat. From Bikenibeu beach to the edge of the lagoon there is a regular increase in the aragonite content from 50 to 84%, whereas on the ocean size of the island, the percentage of aragonite tends to decrease from 65% at the shore to 45% at the reef edge. Low~magnesium calcite constitutes from 3 to 2% of the sediments on the lagoon reef flat but was seldom detected in samples collected on the seaward flat. Thus the aragonite and high-magnesium calcite distri- bution patterns can be considered complementary. ORIGIN OF THE LIME MUDS The quantitative importance of lime mud in the stratigraphic record has De2n emphasized by Matthews (1955) who foundit paradoxical that studies of Recent carbonate sediments general- ly have concentrated on the genesis of the sand size particles rather than on the origin of the fine-grained constituents. In som? major coral reef provinces, carbonate sands are clearly mich more abundant than carbonate silts or mids. Sediments of the Capricorn reef com:lex of Australia, for example, seldom contain more than 2% mud (Maiklem, 1970). According to Maxwell (1968), very little fine carbonate is produced in the Great Barrier Reef system of Australia, and the mud present in sedim=nts of that region is predominan‘ly of terrigenous origin. Elsewhere the deficiency of fine particles and the high proportions of sand and gravel may be explained by removal of the fines into deeper water, as at Pearl and Hermes Reef (Thorp, 1936). Gross et al. (1969) found that particles less than 125 microns in diameter are relatively rare on Kure and Midway Atolls except in the deeper parts of the lagoons. Much of the fines apparently remains in suspension and is transported out of the lagoon, leaving behind the coarse-grained sediment in shallow water. Calcareous oozes are reported in the deepest parts of Kapingamarangi Lagoon, between 50 and 80 m. but McKee (1958) concludes that the mudis not likely derivedprimarily from the residue of sediments occurring in shallower water. The situation at Fanning Island appears quite different in that fine-grained carbonates are unusually abundant. Muddy sediments in the lagoon at Fanning are reported by Roy (1970) who attributes most of the material to physical and biological abrasion or comminution of corals, molluscs, and calcareous red algae. Water in Fanning Lagoon is characterized by a high degree of turbidity, with underwater visibility often limitedto 2 m or less. According to Bakus (1968) the suspended particles are generally between 5 and 15 microns in diameter. Hence the presence or absence of carbonate fines may be controlled more by the degree of current winnowing than by actual production of the grains. This is demonstrated in the case of Palmyra Atoll, at least, by the drastic changes in the lagoon following the construction of inter-island causeways in the early 1940’s. Dawson (1959) reported that reef growth in the lagoon ceased after being deprived of circulating water. The lagoon became murky throughout due to the suspen- sion of fine calcareous sediment, and inshore reef areas which once supported luxuriant 6 coral growth are now deeply covered with carbonate mud. Several possible mechanisms may be important in generating carbonate fines: inorganic precipitation; algal secretion of aragonite needles; and attrition of larger calcareous skeletons by mechanical or biological processes. The first of these, inorganic precipitation, is contro- versial. Fine-grained sediments covering large portions of the Bahama Banks, for example, are believed to be of inorganic origin by som: investigators (Cloud, 1962; Illing, 1954; Newell etal. 1960; Broecker and Takahashi, 1966) while others consider green algae to be the domi- nant source (Lowenstam and Epstein, 1957). Milliman (1967) reports that most of the lagoonal sediment at Hogsty Reef is ‘‘non-skeletal’’ and probably precipitated inorganically. After studying sedimentation on Serrana Bank, Milliman (1969) later postulated that non-skeletal carbonates might form in open lagoon conditions only where biogenic carbonate secretion is lacking. Lowenstam’s (1955) discovery of alga-secreted aragonite needles established an im- portant endemic biogenic source of carbonate fines. Acicular crystals of aragonite, about 2 to 9 microns in length, are producedinlarge numbers by species in the Codiaceae, Dasycladaceae, Nemalionaceae, and Chaetangiaceae. Fine-grained sediments from Bermuda, Jamaica, Kayangle Atoll, Johnston Island, Guam, Ifalik, Eniwetok, and Bikini were found to contain such needles (Lowenstam and Epstein, 1957) and Lowenstam (1955) predicted that quiet water sediments in equatorial regions should carry them wherever the algae are present. Stockman et al., (1967) attribute most of the lime mud in the south Florida area to the skeletal disintegration of calcareous algae. Although experimental destruction of large invertebrate skeletons by boring organisms and abrasion tend to demonstrate the difficulty in comminuting shells to very fine particle sizes (Driscoll, 1970), it appears that physical and mechanical breakdown is an important mechanism of mud formation in some areas. Matthews (1966) described lime muds of British Honduras which contain both calcite and aragonite butare virtually devoid of aragonite needles. His analyses indicate that mechanical comminution in shoal waters and breakage of fragile mollusc and hyaline foraminifera skeletons in the lagoon are important processes in tne production of these muds. Other significant mechanisms include mastication by echinoids and holothurians, and trituration of carbonates by scarid, chaetodontid, acanthurid, and pomacentrid fish (Cloud, 1952). Land (1970) reports that appreciable quantities of carbonate mud may be produced by epibiotic growth of coralline red algae and serpulid worm: on the turtle grass Thalassia. At Tarawa, lime muds are presently accumulating in the extreme southeastern corner of the lagoon in relatively shallow water (6-10 m). These sediments are cream to very light green in color, are extremely plastic, and poorly consolidated. The unusually soft bottom created minor sampling problems in that the sediment grab, which is triggered on contact, would not always ‘‘fire’’. Despite replicated sampling of this area, heart urchins (Brissopsis and Maretia? and sand dollars (Laganum sp.) were the only living macrobenthos recovered. The two spatangoids were considerably more abundant than Laganum, and the significant con- centrations of echinoderm skeletal debris in these sediments indicate that heart urchins are an important in situ source of carbonate (Fig. 6). Up to 80% of the material in the eastern lagoon is found in the <0.074mm size fraction, and examination by both electron microscopy and scanning electron microscopy indicates that the bulk of the fines actually lies in the particle size range of 1 to 10 microns. Although needles, presumably aragonite (Fig. 9) are not uncommon, electron photomicrographs show a variety of grain shapes including fragments of laths and needles, platy and tabular forms, and nondescript particles. Between 60 and 75% of the carbonate and mud consists of aragonite. Low-magnesium calcite amounts to 4-8%, and high-magnesium calcite accounts for the remainder. 7 In much of Tarawa Lagoon, the water is turbid, and this is especially true in the region of lime mud accumulation where underwater visibility is limited to a few feet. The abundance of carbonate fines, the shallowness of the water, and the high degree of turbidity suggest the possibility of inorganic precipitation, although like the ‘‘whitings’’ in the Bahamas (Broecker and Takahashi, 1966), the turbidity is probably due to the suspension and resuspension of previously formed carbonate rather than in situ precipitation by inorganic mechanisms. Nevertheless, carbon and oxygen stable isotope ratio data were obtained to examine this possibility. The isotopic composition of the ‘‘J’’ size fraction (<0.078mm) was determined for four samples in the area of lime mud deposition. 613, and 618. values, with respect to the PDB ‘ C O CO, standard,* are: 2 6 § 13, Oe 18, ore TA-8 + 1.95 - 2.93 TA-9 + 1.70 = 2.07 TA-10 + 1.86 - 2.55 TA-12 tle DS = 209 Admittedly the J size fraction is a mixture of aragonite, HMC, and LMC, each of which may have a different isotopic composition. A plot of 513¢c and 51809 vs. mineralogy, however, indicated no correlation. The average 6180 for the four lime muds in -2.66 °/oo, which corresponds to a crystallization temperature of 28.9° C using the ‘‘paleotemperature’’ equation of Epstein et al. (1953) and assuming precipitation of the carbonate in oxygen isotopic equili- brium. The 180 content of the ambient seawater also enters into this equation, and since this was not determined, specimens of calcareous coelenterates which precipitate skeletal CaCOg in apparent oxygen isotopic equilibrium with surrounding seawater (Weber and Woodhead, 1970) were analyzed: Millepora (N= 16), Tubipora (5), and Heliopora (13). The average 6180 of these 34 specimens is -2.29 (standard deviation 0.17 °/oo) which corresponds to a calculated crystallization temperature of 27.19C. The mean annual temperature of surface seawater at Tarawa, as obtained from monthly temperature distribution maps of the central Pacific, is 28.LCC- While the oxygen isotope ratios are similar to those predicted for inorganically precipitated carbonate, 513¢ data for the lime muds are several permil lower than the theoretical values, and are in the range of carbon isotopic compositions typical of biogenic carbonates. The presence of HMC and LMC in addition to aragonite further suggests that the lime muds in Tarawa Lagoon are derived from the breakdownof skeletal detritus. It is noteworthy that while the occurrence of aragonite needles is indicated by electron microscopy, numerous other grain shapes are also evident. It appears likely that the fines are a mixture of algal needles and particles produced by attrition of larger skeletal debris. On atolls, fine-grained carbonate is generally found to be accumulating only in the deepest portions of the lagoon, e.g. between 50 and 80 m at Kapingamarangi (McKee et al. 1959). According to McKee etal. a ‘‘shallow- water environment favorable for such fine-grained deposits has not been reported from any of the Pacific atolls’’. Tarawa, however, has extensive deposits of carbonate mid which are accumulating in shallow water. This can be attributed to protection of the sediment from current winnowing, which in turn results from the near absence of deep channels connecting lagoon and ocean, and the windbreak provided by the reef islands. *Using the well known § notation, where: 5c —— rE Sane) 1000 X13 /(112 C | Ciianaara in per mil, relative to the PDB standard CO. 489-409 O- 73-5 ACKNOWLEDGEMENTS We are grateful to D. Cudmore, G. Jones, and R.G. Roberts of the Gilbert and Ellice Islands government for their generous support of the field operations, to N. Suhr for the elec- tron photomicrographs, to Dr. E.W. White for providing scanning electron microscope facili- ties, to Dr. D.M. Raup for identification of the echinoids, to Dr. A. L. Guber for independent microscopic examination of many of the sediment samples, and to the U.S. National Science Foundation (J. N. W.) and the Australian Research Grants Commission (P.M. J.W.) for research support. REFERENCES Bakus, G.J., 1968. Sedimentation and benthic invertebrates of Fanning Island, central Pacific. Marine Geol. 6: 45-51. Broecker, W.S. and Takahashi, T., 1966. Calcium carbonate precipitation on the Bahama Banks. J. Geophys. Res, 71: 1575-1602. Cloud, P.E. Jr., 1952. Prelim’nary report on geology and marine environments of Onotoa Atoll, Gilbert Islands. Atoll Res. Bull. 12: 1-73. ae aie a 1962. Environment of calcium carbonate deposition west of Andros Island, Bahamas. U.S. Geol. Surv. Prof. Paper 350: 1-138. Dawson, E.Y., 1959. Changes in Palmyra Atoll and its vegetation through the activities of man, 1913-1958. Pac. Natur. 1: 1-51. Deines, P., 1970. Mass spectrometer correction factors for the determination of small iso- topic composition variations of carbon and oxygen. Jnt. J. Mass Spectrom. Ion Phys. 4: 283-295. Driscoll, E.G., 1970. Destruction of potential fossils by boring organisms and abrasion. Geol. Soc. Amer. Abstr. for 1970 7: 740-741. Emery, K.O., Tracey, J.I. Jr., and Ladd, H.S., 1954. Bikini and nearby atolls, Marshall Islands. U.S. Geol. Surv. Prof. Paper 260A: 1-265. Epstein, S., Buchsbaum, R., Lowenstam, H.A., and Urey, H.C., 1953. Revised carbonate- water isotopic temperature scale. Bull. Geol, Soc. Amer. 64: 1315-1326. Ginsburg, R.N., 1956. Environmental relationships of grain size and constituent particles in some south Florida carbonate sediments. Bull. Amer. Assoc. Petrol. Geol. 40: 2384- 2427. Gross, M.G., Milliman, J.D., Tracey, J.I. Jr., and Ladd, H.S., 1969. Marine geology of Kure and Midway Atolls, Hawaii: a preliminary report. Pac. Sci. 23: 17-25. Illing, L.V., 1954. Bahaman calcareous sands. Bull. Amer. Assoc. Petrol. Geol. 38: 1-95. Land, L.S., 1970. Carbonate mud: production by epibiont growth on Thalassia testudinum. J, Sed. Petrol, 40: 1361-1363. Lewis, M.S. and Taylor, J.D., 1966. Marine sediments and bottom communities of the Seychelles. Tvans. Roy. Soc. London A259: 279-290. 9 Lowenstam, H.A., 1955. Aragonite needles secreted by algae and some sedimentary impli- cations. J. Sed. Petrol. 25: 270-272. Lowenstam, H.A.and Epstein, S., 1957. On the origin of sedimentary aragonite needles of the Great Bahama Bank. J. Geol. 65: 364-375. Maiklem, W.R., 1970. Carbonate sediments in the Capricorn Reef complex, Great Barrier Reef, Australia. J. Sed. Petrol. 40: 55-80. Matthews, R.K., 1966. Genesis of Recent lime mud in Southern British Honduras. J. Sed. Petrol, 36: 428-454. Maxwell, W.G.H., 1968. Atlas of the Great Barrier Reef. Elsevier, Amsterdam. 258 pp. McKee, E.D., 1958. Geology of Kapingamarangi Atoll, Caroline Islands. Bull Geol. Soc. Amer, 69: 241-278. McKee, E.D., Chronic, J., and Leopold, E.B., 1959. Sedimentary belts in lagoon of Kapinga- marangi Atoll. Bull. Amer. Assoc. Petrol. Geol. 43: 501-562. Milliman J.D., 1967. Carbonate sedimentation on Hogsty Reef, a Bahamianatoll. J. Sed. Petrol, 37: 658-676. msoooesosa 1969. Carbonate sedimentation on four southwestern Caribbean atolls and relation to ‘‘odlite problem’’. Bull. Amer. Assoc. Petrol. Geol. 53: 2040-2041. Neumann, A.C., 1965. Processes of Recent carbonate sedimentation in Harrington Sound, Bermuda. Bull. Mar. Sci. 15: 987-1035. Newell, N.D., Purdy, E.G.andImbrie, J.,1960. Bahamianoolitic sand. J. Geol. 68: 481-497. Roy, K.J., 1970. Sedimentation and reef development inturbid-water areas of Fanning Lagoon. Bull. Amer. Assoc. Petrol. Geol. 54: 867. Stockman, K. W., Ginsburg, R.N., and Shinn, E.A., 1967. The production of lime mud by algae in south Florida. J. Sed. Petrol. 37: 633-648. Thorp, E.M., 1936. The sediments of the PearlandHermes Reef. J. Sed. Petrol. 6: 109-118. Weber, J.N., 1968. Quantitative mineralogical analysis of carbonate sediments: comparison of X-ray diffraction and electron probe microanalyzer methods. J. Sed. Petrol. 38: 232-234. Weber, J.N. and Woodhead, P.M.J., 1970. Carbon and oxygen isotope fractionation in the skeletal caroonate of reef-building corals. Chem. Geol. 6: 93-117. 10 TABLE 1: SIZE DISTRIBUTION STATISTICS Depth(m) 13 Ma 0.45 0.33 0.34 0.13 0.17 0.062 0.028 0.009 0.080 0.090 0.12 0.010 0.19 0.20 0.27 0.30 0.22 0.23 0.35 0.46 0.22 2.10 0.63 0.523 0.37 Sk 0.94 0.84 0.82 1.06 1.24 0.62 1.40 0.89 1.68 0.74 1.08 6.40 0.98 1.15 1.01 0.87 0.97 0.87 eii2 1.01 0.92 0.85 0.88 On72 0.82 table 1 (continued) TA- 26 27 0.63 0.71 0.90 0.87 0.80 0.63 0.50 0.56 0.16 0.53 0.36 0.35 1.20 0.72 0.68 0.73 0.30 0.76 0.46 0.22 0.24 0.63 0.39 0.57 0.70 0.35 0.82 1.80 1.54 1.58 1.70 1.79 1.73 1.46 2.27 1.76 2.52 2.60 2.50 2.07 2.05 1.83 2.45 2.83 1.89 2.75 2.45 2.84 2.08 2.95 1.88 1.84 2.67 o2 JEL 12 table 1 (continued) TA= 54 505 0.52 1.82 1.03 35 4.5 0.53 1.63 1.29 56 4 0.63 2.10 0.87 57 1.5 0.22 2.77 1.17 BL- 1 0.40 1.82 1.09 2 0.47 1.88 1.00 3) 0.57 2.06 0.96 L 0.67 ¥593 0.70 5 0.42 1.73 1.15 6 0.32 1.31 1.05 BO- 1 0.84 1.35 0.99 2 1.00 1.83 1.08 3 1.40 = = 4 0.53 1.69 1.45 y) 1.80 2.50 0.60 6 1.48 2.98 0.53 All TA samples are from Tarawa Lagoon except TA-22 which is beach sediment from ~ Bairiki Island. BL = lagoonal reef flat at Bikenibeu Is.; BO = ocean reef flat at Bikenibeu Is.; Md = median diameter in mm; So = Trask sorting coefficient; Sk = — Trask coefficient of skewness. TABLE 2: MINERALOGICAL COMPOSITION OF SIZE FRACTIONS * el I a ee Se TA-l A 87 89 86 83 86 83 80 78 77 68 85 HM 10 9 2p lS set oe 200 19) 28 13 LM 3 2 2 2 1 3 3 2 L Lb 2 TA-2 A 88 89 90 91 88 86 88 81 78 79 87 HMC 9 7 7 7 Shee 3 I es U Aa tay aan 10 LM 3 L 3 2 3 3 3 5 5 L 3 TA-3 A 89 89 89 88 91 8 86 8h 79 78 87 HMC 67 7 8 9 Manele te Mae ON NB RS 10 IM 4 L, 3 5) 2 2 3 L, 5 h 8) TA-4 A SIS 7 OO GON SO SS ° 82) B82... 76 76 83 HMC 13 9 t 8 8 Syd 15 AS os 13 IM 3 L 3 2 3 4 4 3) 6 6 4 TA-5 A 83 89 86 9h 91 83 87 82 83 75 85 HMC Ol, 720 h 6y) 13 Oo ho 135 19 11 IM 3 4 4 2 3 4 4 L, L 6 4 TA-6 A NO” FO) (Ore tes 295i) 96 19h 186". 76) . 73 77 HMC 6020 8 5 9 4 3 5) 208 N6y i AD 15 Ime 4 16 19 3 1 1 1 4 8 8 8 TA-7 A 13) OF Ol aeo. | Ble Ba IT 75) JO 72 HMC 25 25 Oo Cea bedi ia UB Aiea Ws Ueideigees C2 als Wey (aia he Dana 22 IM 862 6 4 6 3 5 5 6 6 7 6 TA-8 A 1M he OM Miia) SOD = = - 6h, 66 EMCI 2O ia Zion 23 Wa 5) Sia - = 29 27 UCM TSU momo NEN ee em 8g ALS) 14 table 2 (continued) TA-9 TA-10 TA-11 TA-12 TA=13 TA-14 TA-15 TA-16 A HMC LMC A 19 16 5 66 33 17 table 2 (continued) TA-17 A 86 8686 me 2a al me. 3 3 TA-18 A 8 88 AMC 14 9 LM 2 3 TA-19 A 83 ©80 HMC 1, 18 IM 3. 2 TA-20 A 86 8688 HM 609Ss«d2100 IM 65 2 TA-21 A 76 8&0 HMC 23) «16 tw OU 4 TA-22 A 19) Aa HMC 81 89 IMC (OO 0) TA-23 A 80 82 HMC 17 16 LMe 63 2 TA-2h A wm (789 HMC 27 9 IMC 2 2 15 16 table 2 (continued) TA=25 TA-26 TA-27 TA-28 TA-29 TA=30 TA=31 TA=32 A HMC LMC A HMC LMC A 22 10 19 16 table 2 (continued) TA-33 TA-34 TA=35 TA-36 TA-37 TA-38 TA=39 TA-L0 A HMC LMC A LMC 88 16 10 83 i) 7h, 20 anh 18 table 2 (continued) TA-42 TA-L3 TA-44, TA=45 TA-46 TA-L7 TA-48 TA-L9 A HMC LMC A LMC 89 10 719 15 83 14 table 2 (continued) TA-50 TA-51 TA-52 TA-53 TA-54 TA-55 TA-56 TA-57 A HMC LMC A LMC 88 10 79 18 er ih & 19 20 table 2 (continued) BL-l A 29 «=L6 HMC 70 0= = (53 IM 1 1 BL-2 A 49 5k HMC 48 46 IMC 3 0 BL-3 A LV 56 HMC 57 843 LMC 2 ub BL-h OA 50 64 HMC 49 32 LMC 1 4, BL-5 A 60 62 HMC 390 3k LMF 61 4, BL-6 A (5 1/5 HMC 20 = 22 EMG 9) 73 BO-l A 66 64 HMC 34 36 LMC 0 0 BO-2 A 56 818 28 eal table 2 (continued) BO-3 =A 55 DAO ne 21 39k 267" 72k 6OR = = 53 HMC. E5r Som 90mm 9 a Oly 153). 295 LO. - L7 LMC 0 0) 0) 0 0 Le) 0 Oo - = 0) BO= 5p A) 330) LOe MeO wemod 57260: 59.50) 52" 1k 37 BO-6 =A IN RGy 25 Pal 5 | ON 5602 55. = = 45 HMC OLS, 5k fog 0) who (39 hh LS = = Py) LMt 6—O 0 0 0 0 0) 0 Or .= = 0 # mm range of size fractions specified in text. A = aragonite; HMC = high-magnesium calcite; LMC = low-magnesium calcite. MEDIAN GRAIN SIZE (mm) DEPTH (m) ov BIKEMAN REEF EDGE 28 29 30 3I 32 33 BK! 2 SAO Ore Ono 10 SAMPLING STATION Figure 2: Median grain size of lagoonal sediments along the sampling traverse from Bonriki (eastern lagoon) to the western reef margin. Location of sampling stations given in Fig. 1. °}X9} UL poljioeds ore ¢ pues V Sassejo 0} suIpuodsei100 Sesuet azIS UTeINn ‘ayUOSere ST yey} AT0SE1"O oeZIS UIvIS YORE JO asejUsoIed ay} JUSsSetdet SaAIND *([ °BI.q ULUMOYS UOT}E}S SuTTdUIeS 0} Spuodserzo0d aAINo MOTAQ Jaquinu) setdures [eotdA} 10} ozis uteis Jo uoljouny & Se UOTIISOdUIOD Teorsojerourpy, °€ A4Nd2.7 He Oe 3 GD) Be te dd 3 = 8. 8 489-409 O- 73 - 6 DY A — comb SLUM UN Ut bbb gst tt sigececers: ITTTTT Da VLA, LL, TA Yj (2QILLIILLLLLLLLLLLL LL AI LID ITLL LLL eee EIT EILLLILILLIIILILT Ws TA 3 pezezzZLLLL 6b! VIL j) VZZZZZA ie errrrrrees VM. Yih eS 7///// eee LL, TA CLLLLL: Mel} YJ}! Witty eovscer 3 | TA VM bree 30 TA | = YU eee - 9 TA ° lean MMMPTIZIZD : 28 4 alBiclolelF[S[4|1 |v] Figure 4: Distribution of total sample aragonite among grain size classes for sediment samples along the TA-10 (eastern lagoon) to TA-28 (western lagoonal reef margin) transect. The height of a column ina histogram indicates the percentage of the total sample aragonite that is in the grain size range designated by the corresponding alphabetic character (millimeter equivalents specified in text). 53 54 55 56 of Figure 5: Systematic changes in the distribution of aragonite among grain size classes for sediments along the TA-53 to TA-57 transect in the northern corner of the lagoon (locations in Fig. 1). Curves B and J indicate the percentage of total sample aragonite that is in grain size classes B (coarse) and J (fine) respectively. 70 60 010) a Mollusc 40 Halimeda 30 20 © Coral “ Echinoderm Dar 2ONSO Sin San sat Pull Smale a7 OnmnO SAMPLING STATION Figure 6: Percentage of four major components (>0.5mm grains) of the sediments in Tarawa Lagoon. Geographic locations of the sampling stations are shown in Fig. 1. “8S -VL a[Cwes WOT} paj0eI}X9 9TEM SapoTVIed [Ty ‘SUTeIS a[SUTS UTYIIM SoT}eI e5 /igpue ep /SN aTqelsrea PoMOYS SN}IA}Ep PITSuUe PUR OSNITOW °(STAQep [B109) N ‘(Dpawmoy) W ‘(opew2oy) J ‘(quewSesy OSNTICW) yy ‘(ou2zsazsiyduy) pc ‘(purzsazsiyquy) | ‘(bAjawouiyog ‘sautds ptoutyoe) Wy ‘(sayeTd TeuoLoo ploutyo9e) H ‘(pu210910D) q ‘(DuU2Ge}S04010H) q ‘(SatnoTds ueTIeUOADTe) q ‘(DUILD910D) OD ‘(osogoursrnyy) q “(Satotsso prorntydo pue proutz9) y ~‘aqordozo1wi uosjoeTa Aq eg pue ‘ig ‘SI JO SISA[eue SnosuRNUIIS Aq poulUIajep syuosUCdUIOD yUeUIpes Jo ASZoTereUTy +2 oAnS2Z DD/4S JLINOOVYEV 3JLINIVD WNISSANOSVW MOT SJLINTV9 WNISSNOSVW HOIH D9/5W *(7X9} UI potfloads sjuajeaArnbe A9zSUlIT]1UW1) Tajoe1vYyo ITIjyaeqevydte Sutpuodsets -100 oy} Aq PaYXUSISApP 9SURI 9ZIS UTVIS JY} UT ST }eY} a}TUOSeAIe oTdUIeS [e}0} 94} Jo adrejUDOIEd 94} SO8}COTPUL WUILISOJSTY & UT UUINTOD & Jo JYSTEY sy, “*T “SI UI pa}eoTPUI SaTdures Jo uoT}eEI07 *SJUSWIIpPaS }eTJ Joo PIeMvaS pUL UOOSET UI UOT}NqII}SsIp ajytuOSere Jo uOosTueduIOD :9 aunsiz ivild 4335u4 GYVMV3SS 1V14 4j33Y¥ IWNOOSV1 Figure 9: Electron photomicrograph of the fines (<0.074mm) from sample TA-8, showing whole and fragmented aragonite needles and laths. The prominent dark colored acicular crystal at lower right is between 1 and 2 microns in length. ; : Ohl { j i) j ‘i j ro 1 Ve a y ut ; ls 7 ‘ j ; } f ‘ cA a q = a 7 ; ann ae < Fy oa : 5 wl if v ae ne, : , ; vee oa, ; ‘ r +a 4 1 uy - gm a F; ag SE ; : i ) hx + A y dg sil 7s rf U } * : _ : ; { > . : ! flex : _ ; - 5. | _ ¢ | De ; i ry H : i a o, : ah ; { + te =~ ¥ ms | =I (eee eo . jinn she Ye te A ~ " s : = Ue td -o é ATOLL RESEARCH BULLETIN No. 158 BIRDS SEEN AT SEA AND ON AN ISLAND IN THE CARGADOS CARAJOS SHOALS by Ro Pocklington, Pon.) Wiltis, and M. Palmiiers Issued by THE SMITHSONIAN INSTITUTION Washimetoniia DCs Ur Ol A December 31, 1972 1. J Aye if v a) , : : ‘ te ‘a ft Ms : - ee ti uli i | 7 4 ; i : . ‘ f Wy, * i. A = i ay = 7 ees ji ‘ i eo ee i att 4 5 : f ho ep Wo “ : i ( ny’ P iG } _ ten oe i = if, ye i eS | a) or ai ' 7 td - iv i 5 os i a } . fueeth } : ; Onn. r 7 : - : ; - ; 4 ee Mt rs iy a i , the / Davy AN a1 Aun K : i iY i a ow y aon : 5 ain y rt a : 4 : male _ 1 . +. 7 a : : : i) - ¥ 1 5 i : ‘ m : t - =i l . y = , Rac a 1 i ; Vead ' ii My Ne *) : i iY 7 q ; Ae , iy ’ fi : : : i ; : Dy na eed set : “ oe fi \ : : H J 4A 7 M if , ¥ re ve : : ; Ww, ene : 1 , me : 1 U ’ } : i ; i : 1 7 1 1 ey . Y Si : 7 - ' { i : J i i i i ' ’ : ' ‘ 7 “ U 7 F ; : Pay - = 1 1 Ss " 1 vw : A : : ; : i 7 ae ; ; . _ } : t : ‘ a - \ ; a x - ‘ Tee, : y i. ‘ : : : ‘s ' rrr z aa 1 ", - ir ot . + - : re 7 an 5 a- gf ‘ ; -oJ 4 >. . . , we, - = he mn he Th Pea, thay - i , eet ae fi : dol SSS Fun : 7 - - 7 2 1 . : , yee a : i : oot : Te : : \ i : tae 7 7 ’ : ’ ey ‘ 3 ‘ ae ne, t eo , - . ; : od a i : : { 7 ~ bs e 2 at | : r 2 a = 7 i an i) A - ", y x A » “a ar: J : 7 / : “my = = : iy ‘ 43, A ea . ” : ae - i " o' i : - 7 + + a 1 Pine - : = 7 = a : : : ) , oe ‘ - a ¥ oe 1 = 1 an ar ea ‘ > =. a a 7 } - Fatt hy ee > Ber ‘ : ae Paine 7 : } N ee a v - : o vie : 7 ; P r Es 4 Sat A AO, ‘y BIRDS SEEN AT SEA AND ON AN ISLAND IN THE CARGADOS CARAJOS SHOALS by R. Pocklington, P.R. Willis and M. Palmieri During a second cruise to the Indian Ocean by the Woods Hole Oceanographic Institution vessel R.V. ATLANTIS II in 1965 as part of the International Indian Ocean Expedition, a return visit waS made to the Cargados Carajos Shoals, where we had previously gone ashore in November, 1963 (Pocklington, 1965). No other vessel during the I.I.O.E. had a similar oppor- tunity to observe the western Indian Ocean at opposite seasons with the same personnel and techniques. The Cargados Carajos are interesting because they are geographically between the Seychelles, where sea-bird species breed mainly in the northern summer (Loustau-Lalanne, 1963), and the Mascarenes, where many sea-birds breedin the southern summer (Newton, 1958). The species list which follows covers records made at sea between Tromelin (15952'S, 54°25'E), Ile du Sud in the Caragados Carajos Shoals (16949'S, 59930'E), Rodriguez, and Mauritius, June 2-7, 1965, inclusive, and ashore on Ile du Sud, June 3. Nomenclature follows Alexander (1955), and names of colors and topography of birds follow Palmer (1952). Puffinus lherminievi | Audubon’s Shearwater One at 18252'S, 62°13'E, 4 June, in the vicinity of Rodriguez. Bailey (1968) saw few more than 50 miles from their breeding grounds, and Gill (1967a) found them common only in the vicinity of the Seychelles, Mauritius and Reunion, but not in the intervening seas, which hints at islands off Rodriguez as possible breeding stations, as suggested by Gill (1967b). Pierodroma spp. Gad-fly Petrels On 4 June at 18°52'S, 62913'E, nine petrels with grey upper-parts, white underparts and a dark leading edge to the upper wing, were seen. They were unlike any of the Petrels and Shearwaters with which we were already acquainted. A possibility exists that these birds were Barau’s Petrels Pterodvoma baraui (Jouanin, 1963), of the existence of which we were un- aware at the time. This bird breeds on Reunion in the southern summ=2r, dispersing late April/early May for parts unknown (Jouanin and Gill, 1967). A recent examination of sub- fossil petrel bones from Rodriguez (Bourne, 1968) indicates that gad-fly petrels did breed there, although they apparently no longer do so (Gill, 1967b). Oceanites oceanicus Wilson’s Storm-Petrel Two storm-petrels, dark with white rumps, seen withthe above-mentioned gad-fly petrels, were believed to be this species, which we recorded with decreasing frequency as we went south. um eee R. Pocklington: Marine Science Branch, Atlantic Oceanographic Laboratory, Bedford Institute, Dartmouth, Nova Scotia. P.R. Willis: 10 Orne Square, Salem, Mass. 01970. M. Palmieri: Woods Hole Oceanographic Institution, Woods Hole, Mass. 20543. (Manuscript received Jan. 1971--Eds.) 2 Phaethon rubricauda Red-tailed Tropic-bird Two were seen between Cargados Carajos and Rodriguez, one at 18°25'S, 62°13'E, the other at 19°09'S, 62°33'E. It has not been recorded in recent times from Rodriguez (Bourne, 1968), nor from Cargados Carajos (Newton, 1958), though Staub and Guého (1968) mention a red tail-feather picked up on Albatros I, Cargados Carajos. It breeds on the cliffs of Gunners Quoin, Mauritius (Newton, 1958) where many were seen 7 June. Phaethon lepturus White-tailed Tropic-bird One adult flew about the ship while the party was ashore on Ile du Sud. No others were seen until we approached Mauritius, 7 June, where they breed (Gill et a/., 1970). Sula dactylatva Blue-faced Booby Three adults were seen close to Ile du Sud, but they were not nesting there, though they do on Ile du Nord (Newton, 1958; Staub and Guého, 1968). Three more were in a mixed flock with Sooty Terns, Sterna fuscata, between Rodriguez and Mauritius, where they breed on Serpent I. (Newton, 1958). Fregata minoy Great Frigate-bird In addition to numbers of unidentifiable frigates among large flocks of terns feeding off Tromelin, where F. minor breeds (Staub, 1970) others were seen between there and Cargados Carajos 16°907'S, 56°05'E, feeding in association with flocks of Sooty Terns. Three all-black males were positively identified among them. No more were seen outside this area. Fregata ariel Lesser Frigate-bird Males of this species, identified by white patches on each side of the abdomen, were seen at sea as we approached Ile du Sud. They were soaring over groups of feeding Noddies Anous spp. and Sooty Terns. No frigates were seen at sea subsequent to our visit to this island. On Ile du Sud, there were ca. 200 nesting pairsin the N.E. and 60 in the S.W. of the island, an increase over the total of ca. 200 pairs nesting in November 1963. The red gular sac of some sitting and flying males was inflated, in contrast to our previous visit, when none of the males showed more than a small shrunken throat-pouch. In the case of the Great Frigate-bird on Tower I. in the Galapagos, Nelson (1968) found that the pouch is inflated for only three or four weeks at the beginning of the breeding season. If this holds true for F. ariel] then this indicates that the beginning of the breeding season was not long past. Staub, who visiged during the latter half of April, 1968, found that ‘‘nearly all the males had their red gular pouch fully extended,’’ and that eggs were already present (Staub and Gueho, 1968). Only one juvenile was seen flying over theisland, which is in accord with the idea that they disperse away from the breeding stations while the adults remain relatively close by (Bailey, 1968). There were no hatched eggs, in contrast to November 1963, when eggs, naked young and downy chicks were simultaneously present, as shown in published photographs (Willis, 1966). Reference to Table I shows that nesting occurs in both northern and southern summer, in con- trast to Ff. minor which breeds only in the southern spring. Also, the duration of the F, ariel breeding cycle is no more than 8-9 months, as found by Gibson-Hill (Palmer, 1962), for F. andrewsi on Christmas I., near Java, and not 15 to 18 months, as found for F. minor on Tower Island, Galapagos (Nelson, 1968), where long before the juveniles from one season were fully independent, the next season began. 3 The female Lesser Frigate-birds were of two types: bluish-gray bill, black eye-ring, preast pure white with a sharp demarkation between the black of the throat and the white of the preast, side of neck and hindneck white; and, pink bill, rose eye-ring, white breast with cinnamon flecking which destroys the sharp demarkation of throat from breast so charac- teristic of the other type, side of neck as breast, the white of the hindneck obscured by black and cinnamon flecking. Both types were sitting on nests. These differences may be due to age or stage of physiological cycle of one form only, but the possibility exists that this represents a stable polymorphism or an overlap of breeding ranges of two previously undescribed forms of this species. Staub confirmed this variation (Staub and Guého, 1968) and noted that ‘‘the eggs of the two varieties seem somewhat different in shape and size,’’ a point that we had not noticed. He also noted red eye-ringed females as being more numerous than the black-ringed variety. Sterna dougalli Roseate Tern About 200 adults were seen ashore onthe sand-pit west of Ile du Sud. There were no young birds as there hadbeenin November 1963. Nesting would appear to be restricted to the southern spring (see Table I) in contrast to the other western Indian Ocean breeding stations of this cosmopolitan species, where nesting is in the northern summer (Watson @? al., 1963). On islands at comparable latitude off Western Australia, nesting in both April-June and November- December has been recorded (Serventy and Whittell, 1962). Sterna fuscata Sooty Tern This species, by far the most frequently encountered bird in the western Indian Ocean (Pocklington, 1967) as also noted by Parker (1970), was seen in hundreds at sea between Tromelin and Cargados Carajos, but was absent as a nesting species from Ile du Sud. A dozen or so were seen flying over the island: they were probably breeding on other islands of the group (Staub and Guého, 1968). Between Cargados Carajos and Rodriguez, and between there and Mauritius, where they breed (Gill et al., 1970) smaller parties of up to 30 were seen, but they were absent from the immediate vicinity of Rodriguez, as also noted by Gill (1967b) in September-October 1964, though they were present in April (Bourne, 1968). Anous stolidus Common Noddy None were seen at sea until we approached the Cargados Carajos reef where parties were feeding in mixed flocks with Roseate and White Terns Gyg7s alba. On Ile du Sud, at least 200 individuals were sitting on eggs and brooding downy young; older chicks and fledglings were running about the island. There were many dead and dying immatures; some were heavily in- fested with ticks, others by seeds adhering to the feathers, making it impossible for them to move. Parker (1970) noted large numbers of dead and dying juvenile Stevna fuscata with Anous stolidus on Goelette Island, Farquhar, apparently starving and Bailey (1968) made a similar observation on Desnoeufs Island, Amirantes. Other adult birds were gathered in groups of 50-60 at the water’s edge, where pairs were observed 7n copula. Thus, all stages of the breeding cycle were in progress. From the data of Table I it appears that breeding is well nigh continuous throughout the year at this latitude. A similar situation apparently existed on Rodriguez according to ancient records (Bourne, 1968) and recently Common Noddies with young of all sizes were observed in mid-July on Sandy I. off Rodriguez (Bourne, 1968), though they were not breeding there in late September/early October (Gill, 1967b). As we crossed the broad fringing reef of Rodriguez, Noddies Anous spp. were the only birds seen. They re- appeared off Mauritius where, on Serpent Island, they nest primarily in October and November (Vinson, 1950). 4 Anous tenuirostris Lesser Noddy As in Novemb:: 1963, this species was nestinz in low bushes Tournefortia sp. on Ile du Sud. It apparently nests continuously there (Table I). On Sandy I., off Rodriguez, there were nests, but no eggs in September/October (Gill, 1967b). The nesting dates on other Western Indian Ocean islands are: Seychelles, throughout the year (Loustau-Lalanne, 1963); Chagos, December/February and, main season, May/August (Loustau-Lalanne, 1962); Mauritius, through- out the year with a peak in September/December (Newton, 1958). Off Western Australia in the Abrolhos Is., hatching is over by mid-November, this being the only breeding season (Serventy and Whittel, 1962). This bird was not seen far out at sea.) Gygis alba White Tern Only seen at sea as we approached Ile du Sud. There were about 500 pairs on the island, some sitting on their solitary egg on branches, on pieces of coral, on the ground and even in the guano-collectors’ huts. Others were brooding downy young or feeding fledglings. It would appear that nesting in this species is continuous throughout the year at this location (Table I) as it is in the Seychelles (Loustau-Lalanne, 1963) and Rodriguez (Bourne, 1968, Gill, 1967b). ACKNOWLEDGEMENTS The authors wish to thank Chief Scientist A.R. Miller, who included this work in a very full ship’s program; Captain E.H. Hiller, officers and men of the Atlantis I; R.G. Munns, who accompanied us as photographer and the late M. Jean Vinson, formerly Director of the Mauritius Institute. The field work was sponsored by N.S.F. Grant GP281, and the paper was written while the senior author was in receipt of financial support from the National Research Council of Canada and the National Science Foundation. SUMMARY A return visit was made in June 1965 to the Cargados Carajos Shoals previously visited in November 1963. These islands are interesting as they lie between the Seychelles, where much breeding is in the northern summer, and the Mascarenes, where many species breed in the southern summer. Records are also given of all species seen at sea between Tromelin, Cargados Carajos, Rodriguez and Mauritius. Lesser Frigate-birds, Fregata aviel, were nesting on the island in Cargados Carajos. Our visit coincided with the beginning of the breeding season. Other nesting birds were Common Noddy Anous stolidus, Lesser Noddy Anous tenuirostris, White Tern Gygis alba for which a continuous breeding cycle at this latitude is inferred. Birds seen at sea include petrels, storm petrels, tropic-birds, boobies, and terns, all in small numbers at this season. REFERENCES Alexander, W.B. 1955. Birds of the Ocean (2nd edition). London:Putnam. Bailey, R.S. 1968. The pelagic distribution of sea-birds in the W. Indian Ocean. Ibis 110: 493-519. Bourne, W.R.P. 1960. The petrels of the Indian Ocean. Sea Swallow 13: 26-39. Jo Seoooeedo 1968. The birds of Rodriguez, Indian Ocean. Ibis 110: 338-344. Gardiner, J.S. and C.F. Cooper. 1907. The Percy Sladen Trust Expedition to the Indian Ocean in 1905. Description of the Expedition: PartII, Mauritius to Seychelles. Trans. Linn. Soc., London Il, Zool., 12(2): 123-129. Gill, F.B. 1967a. Observations on the pelagic distribution of sea-birds in the western Indian Ocean. Proc. U.S.Nat. Mus. 23: 1-33. ---------- 1967b. Birds of Rodriguez Island (Indian Ocean). Ibis 109: 383-390. Gill, F. B., C. Jouanin andR.W. Storer. 1970. Notes on the seabirds of Round Island, Mauritius. Auk 87: 514-521. Jouanin, C. 1963. Un pétrel nouveau de laRéunion ‘‘Bulweria bavaui’’ Bull. Mus. Hist. Nat., 35: 593-597. Jouanin, C. and F.B. Gill. 1967. Recherche du pétrel de Barau, Pterodroma baraui. Oiseau Gtiyiletslas Os Bis alsa). Loustau-Lalanne, P. 1962. The birds of the Chagos Archipelago, Indian Ocean. Ibis 104: 67-73. ---------- 1963. Sea and Shorebirds of the Seychelles. Seychelles:Gov. Printer. Nelson, B. 1968. Galapagos, Islands of Birds. London:Longmans. Newton, R. 1958. Ornithological notes on Mauritius and the Cargados Carajos Archipelago. Proc. Roy. Soc. Arts Sci., Mauritius 2: 39-71. Palmer, R.S. 1962. Handbook of North American Birds. Vol. 1. Yale Univ. Press. Parker, I.S.C. 1970. Some ornithological observations from the western IndianOcean. Atoll Res. Bull. No. 136: 211-220. Pocklington, R. 1965. Birds seen on Coco Island, Cargados Carajos Shoals, Indian Ocean. Ibis 107: 387. =---------- 1967. Observations by personnel of R.V. ATLANTIS II on islands in the Indian Ocean. Sea Swallow. 19: 38-41. Serventy, D. L. and H.M. Whittell. 1962. Birds of WesternAustralia. Perth, Paterson. Staub, F. 1970. Geography and ecology of Tromelin Island. Atoll Res. Bull. 136: 197-209. Staub, F. and J. Guého. 1968. The Cargados Carajos Shoals or St. Brandon: Resources, avifauna, and vegetation. Proc. Roy. Soc. Arts. Sci. Mauritius 3(1): 7-46. Vinson, J. 1950. L’fle Ronde et Vile aux Serpents. Proc. Royal Soc. Arts Sci., Mauritius L322 =0)2'e Watson, G.E., R.L. Zusi, and R.W. Storer. 1963. Preliminary field guide to the birds of the Indian Ocean. Smithsonian Institution, Washington, D. C. Willis, P.R. 1966. Birdwatching at sea. Oceanus 12: 2-6. S961 9S6T SS6T uoIMON ‘uel sunoA sunoA sated sutpee1q F961 S961 €961 SO6I S961 8961 ——" ‘ye ja _ a SS le eee SUL) uoysuUuTTAIOg aradoo9g Ry Ce a2) 3 qne3S ‘qneisg routpiey uo4sUuITYIOd qnej4s "29q@ "AON ‘sny aune ‘adv suno& s33o0 s330 s330 Rsddo sunoA sjsou sjsou s33do Rs3do suno& suno& autos ae sie sido sunoA "y sdd0 4 SodO suno& ATuo sunoA Sutpsos1q Moye sqnpe s3do jou suno& AqTuo sutpso1iq RH Ssdoa sunod you suncoA Aquo s3390 9uI0S sunod RH «sdBe s33o RH sysoU sated ATUo sutjsou sjsou sutpooaiq s}[npe jou Aqyuo sznpe sated sated AyUO __sutpeeaq surtpaeiq s}npe sjsou STVOHS SOLVUVD SOGVDUVO NO ONICYaUd SCUId AO SAUYOOAU “Tr S192 .L SIOJISTA syoW eqje stdhy SstijSOImMus} snouy SNPpI[O}s snouy eyeosny eura4S t[yesnop eusrs4s jotze eyesor gy ToutUL eyedoIy e[ns e[nS eayetAyoep ens snoyroed snutyjnd AP PHNDIX While this paper was in preparation, important accounts of seabirds breeding on other oceanic islands in the tropics were published. These are by Bourne (1971) on the birds of the Chagos Group (5°20' - 7935'S, 71920'-72°40'E), Schreiber and Ashmole (1970) on Christmas Island in the Pacific Ocean (2°N, 157°W) and Harris (1969) on the Galapagos Islands (1950'N - 1920'S, 89°-92°W). We wish to review the records of birds breeding on Cargados Carajos Shoals in the light of this new information. To avoid repetition, the names of the Islands will serve to identify the authors; also Tromelin to indicate Staub (1970). Puffinus pacificus Wedge-tailed Shearwater Occurs in the offshore waters of the Chagos Group during most of the year and reported to breed on some outer islands from November to February. It therefore appears to breed in the same season (southern summer) in the Cagador Carajos (Table 1) as in the Amirantes (Parker, 1970), Chagos and Round Island off Mauritius (Gill et al.,1970). In contrast, on Christmas Island in the equatorial Pacific, few are even present from November to January. The peak period for eggs is June to July and for chicks August to October, i.e. northern summer and fall. Sula dactylatra Blue-face Booby On Christmas Island it lays at all times of the year but mainly from April to October. In the Galapagos it has an annual cycle, with colonies out of phase by up to four months. In the Western Indian Ocean the November to January breeding on Cargados Carajos compares with the October to November peak on Serpent Island (Vinson in Gill et al., 1970), the October peak on Latham Island off the East African Coast, 97km south of Zanzibar (Parker, 1970), and the peak period of reproduction from November to March on Tromelin, to indicate a similar out-of-phase annual cycle. Sula sula Red-footed Booby This bird has been reported to breed in the Chagos Group in both July and September. On Christmas Island, some egg-laying occurs in all months. There are two peak periods from April to May and December to January. In the Galapagos (Tower Island only) it is suggested that the breeding cycle is less than one year to account for the different peak laying months in different years, though eggs have been found in every month. On Tromelin nesting has been noted in August, November and February in different years so the situation there might be comparable. The bird has now disappeared as a breeding species from Cargados Carajos (Staub and Guého, 1968). Fregata minor Great Frigate-bird On Christmas Island most laying is late March to May, with chicks but no eggs September to January. The birds are on individual cycles because of the greater-than-annual cycle of successful breeders. There is more precise synchrony within than between colonies. In the Galapagos there are annual breeding cycles on individual islands but breeding is not in phase on different islands. The available evidence from Cargados Carajos indicates laying from November to January as compared with a late August start on Tromelin, so a situation similar to that in the Galapagos may apply. Fregata ariel Lesser Frigatebird On Christmas Island this bird lays from late April to May. By January no eggs, chicks or adults remain. One definite breeding season during the northern spring and summer is 489-409 O- 73-7 8 implied. In the Cargados Carajos nesting was also observed starting in late April (Staub and Guého, 1968) at the beginning of the cool dry season. Eggs and young are also present from November to January at the start of the warm wet season. On Tromelin, 480km due west, the species was not breeding in late August, so one long breeding period seems less likely than two discontinuous ones at this latitude. Anous stolidus Common Noddy In the Chagos there are apparently two main breeding seasons, May to August and Decem- ber to February but nesting from September through November has also been reported. On Christmas Island eggs have been found throughout the year but fewer from July to November than in other months. In the Galapagos, breeding is continuous but there are only a few eggs from August to October. Anous tenuivostris Lesser Noddy This bird nests in trees in the Chagos in June and November through December. On Christmas Island there is one definite breeding season in the northern spring, laying mainly from April to May. On Serpent Island both A. stolidus and A. tenuivostris nest primarily in October and November (Vinson in Gill et al., 1970). On the basis of these comparative data there may be discontinuities in the breeding of these species on Cargados Carajos during those months, in particular February to May, which corresponds to the northern fall non- breeding season at Christmas Island. This statement supercedes our previous inference about the two noddies (see pp. 3, 4). Gygis alba White or Fairy Tern In the Chagos Group it has been found nesting in trees in all islands in February, May to July, September and October, and December. On Christmas Island substantial numbers of eggs are laid in all months, the minimum being around November. By analogy, continuous breeding at Cargados Carajos is therefore likely. ADDITIONAL REFERENCES Bourne, W.R.P. 1971. The birds of the Chagos Group, Indian Ocean. Atoll Res. Bull. 149: 175-207. Harris, M.P. 1969. Breeding seasons of sea-birds in the Galapagos Islands. J. Zool., Lond. 159: 145-165. Schreiber, R.W. and N.P. Ashmole. 1970. Sea-bird breeding seasons on Christmas Island, Facific Ocean. Ibis 112: 363-394. ATOLL RESEARCH BULLETIN : No. 159 GEOMORPHOLOGY AND VEGETATION OF ILES GLORIEUSES by R. Battistini and G. Cremers Issued by THE SMITHSONIAN INSTITUTION Washanptomee D.C. Ut. Ar. December SIs Oe SesneT10[D saj{ JO uotqeo0T “7 aunbr7 S94jeuwW UI S4NO}UOD SUulUDWGNS yudg uaskeg SESNIdO|O | (o) QUOWOD epudUuO GEOMORPHOLOGY AND VEGETATION OF ILES GLORIEUSES by R. Battistini’ and G. Cremers’ INTRODUCTION The present study was carried out with the assistance of the O.R.S.T.O.M. oceanographic vessel Vauban, a 24.5m trawler. The field party, which spent the three days 28, 29 and 30 January 1971, on the islands, consisted of R. Battistini, geomorphologist, of the Universite de Madagascar, G. Cremers, botanist, of O.R.S.T.O.M., and A. Crosnier, director of the Centre O.R.S.T.O.M. at Nossi-Bé. The Iles Glorieuses are situated north of the Mozambique Channel, at 11°30'S latitude and 47°20'E longitude. The group consists of two islands, Grande Glorieuse and Ile du Lys, located on a large coral platform 17m long and aligned southwest-northeast. The outer slopes of the platform are extremely steep (between 20° and 35° on the south and southeast sides between 75 and 1500m depth), suggesting that the coral has a volcanic basement. No volcanic rocks outcrop, however, and the depth of the basement beneath the coral is not known. There is a meteorological station on Grande Glorieuse, maintained by the Service Météoro- logique de La Réunion, which has operated for about ten years. The mean annual rainfall is 1012mm, and the mean monthly figures are as follows: Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dee ——_S oe ———— —_—_—s—_#éeoromo= MOBS UIDAKD Bile Cet Bot GA). S30) Bot ato Sih AWG HO) The prevailing winds are the Southeast Trades, with 42.3% of the observations in the sectors ENE, E and ESE, except for January and February when the dominant winds are W, WNW and NW (37.9% in January, 38.8% in February). ACKNOWLEDGEMENTS We thank the Centre Océanographique of O.R.S.T.O.M. at Nossi-Bé, and especially its Director, M. Alain Crosnier, for the welcome andassistance given to us during this expedition. We thank also Commandant Fiuric of the Vauban and his crew. M. Poonoosawny, in charge of the meteorological station, and his three colleagues also assisted in this work, and we are glad to thank them here. We thank the Service Météorologique de La Réunion, and particularly M. Malick, for the climatic statistics. Cremers also thanks Dr. J.-L. Guillaumet and A. Rakotozafy for their valuable aid. ‘Laboratoire de Géographie, University of Madagascar. “Laboratoire de Botanique, Office de la Recherche Scientifique et Technique Outre-Mer (O.R.S.T.O.M), B.P. 434, Tananarive, Madagascar. (Manuscript received Dec. 1971--Eds.) GEOMORPHOLOGY OF THE ILES GLORIEUSES by R. Battistini Grande Glovieuse Grande Glorieuse, at the southwest end of the reef, is oval-shaped, with a maximum length of 2300m and width of 1700m. It is a low sandy island, formed during the Flandrian by the accretion of a large number of parallel beach-ridges. The beach-ridge morphology is ob- scured in the east and northeat by dune-formation: on this side of the island there is a large dune field, reaching a height of 12m, covered and fixed by a dense vegetation of Casuarina and other species. The northeast and east coasts are the only ones to have a narrow zone of active dunes, with nebkas and small parabolic dunes, formed by the dominant easterly winds. On the south and southeast coasts the beach crest is a sandy storm ridge, completely covered with a shrubby vegetation of Scaevola and Cordia, and without dunes. Water from storm waves covers the vegetation and reaches up to 10m inland from the beach crest. The west coast, formed by lower flatter beach ridges, also lacks dunes. The centre and western part of the island is occupied by a large but now abandoned coconut plantation. In the south and southwest the Flandrian sand deposits abut against the karst-eroded re- mains of an older reef structure which we correlate with the Karimbolian high stand of the sea (probably about 125,000 yr B.P.); these remnants are named Ile aux Crabes, Rocher Champignon, and Rochers du Cap Vert. They consist of banks of limestone with Tvidacna and with large coral heads in the position of growth, covered at Ile aux Crabes with shelly lime- stones with rolled coral debris, reaching up to about 3m above the level of present highest tides. The limestones are deeply dissected into large lapiés, with tidal notches and over- hanging visors overlooking a low-tide platform, smooth or with enclosed pools, in the same old limestones. These limestone remnants include in their upper parts inclusions, brown to black in colour, some of several cubic metres in size, which appear to have formed in old solution holes and are now consolidated. The upper surfaces of the inclusions are eroded like the rest of the limestones. Bone fragments and pieces of carapace of an unidentified tortoise have been obtained with difficulty from these inclusions. Similar inclusions, but without bones, have been described at Europa (Battistini, 1965, 1966; Berthois and Battistini, 1969). The coast zonation in the west and south is as follows: (1) the beach, which is always sandy. (2) beachrock, of which there are only three outcrops, two on each side of Ile aux Crabes, and one to the southeast of Cap Vert. (3) a platform of reef limestone, 300-500m wide, whichdries almost completely at low water; as at Europa, this is probably largely an eroded pre-Flandrian structure. (4) a rubble zone of small boulders. Beyond the rubble zone, the grooved outer slope had little living coral at the tw» places where it was observed. In shallower water it is covered with banks of Halimeda sand, with, in deeper water, isolated coral heads, which are more numerous between 15 and 35m depth. Ile du Lys Ile du Lys, about 600m long, is muchsmizller than Grande Glorieuse. It is almost entirely formed of reef limestones with Tvidacna and coral heads in the position of growth and by stratified beds of reef debris of pre-Flandrian age. On the north sid2, exposed to the heavy swell, the limestones are eroded along their whole length into lapiés, in places with overhangs 4-6m high. The northwest point is a karst-eroded plateau 3-5m above the level of present highest tides, scattered with large coral heads in the position of growth. The old coral lime- stone is here covered by stratified deposits, dominated by Halimeda debris; some of thes? deposits seem old, but others are recent, and are probably storm deposits cemented to form beachrock, since they contain copper nails from a wreck. The coast zonation on the north side is as follows: ) storm beach overlying the coral limestone. ) high-tide platform with lapis and pools with undercut edges. ) cliff with large pinnacles, with or without visor, 4-6m high, cut in old coral limestone. ) low-tide platform, with large smooth grooves perpendicular to the shore, cut in the same old limestone. (5) shallow channel with some patches of Halimeda sand and marine grasses. (6) rocky outer rim, 0.5-1m in height, consisting of old coral limestone with superficial in- crustation by recent calcareous algae. (7) outer slope, with some large living coral heads. Bedded detrital limestones, with dips of up to 1528 formed of coral and shell debris, out- crop on the east and west sides of the island. These deposits overlie the massive coral lime- stones of the north, but, with the exception of som? outcrops near the leeward point which are probably Flandrian beachrocks, they are placed in the same period (Karim olian). In the centre of the island there is a shallow pool, with outflow to the north, with a vegeta- tion of Pemphis acidula. The pool is tidal, and is reached by storm waves across the shrub- covered storm ridge on its northern side. The whole of the leeward or southern part of the island consists of guano mixed with beds of Halimeda sand, of a yellow to light brown colour, with a vegetation of Tournefortia. This area includes the highest point on the island (11m). A sand spit is forming towards the south on this leeward side, and is colonised by a few patches of grass. The island is inhabited by thousands of rats, which probably arrived either on the wrecked ship the remains of which are visible near the pool outflow in the north, or during a recent attempt at settlement from which a cistern and some other articles remain. There is now only a small colony of terns, preserved from the rats on mushroom rocks of Halimeda limestone at the western end of the island. The Reef flat and Roches Vertes The upper surface of the reef was traversed with an outboard to the south of Ile du Lys and between Grande Glorieuse and Roches Vertes. It is almost entirely sandy, with higher sandy banks drying at low water. The Roches Vertes form a group of four rocks entirely constituted of the same reef limestone, with large coral heads in the position of growth, that outcrops at Ile du Lys and in the south of Grande Glorieuse, of Karim»bolian age. Their upper surface, with small-scale karst erosion, carries a skeletal soil with a low halophytic vegetation of Sesuvium portu- lacastrum, at least on the two southwest Roches. A very large colony of terns survives here. 4 The coral limestone has inclusions of brown and black material, with bone fragments, identical to that described on Grande Glorieuse and also found in the northeast of Ile du Lys, which we interpret as the lithified fill of ancient solution holes. The Roches are dissected into large lapiés and mushroom rocks, standing on a low-tide platform of the same old coral limestone. CONTRIBUTION TO BOTANICAL KNOWLEDGE OF ILES GLORIEUSES by G. Cremers Formation and evolution of the islands The Iles Glorieuses are formed of coral andalgal sand, with outcrops of coral limestone on the west coast of Ile du Lys, at Cap Vert on the southwest coast of Glorieuses, at the small Ile aux Crabes to the south of the main island, and forming the Roches Vertes. The process of development of the islands is most apparent on Glorieuse,. which covers 4 sq km. In the field we observed zones of parallel dunes, especially in the southern part of the island. Air photographs taken by the French Army based at Diego Suarez have confirmed that this is an area of progradation. Conversely in the north-north-west is a zone of erosion. The general tendency is towards an increase in the surface area of the island. This evolution has resulted in a zonation of vegetation as well as of relief. From the periphery towards the centre the main zones are: (1) the beach, (2) the zone of dunes, well-marked on the south and east coasts but eroded away in the north, (3) a shallow depression on the landward side of the dunes, swept by waves overtopping the dunes during periods of high tides and cyclones, the central zone, covering the main part of the island. This has a dense vegetation of shrubs and small trees, with a few taller trees. It is probable that the whole central area was once covered by this vegetation, whichis now confined to the east and south. The west of the island is occupied by a coconut plantation, now abandoned and more or less invaded by other species. Between the plantation and the natural woodland there is a zone of degraded vegetation. Finally the existence of the airstrip has favored the estab- lishment of numerous herbs not found elsewhere. (4 ~~ Botanical history of the islands The first botanical report on the Iles Glorieuses which has been found is that of Coppinger (1882). He found ‘‘a dense vegetation of primary forest’? on Grande Glorieuse, and on Ile du Lys a wooded region with Ficus, Hibiscus and Scaevola, and with Pemphis in the mangrove. The islands have been officially administered by France since 23 August 1892, although a French colonist, M. Caltaux, had already lived there for ten years. Abbott (1893) found 30 species of plants on Grande Glorieuse, and trees 15-20m high in the mangroves on du Lys. The vegetation has clearly been much altered and impoverished since this time. Nicoll (1908) also visited the islands. According to the report of Captain Lebegue on 16 November 1921, the plantation on Grande Glorieuse consisted of 6000 coconut palms, with a population of 17 Seychellois. The settlement was in the northwest part of the island. The vegetation, apart from the plantation, som? maize, and Casuarina, was a stunted serub. The Ile du Lys was uninhabited, but there were numerous traces of attempts at ex- ploiting the guano. A herd of about 200 goats was on this island. A) While it was thought that the islands were uninhabited, the visit of the lightship Marcus Moutet in February 1954 revealed that a colonist from *he Seychelles had lived there for about a dozen years and the plantation had increased to about 15,000 trees. At the tim= of our visitin 1971, there were four men from Réunion, members of the Service Metéorologique de la Réunion living on Grande Glorieuse. The herd of goats had disappeared from Ile du Lys, but had been replaced by a larger number of rats. Vegetation Grande Glorieuse: On this island of 4 sq km, 43 species of plants have been found, some of them Drought by man. These include Cocos nucifera, but also Carica papaya, Flacourtia vamontchii (the fruits of which are eaten), Ricinus communis, and perhaps Catharanthus sp. (1) Dune belt (Plate 13): As already pointed out, there is a well developed zone of dunes in the south of the island, but they have been eroded away in the north. The vegetation consists of a woodland 4-5 m high at the top of the dunes, the width varying with the terrain. This woodland consists essentially of three species of small trees. One with orange flowers, particularly abundant, is Cordia subcordata. The others are Scaevola taccada and Guettarda speciosa. On these trees is a lichen of the genus Rocella and the parasitic vine Cassytha filiformis. The trees are all heliophiles, with leaves at the ends of branches, forming a canopy with an understorey of Achyranthes aspera. Suriana maritima is another shrub scattered through the trees. In the open areas there are several creeping herbs, suchas Launea sarmentosa, Sida ovata, Boer- havia vepens, anda sterile grass which has not beendetermined but which may be Tvricholaena monachme (no. 1356). (2) Depression inland from the dunes (plate 13): In this depression, submerged during high tides, one finds almost exclusively the sedge Fimbristylis abbreviata. The young plant forms a tuft, which increases in diameter as the plant grows; the inner parts die so that the living parts of mature plants form rings (Plates 14 and 15). One finds also certain species from the dune belt, suchas Achyranthes aspera, which is mich more stuntedthan inthe shade, an unidentified grass (no. 1356), another (no. 1394) which may be Dactyloctenium aegyptium, and also Portulaca oleracea which is not found in the interior of the island. (3) Centre of the island: It is the area south of the airstrip which we have attempted to study in greatest detail, in spite of the difficulties of penetrating the vegetation of smzll trees and shrubs. The area to the north has been much influenced by man, with the coconut plantation, the old and the new meteorological stations, the trial airstrip, and the present airstrip at an angle to the first. (a) Natural vegetation (Plate 16): This consists of a dense woodland 2-4m high. The most num=2rous tree is Tournefortia argentea, forming small dome-shaped strongly-branched trees which were found in all stages of flowering and fruiting. The other speciesis Scaevola taccada, each plant covering a smaller area than one of Tournefortia. This species was fruiting, although it was in flower on the dunes. Parasitic Cassytha filiformis is very common on these trees. Several large isolated trees dominate the area: Cordia subcordata and Guettarda speciosa (fruiting). The herbaceous stratum is almost non-existent, except in openareas, where we find the sterile grass (no. 1356) and Fimbristylis abbreviata. (b) Degraded vezetation: Along the airstrip, apart from two trees, Flueggea microcarpa and Colubrina asiatica, one finds mainly prostrate herbs of which the seeds have probably been brought by aircraft. They include Sida rhombifolia, two species of Phyllanthus (nos. 1368 and 1369), one species of Cyperus (no. 1383), and Portulaca oleracea. In the coconut plantation, Casuarina equisetifolia, which was planted along the shore as a windbreak, has multiplied greatly since the plantation was abandoned and is now found through- out. The same is true of the very abundant Ficus sp. (no. 1410) Several other species are widely distributed, such as Fluegga microcarpa, Colubrina asiatica, and tufts of Cyperus sp. (no. 1383). At the edge of the plantation is a big Terminalia catappa, with Stachytarpheta jamaicensis and Ipomoea sp. (nd. 1390) beneath. Several Flacourtia ramontchii have been planted along the paths. In the north of the island, other species are growing at the site of the former meteorologi- cal station and at the cemetery. Human influence is clear in the importation of Carica papaya, Ricinus communis, Gossypium brevilanatum, and probably also of Acalypha sp. (no. 1374), Adenia sp. (no. 1385), Caesalpinia bonduc, Catharanthus sp. (no. 1375), and Ipomoea sp. (no. 1379). (ce) Pool: There is a small tidal pool in the northwest part of the island, which dries completely at low water. Round its margins growthe shrub Suriana maritima, herbs, Sporobolus virginicus, and an undetermined sterile grass which may be Tvicholaena monachme (no. 1378). (d) Beach: Several different species are found on the shore in the north of the island. These are Sporobolus aff. virginicus, Ibomoea pes-caprae, Tribulus cistoides, and Wedelia biflora. Ile du Lys: This much smaller island, a few hundred meters long, is poor in species, with a total of only 8. Overtopping by waves during very high tides may be the reason for this poverty. One can see no zonation in the little vegetation which now exists on the island. Trees of Tournefortia argentea, characterized by their dome-shaped appearance, are found almost everywhere round the periphery of the island. In the higher southern part, however, they are much bigger, reaching a height of 5m and crown diameters of 10-12m (plate 17). The small trees growing amongst them are Thespesia populnea, also in full flower. Behind these trees one finds several trees of Ficus sp. (no. 1403), completely leafless at this time, though on some buds have started to develop. This Ficus is close to no. 1410 of Grande Glorieuse and is not the Ficus marmorata found at Europa I. (Bosser, 1952). In the east there are largeclumpsof Sclerodactylon macrostachyum. Inthe west a sterile sedge (no. 1406), which could not be determined, grows in crevices in the coral rock. In the northwest there is a pool open to the sea: around it and within it there are very numerous shrubs or small trees of Pemphis acidula (photo 19). The centre of the island is formedbya large flat plain. There are two clumps of coconuts, comprising five plants in total. All around, plants of a Boerhavia species (no. 1408), although completely desiccated, form the first stages of vegetation (plate 18). Roches Vertes The Roches Vertes are always emerged, though constantly swept by waves and spray. A patch of Sesuvium portulacastrum has been found. Flora The plant specimens collected during the expedition are deposited in the Herbarium, Laboratoire de Botanique, Centre O.R.S.T.O.M., Tananarive, Madagascar, and som? have been sent to the Herbarium, Muséum National d’Histoire Naturelle, Paris. Grande Glorieuse Acalypha sp. Euphorbiaceae Achyranthes aspera L. Amaranthaceae Adenia sp. Passifloraceae Boerhavia sp. Nyctaginaceae Boerhavia repens L. Nyctaginaceae Caesalpinia bonduc (L.) Roxb. Caesalpiniaceae Carica papaya L. Caricaceae Casuarina equisetifolia L. Casuarinaceae Cassytha filiformis L. Lauraceae Catharanthus sp. Apocynaceae Cocos nucifera L. Palmae Colubrina asiatica (L.) Brongn. Rhamnaceae Cordia subcordata Lam. Boraginaceae Cyperus sp. Cyperaceae Dactyloctenium aegyptium (L.) Willd. (?) Gramineae Ficus sp. Moraceae Fimbristylis abbreviata Boeck. Cyperaceae Flacourtia ramontchii L’Hérit. Flacourtiaceae Flueggea microcarpa Bl. Euphorviaceae Gossypium brevilanatum Hochr. Malvaceae Guettarda speciosa L. Rubiaceae Ipomoea pes-caprae (L.) R. Br. Convolvulaceae Ipomoea sp. Convolvulaceae Launaea sarmentosa (Willd.) Kuntze Compositae Phyllanthus sp. Euphorbiaceae Phyllanthus sp. Euphorpiaceae Portulaca oleracea L. Portulacaceae Ricinus communis L. Euphorbiaceae Rocella sp. Lichenes Scaevola taccada (Gaertn.) Roxb. Goodeniaceae Sida ovata Forsk, Malvaceae Sida rhombifolia L. Malvaceae Sporobolus virginicus Kunth Gramineae Sporobolus aff. virginicus Kunth Gramineae Stachytarpheta jamaicensis (L.) Vahl Verbenaceae Suriana maritima L. Simaroubaceae Terminalia catappa L. Combretaceae Thespesia populnea (L.) Sol. ex Correa Malvaceae Tournefortia argentea L. f. Boraginaceae Tribulus cistoides L. Zygophyllaceae Tricholaena monachme Stapf. & Hubb. (?) Gramineae Wedelia biflora (L.) DC. Compositae Ile du Lys Boerhavia sp. Nyctaginaceae Cocos nucifera L. Palmae Cyperaceae indet. Ficus. sp. Moraceae Pemphis acidula Forst. Lythraceae Sclerodactylon macrostachyum (Berth.) A. Cam. Gramineae Thespesia populnea (L.) Sol. ex Correa Malvaceae Tournefortia argentea L. f. Boraginaceae Roches Vertes Sesuvium portulacastrum (L.) L. Aizoaceae From this list it is apparent that the species found on Ile du Lys and Roches Vertes are not found on Grande Glorieuse, in spite of the small distance between them, and this can be explained by the edaphic differences between the islands. The species found only on Ile du Lys are Pemphis acidula, Sclerodactylon macrostachyum, Ficus sp., a species of sedge, and Boerhavia sp. Sesuvium portulacastyum is only found on Roches Vertes. Two species are not found in Madagascar. Some are pan-tropical. and are found every- where, others are cultivated; in fact these last have a very well-marked distribution and are most numerous on the west coast. However Launea sarmentosa, Scaevola taccada, Tournefortia argentea are only found onthe east coast. Sporobolus aff. virginicus is very similar to the form on the east coast. One species is a Malagasy endemic, Gossypium brevilanatum, and has clearly been brought by man. The two species not known in Madagascar are Wedelia biflova and Sida ovata. The first is found on the shores of the Indian Ocean, in the Comoros and at Juan de Nova I. The second has been collected at Tromelin I. by Paulian in 1953 (unnumbered specimen). According to Hutchinson and Dalziel (1954) this species is distributed in Africa with extensions to Arabia and India. Three species (Colubrina asiatica, Fimbristylis abbreviata, Launaea sarmentosa) are not recorded from Europa or Juan de Nova. No comparison has been made with the islands to the north of Iles Glorieuses. Certain species recorded by Hemsley (1919) have not been found during the present investi- gation. These are: Apodytes mauritiana Planch. (Icacinaceae) =A. dimidiata E. Mey. Distribution: East Africa, Mascarenes, Comoros, Madagascar, Cosmoledo, Astove, Aldabra. Boerhavia diffusa (Nyctaginaceae). Pantropical. Cleome strigosa Oliv. (Capparidaceae). East Africa, Madagascar, Aldabra, Astove. Cucumis trigonus Roxb. (Cucurbitaceae). East Indies. Cyperus rotundus (Cyperaceae). Pantropical and subtropical. Eragrostis tenella (Gramineae). The name of the author was not given by Hemsley, and it is hence not possible to define this species, which may be related to: E, tenella Benth. =E.interrupta Beauv. Africa, Asia, tropical Australia; FE. tenella Nees=E£. pilosa Beauv. Tropical regions; E, tenella Roem. et Schult. = £. plumosa Link. Tropical Asia and Africa; but is more likely: E, tenella (L.) eee - E, amabilis (L.) W.& A. Euphorbia prostrata Ait. (Euphorbiaceae). Pantropical. Ficus aldabrensis Baker (Moraceae). Aldabra, Assumption, Cosmoledo, Astove. = F. reflexa Thunb. Ficus nautarum Baker (Moraceae). Seychelles, Aldabra, Assumption, Cosmoledo, Astove, St Pierre. Hibiscus hornei Baker (Malvaceae) = H. physaloides Guill. et Perr. Tropical and subtropical Africa, Comoros, Madagascar; probably introduced. Ipomoea glaberrima Boj. ex Hook. (Convolvulaceae) =/. macrvantha R.&S. Tropical coasts and islands. Plumbago aphylla Boj. (Plumbaginaceae). Aldabra, Assumjtion, Cosmoledo, Astove, Mada- gascar. Portulaca quadrifida L. (Portulacaceae). Tropical coasts and islands. Sida spinosa L. (Malvaceae). Pantropical. Solanum nodiflorum Jacq. (Solanaceae) =S. nigvum L. Pantropical. Some of our undetermined species may correspond with some of those in this list. Conclusion It is clear from the reports of early visitors that the vegetation of the islands has been transformed. It appears that the vegetation was formerly quite dense: primary forest in 1882; trees 15-20m tall in the mangrove of Ile du Lys in 1893. In 1921 a coconut plantation replaced part of the forest and only shrubby vegetation remained on Grande Glorieuse. On Ile du Lys there is no longer any trace of big trees in the mangrove. If the number of species present has increased, from 30 in 1893 to 48 in 1971, the in- fluence of man is the main reason. Some herbaceous species have been introduced by aircraft following the construction of the airstrip. As Capuron (1966) has suggested for Europa I., since these islands have no economic value, it would seem useful to protect their fauna and flora in order to be able to study their natural development. REFERENCES Abbott, W.L. 1893. Notes on the natural history of Aldabra, Assumption and Glorioso Islands, Indian Ocean. Proc. U.S. Nat. Mus. 16: 759-764. Battistini, R. 1964. L’extreme Sud de Madagascar, étude géomorphologique. Paris: Editions Cujas, 636 pp. Soo booooaS 1965. Note préliminaire sur la morphologie de Vile Europa. Madagascar, Revue de Geographie 6: 37-59. Berthois, L. and Battistini, R. 1969. Etude sédimentologique de Vile Europa. Madagascar, Revue de Géographie 15: 7-52. Bosser, J. 1952. Note sur la végétation des Iles Europa et Juan de Nova. Le Naturaliste Malgache 4: 41-42. Capuron, R. 1966. Rapport succinct sur la végétation et la flore de l’ile Europa. Mém, Mus. Nat. Hist. Natur. A (Zool.) 41: 19-21. Coppinger, R.W. 1883. Cruise of the ‘‘Alert’?: Four years in Patagonian, Polynesian and Mascarene waters (1878-1882). London: W. Swan Sonnenschein, 256 pp. 10 Guilcher, A., Berthois, L., Le Calvez, Y., Battistini, R., and Crosnier, A. 1965. Les récifs coralliens et le lagon de Ile Mzyotte (Archipel des Comores, Océan Indien). Mém. O.R.S.T.O.M. 8: 1-210 (Iles Glorieuses, py. 13-14). Hemsley, W.B. and others. 1919. Flora of Aldabra: with notes on the flora of the neighbouring islands. Kew Bull. 1919: 108-153. Hutchinson, J. and Dalziel, J.M. 1954. Floraof West Tropical Africa. Edition 2, Vol. 1, part I. ---------- 1958. Flora of West Tropical Africa. Edition 2, Vol. 1, part Il. Lebegue. 1921. Les Glorieuses. Bull. Economique de Madagascar, 1921, 4e trimestre: 289-290. Nicoll, M.J. 1908. Three voyages of a naturalist, being an account of many little known islands in three oceans visited by the ‘‘Valhalla’’? R.Y.S. London: Witherby, 246 pp. Perrier de la Bathie, H. 1921. Note sur la constitution géologique et flore des fles Chester- field, Juan de Nova, Europa et Nosy Trozona. Bull. Economique de Madagascar, 1921, 3e trimestre, 31: 170-176. Stoddart, D.R. 1967. Summary of the ecology of coral islands north of Madagascar (excluding Aldabra). Atoll Res. Bull. 118: 53-61. Sa / Roches Vertese° “4 kilometres Submarine contours in metres Evounciien | bkesiGlorteuses,) ine pant atter hydrographic ehart ‘ NEW YA 1 Q METEOROLOGICAL Yl STATION > Champignon = e / / lle aux Crabes i | Karst-eroded remnants of old Old) beach ridges @ reefrock of Karimbolian age Fixed dunes — —,. Reef.rim with grooves Wry Active dunes eo o Boulder zone MOL Storm wave deposits | on inter beach ridge Reef flat of grooved dead coral Sand beach “CD. Shallow sand with aligned sandbanks monnn br beach rock one kilometre Figure 8. Geomorphic sketch-map of Grande Glorieuse Figure 4. Section in the southern part of Ile aux Crabes beet ee : Storm ridge Flandrian Ml Zz : 5m beachridge of BL oes ala SaaS Oo large blocks a a A> ee Sao ners ie et a Shelly limestone ‘ with coral debris branches es) FARS Coral limestone with massive coral ————»> Halimeda lenses Tridacna limestone Massive coral Brown and black limestone solution - hole fill with tortoise remains Figure 5. Section in the reef-limestone outcrop of Cap Vert 489-409 O- 73 - 8 (Tut Ist 7eq@ “yY Aq usye] ANN ey} woiz yde1s0j0yd [etise ue WOIF MATA anbt][qo) SAT np a{[ FO yo eys OtTYydAowosay °9 adnb1 J Ga Quo ssw] Jadu PjO paepouse -ySUDy Ul! JJI]D PHOUSpPUNy — 3 a) = : oes ea) oe spDdey DUO BulAlq a) eel) < a mS ~~ Py ),Ul0d 1SD9 UION snnsneetF oe SN S eneeeuere ° Cea sesspuB pas yoOuyodDeg . 2 ES Se: ye yopdeq 1SEM yynos SSS AD oupn x -41ds pups pupmse) QuUOWSW!| Jadu PJO JO PAWS} SWIOJ D)d : 2 aes ee ee . ee eh nes Oe oles ae epi-Mo} peaocoub upjnBeuy| Dea eee or _ pups mojjDuS © * - yUuDG puDS poeBuawiqns-. OLD METEOROLOGICAL STATION ene) £) LAGOON COCONUT PLANTATION NEW SS METEOROLOGICAL \.’-U. STATION" oo Nee eee AIR STRIP (te aux crabes “| Thicket on coastal dunes f-.]Degraded vegetation in central area Zone behind dunes [[ljeare area in central area WI. Scrub in central area one kilometre Figure 7. Sketch-map showing vegetation distribution on Grande Glorieuse Plate 1. Ile du Lys from the northeast (oblique air photograph by Rese Ba bit staan) Plate 2. Southwest part of Grande Glorieuse (oblique air photo- graph by R. Battistini) Plate 3. South coast of Grande Glorieuse, eroded by Cyclone Félicie. Shrubs of Seaevola taecada and Cordia subeordata (photo R. B.) Plate 4. Vegetation on the southern part of Grande Glorieuse: Scaevola taecada covered with the parasitic vine Cassytha filtformis (Ghote: Renee ) Plate 5, Ile aux Crabes: coral limestone at the base, covered with bedded detrital limestone, and above this a storm ridge of large boulders. Vegetation of Pemphis actdula (photo R. B.) fe on. a Die 2 >. Plate 6. Mushroom rock and outer rubble ridge. On the horizon the rocks of;Cap Vert. (photo 'R. B.) Plate 7. North coast of Ile du Lys. Foreground: the karst-eroded high-tide platform; background: the low-tide platform with large grooves, the channel, and the outer ridge encrusted with calcareous algae (photo R. B.) Plate 8. Ile du Lys: the outer ridge encrusted with calcareous algae (photo R. B.) 7 BO Ni ne PVGEOng Tle du Lys: karst-eroded surface in the extreme northwest, on the bedded Haltmeda limestones. In the background, the west beach (photo R. B.) TEER ORR YET Plate 10. Ile du Lys: basal coral limestone overlain by bedded Haltmeda limestone, eroded into a projecting visor overlooking the low-tide platform. In the background, the outer ridge (photo R. B.) Plate 11. Ile du Lys: the inner pool seen from the southern guano area. Bushes of Pemphis acidula round the pool (photo R. B.) cae oe Plate 12. East coast of Ile du Lys: storm beach overlying old bedded detrital rocks. In the background, a seasonal sandspit (photorR= Be) Plate 13. Grande Glorieuse: depression inland from the dunes, with Fimbrtstylts abbreviata, at the foot of dunes covered with Cordta subcordata, Guettarda speciosa and Scaevola taccada. In the depression one can see traces of lines of dunes now eroded (photo G. Cremers) : eS Plate 14. Young tuft of Fimbristylis abbrevtata on Grande Glorieuse (photo G. Cremers) Plate 15. Old tuft of Fimbristylis abbreviata on Grande Glorieuse (pheto @. Cs) Plate 16. Scrub in the centre of Grande Glorieuse, with Tournefortta argentea and Seaevola taecada Goose! 1G aCe) Plate 17. Zone of Tournefortia argentea on the south coast of Ile du iyis (photo, Ger Ca) Plate 18. Central area of Ile du Lys, with completely dessicated Boerhavta sp. In the background, dome-shaped Tournefortia argentea (photo G. C.) Plate 19. lle du Lys: small pool open to the sea, with numerous bushes of Pemphis actdula (photo G. C.) ou é 7 a if 4, ; . 5 ATOLL RESEARCH BULLETIN No. 160 REEF ISLANDS OF RAROTONGA by D. R. Stoddart LIST OF VASCULAR FLORA INP Ig ING WOSsloeiess Issued by THE SMITHSONIAN INSTITUTION WEislanlavenoas D6 (Gog) Worsia Jvc December *Ssik, wWGire nn fe) fe) je} o : ie h Ni | ‘ll E a et | CF & \i REEF ISLANDS OF RAROTONGA by D.R. Stoddart INTRODUCTION Remarkably little is known of the coralreefs of Rarotonga, southern Cook Islands. Cross- land (1928a, 616-619) and Davis (1928, 407-408) gave brief descriptions following short visits, and remarks on reef structure in the context of the geological history of the island have been made by Marshall (1908, 1912, 1930) and later workers (Wood, 1967). A marine biology party from the Cook Bicentenary Expedition to the Southwest Pacific worked on Rarotonga from 21 to 27 August 1969. The party consisted of Dr. H.G. Vevers (Zoological Society of London), Dr. P.E. Gibbs (The Marine Laboratory, Plymouth), and the author. After a reconnaissance of the coast of Rarotonga, work was concentrated at Ngatangiia Harbour on the east side of the island, though some collections were also made at other locali- ties. Subsequently the Ngatangiia area was also visited by Prof. W.R. Philipson (Department of Botany, University of Canterbury, Christchurch, New Zealand), during floristic studies carried out on the same expedition, and some of his results are used here. The present paper gives a general description of the reefs and environment of Rarotonga, a detailed account of the reef islands of Ngatangiia, and a list of the vascular flora of the islands determined by Dr. F.R. Fosberg (Smithsonian Institution). Other results of the Ngatangiia work will appear elsewhere. A preliminary account of coral reef studies in the Cook Islands during the 1969 expedition hasalready been published (Gibbs, Stoddart and Vevers, 1971). REEF GEOMORPHOLOGY Rarotonga (latitude 21912'S, longitude 159°46'W) is anisolated volcanic island, now deeply dissected by erosion, with a maximim altitude of about 640m. It rises from the ocean floor at a depth of about 4000m (Summerhayes and Kibblewhite, 1967), at which depth the volcanic cone is 45-55km in diameter (Figure 1). The present island has maximum dimensions of 11.5 x 8km. The exposed volcanics consist of basaltic and phonolitic eruptives; the latter have been dated radiometrically at 2.3-2.8 million years, indicating a Pliocene age for the upper- most lavas of the cone (Tarling, 1967). The volcanics are surrounded at the coast by a low apron of gravels and sands (Fig. 2). The gravels (Nikao Gravels) were deposited first and slightly cliffed before the deposition of the more recent Aroa Sands (Wood, 1967). Swamp deposits in places occupy gaps between the gravels and the beach ridges of the Aroa Sands, particularly along the south and west coasts. Small areas of elevated reef limestone outcrop in places round the coast, and may be extensive beneath the Aroa Sands; their significance will be discussed later. (Manuscript received Oct. 1971--Eds.) 489-409 O- 73 - 9 The island is surrounded by a fringing reef about 400m wide along the south and 200m wide along the west and north coasts. In Ngatangiia Harbour the reef edge lies about 1km from the coast. The reef edge is continuous except for steep-sided narrow inlets at Avatiu and Avaroa on the north coast, Ngatangiia on the east, and several places on the south. The reef flats are planed-rock features, rarely carrying more than 1.5m of water even at high tide; they are covered with sand sheets, and growing corals are not common. Marine phanerogams are absent: These flats are distinguished, especially at Ngatangiia Harbour, by extremely large populations of holothurians, with densities of Holothuria atva reaching up to 10 per sqm. An algal ridge is only weakly developed onthe reefs, though Marshall (1930) states that it is higher in the south than the north. Crossland (1928a, 616-619) considered that the reef was originally a barrier reef converted to a fringing reef by infilling of the lagoon by the coastal sands and gravels. He said little of the composition of the reef, except to note the existence of large brain corals and also of Porites heads in the boulder beach at Avarua (Crossland, 1928b, 721). Davis (1928, 407-408) rather misleadingly referred to Rarotonga’s ‘‘close-set and little-eroded barrier reef, from a quarter to a half mile wide, now about 15 feet above sea level, enclosing a narrow lagoon flat or Swamp’’; he considered that the amount of sediment produced during dissection of the island, both before and during reef growth, required considerable subsidence for its disposal, and that this overall sinking of the island was to be distinguished from the more recent 5m uplift of what he termed the barrier reef, and the subsequent infilling of its lagoon. Ngatangiia Harbour on the west coast forms the most pronounced coastal indentation. Rarontonga’s largest river, the Avana, flows into the harbour and discharges by the deep reef ‘gap north of Motutapu (Fig. 3). Outcrops of elevated reef limestone are found on both sides of the harbour entrance. The Harbour islands, from Motutapu southwards, continue the general trend of the coast (though Taakoka is volcanic, not detrital). Wood (1967) maps the detrital islands as Aroa Sands, and both Marshall (1930) and Wood (1967) draw attention to the absence of sands and gravels on the mainland coast of Ngatangiia Harbour, where volcanic rocks reach the sea. Marshall (1930, 19-20) proposed that the detrital islands represented fragments of a formerly continuous beach ridge breached by hurricanes, with as a result the sea flooding the swampy former back-ridge area. ENVIRONMENT Rarotonga lies in the South-east Trade Wind belt. There are no climatic data for the Ngatangiia area, on the windward, wetter side of the island. Avarua, on the leeward side, has a mean annual rainfall of about 2050mm, though with considerable variation between years. Most of the rain falls during March, Apriland May; and the driest months, with less than 100mm average, are July and September. Mean annual temperature is 23.6°C. Highest recorded temperature near sea level at Avarua is 33.5°C, andlowest 8.9°C (Marshall 1930; Grange and Fox 1955). The island is frequently affected by storms of hurricane intensity, generally approaching from the northwest. Major recorded storms are those of December 1831, December 1842, March 1846 (especially damaging at Ngatangiia), Decem%er 1848, March 1926, March 1943, January 1944, January 1946, December 1948, and September 1950 (Gill, 1885; Marshall, 1930; Hutchings, 1953), but this list is clearly incomplete. Because of the open-ocean situation, storm surges associated with hurricanes are not important; most of the effects are from wind and wave activity. Tsunamis also occasionally occur. Those of May 1960 resulting from the Chilean earthquakes reached the comparatively low height in inlets at Rarotonga of 1.5m above normal sea level, partly because the tide was low at the time and normal water level was below the outer reef edge (Keys, 1963). Both hurricanes and tsunamis, however, are likely to have been significant controls in the development of the reef islands. 3 Tides are semi-diurnal, with a rather pronounced diurnal inequality at and following neaps. According to predictions prepared by the Hydrographic Department, Ministry of Defence, London, for the expedition, in 1969, the range at springs is 0.85m and at neaps 0.33 m. DESCRIPTION OF THE ISLANDS Motutapu (Fig. 4) Motutapu is the northernmest and largest of the Ngatangiia islands: it is 600m long, 360m wide, with an area of 11 ha. The northern part of the island consists of a rough makatea or elevated reef limestone, well cemented, with a local relief of up to 1m though with a rather subdued erosional topography (Plate 1); it is cliffed and undercut at intertidal levels, and its upper surface stands at 2-3m above the sea (Plate 2). A small patch of similar rock outcrops further south on the seaward shore, and much of the northeastern part of the island is presum- ably underlain by this limestone. At the north point the makatea directly overlooks the deeper water of the harbour entrance, and similar rock outcrops on its northern side. Apart from the makatea the island is a simple cay, with a gravel and cobble ridge and intertidal storm rubble forming a beach up to5m high on its seaward side (Plate 3), anda wide intertidal expanse of fine sand and silt forming an Uca-dominated flat on its leeward side. The seaward beaches are 25-30m wide; those on the lee side are narrower as well as lower. All are aggrading except for a cliffed sector of cobble beach on the north coast. The vegetation of the makatea differs from that of the rest of the cay. Sesuvium portu- lacastrum covers the floors of potholes, with a low scrub of Wedelia biflova and Capparis cordifolia. Heliotropium anomilum and [Ipomoea pes-caprae are also present. Most of the characteristic strand plants of the sand cay are absent. On the seaward coast of the sand cay section there is a long narrow zone of Scaevola taccada, with some scattered bushes of Tournefortia argentea anda patch of Lantana camara. Outpost vegetation is restricted to small areas of Vigna marina and Cassytha filiformis. The Scaevola, in places up to 4m tall, is replaced inland by a dense woodland of Hibiscus tiliaceus 6-8m tall, with some Guettarda, Morinda, Casuarina, and occasional conspicuous tall coconuts. The interior of the cayis occupied by a higher woodland, dominated by coconuts, with Hernandia sonora, Leucaena insularum, Morinda citrifolia, Carica papaya and other spe- cies. On the low-lying, partly waterlogged lee shore there are areas of grass and sedge marsh dominated by a sterile grass (possibly Paspalum) and inhabited by Uca. Oneroa (Fig. 5) Oneroa, south of Motutapu, is slightly smaller; it lacks the makatea but is otherwise similar in topography and vegetation. The island is 500 m long and 200-250m wide, with an area of 10.6ha. The seaward beach (Plate 4) consists of sand, gravel and cobbles, with broken coral rubble at its foot and strewn acrossits surface; the coarser sediments are more common in the south- east. The beach is 20m wide and 2-3m high. The leeward beach is low and sandy, with a sand spit 100m long extending lagoonward in the north. This spit encloses, as at Motutapu, a wide intertidal flat of sand and fine gravel with large num»ers of Uca. Cemented rubble forms a low shelf at the foot of the seaward beach towards the north, but otherwise there is no beach- rock. The main vegetation of the island is a2 mixed woodland dominated by Casuarina, broadleaf trees and coconuts. Hzbiscus tiliaceus is an important component, together with Hernandia sonora, Pisonia grandis, Morinda citrifolia and Leucaena insularum. Several species of ferns 4 form a ground cover. Casuarina reaches the shore in several places, and is locally being undermined by beach erosion. Scaevola taccada forms an interrupted beach-crest scrub up to 4m tall, with some Tournefortia and Guettarda, on the seaward shore. Inland from the narrow Scaevola belt is a zone oftaller Pzsonia and Hernandia woodland 40-50 m wide, largely growing on coarse beach-crest coral rubble with no other ground cover. At the abrupt inner edge of the rubble spread, on sand and fine gravel, the vegetation changes to a mixed woodland dominated by Casuarina with a ground cover of fernsandgrasses. The Scaevola belt is wider and more continuous on the lagoon shore, again with occasional Tournefortia; Sophora and Suriana are represented by isolated bushes. Outpost species on the beach outside the Scaevola zone include Cassytha, Cenchrus and Triumfetta. Koromiri (Fig. 6) Koromiri, the smallest of the cays (320mlong, 120m wide, area 3ha) strikingly resembles Oneroa in its topography, with a prominent northern sand spit. The beaches are narrower, however, and on the southeast coast cliffed and retreating. The island is distinguished by its relict beachrock extending up to 120m seaward of the cay. Apart from patches of Hernandia, Hibiscus, Morinda and Guettarda woodland, the island is covered with Casuavina woodland. Beach-crest scrub of Scaevola taccada, with some Tournefortia argentea, is most continuous on the lagoon shore, where Cassytha is abundant. Outpost vegetation is very limited, with some Vigna marina and Triumfetta procumbens. Taakoka Taakoka, the southernmost island, set well back from the reef edge, is not a sand cay but a low hill of basalt forming an island half the size of Koromiri. The island may be described in two parts. First, a central plateau, consisting of a jumble of large angular and sub-rounded basalt blocks, encrusted with lichens and covered with bryophytes, and surrounded on its sea- ward margins by large spreading trees of Barvingtonia asiatica 10-15m tall. In the centre of the plateau there are some 50 coconut trees 15-25 m tall, with Morinda, Hernandia and ferns. The plateau stands about 6m above sea level. Second, there is a “‘tail’’ lagoonward of the plateau, of basalt blocks up to 1m long. This has an outer zone of Casuarina up to 20m tall, with Vigna marina and Ipomoea macrantha beneath, and an inner zone of Hernandia, 10mtall, with Morinda, coconuts and ferns. Scaevola grows abundantly among the basalt blocks of the seaward coast, with Wedelia biflora, Vigna marina and Ipomoea pes-caprae. Along thenorthandsouth shores Barringtonia approaches close to the beach, with thickets of Hibiscus tiliaceus and outpost Vigna marina and Ipomoea macrantha. Several species are clearly introduced, though the island is not inhabited. These include not only the coconuts, but cultivated Hzbzscus and Hippeastrum, anda single banana plant. It is clear that Taakoka, so different physiographically, has little in common with the other three islands, and is .nly linked with them by proximity. Along the north shore there is a thick outcrop of beachrock, consisting of angular basalt cobbles ina red clayey matrix which is presumably a decomposition product of the basalt. The beach sands, as on the other islands, are, however, reef-derived. DEVELOPMENT OF THE ISLANDS From these surveys of the cays, it is not possible to confirm Marshall’s proposal (1930, 19-20) that the islands form part of a formerly continuous ridge of Aroa Sands fragmented by ' storm action. This could be so, but equally the islands have all the characteristics of ordinary sand cays, and they show no obvious signs of major erosion or dissection. Nor is there any evidence between the islands, e.g. in relict beachrock, of form2rly more continuous land. The islands, other than Taakoka, consist of successive increments of storm--deposited rubble and cobbles of reef origin on the seaward beaches, with infill and spit-growth of fine gravel and sand to leeward. It seems likely that these processes of aggradation are still active. The islands in their present form are not therefore directly related to the occurrence of raised reefrock remnants in the makatea of Motutapu and the mainland coast. There are also two smsll islands on the northern reef flats of Rarotonga: one near Avatiu has been much altered by land reclamation on the reef flat, but the other, Motu Toa, has probably also originated by sediment accretion to form a discrete island on the reef flat. Some data are available on the absolute chronclogy of late Pleistocene and Holocene events at Rarotonga. Schofield (1970) has published C14 gates on makatea samples: a sample from Ngatangiia Harbour, elevation 3.05m, is datedat 28,200 4+ 850 yr B.P.,and one at Te Ara Vaka, elevation 1.83m, at more than 48,900 yr. A sample from 3.2m elevation at Matavera could not be dated. Schofield suggests that the reef from which these samples were taken (equivalent to the Motutapu makatea) was formed during an interstadial at 32,000—35,000 yrB.P.; it is per- haps more probable, in view of the difficulty of dating such material, that it correlates with the makateaof Mangaia, for which Veeh (1966) obtained a uranium-series age of 110,000 + 50,000 yr. There is somes evidence from Rarotonga for much more recent high stands of the sea. A raised reef at Avarua on the north coast, at 1m above present low water level, with corals in the position of growth, has been datedat 2030 + 60 yr B.P., and soil samples from beach ridges of Aroa Sands age have been dated at1235 + 57, 2470 + 63, and 3510 + 50 yr B.P. (Wood 1967; Schofield 1970). Schofield also reports a beachrock at Titikaveka at 1m bove present high water level. He suggests that these recent dates indicate sea stands at +2 and +1m above the present, and that the Aroa Sands beach ridges, which rise to about 8m above present sea level, were formed during these higher stands. Thereis no evidence of such stands in the topography of the Ngatangiia reef islands, and indeed large parts of the islands would be subm=2rged with such higher sea levels. It should be noted that the much older Motutapu makatea is tidally notched at present intertidal levels, and there are no apparent signs of higher notches. The Ngatangiia islands can thus be assumed, from their own characteristics, to have had a simple aggradational history, punctuated by hurricane events, accumulating on reef flats partially formed by the erosion of older elevated reef limestone. It is possible that they are younger than the sea-level events described by Schofield, or that the evidence of such events in the phySiography of the islands has been erased by later storm action. SUMMARY OF VEGETATION The vegetation of the islands is relatively simple. Essentially it comprises a broadleaf woodland of Hibiscus, Guettarda, Morinda, Pisonia and Hernandia, with coconuts and Casuarina, the latter apparently spreading at the expense of broadleaf trees; a beach-crest scrub of Scaevola taccada with some Tournefortia argentea; and an outpost or pioneer strand vegetation, very patchily developed, of Tvzum/fetta, Ipomoea, Vigna and other species. The Motutapu makatea has its owndistinctive vegetation of Wedelia biflova, Ipomoea, Heliotropium, and Capparis cordifolia. IWl-drained leeward sand flats have a low vegetation of grasses and sedges. This restricted range of vegetation types reflects the limited range of habitats and the 6 small size of the islands, as well as floristic poverty. Some types, notably Barringtonia woodland, are found only on Taakoka and not on the sand cays. There iS 10 mangrove vegetation, and sea-grasses are absent, not only on Rarotonga but throughout the Cook Islands. Some species, especially littoral shrubs, are surprisingly rare, particularly Suriana maritima, which is extremely common on Aitutaki on similar cays. Wilder (1931) recorded this species as occurring only on Motutapu; it was not seen there in 1969, but a single specimen was growing on Oneroa. ACKNOWLEDGEMENTS I thank the Royal Society of New Zealandand the Royal Society of London for the opportunity to take part in the Cook Bicentenary Expedition. Our work was supported by the Cook Island Government through the Premier, Hon. Albert Henry. The late Mr. L. Peyroux acted as Ex- yedition Liaison Officer with the Premier’s Office. Iam grateful to the Rev. Bernard Thorogood and other members of the Cook Islands Library and Museum Association; to Mr. Dawson Murray of Teriora College, Rarotonga; to Prof. W.R. Philipson for making some of his own results available; to Dr. F.R. Fosberg for making the plant identifications; and to Dr. H.G. Vevers and Dr. P.E. Gibbs for assistance in the field. REFERENCES Crossland, C. 1928a. Coral reefs of Tahiti, Moorea and Rarotonga. J. Linn. Soc. Lond. 36: 577-620. Crossland, C. 1928b. Notes on the ecology of the reef-builders of Tahiti. Proc. Zool. Soc. Lond. 1928: 717-735. Davis, W.M. 1928. The coral reef problem. Amer. Geog. Soc. Spec. Pub. No. 9: 1-596. Gibbs, P.E., Stoddart, D.R., and Vevers, H.G. 1971. Coral reefs and associated commniinities in the Cook Islands. Bull. Roy. Soc. New Zealand 8: 91-105. Gill, W.W. 1885. Jottings from the Pacific. London: The Religious Tract Society. 248 pp. Grange, L.I. and Fox, J.P. 1953. Soils of the Lower Cook Group. New Zealand Soil Bureau Bull. n.s, 8: 1-55. Hutchings, J.W. 1953. Tropical cyclones in the southwest Pacific. New Zealand Geographer 9: 37-57. Keys, J.E. 1963. The tsunami of 22 May 1960 in the Samoa and Cook Islands. Bull. Seis. Soc. Amer. 53: 1211-1227. Marshall, P. 1908. Geology of Rarotonga and Atiu. Trans. New ZealandInst. 41: 98-100. Marshall, P. 1912. Coralreefs of the CookandSociety Islands. Rept. 13th Meeting Australasian Assoc. Adv. Sci. (Sydney 1911): 140-145. Marshall, P. 1930. Geology of Rarotonga and Atiu. Bull. Bernice P. Bishop Mus. 72: 1-75. Schofield, J.C. 1970. Notes on late Quaternary sea levels, Fiji and Rarotonga. New Zealand J. Geol. Geophysics 13: 199-206. 7 Summerhayes, C.P. 1967. Bathymetry and topographic lineation in the Cook Islands. New Zealand J. Geol. Geophysics 10: 1382-1399. Sumimerhayes, C.P. and Kibblewnite, A.C. 1967. Rarotonga provisional bathymetry. New Zealand Oceanographic Institute Chart, Island Series, 1: 200,000. Tarling, D.H. 1967. Som: paleomagnetic results from Rarotonga, Cook Islands. New Zealand J. Geol. Geophysics, 10: 1400-1406. Veeh, H.H. 1966. Th239/y238 and 234/238 ages of Pieistocene high sea level stand. J. Geophys. Res. 71: 3379-3386. Wilder, G.P. 1931. Flora of Rarotonga. Bull. Bernice P. Bishop Mus. 86: 1-113. Wood, B.L. 1967. Geology of the Cook Islands. New Zealand J. Geol. Geophysics 10: 1429- 1445. ir es) Ae pp iam sogyeagene * Joe Bess : ae ‘ AY ihe oe ra pine 9 yet. crivikten 4) Lene way ald iar oy LIST OF VASCULAR PLANTS by F.R. Fosberg Plants collected in 1969 and determined by Dr. F.R. Fosberg, together with those collected on the cays by Prof. W.R. Pnilipson, comprise 2 bryophytes (not determined), 1 lichen (not determined), and 49 species of vascular plants, including 4 species of ferns. Some earlier records, usually without precise location, are given in the floras of Cheeseman (1903) and Wilder (1931). One species recorded on Motutapuby Wilder, Myoporum sandwicense A, Gray, was not collected in 1969. Of the plants in the following list, excluding ferns, 26 species are recorded from Motutapu, 17 from Oneroa, 14 from Koromiri, and 18 from Taakoka. Seven of the Taakoka species, plus one fern, are recorded only from that island and not from the three reef islands, bringing the number of species from the latter down to 41. The list is interesting for its omissions, compared with other Cook Island reef islands, particularly those of Aitutaki. There are no pandans; Pemphis acidula, elsewhere abundant, is missing; and there are no species of Euphorbiaceae, though on similar islands on Aitutaki Euphorbia chamissonis iS an important component of the vegetation. Some aspects of the floristics of Rarotonga are discussed by Philipson (1971). Almost complete sets of the specim2ns cited are deposited in the U.S. National Herbarium, Washington, D.C. and in the herbarium of the Botany Division, Dept of Scientific and Industrial Research, Christchurch, New Zealand. POLY PODIACEAE Asplenium nidus L. Oneroa: Stoddart 2118, Koromiri: Stoddart 2154. Davallia solida (F. f.) Sw. Oneroa: Stoddart 2119, Taakoka: Stoddart 2167. Nephrolepis hirsutula (Forst. f.) Presl Taakoka: Stoddart 2169. Polypodium scolopendria Burm. f. Oneroa: Stoddart 2121. Koromiri: Stoddart 2157. Taakoka: Stoddart 2168. GRAMINEAE Cenchrus echinatus L. Oneroa: Stoddart 2158. Lepturus repens R. Br. Taakoka: Philipson 10349. Stenotaphrum secundatum (Walt.) O. Ktze. Oneroa: Philipson 10354, Thuarea involuta (Forst.) R. Br. ex R. and S. Motutapu: Stoddart 2135. 10 CYPERACEAE Fimbristylis cymosa R. Br. Motutapu: Phillipson 10364. PALMAE Cocos nucifera L. Motutapu: Stoddart, sight record. Oneroa: Stoddart, sight record. Koromiri: Stoddart, sight record. Taakoka: Stoddart, sight record. AMARYLLIDACEAE Hippeastrum puniceum (Lam.) Voss Taakoka: Stoddart 2159. M'JSACEAE Musa sapientum L. Taakoka: Stoddart, sight record. CASUARINACEAE Casuarina equisetifolia L. Motutapu: Stoddart, sight record. Oneroa: Stoddart 2115. Koromiri: Stoddart, sight record. Taakoka: Stoddart, sight record. PIPERACEAE Peperomia leptostachya H. and A.? Motutapu: Stoddart 2140. Peperomia pallida var. Oneroa: Philipson 10360. NYCTAGINACEAE Pisonia grandis R. Br. Oneroa: Stoddart 2114. AIZOACEAE Sesuvium portulacastrum L. Motutapu: Stoddart 2105; Philipson 10365. 11 PORTULACACEAE Portulaca lutea Sol. ? Motutapu: Stoddart 2106. LAURACHEAE Cassytha filiformis L. Motutapu: Stoddart, sight record. Oneroa: Stoddart 2117. Koromiri: Stoddart, sight record. HERNANDIACEAE Hernandia sonora L. Motutapu: Stoddart 2137. Oneroa: Stoddart 2124. Koromiri: Stoddart 2148. Taakoka: Stoddart 2164. CAPFARIDACEAE Capparis cordifolia Lam. Motutapu: Stoddart 2141; Philipson 10361. Koromiri: Stoddart 2155. LEGUMINOSAE Canavalia sericea A. Gray Motutapu: Stoddart 2133. Leucaena insularum (Lam.) Dan. Motutapu: Stoddart 2139. Oneroa: Stoddart, sight record. Mucuna gigantea (Willd.) DC, Taakoka: Philipson 10351. Sophora tomentosa lL. Oneroa: Stoddart 2127. Vigna marina (Burm.) Merr. Motutapu: Stoddart 2112. Koromiri: Stoddart 2146, Taakoka: Stoddart 2165. SURIANACEAE Suriana maritima L. Oneroa: Stoddart 2123. RHAMNACEAE Colubrina asiatica (L.) O. Ktze. Oneroa: Stoddart 2131; Philipson 10356. 12 TILIACEAE Triumfetta procumbens Forst. Oneroa: Stoddart 2132. Koromiri: Stoddart 2147. MALVACEAE Hibiscus sp. (cultivated variety) Taakoka: Stoddart 2161. Hibiscus tiliaceus L. Motutapu: Stoddart 2104. Oneroa: Stoddart 2130. Koromiri: Stoddart 2150. Taakoka: Stoddart 2166. CARICACEAE Carica papaya L. Motutapu: Philipson 10363. LECYTHIDACEAE Barringtonia asiatica (L.) Kurz Taakoka: Stoddart 2172. MYRSINACEAE Ardisia etliptica Thunb, Motutapu: Stoddart 2142. CONVOLVULACEAE Ipomoea macrantha R. and 5S. Taakoka: Stoddart 2163. Ipomoea pes-capvae subsp. byasiliensis (L.) v. Ooststr. Motutapu: Stoddart 2108. Taakoka: Stoddart, sight record. BORAGINACEAE Heliotropium anomalum H. and A. Motutapu: Stoddart 2113. Tournefortia avgentea L. f. Oneroa: Stoddart 2128. Koromiri: Stoddart 2152. 13 VERBENACEAE Lantana camara L., E Oneroa: Stoddart 2122. Lantana camara var. aculeata (L.) Mold. Moiutapu: Stoddart 2110. Taakoka: Stoddart 2160. Koromiri: Philipson 10352. Stachytarpheta urticifolia Sims Koromiri: Stoddart 2153. Vitex trifolia L. var. bicolor (Willd.) Mold. Motutapu: Philipson 10362. RUBIACEAE Guettarda speciosa L. Motutapu: Stoddart 2136. Oneroa: Stoddart 2116. Koromiri: Stoddart 2151; Philipson 103852. Morinda citrifolia L. Motutapu: Stoddart 2144, Oneroa: Stoddart 2126. Koromiri: Stoddari 2149. Taakoka: Stoddart 2170. GOODENIACEAE Scaevola taccada (Gaertn.) Roxb. Motutapu: Stoddart 2145. Oneroa: Stoddart 2129. Koromiri: Stoddart, sight record. Taakoka: Stoddart 2171. COMPOSITAE Bidens pilosa L. Koromiri: Stoddart 2156. Elephantopus mollis HBK. Motutapu: Stoddart 2143. Emilia sonchifolia (L.) DC. Motutapu: Stoddart 2134. Sonchus oleraceus L. Motutapu: Stoddart 2111, 2138. Wedelia biflova (L.) DC. Motutapu: Stoddart 2107. Taakoka: Stoddart 2162. 14 REFERENCES Cheeseman, T.F. 1903. The flora of Rarotonga, the chief island of the Cook Group. Trans. Linn. Soc. London II. Bot. 6: 261-313. Philipson, W.R. 1971. Floristics of Rarotonga. Bull. Roy. Soc. New Zealand 8: 49-54, Wilder, G.P. 1931. Flora of Rarotonga. Bull. Bernice P. Bishop Mus. 86: 1-113. oTesow ydeisoj0ud Ate worlz paonpeir Aydeiso0do} $(/961) poom 4eqFy “e8u0j01eYy FO ABojoaD *g aunbry SOINVIIOA 3N390Id es] S13AVYE9 OVXIN ISS sanvs vouv [a] sy SS _VvOUvAV vAV>~ sisod3aa dwwms Ej eS WNIANTIVY WV34Ls [ ~] —S SS iva ~ ~ WS NES nlLwav ~< we ees SIYLIW NI SLHOIFZH IIVSOW OLOHAYIY WOYF GIY9 FYLIWOTIN DOL NOW SL vl €i Al Figure 38. Reef islands of Ngatangiia Harbour. For location, see Figure 2 MOTUTAPU metres Re COCONUT WOODLAND o a < a a < O < Ww {e) = ce) 2 é ) Ww (a) Ww = [> | HIBISCUS WOODLAND WITH GUETTARDA AND MORINDA = SCAEVOLA WITH TOURNEFORTIA [« ] PASPALUM MARSH 489-409 O - 73 - 10 (ee CASUARINA (ea INTERTIDAL SAND (UCA FLAT) mow UNDERCUT SHORELINE Figure 4. Motutapu ONEROA metres WITH TOURNEFORTIA ES HIBISCUS WOODLAND =| SCAEVOLA TACCADA ca CASUARINA WOODLAND WITH SCAEVOLA < z a < =) Ww < ) a Zz < Ww > ) 2) a Ee ae & = [a) Zz \ 7) <3 | D| DPS, Pi ost AW aie yn ih, Up ‘A Lp S————__qulog npund1uDW ae <7 (9n ‘d ‘[961) JUewjzedeq ITYder8o1phy :ejep Fo aoinog ‘s}ouy ut paeds purM uvell sinsty Aamo, pue sues a8equaoied smoys esin8ty teddq ‘(ejep azeak ¢ ‘sanoy QF€/T Ie SsuoT}eAAasSqO JO aBeqUedi1ed) uequeg IOJ SeSOI putmM ATYWUOM ° aunb27 INCHES 3D) 3© 29 2O y GG ml a OY, 93 a GZ can vaya AA W J Figure 4, MM 900 800 700 SOO SOO 400 SOO SX SQ QI IONNHOAAAAAY\i RSS 4 EM AWeeMireee ean Ag eS @ > N° D Mean monthly rainfall, monthly extremes, and monthly quartile ranges for 62 years of record at Pamban. Source of data: Meteorological Department, Government of India (1891-1954). [e] Individual Thespesia FHA Thespesia Woodland [a] Borassus = Acacia (i Grasses and Herbs V4 Pemphis acidula [A] coconut Woodland Sand MUSAL TIVU OR HARE ISLAND metres 222777772 Figure 5, Musal Tivu or Hare Island. This and the following figures based on prismatic compass and pacing surveys pueTsSI ysom °9 aanb17 Eeeeneerared! OOL Seujow O pups jppiyequy fo] PUDS snjoqouods k Les} TSI 2S S]inn SqUSH DUD =sossDUS TT DUaZ}IUWNA YIM DIUUddIAY [eo] T[neueW +7 aanb1z siyudwad ZZ puds sassDU SSDUD PUD DpaDNs Disedsau xKaJIUIdS WNniAnses DINPIDD siudwad puainBnug DIUUdDJIAY | CH EA ZZ Le] [2] pue[S] MeN °9 aanb17 (eee erel OO! S24}9W O QNV1SI MAN > fr ATOLL RESEARCH BULLETIN No. 162 ISLAND NEWS AND COMMENT Issued by THE SMITHSONIAN INSTITUTION Washington, D. C., U. S. A. December 31, 1972 ISLAND NEWS AND COMMENT Readers will notice a neater format in this Bulletin. It is not a permanent improvement but is the result of very welcome, though temporary, help from the U.S. Air Force (AF/LGFC, Hq USAF). We owe the nice appearance of this issue to Mrs. Martha M. Green, of that Air Force unit, who did the final typing; our best thanks to her. This help bailed us out of a diffi- cult situation for this issue. The next one will appear thanks to the Bureau of Sports Fisheries and Wildlife. After that we will probably be in trouble again. If any of our audience has any ‘ideas on how we can get camera-ready copy typed on an “‘editing typewriter’’ or ‘‘word- processing machine,’’ with no funds to pay for it, we would appreciate suggestions. We simply cannot handle the burden of repeated proof-reading of work done on an ordinary machine. We hope to get the production of the ARB back ona stable basis soon, but have had some bad breaks recently. Since the last Island News and Comment appeared, the Island Bibliography Supplement, by Sachet and Fosberg, has been published (see under Publications). A leaflet has been sent to ARB readers to this effect. As there will be no further printed supplements to this volume, the section on Publications, Briefly Noted Items, will include lists of current publications on islands and reefs. Mrs. Linda Smithhas volunteered to compile these lists for marine subjects and Dr Bryce G. Decker for terrestrial ones. Reprints and references sent to us will make this task easier and the lists more complete. We are always glad to receive good reviews of appropriate works within our scope, as well as newsitems, comments on pertinent subjects, and short original papers. NEWS PALAEOECOLOGY OR REEFS: The Palaeontological Association organized a one-day symposium at Edinburgh on 9 September 1971 as part of the joint meeting of British Geological Societies. Papers covered sooth modern and ancient reefs. In the former class, several of the papers were based on work either in the Seychelles or at Aldabra. Thus Dr E.Drew spoke on ‘‘Reef front phenomena of Aldabra: an ecological interpretation;’? Dr J. Taylor on ‘‘Habitat complexity and faunal diversity on Recent reefs;’? Mr. B.R. Rosen on ‘‘Ecological stratifica- tion in coralline environments;’’ and Dr. C.J.R. Braithwaite ‘on ‘‘Ecology, palaeoecology and reef recognition.’?’ Two papers drew on recent work at Bermuda. Dr P. Garrett considered “‘Bermuda lagoon reefs and their palaeoecological significance,’’ and was concerned with the ways that small coral colonies develop into micro-atolls and then into patch reefs or, as he termed them, mesa-reefs. He stressed the importance of biological erosion concurrent with growth, and showed that a large percentage of the volume of patch reefs was open cavity. Dr T.P. Scoffin also drew on West Atlantic experience in interpreting the ‘‘Conditions of growth of the Wenlock reefs of Shrophire.’”’ The fossil reefs covered a wide range: Palaeozoic (Dr C.T. Scrutton), Mesozoic (Dr F.M. Broadhurst and Dr I.M. Simpson on Derbyshire; Mr. J. Miller on west Yorkshire; Dr D.B. Smith on the Permian in Durham and in the Guadeloupe Mountains of Texas), and Coenozoic (Dr E.B. Wolfenden and Dr D.D. Bayliss on the Libyan Palaeocene). Dr G. Farrow also spoke on rudist reefs. The often heated discussion revealed a sharp difference of view between the ecologists and the palaeoecologists. The latter were content with a catholic interpretation of the term reef; the former, impressed by the thinness of many modern reefs and the fact that coral com- munities are not necessarily reef structures nor are reef structures necessarily covered with coral communities, wanted a more precise definition which they were themselves, however, unable to succinctly provide. Brian Rosen summarized the position lucidly when he pointed out that modern reefs are growing in highly atypical immediately post-glacial conditions, and it is hence difficult to extend concepts of reef structure derived from the study of modern reefs to those in the past. Dr Braithwaite considered, however, that there were certain criteria of scale, zonation and structure (in the sense of frame-building) that must be satisfied before the term reef could legitimately. be used. It was noticeable that some of the palaeoecologists used terms such as “reef flat,’? derived from contemporary reefs, without considering the extent to which these forms owed their characteristic features to the highly unusual conditions of the last several thousand years. The discussion ended rather inconclusively, with the palaeoecologists on the whole unconvinced that the issue of reef definition was a real one, and the ecologists unable to provide a convincing definition of what they considered to be a reef, In summing up Sir Maurice Yonge drew attention to the need for further work on corals as animals. He noted recent work on the role of zooxanthellae in coral nutrition, on competi- tion between corals, and on the testing of species in corals by immvnological methods, and he stressed the need for long-term study of population fluctuations on reefs, possibly on islands set aside as laboratories by the Islands for Science Programme. One was left with the feeling that more palaeoecologists should follow the lead given by Newell twenty years ago and study modern reefs as well as ancient ones - - but that comment comes from a modern reef-man, not an ancient one; the latter might well reverse it. Diahass Apropos of some of the remarks reported from this symposium it may be well to remind readers that regardless of the supposedly atypical conditions under which modern reefs are growing, reports on- the cores and cuttings from the various deep drillings on atolls and barrier reef islands, some going back to the early Eocene or even late Cretaceous, do not indicate any startling difference or new characteristics introduced in the structures produced during the last few thousand years. A survey of fossil reefs made some years ago by Ted Ballard indicated that the great preponderance of reef growth throughout geological time has taken place during periods of transgression (personal communication). Ona geological scale, at least, that is what post-glacial time has been. F.R.F. NEW INDIAN OCEAN RADIOCARBON DATES: D. P. Agrawal, S.K. Gupta and S. Kusumgar (‘Tata Institute. radiocarbon date list VIII’’) report some new dates from reef areas in the Indian Ocean in Radiocarbon 13: 84-93, 1971. Samples collected by S8.G. Patil from Minicoy are dated as follows: TF-1017, coral, depth 3m, 1575 + 85 BP; TF-1022, coral, depth 0.9m, modern. Several samples from the Gujarat coast and the Rann of Cutch, mostly from coral samples, are dated between ca 4500 and 6500 BP. Other samples dated come from coastal plain and continental shelf environments, the latter including one sample, TF-969, in 96m of water off Bombay, of coral dating atonly 140 + 90 BP. We look forward to the papers which will place these dates in their geological framework. SCOTT REEF, SAHUL SHELF, AUSTRALIA: The northwestern Australian reefs are among the least known in the world, apart from a reconnaissance report by C. Teichert and R.W. Fairbridge (Some coral reefs of the Sahul Shelf, Geog. Rev. 38: 222-249, 1948). In June 1971 Australian newspapers carried reports-of deep drilling on Scott Reef, 14005'S, 121°50'E, immediately south of Seringapatam Atoll, carried out by a consortium including Woodside Oil NL, BOC of Australia, Mid-Eastern Oil NL, Shell Development (Australia), BP Development Australia, and California Asiatic Oil Co. Gas flow rates from Scott Reef No 1 well have been 3 reported as: 18 million cuoic feet of wet gas a day from the interval 14,105-14,124 ft; 11 million cubic feet from 14,240-14,330 ft; and 9.8 million cubic feet from 14,370-14,390 ft. Exploitation of this gas field is being considered, and it is possible that a land-based produc- tion installation could be based on a small island on Scott Reef. The lagoon with a depth of 20 fathoms could be used as an anchorage. In view of the lack of knowledge about these reefs, it is to be hoped that these developments will provide an opportunity for thorough faunal and floristic studies of Scott Reef (including land areas) before development begins. These details of the Scott Reef well appeared in The Australian for 9 and 16 June 1971. TONGA: The Oct. 7 issue of the Tonga Chronicle announces the initiation of oil drilling with great fanfare at Ma’ufanga, Tongatabu. It is with mixed feelings that we read this. We are, of course, Sympathetic to the Tongans’ desire for prosperity, and we are interested in what will be learned about the subterranean structure of Tongatabu. On the other hand, we have seen what oil strikes have done to other societies, including some regional branches of our own, and we would be most unhappy to see one of the few still truly viable Polynesian cultures destroyed by too sudden prosperity and ‘‘development.’’ We hope that, at least, the agreement between the Tongan Government and the Tongan Oil Participants calls for full publication of the scientific results of the drilling, that cores will be taken, and that provision is being made for their scientific examination and study. We are happy to offer the pages of the ARB for publication of any scientific observations that result from “his enterprise. SEYCHELLES: The Government of the Seychelles has just published a ‘‘white paper’’ setting forth the current government policy on conservation matters in the Archipelago. This 10- page document is obtainable from the Seychelles Ministry of Agriculture, Natural Resources and Marketing, Victoria, Mahé, Seychelles, for 3 shillings. The paper summarizes the present conservation legislation and explains the plans of the government to implement these regulations and to introduce additional provisions where neces- sary. The plans are largely based on a report by Mr. J. Procter, who spent some months studying the situation in detail. The government has, in most respects, accepted the recom-- mendations of the Procter report. We need not go into any detail on what is planned, as anyone interested will certainly send for the document itself. It is a satisfaction to be able to congratulate the Seychelles Government on their plans for the protection of what is left of the magnificent land fauna and flora of the islands, and especially to commend their being ahead of most of the rest of the world in establishing marine parks. The intent to prohibit spear-fishing is an encouraging development that could well be emulated elsewhere. The recent opening of the new international jet airport on Mahé makes this firm set of policies on conservation especially timely. In order to support a lasting tourist trade, the Seychelles will have to have, over the long pull, something to offer beside novelty and remote- ness. The natural features, both marine and terrestrial, provide this. To properly utilize them, an active program to study and make known the island natural history is essential. There is a rumor that Mr. Procter will return to the Seychelles as conservation officer. If true, this will give him an opportunity to help implement this program for which he is largely responsible. SOOTY TERN STUDIES IN THE SEYCHELLES: For some time concern has been expressed about the exploitation of Sooty Tern eggs in the Seychelles, the scale of which has led to a reduction in the sizes of some colonies (see, for example, M.W. Ridley and R. Percy: The exploitation of sea birds in Seychelles, Colonial Res. Stud. 25: 1958). Dr C.J. Feare, of the University of Aberdeen, has now been given a grant by the Natural Environment Researcn Council for a three-year study of the problem, beginning in 1972. The aims of the study are to collect basic data on the breeding output of the birds, and to lay the foundations for longer- 4 term work which will permit the construction of mathematical models of the population dynamics of the species in the Indian Ocean. From the results a rational plan of exploitation will be formulated. During the first season the general breeding biology will be studied in order to discover the causes and extent of natural losses within a colony, and how these vary in different parts of the colony and at different timesinthe season. The second year will be devoted to observing the effects of experimental removal of eggs on the overall output of the colony. The results of these investigations will provide a basis for the work of the third season, in which Sooty Tern colonies in the Seychelles will be counted, and recommendations made for the modification (if necessary) of existing regimes of commercial exploitation, in order that the futures of both the tern colonies and the egg industry will be assured. DUCIE ATOLL: As a part of his long-term investigation of the marine mollusk fauna of Polynesia, Dr Harold Rehder, on Jan. 13-15, 1971, visited little known Ducie Atoll, between Pitcairn and Easter Islands, perhaps the most remote and isolated of all coral atolls. Notable was the fact that the forest on Ducie is composed of only one species of tree, Tournefortia argentea. Rehder gave a lecture on the impoverished biota of this atoll at the National Museum of Natural History on Feb. 10, 1972. We hope to be able to offer our readers a description of Ducie by Dr Rehder in the near future. FANNING ISLAND EXPEDITION: Under the auspices of the Hawaii Institute of Geophysics, a group of 26 scientists and students from the University of Hawaii and the B.P. Bishop Museum spent over three weeks on Fanning Island doing scientific investigations of great diversity. This atoll was the site of one of your editors’ (F.R.F.) introduction to coral atolls, some 38 years ago, so this expedition is of extra special interest to him. The leader of the party was the eminent geologist, Prof. Keith E. Chave. Support was provided by the National Science Foundation. That the visit was a most productive one is shown by the massive report, HIG-70-23, published in November 1970, and a fine series of papers in the April 1971 issue of Pacific Science (25: 188-289). The papers in the report and in the Pacific Science number dealing with land aspects have been listed and annotated in the Supplement to Island Bibliography, recently published. The marine papers, by far the majority, are listed in the ‘‘Briefly noted items’’ section at the end of the present Island News and Comment. The University and Keith Chave are to be congratulated on this achievement. MEETINGS: The Regional Symposium on Conservation of Nature - - Reefs and Lagoons, organi- zed and hosted by the South Pacific Commission in collaboration with the International Union for Conservation of Nature and Natural Resources, took place in Noumea, Aug. 5-14, 1971. During the first days, the territorial representatives describedand discussed the problems of their territories, of which unfortunately only a limited number were represented. It became quickly apparent, then and later in the more general Symposium starting on Aug. 9, that lagoon pollution, reef destruction and other conservation problems are not uncommon in Pacific island territories and are in fact rapidly becoming alarming in many of them. Twenty-one resolutions on the protection of reefs and lagoons and on more general topics were adopted and addressed to the SPC and governments and administrations concerned. They are published as a supplement to IUCN Bulletin 2(21): Oct.-Dec. 1971. A large number of mimeographed background papers, and draft proceedings, were available at Noumea. Planned utilization of the lowland tropical forests. _A symposium sponsored by the Indonesian Institute of Science (LIPI), the Regional Center for Tropical Biology (BIOTROP) and Unesco was organized by the Standing Committee on Botany of the Pacific Science Association at ) Tjipajung Indonesia, Aug. 12-14, 1971. It included one session on Pacific islands forest conditions. A report on this meeting, by Prof. D. Mueller-Dombois, appears in Nature and Resources, a newsletter published by Unesco, 7(14): 18-22, Dec. 1971. Twelfth Pacific Science Congress, Canberra, Aug. 18-Sept. 3, 1971. The Noumea and Tjipajung symposia were pre-congress meetings and many of the participants went on to Canberra where the Australian National Academy organized the congress at the Australian National University. A report on the Congress is available in the Pacific Science Association Informa- tion Bulletin 23 (3-5): Oct. 1971. This includes portraits of the recipients of honors and awards announced at the opening session: Dr Sarwono Prawirohardjo and Sir Maurice Yonge were made Honorary Fellows of the Association, Dr Carl L. Hubbs received the Shinkishi Hatai Medal and Dr F. Raymond Fosberg the Herbert E. Gregory Medal for distinguished service to science in the Pacific. The 13th Pacific Science Congress will be held in Vancouver in 1975, at the invitation of the National Research Council of Canada. M.-H.S. The International Symposium onCoralReefs (see ARB 148: 8) sponsored by the Great Barrier Reef Committee with the Committee for International Symposia on Corals and Coral Reefs is to be held at Heron Island and other locations on the Great Barrier Reef, tentative dates are now 29th May-10th June 1973. Address inquiries to Dr. G.R. Orme, Dept. of Geology and Mineralogy, University of Queensland, St. Lucia, Q4067, Australia. PLANTS OF OCEANIA: Botanists interested in Pacific island floras will be glad to learn that the vast accumulation of unmounted island collections in the Paris herbarium, long in storage, are being sorted to family and made accessible to visiting botanists. Mr. N. Hallé, Sous-Directeur, Laboratoire de Phanérogamie, has undertaken the immense task of sorting through this accumulation of 150 years. He is willing to make available to serious workers material in any family, providing the botanists intending to visit the Paris herbarium will inform the Directeur of the Laboratoire de Phanérogamie, 17 rue Buffon, Paris V% France, of their planned visit and their needs sufficiently in advance. ISLAND SCIENCE NEWSLETTERS: A number of informal serial publications have been established that deal wholly or in part with scientific affairs on islands or with matters of interest to island scientists. Several of these will be listed, with their sponsoring organiza- tions and addresses and editors if known. We will not usually repeat news items that appear in these newsletters, unless they seem of unusual importance to our readers. The Elepaio. Hawaiian Audubon Society, eds. Miss Charlotte Hoskins and Miss Unoyo Kojima, P.O. Box 5032, Honolulu, Hawaii 96814, U.S.A. Newsletter of The Hawaiian Botanical Society, c/o Dept. of Botany, University of Hawaii, Honolulu, 96822, ed. Russell K. LeBarron. Aldabra Newsletter. The Royal Society, 6 Carlton House Terrace, London, S.W.1, England. Requests to be placed on the mailing list should be sent to Dr. D.R. Stoddart, Dept. of Geography, Downing Place, Cambridge CB2 3EN, England. He also keeps a mailing list of those persons interested in exchanging publications, especially in the fields of terrestrial and marine ecology of tropical oceanic islands and coral reefs, for reprints of papers on Aldabra forming a series entitled Contributions from the Royal Society Aldabra Research Station. Association for Social Anthropology in Oceania Newsletter, ed. Bob Kiste, Center for South Pacific Studies, University of California, Santa Cruz, Calif. 95060, U.S.A. The Palaeontological Association Circular, ed. Dr. J.A.E.B. Hubbard. Geology Dept., Kings College (London University), Strand, London, W.C. 2, England. 6 Environmental Sciences Quarterly Newsletter, Office of Environmental Sciences, Smithsonian Institution, Washington, D.C. 20560, U.S.A. CITRE Newsletter, ed. Stephen Smith, Smithsonian Institution, Washington, D.C. 20560, U.S.A. An Environmental Newsletter, Caribbean Conservation Association, c/o Caribbean Research Institute, College of the Virgin Islands, St. Thomas, U.S. Virgin Is. 00801. Pacific Island Program Bulletin, Dept. of Anthropology, University of California, Los Angeles, Calif.. 90024, U.S.A. (This bulletin ceased publication with no. 15, Oct. 20, 1971). News and items of current interest are also reported in the Pacific Science Association Information Bulletin, Miss Brenda Bishop, ed., Bishop Museum, P.O. Box 6037, Honolulu, Hawaii 96818, and Cahiers du Pacifique usually include sections on News, Meetings, and Congresses, and bibliographies. No. 15, Sept. 1971, received in Washington in early 1972, includes such a section of ‘‘Nouvelles du Pacifique’’ of 74 pages, by J. Plessis. Address for the Cahiers: Fondation Singer-Polignac, 43 Ave. Georges-Mandel, Paris 16e. ISLAND BIOLOGICAL STATIONS: West Indies Laboratory: In July 1971 classes opened at the new West Indies Laboratory of Fairleigh Dickinson University at Teague Bay on St. Croix Island, U.S. Virgin Islands. Starting with a magnificent physical plant, though modest in size, including laboratory and library buildings, dining hall, living quarters for students, staff, and visiting research workers and faculty, a pier and boat house, shop, sewage treatment and recycling plant, and warehouse, this tropical station has immediately taken its place as one of the finest combination teaching and research facilities in the Caribbean. The director is Prof. H. Gray Multer, the resident scientist is Dr John Ogden, and the manager is Mr. Lowell Bingham. Courses in geology, ecology, zoology, botany and marine biology are offered. During periods when no regular classes are scheduled the laboratory is available for use by classes from other universities by arrangement with the director. Arrangements also may be made for use of the facilities by visiting investigators. A brochure isavailable on request. The address of the station is West Indies Laboratory, P.O. Annex Box 1010, Christiansted, St. Croix, U.S. Virgin Is. 00820. University of Guam Marine Laboratory: The new marine science facility of the University of Guam opened on January 15, 1971, in a well-equipped building, beautifully situated on Pago Bay, east coast of Guam, just below the University campus. Flowing sea-water, boats, diving gear and other equipment for marine biological studiesare available. Prof. Lucius G. Eldredge is director and will supply information on request. The address is Marine Laboratory, University of Guam, P.O. Box EK, Agana, Guam, 96910, U.S.A. The University Marine Biological Station, Millport, Isle of Cumbrae, U.K.: This station, form- erly known as the Marine Station at Millport, is now functioning under new auspices, con- trolled by the Universities of London and Glasgow. Its objectives are teaching and research in marine biology. The director is Prof. Norman Millett, formerly Professor of Zoology at Bedford College, University of London. Besides courses offered by the station, accommoda- tion will be provided for outside classes accompanied by their own teachers. Royal Society Research Station, Aldabra: The Royal Society has announced the appointment of Dr David Wood as the new director of the station. The lease of the island, formerly held by H.Savy and Co., Ltd., of the Seychelles, has been assigned to the Royal Society. The purchase of this lease was in large part made possible by a donation from Mr. Christopher Cadbury. Detailed news on Aldabra research activities will be found inissues of the Aldabra Newsletter and in press releases from the Royal Society, 6 Carlton House Terrace, London, S.W. 1, U.K. 7 Antenne de Tahiti: The Muséum National d’Histoire Naturelle of Parishas recently established an outpost or ‘‘antenna’’ in French Polynesia. There will be a small station on Moorea, and research plans for 1972 include studies on Rangiroa, Bora-Bora, and Taiaro Atoll. The director is the very active malacologist, Dr Bernard Salvat of the Muséum (55 Rue de Buffon, Paris 5), and the mailing address of the station is B.P. 562, Papeete, Tahiti, French Polynesia. RECENT DEATHS: Edwin C, Allison: Ned Allison died suddenly on Jan. 1, 1971. He was struck down while guiding students on a fieldtrip, near Caborca, Sonora, Mexico. His work was mostly in paleon- tology, but he also collected and studied living forms, especially mollusks. He took part in the 1958 Scripps Institution expedition to Clipperton Island and published several papers on the mollusks with L.G. Hertlein. He was a gentle and patient colleague in the field, and generous in exchanging information or photographs later. Of some comfort to his friends and colleagues, as well as his family, is the fact that San Diego State College has announced that there will be an Edwin C. Allison Center for the Study of Pacific faunas in the Department of Paleontology where he taught. Richard J. Russell: Dean Russell died on September 17, 1971, at Baton Rouge, Louisiana at the age of 75. Coastal geology and geomorphology thus lost one of its most active and distin- guished leaders. Not only through his own research did he advance his science, but even more through the students and younger colleagues that he helped and encouraged. His Coastal Studies Institute at Louisiana State University is one of the most important centers for the study of coastal geomorphology. We sincerely hope that, even without Russell’s leadership, it will continue to be outstanding and productive in this fascinating field of research. SHORT PAPERS NOTES ON THE HERPETOFAUNA OF KUME-JIMA AND O-JIMA, RYU KYU ISLANDS by Clifford Ray Johnson Department of Zoology, The University of New England Armidale, N.S.W. Australia Kume-jima, the westernmost island of the Okinawa Group, covers 26 sq. miles and lies 55 miles W of Okinawa in the East China Sea. Like most of the Ryu Kyu Islands, it is sub- tropical with a mean annual temperature of 72°F, average humidity of 80%, and a mean annual rainfall from 51 to 91 inches. O-jima lies 1 mile S of Kume-jima and has an area of about 1 sq. mile. The islands appear to be of volcanic origin with a shoreline composed of disintegrated corals and conglomerates. Kume-jima is forested, like Okinawa, in montane regions and culti- vated along the coastal areas. O-jima is entirely cultivated. Both islands were by-passed by World War II. The herpetofauna of Kume-jima, although one of the larger islands of the Okinawa Group, is very poorly known. During March 1965, I visited Kume-jima and O-jima and made small collections of reptiles and amphibians over a period of five days. The only previous mentions of Kume-jima in the literature are by Koba (1959) and Okada (1966). 8 Rana limnocharis Boie (in Weigmann) Four specimens were collected near Gima in a cultivated field. They appeared similar to specimens from Okinawa where this is the most abundant species of ranid (Johnson, 1969). Rana namiyei Stejneger A single specimen, 85 mm snout-vent length, was capturednear a rice field close to Gima. This species on Okinawa is usually restricted to montane areas, but appears to be more eurytopic on Kume-jima. Eumeces margenatus (Hallowell) One specimen was collected at the edge of a cultivated field in dense brush near Gima. Another was seen in a similar habitat near O-village, O-jime. None were seen along the beaches, although they were often found in such habitats on Okinawa. Ateuchosaurus p. pellopleurus (Hallowell) One specimen, 42mm snout-vent length, was collected near Gushikawa in a grass field. Takydromus smaragdinus Boulenger Three specimens were collected ia thick grass adjacentto a cultivated field near Gima. Trimeresurus okinavensis Boulenger One specimen was seen in a grass field on Mt. Uegsuka. According to the natives these snakes are common throughout the island. Other species of Trimeresurus may also occur on the island. The following is the most complete check-list to date of the herpetofauna of Kume-jima based upon my collection and those of Koba (1959). Order Salientia Microhyla ornata (Duméril and Bibron) Rhacophorus viridis (Hallowell) Rana limnocharis Boie (in Weigmann) Rana namiyei Stejneger Order Sauria Eumeces margenatus (Hallowell) Ateuchosaurus p. pellopleurus (Hallowell) Takydromus smavagdinus Boulenger Order Squamata Trimeresurus okinavensis Boulenger ACKNOWLEDGEMENTS I wish to thank Cdr. William F. Russell, CEC, USN, for his cooperation during field opera- tions and Rev. and Mrs. Kanard for extending their hospitality during my stay on Kume-jima. Transportation was arranged by the U.S. Navy. Dr Harold F. Heatwole, The University of New England, reviewed the manuscript. LITERATURE CITED Johnson, C.R. 1969. Herpetofauna of Okinawa, Ryu Kyulslands. Herpetologica 25(3): 206-210. Koba, K. 1959. Herpetofauna of the Amami Group of the Loo Choo Islands (III). Mem. Fac. Educ. Kumamoto Univ. 7: 187-202 (in Japanese). Okada, Y. 1966. Fauna Japonica: Anura (Amphibia). Biogeogr. Soc. Japan, Tokyo, 1-234, 24 plates. LAYSAN ALBATROSS AS CARRIER OF FLOATING DEBRIS TO LAND by Miklos D.F. Udvardy Sacramento State College, Calif. Upon reading Kenyonand Kridler’s interesting note (Laysan albatrosses swallow indigestible matter, The Auk 86: 339-343, 1969), I found that my own observations on Laysan might modify the impression these authors gave about the carrying ability of albatrosses and the hydrography of that island. During two expeditions to Laysan Island (see notes in Elepaio 20: 16, 1959; 22: 43, 1961, and in ARB 103: 1964) my own experience with juvenile and adult albatross skeletons was by and large similar to that of Kenyon and Kridler: the carcasses and skeletons we found were also lined with pumice, armoured fish, kukui nuts and other indigestibles though plastic arti- facts were not observed by us. Kenyon and Kridler note that the ‘‘Lagoon’’ of Laysan Island is not connected by any channel with the sea (i.e. it is, in reality, a lake) and therefore they conclude that the plastic and pumice they found deposited at high water line of the lagoon must have originated from contents of albatross carcasses. But this conjecture needs to be modified: some, but not all, the former floatsam on the perimeter of this lagoon originates with the albatrosses. An undeterminable portion of the debris--and certainly all large-sized pieces, too big for albatrosses to swallow--should rather be assumed to have been brought in by the huge waves of winter storms which break through the flat coral sand area that girdles the lagoon on the south and which is only at a few places reinforced by low ridges or patches of phosphate rock. This area is devoid of a protective beach crest and here and northward up to the edge of the lagoon we found in June 1959 dozens of Japanese fishnet floats: heavy glass balls of 80 and 90mm, or even more, in diameter, certainly unfit for albatross consumption. It is safest to assume that these glass floats were brought in by wave action. Then, other floatsam is likely to enter the lagoon the same way and the albatrosses are not the only carriers to blame for all deposited foreign material. MOROTIRI (BASS ROCKS) AUSTRAL ISLANDS by F.R. Fosberg Smithsonian Institution Morotiri or Bass Rocks is a small cluster of 4 rugged volcanic rocks and a number of stacks that form the southeast extremity of the Austral Island Group, in French Polynesia. They are located about 46 miles east by south of Rapa, at about 28°9S, 143930‘W. As nothing of a general nature seems to have been published on their natural history, it may be worthwhile to publish notes made on a short visit on July 22, 1934, when Harold St. John, Elwood C. Zimmerman and I landed on the largest of the rocks and collected what could be found and 10 reached in a couple of hours of scrambling and climbing over guano-covered ledges and cliffs. The highest of the rocks is about 100m high and not much more than that wide, several times as long. The rocks were practically without a real covering of vegetation. A sparse growth made up of Cyperus, Bidens, Digitaria and Portulaca was present on the non-perpendi- cular slopes and ledges, almost luxuriant in favorable spots. In the crevices in the rocks Asplenium, Nephrolepis, and Cheilanthes formed tufts. Solanum nigyum was present hereand there. Lyciwm was seen on the lowest slopes, anda prostrate Euphorbia formed mats on one slope on one end of the island. A sterile rosette of Sonchus was seen. Some of the rocks were sparsely covered with lichens. A sterile moss was occasional around seeps. These seeps were evidently highly charged with lime or some other substance that crystallized out around the cracks. Under the plants and stones was a remarkably large fauna of insects, spiders, centipedes, and isopods. Of the last, 3 species occurred under stones anda species of Ligia ran around over rocks, collecting in numbers under overhanging rocks. At least 1 species of centipede, 4 or 5 of spiders, 1 or 2 of ticks, 2 of mites, 1 of Lepisma, 1 of Machilis, 2 of Collembola, 1 cricket, 1 or 2 of ants, 1 lygeid, 4 species of Rhynchogonus, and 1 other weevil all lived under stones and plants. Two or 3 kinds of flies and 2 of moths were seen flying. A louse fly (Olfersia sp. ?) was abundant on shearwaters. Birds were more than abundant, but nearly all belonged to a species of gray tern and 2 or 3 of shearwater. A couple of white-tailed tropic birds were seen. The shearwaters were nesting and eggs and young were abundant on small ledges and between tufts of sedges. Bur- rowing would be difficult here. The rock is composed of more or less bedded basalt, forming a high core in the center, and the two ends are capped with tilted beds of what is probably tuff (described as ‘‘apparently sedimentary rock’’ in my notes at the tim?). No coral was seen. Encrusting calcareous algae colored the lower rocks. Non-caleareous algae were reasonably abundant but badly beaten to pieces by wave action. The waves, even at the ‘‘quiet’’? period of our visit, were several meters high, making landing hazardous, to say the least. Acorn barnacles and chitons were seen on the lower rocks. Fish of many kinds were abundant in the area and many were caught while the ship cruised around the area near the rock while we worked. SOME LAND BIRD MIGRANTS IN THE WESTERN INDIAN OCEAN by C.W. Benson Mr. H.H. Beamish has shown me a colour slide of a bird photographed by him in November 1970 on African Banks, Amirante Islands, at ca. 5°S, 53°E (for an account of the geography and ecology see Stoddart & Poore, ARB136: 187-191, 1970). Undoubtedly the bird is a Phylloscopus species, and on geographical grounds by far the most likely is the Willow Warbler, P. trochilus, from which the bird onthe slide was indistinguishable. On the African mainland this species is abundant during the palaearctic winter, even reachingasfar south as South Africa (Mackworth- Praed& Grant, African handbook of birds, Ser.I, Vol. 2: 1955). According to the same authors, the Chiff-Chaff, P, collybita, not certainly distinguishable on the evidence of this slide from P, trochilus, migrates almost as far south as the equator, but its occurrence on African Banks seems extremely unlikely. As far as I am aware, this is the first record of a palaearctic breeding sylviid from any island in the western Indian Ocean south of the equator. Indeed 11 Ripley and Bond (Smiths. Mise. Coll’ns. 151(7): 1966) do not even give any such record from Socotra, although they quote single old records each of P. collybita and the Whitethroat, Sylvia communis, from Abd-el-Kuri, between Socotra and Cape Guardafui. Among records of birds received by the Royal Society from J.A. Stevenson on Aldabra, copies of which were transmitted to me by D. Griffin, the following from West Island should be mentioned: Eurystomus glaucurus, Broad-billed Roller: One seen at 15:00 hrs. on 20 October 1970. Benson and Penny (Phil. Trans. Roy. Soc. B260: 517, 1971) give but few records from Aldabra and neighbouring islands. Oenanthe oenanthe, European Wheatear: One seen at 15:00 hrs. on Wed. 27 January 1971, with a note that there had beena cyclone the previous week. What was presumably the same in- dividual was seen again on 1 and 5 February 1971. Benson and Penny (op. cit.: 519) suggest that this species may winter regularly on Aldabra in very small numbers. HURRICANE LAURA, WITNESSED IN BRITISH HONDURAS by Arnfried Antonius Smithsonian Institution Hurricane Laura was first reported on Sunday, November 14, 1971, in the morning. It was then a mere tropical storm near Swan Island, but in the afternoon it reached hurricane force and was baptized Laura. During the following days Laura first travelled N in the direc- tion of Cuba, then turned W and finally S along the coast of the Yucatan Peninsula. It entered British Honduras territory in the early morning of November 20. By about mid-afternoon the hurricane had passed Belize, with wind speeds about 70 knots and thus not doing any harm. In Stann Creek though, things were slightly different. Belize and Stann Creek lay both on the western rim of the hurricane, but as it moved more and more southwesterly, the eye came closer to Stann Creek. Winds blew during the day first from the W, then NW, N, NE, and in the afternoon from the east. The eye was closest then. The winds came from the open sea and reached at least 80 knots. About half the harvest was lost in the extensive Citrus plantations east of Stann Creek, and wave action removed the longest wooden pier of the area. On Glover’s Reef, winds first came roughly from the W with speeds up to 70 knots. Wave action piled up large heaps of Thalassia and algae along the lagoon side of the cays without doing damage. The eye passed around noon, creating a two-hour lull. The most violent stroke came after this, with sudden wind forces of 80-100 knots from the NE; this lasted only a few minutes and calmed down to 70-80 knots shortly afterwards, but during this brief time some damage was done to the facilities of a diving resort on Long Cay. The winds sank a 35-foot vessel, blew down one of eight existing cottages, and removed half the tin roof of another hut. Very few coconut trees fell. Underwater, on the eastern and northeast fore-reef slope, broken trunks of Acropora palmata up to 20cm in diameter could be observed here and there, as well as large colonies of Acropora palmata and Diploria strigosa turned upside down. However, only 2m to the right or left of them, it was possible to find much younger colonies, very fragile and yet completely unharmed. Therefore, the character of the turbulence must have varied considerably within a space of a few meters. In the evening, Hurricane Laura was in the Monkey River area and it dissolved and disappeared during the night south of Punta Gorda. Hurricane Laura was at least 100 miles in diameter, slow moving and of moderate force. 12 In its wake, Laura dragged along the windy, cool and rainy weather of a proper ‘‘Norther.’’ The heavy rainfall during the night and during November 21 caused extended inundation in the flat coastal areas of British Honduras and made some of the most important roads impassable. A zone several hundred meters wide of brown muddy water along the British Honduras coast made the freshwater influx clearly visible, the major rivers causing extensive protrusions of this discoloration to almost half way across the barrier reef lagoon. In conclusion, it can be stated that Hurricane Laura was an unusually late Hurricane, followed by an unusually early ‘‘Norther.’’ Fortunately, the overall effects of both on British Honduras can be considered negligible. PUBLICATIONS REVIEWS: Westoll, T.S. and Stoddart, D.R. (organizers), 1971. A discussion of the results of the Royal Society Expedition to Aldabva 1967-68, Philosophical Trans. Royal Soc. London, ser. B, 260 (836): 654 p. £19.00, $50.00. It is a curious fact that two of the most intensively studied atolls in the world were investigated for entirely different reasons. Bikini, in the Pacific, was investigated twenty-five years ago to evaluate the destructive power of atomic bombs. Now, Aldabra, in the Indian Ocean, is being given similar treatment in a laudable attempt to save it from the effects of encroaching civilization. Aldabra, an atoll that rises from the deep sea, is the last undisturbed elevated reef island in the Indian Ocean. Its continued existence in this category was threatened in 1965 when the British Ministry of Defence announced a plan to construct a Royal Air Force Staging-Post there for planes flying to the Far East. In addition to a landing strip, the plan called for the construction of a harbor for tankers, a road, anda radio transmitting station. Realizing that such construction- would probably have a devastating impact on the island and its unique fauna, the Royal Society, on behalf of scientific and conservation organizations, dispatched in 1967 a scientific expedition to study the island. In that same year the military plans were abandoned for financial reasons but the Society wisely continued its studies in the hope that the establish- ment of a scientific research station on the island would give protection to the area in the foreseeable future. The present volume presents Aldabra’s case in a most impressive manner. In September 1966 D.R. Stoddart and C.A. Wright had accompanied the survey party that went out to reconnoiter the island as a potential staging-post and radio station site. Stoddart was named a member of the Aldabra Research Committee set up by the Royal Society to work with personnel from the Smithsonian Institution, the U.S. National Academy of Sciences and the University of the Witwatersrand in a lengthy research program. Stoddart became the over- all expedition leader and is the author, or co-author, of a number of the papers in the present volume. To date, more than forty persons have participated in the expeditions to Aldabra. The biota of Aldabra is an interesting one that is intermediate between that found on sea level atolls and that of the high islands. It has many remarkable features. It supports, for example, an estimated total of 100,000 Giant Land Tortoises; it is the land base for several kinds of marine turtles; the largest colony of frigate birds in the Indian Ocean (39,000) breeds there. The island is the home of several distinct species and subspecies of land birds. One of these, a new warbler, was discovered during the present investigation; another is the rail, Dryolimnas, This is the last of the flightless birds of the western Indian Ocean but at least 1000 still survive on Aldabra. 13 Aldabra is a very smallisland. The overall area is only 365 square kilometers, the land area about 40 percent of the total. Bikini, with an overall area of more than 700 square kilo- meters, is nearly twice the size of Aldabra, but Bikini’s land area is only about 10 percent of its total. Aldabra was probably discovered during the earliest years of the 16th Century but it remained uninhabited by man for about 400 years. Scientific studies of several sorts have been made at Aldabra over a period of nearly 100 years and many of the early visitors made large collections of plants, insects and birds. An ecological survey of Aldara and nearby istands was made by Fryer in 1903 and Aldabra becam: one of the better known reef islands of the Indian Ocean. Not much additional work was done in the sixty years that followed - - not until the activity that grew out of the recent crisis. The present quarto volume with more than 650 pages and many excellent illustrations is an elaborate and satisfying treatment of many aspects of Aldabra and its life. It consists of two dozen reports. These give many data that support a series of earlier papers that presented the bare essentials of what came to be known as ‘‘The Aldabra Affair.’’ A summary of earlier scientific studies at Aldabra and neighboring islands is followed by a series of reports dealing with the physical environment - - the geomorphology, climate, tides, and shallow water environments. These studies set the stage for reports that deal with the existing flora (ground covers, mangroves, terrestrial and freshwater algae) and fauna (invertebrate and vertebrate). Among the invertebrates, special treatment is given to the ostracodes, the freshwater snails, insects and decapod crustaceans. Among the vertebrates, the tortoises, sea turtles, and birds (land birds, migrant waders and sea birds) are fully treated and a report on bats is included. Both reptiles and birds were examined for blood parasites. Some of the reportsare shortand admittedly preliminary, but all are well organized and offer a sound foundation for future work. In addition to the subjects already mentioned, there is a special report on the origin and distribution of the flora of Aldabra and an analytical study of the evolution of terrestrial faunas in the western Indian Ocean. It is surprising and certainly most gratifying that Aldabra has remained almost completely undisturbed during a lengthy and not entirely uneventful history. Commercial exploitation has been only sporadic. Unlike its three neighboring islands, Assumption, Cosmoledo and Astove, Aldabra has never been the site of serious mining operations for guano. The fact that rainfall on Aldabra is small and erratic and that areas of sandy soil are very limited has forestalled development of coconut growing. The fisheries potential is not great in the lagoon or in the waters outside. The Giant Tortoises make a profitable export but this traffic is controlled. Aldabra is the largest of a group of four islands. It has a maximum height of 8.8 meters but still retains its atoll configuration --a rim encloses a shallow lagoon with three entrances, two of which are deep channels. Large parts of the rim consist of jagged cavernous lime- stone (champignon) that is covered by dense scrub. Small areas (14 percent of the land area) at the eastern end are smoother and flatter (platin); they support a more open vegetation and furnish a home for the tortoises, frigate birds, and other species. Solution surfaces and residuals are described in fair detail. Some are satisfactorily explained but others remain problematical. There is wide lateral facies variation in the elevated lime- stones and it is not possible at present to make island-wide stratigraphic correlations. Thus, the chronological framework of the limestones remains uncertain. Some Carbon-14 determina- tions have been made on Tvidacna and Chama shells in the limestone; more are awaited. 14 The report on geomorphology is a joint effort involving four investigators. It discusses the topography and morphology of Aldabra itself, but these discussions are preceded by a wide-ranging summary of the regional setting that prepares the reader for a sketch of the probavie evolution of the island. This is an interesting story but the authors recognize that it is by no means a final account. D.R. Stoddart, the senior author of the report on geomorph- ology, has informed this reviewer (written communication March 27, 1971) that subsequent work points to a much more complex picture. Studies of the limestone outcrops have revealed half a dozen transgressions and regressions and a large land fauna. Uranium dates at 125,000 years have also been obtained; these markedly older than the equivalent c14 gates. The report on geomorphology deals only with the surficial features of Aldabra. The in- vestigators hope, however, that, following additional geophysical work (magnetic, gravity and seismic) they can carry out a drilling program that will determine the nature and age of the atoll’s foundation and reveal the major steps in its geologic history. The geophysical surveys will not injure the existing environment but the drilling would present a hazard and probably should be done on nearby Assumption Island. Deep drilling could establish a standard strati- graphic section for the western Indian Ocean comparable to that set up for the western Pacific by drilling in the Marshall Islands. The work done to date has resulted ina fairly complete inventory of the fauna and flora existing on Aldabra. This includes a determination of the importance of exotic elements present. The time has now arrived to draw up a program for a Research Station that will preserve the rich values now in existence while studying the structure and functioning of the ecosystem. Future investigations will include dredging in the waters surrounding the atoll but this will in no way injure the existing environment. The threat of exploitation has diminished because of economic reasons but it could be reactivated ata later date. Meanwhile, the excellent preliminary investigations are continuing and it is hoped that they will furnish any additional information that may be needed by the con- servationists in their effort to save undisturbed Aldabra as a nature reserve and scientific research station. Harry S. Ladd Basilewsky, P., ed. 1970. La faune terrestre de I’tle de Sainte-Héléne. Ann. Mus. Roy. Afr. Centr., Zool., 181: 1-227. This informative volume is the first inananticipated series stemming from two expeditions (1965-66) dispatched to St. Helena by the Musée Royal de l’Afrique Centrale of Tervuren, Belgium, to intensively collect the land and fresh water fauna of this ecologically devastated tropical south Atlantic island. Especially arresting is a series of excellently reproduced photo plates of a barren, weedy landscape, and tiny vestiges of native forest. Four introductory chapters, with maps, afford useful background on the present status of the island and its biota: Geography and climatology by N. Leleup; ‘‘Geological history of St. Helena in relation to its floral and faunal colonization’? by Ian Baker; Vegetation by J. Decelle; a history of faunal exploration and study by P. Basilewsky and P.L.G. Benoit; and an account of the Museum’s expeditions to the island with notes on collecting stations by Basilewsky and Decelle. Animal groups systematically treated are: Vertebrates, largely sea birds, by Basilewsky; and eight orders of insects by specialists of several nationalities. Many new records and several new insect species attest the heretofore scant knowledge of a fauna already much depleted. The insect groups treated are Collembola, Diplura, Thysanura, Odonata, Blattaria, Isoptera, Orthoptera, and Dermaptera. Bryce Decker 15 Yoshino, M.M., ed. 1971. Water balance of MonsoonAsia - -a climatologicalapproach. Honolulu, University of Hawaii Press, 1-308. $16.00. This handsomely produced volume comprises fifteen contributions, by as many Japanese authors, onthe water balance and related atmosphe- ric phenomena of Monsoon Asia andadjacent regions including the islands of Indonesia, northern Indian Ocean and the west and southwest Pacific. Yoshino’s introductory review of water balance problems and historical background is followed by four sections that take up sources and transfer of water vapor from season to season; distribution of precipitation, cloudiness and precipitable water during the summer monsoon; secular variation of precipitation and climatic change; and finally applications of several climatic classifications. There are many excellent maps and diagrams, the English is unstilted, and the bibliographies will be of interest for the access they afford to Chinese and Japanese literature on climate and weather. Bryce Decker Veevers-Carter, W.D. 1970. Island Home. 1-345, Random House, New York. Most of us know coral atolls from expeditions and short visits only. We hurry frantically to collect what we can and write what notes we can in the short time we have. We miss some of the essence of life in these microcosms. Our friend Wendy Veevers-Carter lived on one tiny 65 acre coral island, Remire, of the Amirante group in the Western Indian Ocean, for three years. One might expect that she had infinite leisure to soak up the essence of island life - - but the book gives quite the opposite impression. She was so continuously busy that one wonders when she had time to write the book. Essentially this is an account of human relations, of an effort to understand and deal with the entirely different pattern of behavior and set of mores and ethics of the Seychellois laborers. To the reader’s continual great surprise the author pulled it off astonishingly well. And in telling the story she also succeeds in giving an idea, at least, of the coral island environment. The book is fascinating reading and recommended to anyone who has ever had the yen to run away and live ona tropicalisland. F.R.F. Wodzicki, K. and: Laird, M. 1970. Birds and bird lore in the Tokelau Islands. Notornis 17: 247-276. This small group of atolls is well on its way to becoming one of the better- known of atoll archipelagoes, largely due to the investigations and publications of the authors of this paper and their colleagues in the rat and mosquito control projects carried out between 1958 and 1970. The present paper lists the bird species observed and collected and provides one to several paragraphs of observations of various sorts on each, largely descriptive and ecological, with native names and data on use of the birds as food. Several native folk tales about birds are placed on record. Occurrence, behavior, zoogeography and conservation problems are discussed. A bibliography of 44 items (not all on Tokelau birds) is provided, as well as maps of the atolls with place names. F.R.F. Bloom, A. L. 1970. FPaludal stratigraphy of Truk, Ponape and Kusaie, eastern Caroline Islands, G.S.A, Bull. 81: 1895-1904. The author interprets the swampy coastal plains, with their 2-3m of peat, as resulting from shore line progradation during a period of rising sea-level, slowing down about 4100 years ago. These flat benches have been regarded by many workers as evidence of a recent 1-2m emergence, which has been considered to fit nicely with 2m elevated reefs and notches in various parts of the world. The facts reported in Bloom’s paper add to a growing body of evidence contrary to this presumed eustatic fall in sea level. To reconcile these contrary indications, and account for the biological phenomena, which have been hiterto explained in terms of eustatic 6 foot and 11 foot benches, will be the next task of geomorphologists interested in coral reefs and islands. Nine samples of highly organic sedi- ments yielded radiocarbon dates between 1000 and 6500 years B.P. for deposition of the peat they represent and were from depths of 5 to over 20m inthe swamps. F.R.F. 489-409 O - 73 - 13 16 Tsuda, R.T. 1971. Status of Acanthaster planci and coral reefs in the Mariana and Caroline Islands, June 1970 to May 1971. Univ. Guam Marine Lab, Techn, Rept. 2: 1-27. This bro- chure is a collection of resurvey reports of the crown-of-thorns starfish situation on the reefs of Truk, Kapingamarangi, Saipan, Tinian, Aguijan, Guam, Rota, Yap, Palau, Ponape, Ant, and the atolls of the central Carolines. These are mostly areas examined by the Westinghouse teams in 1969, and data are being assembled for comparison. Maps and photos accompany the reports. In some areas control measures are being undertaken with, apparently, some success (at least temporarily). Roy Tsuda is to be complimented on getting these informative reports out so promptly and satisfactorily. F.R.F. Lemon, FE, R., etal. 1969. Biology and ecology of nitrogen, Proceedings of a conference. 1-166, Nat. Acad. Sci., Washington, D.C. This book contains nothing onislands, but summarizes briefly much of what would be needed to start investigating the nitrogen ecology of coral and other islands. It is flaweda bit by some rather naive taxonomy of certain of the higher plants discussed, but presumably the taxonomy of nitrogen-fixing bacteria is better. One is led to wonder just how adequate the treatment of nitrogen biology and ecology is, since no mention is made of the role of blue-green algae in nitrogen fixing. At least the book is a good place for the beginner to start. Some of the articles have substantial bibliographies. F.R.F. Balgooy, M.M.J. van, 1971. Plant geography of the Pacific, Blumea Suppl. 6: 1-222. Plant geography is one of the most controversy-ridden of sciences. Thismaybe, partly, because with an amorphous and diffuse mass of information of every degree of reliability (or unre- liability), plant geographers tend to choose, usually arbitrarily, very diverse techniques, parameters, and assumptions for the organization of their data. Another reason may be that after the data are organized they are not very interesting or even significant in their own right. The interest is added by what the author of the system does with them, usually by what speculations he introduces, based on them. This admirable book is no exception to any of the above, though the author has been un- usually careful in his selection of the bases for his organizational framework and unusually conservative in his speculations. One is tempted to quarrel with him on a considerable number of points, some large, most small. Many of these derive from his selection, justified at great length, of the genus as the basic unit dealt with in his investigation. A brief review is not the place to debate the propriety of sucha choice. Granted this selection as a valid basis for an analysis, Balgooy has done a superb job. He has, moreover, placed Pacific botanists in his debt for compiling the enormous mass of data presented in this volume, and for determing in a generally convincing way the ‘‘distribution types’’ to which all recognized Pacific phanerogam genera belong. One could wish he had used the term Austral for type 7 rather than the misleading ‘‘Subantarctic’’, but this is a minor objection. He has also analyzed and summarized in a thoroughly satisfactory manner all important previous essays of phytogeographicalanalysis of Pacificfloras. This, in itself, is an enormous- ly useful accomplishment. It also enables us to place his own phytogeographic scheme and conclusions in a far more satisfactory perspective. Our opinion is that the picture presented is a substantial advance over its predecessors. It has shortcomings, or course. These will be remedied only after a vast amount of additional collecting and taxonomic research has been done on Pacific plants, and when a means is found to weave into this picture considera- tions based on species and groups of species, as well as on genera. We also do not believe that such a scheme can be very sound which does not take into account the geological setting and what is known of the paleogeography of the Pacific. We cannot refrain from a few comments on Balgooy’s findings on dispersal classes and conclusions on how the island floras originated. There may be some virtue in confining assign- ment of genera to the five functional dispersal classes to cases where the mechanisms involved have actually been seen in operation. However, the inevitable infrequency of such observations 17 naturally throws most of the genera into his two catchbasket classes, those where he has no opinion on dispersal mechanisms and which are only distinguished by small versus large diaspores. The chance of actually observing a diaspore less than 3mm in diameter being dispersed by a typhoon is remote, indeed. This does not say anything about the probable frequency of such events, however. We foundno mention of the role of either typhoons or “‘jet- streams’”’ in the discussion of dispersal. Without considering these agencies, in our opinion, no general conclusions on the relation of dispersal to origin of island floras are likely to mean much. F.R.F. OTHER PUBLICATIONS: Tethys Supplements. The Station Marine d’Endoume, of the University of Marseille, had been publishing the works of its researchers in a series of Recueil des Travaux..., which was superseded in late 1969 by a quarterly journal, Téthys. The supplements to the Recueils, devoted to the works of the Station marine de Tuléar (SW Madagascar) - - which the Endoume Station was largely instrumental in creating and equipping - - have been replaced by a ‘‘tropical Indian Ocean’”’ series appearing as supplementsto Tethys. Two of these were published in 1971. No. 1 includes a list of the contents of the 10 supplements (1962-1970) to the Recueils, with annotations in French and English, and a list, also annotated, of 75 papers on the southwest Indian Ocean published or in press in other journals. The rest of this supplement includes papers on marine life and geology of the Tuléar area and of Réunion Island, where the Endoume Station has a branch station or ‘‘Antenne.”? Téthys, supplement 2, is an attempt to define terms used in coral reef morphology as exemplified by the reefs of the Tuléar region. It is presented in parallel French and English columns and is generously illustrated with diagrams and photos. Both supplements are handsomely printed. M.-H.S. Bibliographies: We have just received two more of Noel L.H. Krauss’ excellent little island bibliographies, these on Ontong Java and Rennell and Bellona, all fascinating islands among the westernmost outposts of the Polynesian culture. The Ontong Java booklet has 112 items with dates ranging from 1756 to 1969. The Rennell-Bellona bibliography has 168 items. They are published as nos. 3 and 4 of Pacific Island Studies. This series is issued and deposited in selected libraries by Mr. N.L.H. Krauss, 2437 Parker Place, Honolulu, Hawaii 96822, U.S.A. The Hawaii Institute of Marine Biology issued as its Technical Reports 20 and 21, an annotated bibliography of Kaneohe Bay [Oahu, Hawaii| and a bibliographic species list for the biota of Kaneohe Bay, both by Joleen Aldous Gordon and Philip Helfrich. Island Bibliographies Supplement, by M.-H. Sachet and F.R. Fosberg, is available from the National Academy of Sciences for $10.50. A circular announcing its publication has been sent to those on the mailing list of the Atoll Research Bulletin. Offset reproductions of the original volume, Island Bibliographies, by M.-H.-Sachet and F.R. Fosberg, may be obtained from the National Technical Information Service, Springfield, Va. 22151, U.S.A., for $6.00. Ask for document AD-738566. If you have already tried to get the reproduction of the original volume from NTIS with unsatisfactory results, try again. The difficulties seem to have been corrected and we are assured that there will be no further problem. From the eminent Pacific historian and documentalist, Professor H.E. Maude, we have received copies of his opening andclosing addresses to the Australia Unesco Seminar on Source Materials related to Research in the Pacific Area, held in Canberra, 6-10 Sept. 1971. These papers, Pacific Documentation: An Introductory Survey, and Pacific Bibliography, were not written for publication, but we certainly hope they will appear in Proceedings of this small and select Seminar, or elsewhere, without being doctored up too much, at least without loosing any of the wit and gentle sarcasm of our friend’s spoken words. 18 Bricker, O.P., ed., 1971. Carbonate cements. 1-376, The Johns Hopkins Press, Baltimore and London. This remarkable volume reports a symposium held in Bermuda in 1969. A copy for review for the next ARB Island News and Comment number has just come to hand. Stoddart, D. R. and Yonge, M., eds., 1971. Regional variation in Indian Ocean Coral Reefs. 1-572 (Zoological Society of London Symposia 28), London and New York. We have also just received a copy of this for review for the next News and Comment number (see ARB 148: 7-8). Bablet, J.P., and Cayet, O., eds., 1972. Le monde vivant des atolls. Pub. Soc. Océanistes no. 28: 1-148, Mlustrated guide toatoll naturalhistory based on the Tuamotus. To be reviewed. Bakus, G.J. 1969. Some effects of sedimentation on benthic invertebrates of atoll lagoons. Mem. Simp. Intern. Lagunas Costeras. UNAM-UNESCO 1967, Mexico 503-504. Abstract. Men- tions experiments on sediment deposition on sponges and ascidians in Fanning Island lagoon. Concludes that sedimentation plays an important role in determining which species survive, but that most species were affected adversely. Barthel, K.W., Janicke, V., and Schairer, G., 1971. Studies on the coral reef complex of Lat- sacker near Neuburg a.D (Lower Tithonian, Bavaria)., N. Jahrb. Geol. Palaont. Mohatsheft 1971 (1): 4-23, This is a descriptive account ofanupper Jurassic reef of corals and mollusks, with sediment-trapping algae, and showing effects of what are probably clionid sponges. This reef rests on older sponge reefs. Expédition Frangaise sur les récifs covalliens de la Nouvelle-Calédonie, Vol. 5: 1-307, 1971, is devoted entirely to a memoir on the scleractinian corals of French Melanesia (New Caledonia to New Hebrides), by J.-P. Chevalier, beautifully illustrated with line drawings and 38 plates of photos. See ARB 148: 29, 1971. Journal of the Marine Biological Association of India, vol. 11 (1&2), was issued in April, 1971 and is dedicated to Dr. Santappan Jones. It contains several papers on coral reefs and cays, corals and other marine organisms. Caribbean Marine Biological Institute, Curvagao.... Collected Papers, 6(84-97): 1969-1971. The individual papers that seem likely to be of interest to ARB readers are listed as appropriate below, with references to the original places of publication. Captain Cook’s Florilegium. An item in The Times (London), January 17, 1972, announces the imminent appearance of this title, the book being printed by the Royal College of Art from copper plates engraved in 1780. After 200 years’ delay, the plates are being used to illustrate plants collected by Joseph Banks on the Endeavour voyage, 1768-1771. Only 120 copies will be made. B.D. Zinderen Bakker Sr., E.M. van, Winterbottom, J. M., and Dyer, R.A., eds. 1971. Marion & Prince Edward Islands. Cape Town, A.A. Balkema, xi, 427 pp. Marion and Prince Edward Islands are sub-Antarctic rather than tropical, but the volume is sucha good example of a description and interpretation of an island ecosystem that we wish to call our readers’ attention: to it. It presents the results of the South African Biological and Geological Expedition, 1965- 1966, to these bleak volcanoes situated southeast of Cape Town, and is an expedition report in the classic manner. The photos, alone, are worth the reader’s time. F.R.F. BRIEFLY NOTED ITEMS: Coral Islands of the Western Indian Ocean: ARB 136, on Coral Islands of the Western Indian Ocean, included a series of tables listing the number of species of insects in various orders recorded from each island in the publications of the Percy Sladen Trust Expedition (Farquhar, 19 table 2; Cosmoledo, table 7; Astove, table 11; Assumption, table 14; Desroches, table 16; Remire, table 18). Since the Bulletin went to press, some additional records have been found in the literature, inpapers overlooked whenthe Bulletin was in preparation. These records are noted below, and the papers themselves shouldbe added to the lists of references on each of the islands. Farquhar Diptera 1 species Bezzi 1923 Cosmoledo Diptera 1 Austen 1920 Astove Hemiptera 1 Green and Laing 1921 Diptera 1 Austen 1920 Assumption Hemiptera il Green and Laing 1921 Diptera 1 Bezzi 1923 Desroches Diptera 1 Austen 1920 1 Bezzi 1923 Remire Coleoptera 1 Fleutiaux 1922 Thysanura 1 Carpenter 1916 The references are as follows: Austen, E.E. 1920. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, and in 1907-1909, under Mr. J. Stanley Gardiner, M.A. Diptera: Tabanidae. Bull. Ent. Res. 11: 43-45. Bezzi, M. 1923. Diptera, Bombyliidae and Myiodaria (Coenosiinae, Muscinae, Calliphorinae, Sarcophaginae, Dexiinae, Tachininae), from the Seychelles and neighbouring islands. Parasitology 15: 75-102. Carpenter, G.H. 1916. The Apterygota of the Seychelles. Proc. Roy. Irish Acad. B, 33: 1-70. Fleutiaux, E. 1923. Coleoptera: Melasidae et Elateridae des Séchelles et des iles voisines. Trans. Ent. Soc. London 1922: 398-436. Green, E.E. and Laing, F. 1921. Coccidae from the Seychelles. Bull. Ent. Res. 12: 125-128. D.R.S. Additional items on Chagos: Feuga, Jean. 1946. L’Emden, croiseur corsaire. Paris, Editions Charcot, 173 pp. Diego Garcia pp. 117-145. Fry, H.T. 1967. Early British interest in the Chagos Archipelago and the Maldive Islands. Mariner’s Mirror 53: 343-356. Spray, W.A. 1970. British surveys inthe Chagos Archipelago and attempts to form a settlement at Diego Garcia in the late eighteenth century. Mariner’s Mirror 56: 59-76. D.R.S. Old items ommitted from Atoll Bibliography and Supplement: Anon. 1830. Some account of the Cocos or Keeling Islands: and of their recent settlement. Gleanings in Science (Calcutta) 2(22): 293-301; reprinted in Jour. Malay. Br. R. Asiatic Soc. 25(4): 174-191, 1952. Jagt, H. van der 1831. Beschrijving der Kokos- of Keeling-eilanden. Verh. Batav. Gen. v. Kunsten en Wetenschappen, Batavia 13: 293-322; translated in Jour. Malay. Br. R. Asiatic Soc. 25(4): 148-159, 1952. Nesbit, J.C.1859. Thehistory and properties of the different varieties of natural guanos. London, Rogerson and Tuxford, 1-50+ 2. (listedin Atoll Supplementas not seen). Analyses of guano from Pedro Keys, Swan I., Baker I. (WestIndies), Bird I. (St Vincent), Sombrero, Jarvis. Ross, J.C. 1836. On the formation of the oceanic islands in general, and of the coralline in particular. Singapore Free press, 2 June 1836; reprinted in Jour. Malay. Br. R. Asiatic Soc. 25(4): 251-260, 1952. D.R.S. Some recent doctoral theses on coral reef topics: Barnes, D.J. 1971. A study of growth, structureandform in modern coral skeletons. Univer- sity of Newcastle upon Tyne, School of Physics, Department of Geophysics and Planetary Physics. Ph.D. thesis, 180 pp. *Buchanan, H. 1970. Environmental stratigraphy of Holocene carbonate sediments near Frazers Hog Cay, British West Indies. Columbia University, Ph.D. thesis, 241 pp. 20 *Chase, C.G. 1970. Tectonic history of the Fiji Plateau. University of California at San Diego, Ph.D. thesis, 95 pp. *Conaghan, P.J. 1968. Marine geology of the southern tropical shelf, Queensland. University of Queensland Ph.D. thesis, 508 pp. *Freeland, G.L. 1971. Carbonate sedimentation in a terrigenous province: the reefs of Veracruz, Mexico. Rice University, Ph.D. thesis, 367 pp. *Garrett, P. 1971. The sedimentary record of life on a modern tropical carbonate tidal flat, Andros Island, Bahamas. Johns Hopkins University, Ph.D. thesis, 259 pp. *Lang, J.C. 1970. Inter-specific aggression within the scleractinian reef corals. Yale University, Ph.D. thesis, 177 pp. *Roberts, H.H. 1969. Recent carbonate sedimentation, North Sound, Grand Cayman Island, British West Indies. Louisiana State University, Ph.D. thesis, 118 pp. Till, R. 1968. Some aspects of the geochemistry of recent Bahaman carbonate sediments from the Bimini lagoon. University of Sheffield, Ph.D. thesis, 166 pp. *Upchurch, S.B. 1970. Sedimentation on the Bermida platform. Northwestern University, Ph.D. thesis, 243 pp. *Vacher, H.L. 1971. Late Pleistocene sea-level history: Bermuda evidence. Northwestern University, Ph.D. thesis, 186 pp. *Ward, W.C. 1970. Diagenesis of Quaternary eolianites of N.E. Quintana Roo, Mexico. Rice University, Ph.D. thesis, 243 pp. *Available in microfilm or xerox form from University Microfilms. D.R.S. Island papers, terrestrial, compiled by Bryce G, Decker Baltzer, F., and Lafond, L.-R. 1971. Marais maritimes tropicaux. Rev. Géogr. Phys. Géol. Dynam. 8(2): 173-196. Considers edaphic regimes of mangrove and other littoral marsh types. Beck, H. 1970. Germania in Pacifico. Der deutsche Anteil an der Erschliessung des Pazifischen Beckens. Akad. Wiss. Lit. Mainz Abh. math.-nat. Kl. Jahrg. 1970(3): 233- 327. This concise historical account of the contributions by Germans to knowledge of the Pacific area will be welcomed for its extensive bibliography; includes con- temporary works. Blanc, C.P. 1971. Les reptiles de Madagascar et des fles voisines. Annls. Univ. Madagascar 8: 95-178. Important catalogue and discussion for western Indian Ocean islands, including reef islands. DRS. Brownlie, G., and Philipson, W.R. 1971. Pteridophyta of the southern Cook Group. Pac. Sci. 25(4): 502-511. Includes published and recent unpublished records. Bryan, W.B. 1971. Coral Sea drift pumice stranded on Eua Island, Tonga, in1969. Bull. Geol. Soc. Am. 82: 2799-2812. Also includes data from One Tree Island, Great Barrier Reef; unaware of Sachet, 1955. DRS. Bullard, E.C. 1957. Gerald Ponsonby Lenox-Conyngham 1865-1955. Biog. Mem. Fellows Roy. Soc. 3: 129-140. Author of paper on Great Barrier Reef, Geog. Jour. 70: 1925. Challinor, D., and Wingate, D.B. 1971. The struggle for survival of the Bermuda cedar. Biol. Conserv. 3: 220-222. Connell, D.W. 1971. The Great Barrier Reef Conservation issue--a case history. Biol. Conserv. 3: 249-254. Coulson, F.I.E. 1971. The geology of western Vanua Levu (an explanation of Vanua Levu Sheets 3&9). Geol. Surv. Dept. (Fiji) Bull. 17: 1-49. Multi-color maps in endpocket; 1: 50,000. 21 Cowan, C.F. 1970. The insects of the Coquille voyage. Jour. Soc. Bibl. Nat. Hist. 5: 358-360. Clarifies bibliography and dates of portions of Duperrey’s Voyage autour du Monde dealing with insects. FRF. Dent, D.W., and Preedy, B.H. 1970. Structure of the troposphere over Gan. Met. Mag. 99: 304-313. DeRoy, T. 1972. Giant tortoises on a volcano. Pac. Disc. 25(2): 14-20. Informative popular account of tortoise behavior on Isabella (Albemarle) I.; excellent photos. Draeger, R.H., and Lee, R.H. 1953. Meteorological data Eniwetok Atoll. Bethesda, Md., Naval Medical Research Institute, National Naval Medical Center, Memorandum Report 53-8 related to Project NM 006 012.01: 51-71. Tabulation of 1949-50 data, Japtan Islet. DRS. Eliasson, U. 1970. Studies in Galapagos plants VII: Chromosome numbers of some endemic species. Botaniska Notiser 123: 149-154; (IX). New taxonomical and distributional records, 346-357. Fisher, M.L. 1970. The albatross of Midway Island: anaturalhistory of the Laysan albatross. 1-164, Carbondale, Southern Illinois Univ. Press; London, Feffer and Simons. $5.95. Gladwin, T. 1970. East is a big bird: navigation and logic on Puluwat Atoll. i-xviii, 1-242, Cambridge, Harvard Univ. Press. $9.95. Goff, M.L. 1971. New records of chiggers (Acarina, Trombiculidae) from the northwestern Hawaiian Islands. Jour. Med. Ent. 8(4): 456. Pearl and Hermes Atoll and Laysan I. Gould, S.J. 1969. Character variation in two land snails from the Dutch Leeward Islands: geography, environment and evolution. System. Zool. 18(2): 185-200. Heatwole, H. 1971. Marine-dependent terrestrial biotic communities on some cays in the Coral Sea. Ecology 52: 363-366. Bird and invertebrate communities on non-vegetated cays on the Great Barrier Reef, related tomarine communities adjacent. FRF. Herbst, D. 1971. A new Euphorbia (Euphorbiaceae) from Hawaii. Pac. Sci. 25(4): 489-490. E. haeleeleana from Kauai. Lee, B.K.H., and Baker, G.E. 1972. An ecological study of the soil microfungi in a Hawaiian mangyove Swamp. Pac. Sci. 26(1): 1-10; Environment and the distribution of microfungi in a Hawaiian mangrove swamp: 11-19. Lewis, D. 1971. ‘“‘Expanding’’ the target in indigenous navigation. Jour. Pac. Hist. 6: 83-95. Discussion of birds, clouds, swell patterns, horizon brightness, deep phosphorescence and other indicators of proximity of land in the Pacific, based on interviews with islanders during a west Pacific cruise in 1969. DRS. Nelson, J.B. 1971. The biology of Abbott’s booby Sula abbotti. Ibis 113: 429-467. Christmas I., Indian Ocean. Odum, H.T. 1971. Environment, power and society. 1-331, New York, London, Sydney, Toronto, Wiley-Interscience. Facing p. 104 is an ecological energy diagram for Lamotrek Atoll, Caroline Is., utilizing data from Alkire, 1965. 22 Ono, M. 1967. Chromosome number of Scalesia (Compositae), an endemic genus of the Galapagos Islands. Jour. Jap. Bot. 42(12): 353-360. See also Eliasson, 1970, and Ono, 1971 for further counts. 2SS5SS5EhR5e 1971. Chromosome number of Scalesia ... (2). Jour. Jap. Bot. 46(11): 327-334. Posnett, N.W., and Reilly, P.M. 1971. Bahamas. Foreign and Commonwealth Office Land Resources Div. Overseas Development Administr., Surbiton (England), Land Resource Bibliography No. 1: 1-74. Ramage, C.S. 1970. Monsoon meteorology. 1-304, New York and London, Academic Press. $15.00. Not seen. Rolle, F.J. 1966. Notes on birds from some West Indian islands. Stahlia 7: 1-3. Islets off east coast of Puerto Rico, and Isla de Aves, Lesser Antilles. Pineros I., Cabeza de Perros I., Palominos I., Palominitos I., Cayo Obispo. FRF. Scheer, G. 1960. Eine neue Rasse des Teichreihers Ardeola grayii (Sykes) von den Malediven. Senck. Biol. 41: 143-147. Describes Maldivian race of Indian pond heron; notes on occurrence and ecology. Schofield, J.C. 1971. Note on high sea-level evidence from Lau Islands, southwest Pacific. N.Z. Jour. Geol. Geophys. 14: 240-241. Tabulation of data from Ladd and Hoffmeister, 1945; main clusterings at ca 550-600, 300-350, 250-275, 200-230, 170-185, 110-130 and 70 ft. DRS. Smith, A.C. 1971. Studies of Pacific island plants, XXII; new flowering plants from Fiji. Pac. Sci. 25(4): 491-501. Describes 12 new species in 6 families. Vogl, R.J., and Henrickson, J. 1971. Vegetation of an alpine bog on east Maui, Hawaii. Pac. Sci. 25(4): 475-483. Floristic list, notes on occurrence, soils, comparison with other Hawaiian mountain bogs; photos. Wagenaar Hummelinck, P., and Roos, P.J.1969. Eennatuurwetenschappelijk onderzoek gericht op het behoud van het Lac op Bonaire. Nieuwe Westind. Gids 47: 1-28, 55 plates. A scientific survey aimed at the preservation of this largest salt lagoon in the Netherlands Antilles. AY eS Weber, D. 1971. Pinta, Galapagos: uneileasauver. Biol. Conserv. 4(1): 8-12. Native vegeta- tion under severe pressure from goats introduced in 1959; maps; photos. Westermann, J.H. 1968. Om het voortbestaan van de flamingo’s van Zuid-Bonaire 1957-1968. Nieuwe Westind. Gids 46: 195-231. Conservation of flamingos on South Bonaire. Island and reef papers, marine, compiled by Linda Smith Baldwin, W.J. 1972. A new genus and new species of Hawaiian gobiid fish. Pac. Sci. 26(1): 125-128. Benjamin, G.J. 1970. Diving intothe blue holes ofthe Bahamas. Nat. Geog. Mag. 138: 347-363. Holes to 230 feet deep. Bruce, A.J. 1972. A report ona small collection of pontoniid shrimps from Fiji... Pac. Sci. 26(1): 63-86. 23 Bryan, W.H. 1948. H.C. Richards memorial address. Proc. R. Soc. Queensland 59: 141-150. Biographical; bibliography. Campbell, A.C., and Ormond, R.F. 1970. The threat of the ‘‘Crown-of-thorns’’ starfish (Acanthaster planci) to coral reefs in the Indo-Pacific area: observations on a normal population in the Red Sea. Biol. Conserv. 2: 246-251. Coomans, H.E. 1970. Volksnamen voor weekdieren op de Nederlandse Antillen (Vernacular names of molluscs in the Netherlands Antilles). Natuurwet. Werkgroep Ned. Antillen 19: 158-186. Alphabetical and systematic lists. Creutzberg, F., et al. 1969. Speciaal Koraalrifnummer. Stinapa No. 4: 1-40. Dana, T.F. 1971. On the reef corals. of the world’s most northern atoll (Kure: Hawaiian Archipelago). Pac. Sci. 25: 80-87. Faure, G., and Montaggioni, L. 1971. Les récifs coralliens Sous-Le-Vent de 1’tle Maurice (Archipel des Mascareignes, Ocean Indien): morphologie et bionomie dela pente externe. C.R. Acad. Sci. Paris 273,D: 1914-1916. Franzisket, L. 1970. Zur Okologie der Fadenalgen im skelett lebender Riffkorallen. Zool. Jahrb., Abt. fur allg. Zool. und Physiol. der Tiere 74: 246-263. Gallagher, B.S., et al. 1971. Tides and currents in Fanning Atoll lagoon. Pac. Sci. 25(2): 191-205. Gardiner, J.S. 1932. The Great Barrier andthe formation of coral reefs. Nature 129: 748-749. Gerlach, S.A. 1961. Ueber Gastrotrichen aus dem Meeressand des Malediven. Zool. Anz. 167: 471-475. Gordon, D.C., Jr. 1971. Organic carbon budget of Fanning Island lagoon. Pac. Sci. 25(2): 222-227. Gordon, D.C., Jr., Fournier, R.O., and Krasnick, G.J. 1971. Note on the planktonic primary production in Fanning Island lagoon. Pac. Sci. 25(2): 228-233. Gosline, W.A. 1971la. Functional morphology and classification of teleostean fishes. 1-208, Honolulu, Univ. Hawaii Press. A general treatise on fishes, not emphasizing island fishes, but very likely of interest to many of our marine biologist readers. FRF. ===------- 1971b. The zoozgeographic relationships of Fanning Island inshore fishes. Pac. Sci. 25(2): 282-289. Guinther, E.B. 1971. Ecologic observations on an estuarine environment at Fanning Atoll. Pac. Sci. 25(2): 249-259. Guppy, H.B. 1886. The coral reefs of the Solomon Islands. Nature 35: 77-78. Henry, D.P. 1958. Intertidal barnacles of Bermuda. Jour. Mar. Res. 17: 215-234. Hoek, C. van den. 1969a. Algal vegetation-types along the open coasts of Curacao, Netherlands Antilles. I, I. Proc. Koninkl. Nederl. Akad. van Wetensch. C, 72(5): 537-577. 24 Hoek, C. van den. 1969b. Notes on Cladophora (Chlorophyceae). 1, U, Jour. Phycol. 5(2): 128-134, 134-136. Holeman, J. and Kohn, A.J. 1970. The identity of Conus mappa (Lightfoot), C. insularis Gmelin, C, auvantius Hwass in Bruguiére, and Hwass’ saniraspeenic taxa of C, cedonullz. Jour. Conch. 27: 135-137. Johannes, R.E., and Wiebe, W.J. 1970. Method for determination of coral tissue biomass and composition. Limnol. Oceanog. 15: 822-824. Kay, E.A. 1971. The littoral marine molluscs of Fanning Island. Pac. Sci. 25(2): 260-281. Kohn, A.J. 1970. Food habits of the gastropod Mitra litterata Lamarck: relation to trophic structure of the intertidal marine bench compniinity in Hawaii. Pac. Sci. 24(4): 483-486. Se et a ate tl 1971. Diversity, utilization of resources, and adaptive radiation in shallow- water marine invertebrates of tropical oceanic islands. Limnol. Oceanog. 16(2): 332-348. Kuhlmann, D.H.H. 1970a. Studien uber physikalische and chemische Faktoren in kubanischen Riffgebieten. Acta Hydrophysica 15(2): 105-152. Much more extensive discussions of Cuban coral reef topics in this and other items appear in Kuhlmann, 1971b. English summary. ---------- 1970b. Die Korallenriffe Kubas. I. Genese und Evolution (The coral reefs of Cuba. I. Genesis and evolution). Int. Revue gesamte Hydrobiol. 55(5): 729-756. English summary. Boone a soe 1971a. Die Korallenriffe Kubas. II. Zur Okologie der Bankriffe und ihrer Korallen (The coral reefs of Cuba. II. On the ecology of the bank reefs and their corals). Int. Revue gesamte Hydrobiol. 56(2): 145-199. English summary. ---------- 1971b. Die Entstehung des westindischen Korallenriffgebietes (The origin of the West Indian field of coral reefs). Wiss. Zeitschrift Humboldt-Univ. Berlin, Math.- Nat. R. 20(4/5): 675-695; Zur Methodik der Korallenriffuntersuchung (On the methods of studying coral reefs): 697-705; Uber einige physikalische und chemische Faktoren in kubanischen Korallenriffgebieten (On some physical and chemical factors in Cuban coral reef areas): 707-719; Untersuchungen zur Okologie und Entstehung kubanischer Bank-Riffe (Studies on the ecology and the origin of Cuban bank reefs): 721-775. English summaries. Lawrence, J.M. 1972. Carbohydrate and lipid levels in the intestine of Holothuria atra (Echinodermata, Holothuroidea). Pac. Sci. 26(1): 114-116. Lewis, J.B., et al. 1968. Comparative growth rates of some reef corals in the Caribbean. McGill Univ. Marine Sciences Manuscript Rept. 10: 1-26. ---------- 1969. Latitudinal differences in growth rates of some intertidal marine molluscs in the Caribbean. McGill Univ. Marine Sciences Manuscript Rept. 12: 1-89. Marcus, E., and Marcus, E. du Bois-Reymond. 1970. Opisthobranchs from Curagao and faunistically related regions. Studies on the fauna of Curagao and other Caribbean islands 33(122): 1-129. Maxwell, W.G.H. 1970. Deltaic patterns in reefs. Deep Sea Res. 17: 1005-1018. 25 McGill University. 1970. Annual Report, Marine Sciences Centre: 1-40. Scope of studies includes Caribbean. Meyer, D.L. 1969. Functional morphology and living habits of shallow water unstalked crinoids of the Caribbean Sea. Abstract of paper presented at annual meeting, Geol. Soc. Amer., Atlantic City, Nov. 12, 1969. soon soSeseo 1971. The collagenous nature of problematical ligaments in crinoids (Echinoder- mata). Marine Biology 9(3): 235-241. Newell, N.D. 1971. An outline history of tropical organic reefs. Am. Mus. Novit. 2465: 1-37. A paleobiologist’s view of the evolution of coral reef ecosystems as interpreted from the unusually complete fossil record. Ohata, C.A., et al. 1972. Diurnal rhythm of body temperature in the Hawaiian monk seal (Monachus schauinslandi), Pac. Sci. 26(1): 117-120. Pearson, R.G., and Endean, R. 1969. A preliminary study of the coral predator Acanthaster planci (L.) (Asteroidea) on the Great Barrier Reef. Dept. of Harbours and Marine (Qld) Fisheries Notes 3(1): 29-55. Qasim, S.Z. 1970. Some characteristics of a Trichodesmium bloom in the Laccadives. Deep-Sea Res. 17: 655-660. Renaud-Mornat, J.C., Salvat, B., and Bossy, C. 1971. Macrobenthos and meiobenthos from the closed lagoon of a Polynesian atoll, Maturei Vavao (Tuamotu). Biotropica 3: 36-55. Gives brief description of atoll, quantitative sampling and analysis of lagoon fauna and sediments; map; diagrams; photos; comparison with Mururoa Atoll. Roy, K.J., and Smith, S.V. 1971. Sedimentation and coral reef development in turbid water: Fanning Lagoon. Pac. Sci. 25(2): 234-248. Salvat, B., and Ehrhardt, J.P. 1970. Mollusques de Vile Clipperton. Bull. Mus. Hist. Nat. Paris 42(1): 223-231. Smith, D.P. 1969. Daily migrations of tropical sea urchins. Am. Zool. 9: 1075. Abstract of motion picture. Curacao. Smith, S.V. 1971. Factor analysis: a tool for environmental studies. Mar. Tech. Soc. Jour. 5(6): 15-19. Smith, S.V. et al. 1971. Flux of suspended calcium carbonate (CaCOg), Fanning Island lagoon. Stacey, C.C.D. 1971. Coral islands: their origins and morphology, with reference to the Maldive Islands. Don (Sheffield Univ. Geog. Soc.) 14: 31-33. Worthless discussion of major reef theories as they apply to the Maldives mainly based on Davis, The coral reef problem, 1928; based on a visit to Gan but with no field observations of any value. DRS. Stock, J.H. 1970. Notodelphyidae and Botryllophilidae (Copepoda) from the West Indies. Studies on the Fauna of Curagao and other Caribbean islands 34(123): 1-45. 26 Stringfield, V.T., and Legrand, H.E. 1971. Effects of karst features on circulation of water in carbonate rocks in coastal areas. Jour. Hydrol. 14: 139-157. Andros blue holes. Summerhayes, C.P. 1971. Lagoonal sedimentation at Aitutaki and Manuae in the Cook Islands: a reconnaissance survey. N.Z. Jour. Geol. Geophys. 14: 351-363. Tixier-Durivault, A. 1957. Les Alcyonaires du Muséum. 1. Famille des Alcyoniidae - - IV. Genre Lobophytum (fin.) Bull. Mus. Hist. Nat. Paris Il, 29: 106-111. L. venustum n. sp. from Aldabra. Tsuda, R.T., Larson, H.K., and Lujan, R.J. 1972. Algal growth on beaks of live parrotfishes. Pac. Sci. 26(1): 20-23. Uchupi, E., et al. 1971. Structure and origin of southeastern Bahamas. Bull. Am. Ass. Petr. Geol. 55: 687-704. Van Dorn, W.G. 1970. Tsunami response at Wake Island: a model study. Jour. Mar. Res. 28: 336-344. Vastano, A.C., and Reid, R.O. 1970. Tsunami responses at Wake Island: comparison of the hydraulic and numerical approaches. Jour. Mar. Res. 28: 345-356. Veevers, J.J. 1969. Palaeogeography of the Timor Sea region. Palaeogeog. Palaeoclim. Palaeoecol. 6: 125-140. Deep well dug on Ashmore Reef, Timor Sea: 2376m carbonates back to Upper Cretaceous, mostly shallow water. Womersley, H.B.S. and Bailey, A. 1970. Marine algae of the Solomon Islands. Phil. Trans. R. Soc. London B, 259: 257-352. Yonge, C.M. 1962a. Dr. A.P. Orr. Nature 196: 719. Member Great Barrier Reef Expedition 1928-1929. =--------- 1962b. Thomas Alan Stephenson 1898-1961. Biogr. Mem. Fellows R. Soc. London 9: 137-148. Obituary. U. S. GOVERNMENT PRINTING OFFICE: 1973 O - 489-409 aos AFS/ NOS. 163-164 Sr December 31, 1972 ATOLL RESEARCH BULLETIN 163. The Natural History of Gardner Pinnacles, Northwestern Hawaiian Islands by Roger B. Clapp 164. The Natural History of Kure Atoll, Northwestern Hawaiian Islands by Paul W. Woodward Issued by THE SMITHSONIAN INSTITUTION Washington, D.C., U.S.A. ATOLL RESEARCH BULLETIN NOS. 163-164 163. The Natural History of Gardner Pinnacles, Northwestern Hawaiian Islands by Roger B. Clapp 164. The Natural History of Kure Atoll, Northwestern Hawaiian Islands by Paul W. Woodward Issued by THE SMITHSONIAN INSTITUTION with the assistance of The Bureau of Sport Fisheries and Wildlife U.S. Department of the Interior Washington, D.C., U.S.A. December 31, 1972 ACKNOWLEDGMENT The Atoll Research Bulletin is issued by the Smithsonian Institution as a part of its Tropical Biology Program. It is co- sponsored by the Museum of Natural History, the Office of Environ- mental Sciences, and the Smithsonian Press. The Press supports and handles production and distribution. The editing is done by the Tropical Biology staff, Botany Department, Museum of Natural History. The Bulletin was founded and the first 117 numbers issued by the Pacific Science Board, National Academy of Sciences, with financial support from the Office of Naval Research. Its pages were largely devoted to reports resulting from the Pacific Science Board's Coral Atoll Program. The sole responsibility for all statements made by authors of papers in the Atoll Research Bulletin rests with them, and statements made in the Bulletin do not necessarily represent the views of the Smithsonian nor those of the editors of the Bulletin. Editors F. R. Fosberg M.-H. Sachet Smithsonian Institution Washington, D. C. 20560 D. R. Stoddart Department of Geography University of Cambridge Downing Place Cambridge, England ATOLL RESEARCH BULLETIN No. 163 THE NATURAL HISTORY OF GARDNER PINNACLES, NORTHWESTERN HAWAIIAN ISLANDS by Roger B. Clapp Issued by THE SMITHSONIAN INSTITUTION with the assistance of The Bureau of Sport Fisheries and Wildlife U.S. Department of the Interior Veisosiaecoml. Ds Co, We So No Decembenm sia Oe i si Rg Y Bote act ayer ‘ } Veil) tds a ee i 1 {) = ¥ Pha Dae ny aD . iit “ 5 7 oy a» y i i - ‘ f f = Pe eT , ee M _ UN ch ue ty * 5h eta Pt ay i t ‘ ye fs ay ip i ca Sea i : * gl = Hens e fe “om 1 * 1 Es ¥ ie y ‘ a om ee fy? : ae Lube j ay Tare the: } a ® ees x iy ( a] i » ' rea ea | . ates | | ete A eee Ae tae ‘ 1 an a i i i y ’ r r 0 ‘ j 2 mtg x E R , y 7 b 3 rt i ie a ‘ pi : ; hi , a x + J | A r ¥ ¥ ve } iv ¥ ” ' i J i = = to iy a ‘ gn nee 7 i an ; At ; jt i x : : aro ool? ed) - - , " . , , , 1 ' ary h Ss i u yi eat F i ' N P h : { ( 1 ‘ é - ‘ : Le «s iF i = . - 4 ais a \ F L) i © i ie i : ny f ies . ay 7 os an I af ‘ a ' ’ if Ai i . . | rf - A | i ; nt E if r * : . ' . ‘ - i E x “ i , , . t - K i , ta = i . a . i 2 i i * iL i . . vy 7 v - ¥ ry wy j i 4 a n ae | } 4 ar x ) 7 ies oF pin] = i ‘ ri 4 + res) é ee no, i Z Fe vi = ¥, ‘ y 4 ‘ ; Es : Q es Th : ote 2 1 pr Z ¥ oS . u 2 tig sf ' r yy = ‘ a - F ; Ss 4 7 : f " ft. 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S100 GOO ODIO mIOId O10 6.8100 6 6.6.6 WMEISCMe GMO ve cre telepcuslicvel cllelel'sl elles p00 oO OOOO OO OOOO Ob OOOUOOOO OOS COOLOBVo oon nooo FOO DOOD OOOO ODDO DOOR OOD OOOO ODO OO OOODGDOOOON BOW Myers tenonderorencieneneicnelevelcionelcncloncleiellelsielicnarciorciorclle/enel oleic! +) elieltetc WSiPCSIOIACS WWI. 6 OOO 0 DOO COOGO ODO DOO DOADOH OO OOOO 0O0D0O00dO ROLLS Ciael MEMES 5 oo ooo COC COON DODD GOD OOO U OOO ODOddS BHBISCUST Te Weveee ete’ chelewencl one ietiane over ciereicienecioucbelehe. sictesienetcrse her cte'e.els COMPOOSUGLOM Oi WAS JAVA. Soa cdoodnon600C Gado OUbb0O Historical Changes) in the Size of Populations....... BNAGUNSS 5 SaSROHOGb'D 00 O60 OOOO DOD DH OO OCU OOO GoD Oe GoD SOC CHIMCTAS sepMeded ati leyeuenoley erelotetel el etekelelcheieioieheicrsliey slo) Coy Ws So uNe Decienbernsl One TABLE OF CONTENTS Table of Contents Figures Tables Introduction History 19th Century 20th Century Description Vegetation Climate Scientific Visits Biological Surveys Other Scientific Visits Kure Avifauna Introduction Breeding Species Common Non-breeding Species Rare Non-breeding Species Accidental Species Populations Annual Population Cycles Breeding Cycles Factors Affecting Nesting Success Polynesian Rats Storms High Tides Rain Human Disturbance Great Frigatebird Predation seals Dogs Starvation Miscellaneous Factors Movement Species Accounts Black-footed Albatross, Diomedea nigripes Laysan Albatross, Diomedea immutabilis Black-footed x Laysan Albatross, Diomedea nigripes x immutabilis Northern Fulmar, Fulmarus glaciali: glacialis rodgersii Bonin Petrel, Pterodroma h. hypoleuca Kermadec Petrel, Pterodroma neglecta Murphy's Petrel, Pterodroma ultima Bulwer's Petrel, Bulweria bulwerii Wedge-tailed Shearwater, Puffinus pacificus sooty Shearwater, Puffinus griseu griseus Christmas Shearwater, Puffinus nativitatis Leach's Storm Petrel, Oceanodroma leucorhoa iv Sooty Storm Petrel, Oceanodroma tristrami Red-tailed Tropicbird, Phaethon rubricauda White-tailed Tropicbird, Phaethon lepturus lepturus Blue-faced Booby, Sula dactylatra Brown Booby, Sula Leucogaster Red-footed Booby, Sula sula Great Frigatebird, Fregata minor minor Lesser Frigatebird, Fregata “ariel Black-crowned Night Heron, Nycticorax nycticorax (hoactli?) Emperor Goose, Philacte canagica European Widgeon, Mareca penelope Pintail, Anas acuta Tufted Duck, Aythya fuligula Bufflehead, Bucephala albeola Peregrine Falcon, Falco peregrinus (pealei?) Dotterel, Eudromias morinellus American Golden Plover, Pluvialis dominica Black=-bellied Plover, Squatarola | squatarola Ruddy Turnstone, Arenaria interpres Pintail Snipe, Capella stenura Common Snipe, Capella gallinago delicata Bristle-thighed Curlew, Numenius tahitiensis Wood Sandpiper, Tringa glareola Wandering Tattler, Heteroscelus incanum Lesser Yellowlegs, Totanus flavipes Sharp-tailed Sandpiper, Erolia acuminata Pectoral Sandpiper, Erolia melanotos Sharp-tailed or Pectoral Sandpiper, Erolia acuminata or melanotos Dunlin, Erolia alpina sakhalina Long-billed Dowitcher, Limnodromus scolopaceus Western Sandpiper, Ereunetes mauri Bar-tailed Godwit, Limosa lapponica Ruff, Philomachus pugnax Sanderling, Crocethia alba Red Phalarope, Phalar Phalaropus - fulicarius Ring-billed Gull, Larus delawarensis Herring Gull, Larus argentatus vegae Slaty-backed Gull, Larus schistisagus Glaucous-winged Gull, Larus glaucescens Glaucous Gull, Larus hyperboreus Black-legged Kittiwake, Rissa tridactyla Black Tern, Chlidonias niger Arctic Tern, Sterna paradisea Sooty Tern, Sterna fuscata Gray-backed Tern, Sterna lunata Brown Noddy, Anous stolidus Black Noddy, Anous tenuirostris Page Sve White Tern, Gygis alba Horned Puffin, Fratercula corniculata Short-eared Owl, Asio flammeus (flammeus? ) Skylark, Alauda arvensis pekinensis Barn Swallow, Hirundo rustica Water Pipit, Anthus spinoletta japonicus Red-throated Pipit, Anthus cervinus House Sparrow, Passer domesticus Snow Bunting, Plectrophenax nivalis townsendi Sand Island Avifauna Specimens Kure Reptiles Species Accounts Hawksbill Turtle, Eratmochelys imbricata Green Sea Turtle, Chelonia mydas Stump-toed Gecko, Gehyra m mutilata House Gecko, Hemidactylus frenatus Kure Mammals Species Accounts Squirrel Monkey, Saimiri sp. Polynesian Rat, Rattus exulans Goose-beaked Whale, Ziphius cavirostris Sperm Whale, Physeter catadon Hawaiian Spinner Dolphin, Stenella roseiventris Bottle-nosed Dolphin, Tursiops truncatus Hawaiian Monk Seal, Monachus schauinslandi Domestic Dog, Canis familiaris Pig, Sus scrofa Acknowledgments Literature Cited vi D= 30 p-4. =). . FIGURES Green Island, Kure Atoll. Vegetation of Green island, Kure Atoll. Aerial view of Green Island, Kure Atoll, looking north, 6 April 1966. Aerial view of Green Island, Kure Atoll, looking south, 27 February 1968. Aerial view of building complex on Green Island, Kure Atoll, 13 October 1964. -North antenna field from north tower, Green Island, Kure Atoll, 1 July 1968. Blue-faced Boobies in north antenna field, Green Island, Kure Atoll, 1 July 1968. Looking towards west point from radar reflector, Green Island, Kure Atoll, 1 July 1968. Breeding Red-footed Boobies in central roost, Green Island, Kure Atoll, 1 July 1968. South beach, Green Island, Kure Atoll, 1 July 1968. Edge of runway, Green Island, Kure Atoll, 1 July 1968. Looking towards north point from north tower, Green Island, Kure Atoll, 1 July 1968. Sooty Tern colony in south antenna field, Green Island, Kure Atoll, 1 July 1968. Generalized annual population cycles of breeding species at Kure Atoll. Generalized annual population cycles of non-breeding Species at Kure Atoll. Generalized breeding cycles of avian species at Kure Atoll. Location of islands or atolls in central Pacific where Kure-banded birds have been recaptured or vice-versa. Page 2 16 aN 18 19 20 20 21 a 22 22 eB) eB) 69 70 (fal 83 WIS-1. Cseir Run le. RnB er. 1SUEUE=3} BFB-1. BFB-2. BHB Sl BFB-4. Locations of birds banded at Kure Atoll and recovered at sea or outside the central Pacific. Breeding cycle of Black-footed Albatross on Green Island, Kure Atoll, 1964-65. Distribution and abundance of breeding Black-footed Albatross on Green Island, Kure Atoll, May 1967. Breeding cycle of Laysan Albatross on Green Island, Kure Atoll, 1964-65. Distribution of breeding Laysan Albatross on Green Island, Kure Atoll, May 1967. Number of Bonin Petrels captured each night in two study areas, 11 February - 14 April 1969, on Green Island, Kure Atoll. Counts of Wedge-tailed Shearwaters in the central plain on Green Island, Kure Atoll, 1967-68. Distribution of breeding Christmas Shearwaters and Sooty Storm Petrels on Green Island, Kure Atoll. Annual population cycle of Red-tailed Tropicbirds flying over Green Island, Kure Atoll, 1964-69. Breeding cycles of Red-tailed Tropicbirds on Green Island, Kure Atoll, 1964-66. Number of Red-tailed Tropicbird nests in which first egg was laid each semi-monthly period in study area at west end of runway on Green Island, Kure Atoll, 1964-67. Breeding cycles of Blue-faced Boobies on Green Island, Kure Atoll, 1964-69. Number of Blue-faced Booby nests in which first egg was laid eacn semi-monthly period in north antenna field, Green Island, Kure Atoll, 1964-69. Number of Blue-faced Booby nests in which first eggs were laid each semi-monthly period on Green Island, Kure Atoll, 1964-69. Number of Blue-faced Booby nests in which first eggs hatched each semi-monthly period on Green Island, Kure Atoll, 1964-69. ‘abal Page 84 93 i 105 LO7 116 12h 131 138 139 WD, 156 157 158 158) Viral aeal, BFB-5. BB-1. BB-2 ° BB-3. BB-4. RFB-3. Gla: GF-2. AGP=ir RE=1. BICG=1.. WTa-1l. Sand-lL. ro LEA e Distribution of breeding Blue-faced and Brown Boobies on, Green’ listland,) Kure “Atoll. Breeding cycles of Brown Boobies on Green Island, Kure Atoll, 1964-69. Number of Brown Booby nests in which first egg was laid each semi-monthly period in north antenna field, Green Island, Kure Atoll, 1964-68. Number of Brown Booby nests in which first eggs were laid each semi-monthly period on Green Island, Kure Atoll, 1964-68. Number of Brown Booby nests in which first eggs hatched each semi-monthly period on Green Island, Kure Atoll, 1964-68. Breeding cycles of Red-footed Boobies on Green Island, Kure Atoll, 1965-69. Number of Red-footed Booby nests in which first egg was laid each semi-monthly period on Green Island, Kure ATole MOGs). Distribution of Red-footed Booby breeding and roosting areas on Green Island, Kure Atoll, 1967-68. Breeding cycles of Great Frigatebirds on Green Island, Kure Atoll, 1965, 1968, 1969. Distribution of Great Frigatebird breeding areas on Green Island, Kure Atoll, 1967-68. Annual cycle of American Golden Plovers on Green Island, Kure Atoll. Annual cycle of Ruddy Turnstones on Green Island, Kure Atoll. Annual cycle of Bristle-thighed Curlews on Green Island, Kure Atoll. Annual cycle of Wandering Tattlers on Green Island, Kure Atoll. Annual cycle of Sanderlings on Green Island, Kure Atoll. Breeding cycles of Sooty Terns on Green Isiland, Kure Atoll, 1965-69. Page 164 MS) 174 176 177 195 196 199 2le 213 226 230 239) 238 243 253 SHES} ST-)., (Grek, GBI-e. (CASHES) 6 Tela BN-2. Moa. Distribution of Sooty Tern breeding areas on Green Island, Kure Atoll, 1964-65. Distribution of Sooty Tern breeding areas on Green Island, Kure Atoll, 1966-67. Distribution of Sooty Tern breeding areas on Green Island, Kure Atoll, 1968-69. Number of Gray-backed Tern eggs laid each semi-monthly period on Green Island, Kure Atoll, 1967-69. Breeding cycles of Gray-backed Terns on Green Island, Kure Atoll, 1965-69. Distribution of breeding Gray-backed Terns on Green Island, Kure Atoll, 1964-69. Breeding cycles of Brown Noddies on Green Island, Kure Atoll, 1965-69. Number of Brown Noddy eggs laid each week at north point study area on Green Island, Kure Atoll, 1967-68. Distribution of breeding Brown Noddies on Green Island, Kure Atoll, 1967. Main Black Noddy roosting area and White Tern breeding sites on Green Island, Kure Atoll. Number of White Tern eggs laid each semi-monthly period on Green Island, Kure Atoll, 1967-68. Population size of Polynesian rats in a 6.94 acre study area on Green Island, Kure Atoll, March 196)- May 1965. Number of Hawaiian Monk Seal pups born each semi-- monthly period at Kure Atoll, 1964-65. ix Page 259 256 257 264 265 268 274 275 e77 285 289 301 310 C=3). c-4., POP= acres invextent. it is oriented along a north to south axis. Surrounding this plain of low herbaceous plants is tall, dense Scaevola. A series of paths runs through this vegetation. Figures Dl-]1 show island scenes and should convey to the reader a better impression of Kure than could ever be expressed in words. To the west of Green Island are three sandbars known eolllectively as Sand island. “These islets are variable in size and shape throughout the year. Generally they follow the pattern of the north and west points of Green Island. The largest one may be as long as 400 meters and as wide as 20 meters. Birds breed only on this largest islet. During the winter these islets may wash away. VEGETATION Christophersen and Caum (1931) reported 13 species of vascular plants collected at Kure Atoll by the Tanager Expedition. In 1959 Clay (1961) found an additional 6 species and Lamoureux (1961) reported another 23 in 1961. During POBSP studies four previously unreported species were found; thus 46 species of vascular plants have been recorded at Kure Atoll (Table P-1). The majority of these species was intentionally or accidentally introduced during the construction of the LORAN station. 16 ‘TTOV emmy “pueTsT user) FO uotTzeqeseA “*T-A eanNSTW DUIONSHD @ wopoud> Yj, wWNUDIOS BENING] ‘DIADYIA0g ‘SIISOIDDIA VW), i DUISAGIAA »-) Be ent EE D[OASDIG wits Li TeTOT SIO ‘yderso0qj0ud paren) 98809 °*S°n "O96T Trady 9 ‘ugzou SuTYOoT ‘*TTOV eany “pueTsI user) FO MeaTA TeTtey I a OMT AT 18 "996T Atenazqeq 12 ‘ydersoyzoud AAeN °S°N TetoT sro ‘ygnos SuTYOoT ‘TToVy emmy “pueTs—T user) JO meta Tel ry 3 qd smst aT 19 qT 96T 2840900 €T 4 Toy emmy 6 "ydersoqoyd AAeN *S°N Tetot sso pueTSsT user) uo xeTdmoo SutpTiInq JO MeTA TeTtey “fel 20 Figure D-4. Figure D-5. North antenna field from north tower, Green Island, Kure Atoll, 1 July 1968. POBSP photograph. Blue-faced Boobies in north antenna field, Green Island, Kure Atoll, 1 July 1968. POBSP photograph. lL Figure D-6. Looking towards west point from radar reflector, Green Island, Kure Atoll, 1 July 1968. POBSP Photograph. Figure D-7. Breeding Red-footed Boobies in central roost, Green Island, Kure Atoll, 1 July 1968. POBSP photograph. Figure D-8. Figure D-9. South beach, Green Island, Kure Atoll, 1 July 1968. POBSP photograph. Edge of runway, Green Island, Kure Atoll, 1 July 1968. POBSP photograph. — Figure D-10. iBalexbage) 1D ALIL 5) Looking towards north point from north tower, Green Island, 1 July 1968. POBSP photograph. Sooty Tern colony in south antenna field, Green Island, Kure Atoll, 1 July 1968, POBSP photograph. % a4 Christophersen and Caum (1931) described Green Island in 1923 as covered with a dense, almost impenetrable growth of Scaevola frutescens..., which averages 5 to 6 feet in height, except in small areas, generally on the tops of the sand hills, where it is only about waist high and fairly open. In these openings and along the outer rim of the thicket are a few other plants, principally the tall bunch-grass, Eragrostis variabilis, and the creeping Boerhavia diffusa. Toward the east central part of the islet is a large, open plain, probably 20 to 25 acres in extent, entirely surrounded by the tall Scaevola frutescens....With the exception of a few "islands" of Scaevola frutescens, scarcely any vegetation in this central plain was over 2 feet high, and most of it was considerably under that height. Achyranthes splendens, Tribulus cistoides, and Lipochaeta intergrifolia were found only on the plain, while Eragrostis variabilis, Boerhavia diffusa, Lepidium owaihiense and Solanum nelsoni grew on the central plain and in open areas among the Scaevola. Two clumps of Cenchrus agrimonioides were found, one among Scaevola and one on the edge of the central plain. Other plants were found as follows: Eragrostis whitneyi grew sparingly on the outer edges of the dunes at the northwest corner; Lepturus repens at one spot among Scaeyola; Ipomoea indica on Scaevola; and Sicyos hispidus on Scaevola ielanae in the central plain and on the inner edge of the scrub. In October 1959 Clay reported a vegetative condition similar to that recorded in 1923. However, he found Cynodon dactylon, Casuarina equisetifolia, Pluchea odorata, and Verbesina encolioides growing near the radar reflector and surmised that seeds of these plants were brought on equipment from Midway Atoll in 1955 when the reflector was built. Two other new species were reported: a young Tournefortia argentea growing on the windward shore and Solanum nigrum on the central plain. By the time Lamoureux visited Kure in September 1961 the island had been greatly altered by the construction of the LORAN station. As a result, there were more open areas: along the runway, around the building complex and roads and through the Scaevola on the south side of the runway (the remains of the albatross paths built in 1959). He : norted (1961) the follow- ing changes in these areas: oy) a. Margins of roads and runway. Boerhavia diffusa has covered most of the margin of the runway, with Tribulus cistoides and some weedy grasses occurring as scattered indi- viduals. Along the roadsides Boerhavia is somewhat less abundant with a larger pro- portion of weeds. b. Albatross runways. Covered with Boerhavia. ce. Clearings around living quarters. Gnaphalium sandwicensium is extremely abundant in areas where Cynodon dactylon has not been planted. Euphorbia glomerifera, Emilia javanica, Conyza bonariensis, and Eragrostis amabilis were found only here’. d. Clearing around LORAN Tower. A series of cleared strips a few meters wide radiate out from the base of the tower to the guy-wire anchors. These strips cut through most of the eastern part of the central plain. Boerhavia diffusa, Lepidium owaihiense, Lipochaeta inte- grifolia and Solanum nigrum are moving into these areas. Twenty-three new species were found on this trip. Only one, Phyllostegia variabilis, was native and had probably been overlooked by earlier observers. Cocos nucifera, Codiaeum sp., Hibiscus sp., Thespesia populnea, Terminalia catappa, Nerium oleander, and Helianthus annuw were cultivated around the buildings while the other new species were found in disturbed areas and were undoubtedly accidentally introduced to the island during construction work. Scaevola was still the most abundant plant and the central plain was still present. During POBSP studies the vegetation on Green Island re- mained similar to that found by Lamoureux, although much of it had grown taller. The following is an account of the vegetation on Green Island in 1968 and applies to most years POBSP personnel worked on the island. Seaevola taccada covers most of the island (Fig. V-1) except for the original central plain and man-made open areas. The extent of these areas varies. For example, after the station was built, most of the area under the LORAN tower and its sup- porting guywires was devoid of Scaevola. By 1963, when POBSP Studies began, most of this area, except for that portion of the central plain where Scaevola has never grown, was. again covered with this species. Given enough time, the majority of these man- made areas would again by covered with Scaevola. 26 There are three types of open areas--the central plain, the area around the buildings, and the other man-made areas. On the central plain, the dominant species are Eragrostis variabilis, Boerhavia diffusa, Lepidium owaihiense, Tribulus cistoides, Solanum nelsoni, Sicyos hispidus, and Verbesina encelioides. Cenchrus echinatus, Achyranthes splendens, Ipomoea indica, Solanum nigrum and Lipochaeta integrifolia also grow there. Around the buildings Cynodon dactylon was planted as a lawn grass and is now the dominant ground cover; most native species are rare or lacking. Cocos nucifera, Casuarina equisetifolia, Hibiscus sp., Thespesia populnea, Terminalia catappa, Nerium oleander, and Pluchea odorata were planted as ornamentals. In the other open areas the commonest species are Eragrostis variabilis, Boerhavia diffusa and Gnaphalium sandwicheum. Elusine indica, Tribulus cistoides, Solanum nelsoni, Solanum nigrum, Sonchus cleraceus, and Verbesina encelioides are also common. Besides Scaevola, Casuarina equisetifolia and Tournefortia argentea are the major shrub or tree species on the island. The former species is present in three groups around the barracks and in one group along the southwest beach near the radar re- flector. These trees are about 20 feet tall. Great Frigatebirds, Red-footed Boobies, and Black Noddies commonly roost in them but no birds nested in them during POBSP studies. Tournefortia is found at the north end of the island (4 trees about 10 feet high), northwest of the central plain (1 tree about 10 feet tall), south of the runway (1 tree about 10 feet tall), and along the southwest end of the runway (3 trees 3 to 4 feet tall) (see Fig. V-1). Red-footed Boobies and Great Frigatebirds breed in the northern- most tree along the northeast beach, while the others are utilized only for roosting. Scaevola provides roosting areas, breeding areas, and/or shelter for most avian species. Verbesina encelioides is the most important of the introduced Species to the island's avifauna. First found in October 1959 growing near the radar reflector, it is now widespread and grow- ing in most of the open areas, mainly the central plain and near the radar reflector. Growing in dense stands and up to 4 feet tall, it is a potential danger to the island ecosystem, especially its ground nesting birds and native vegetation. Birds have dif- ficulty moving through Verbesina stands and when the plant becomes dense enough are no longer able to breed where it grows. Presently the breeding habitat of the Blue-faced Boobies is threatened in such a manner. Other plants cannot grow under it, so native plant species in the central plain are also threatened. The range of this species continues to expand rapidly on the island. ai Other species of importance to the birds include Boerhavia diffusa, Tribulus cistoides, and Solanum nelsoni. These plants are used as nesting materials by Red-footed Boobies and Great Frigatebirds. There is a definite seasonal growth and reproductive period for plants, mainly from June through October. In summer, for example, Tribulus and Boerhavia cover the cleared breeding sites of the Blue-faced Booby and probably prevent the building of nests at this time. There is little growth from November through March. During this period wind and Sand may scour and defoliate Scaevola, primarily on the wind- ward side and to a lesser extent elsewhere on the island. Table P-l. Vascular plants recorded at Kure Atoll by various people. Christophersen and Caum Clay Lamoureux POBSP Pandanaceae Pandanus sp. x Gramineae Brachymenium sp. Cenchrus agrimonioides Macha viele laysanensis x Cenchrus echinatus L. Choloris inflata Link Choloris virgata Swartz Cynodon dactylon (L.) Pers. x Digitaria sanguinalis (L.) Scop. x Digitaria henryi Rendle xs Eleusine indica (L.) Gaertn. : x x Eragrostis amabilis (L.) W&A. x Eragrostis variabilis (Gaud.) Steud. x x x x Eragrostis whitneyi var. * a a mM PM OM Caumii Fosb. x x m x Lepturus repens (Forst. ) Ig 1Bne x x x xX Setaria verticellata (L.) Beauv. X x Cyperaceae Cyperus rotundus L. x x 28 Table P-l. (continued) Christophersen and Caum Clay Lamoureux POBSP Palmae Cocos nucifera L. x x Casuarinaceae Casuarina equisetifolia x x x Amaranthaceae Achryanthes splendens var. reflexa Hbd. x x x Amaranthus spinosus L. x Nyctaginaceae Boerhavia diffusa L. x x Xx XK Caryophyllaceae Spergularia marina (L.) Griseb. x x Cruciferae Lepidium owaihiense C. and §S. 5 x Ke x Lepidium virginicum L. x Zygophyllaceae Tribulus cistoides L. x x xX x Euphorbiaceae Codiaeum sp. x Euphorbia glomerifera eaRS, ; Wheeler x x E E __horbia hirta Gre x Malvaceae Hibiscus sp. x Thespesia populnea (L.) Sol. x Combretaceae Terminalia catappa L. x Apocynaceae Nerium oleander L. x x Convolvulaceae Ipomoea indica (Burm. ) Merr. x x x xX 2S) Table P-l1. (continued) Christophersen and Caum Clay Lamoureux POBSP Boraginaceae Tournefortia argentea bg 6 3 xX x x Labiatae Phyllostegia variabilis Bitter X x Solanaceae Solanum nelsoni Dunal x X xX x Solanum nigrum L. x X x Cucurbitaceae Sicyos hispidus Hbd. x xX x x Goodeniaceae Secaevola taccada x x x x Compositae Conyza bonariensis (L.) Cronq. xX x Emilia javanica (Burm. ) Rab. xX x Gnaphalium sandwicheum Gaud. x x Helianthus annuw L. x Lipochaeta integrifolia Gray X X x x Pluchea odorata (L.) Cass x x Sonchus cleraceus L. x 2 Verbesina encelioides (Cav.) B. and H. x x x CLIMATE The climate at Kure is subtropical marine and is influenced by two main air masses and by the surrounding ocean where surface variables are relatively small. For example, sea surface tempera- tures vary from a low of 69°F. in February to a high of 80°F. in August and September, and salinities from 35.2%. in April to August and 35.0%0in November to February (Seckel, 1962). Most of the year Kure is in the path of the northeast trade winds from the Pacific subtropical anticyclone, but in the winter 30 it is on the southern edge of the Aleutian low, with resultant westerly winds. Table C-1* lists the wind direction at Kure for the years 1961 to 1968. From November to February winds were variable, with a strong westerly component which in some years was not well-developed. Winds were again variable from March to June and September to October, but with a major easterly component. The northeast trade winds were best developed in July and August when almost all winds came from the east. Windspeeds ranged from O to 8 on the Beaufort Scale (0 to 46 knots), with a yearly average of about 2 ( 4 to 7 knots). From 1961 to 1968 average semi-monthly windspeeds (Table C-2) varied from 1.5 tc 4.0 (2 to 18 knots). Although there was no consistent yearly pattern, there was a tendency for the strongest winds to occur in winter and early spring, and the weakest winds in summer. In 1967 windspeed varied little throughout the year. Although there is yearly variation in the average semi- monthly temperatures (Tables C-3 and 4), the pattern of tempera- ture change throughout the year is the same. The average maximum varies about 20°F., from 65°F. to 85°F., and the average minimum varies about 17°F., from 58°F. to 75°F. The absolute variation was SO7E. to. 99mm. From December through April the average semi-monthly maximum is about 70°F., in May, June, and November between 70°F. and 80°F., and from June through October 80°F. to 90°F. The average semi-monthly minimum is between 60°F. to 65°F. from December through May, between 70°F. and 75°F. in November. January and February are usually the coldest months and July, August, and September the warmest ones. In 5 of the 7 years of POBSP study the average temperature began to increase in May, in one year in Jume, and the other in April. It began to decrease in September in # years and in October in 3 years. Since rainfall data were not collected at Kure, the Midway Atoll data were used (Table C-5). As with other climatic para- meters, rainfall was variable from year to year. Some years the wettest months were in the summer, in other years in winter. At Midway rainfall averages about 40 inches annually. On the average it rains 12 days each month. March is the driest month and December, January, and February are the wettest months. Heavy rainfall is most common; light rain and drizzles are almost unknown. Rain occurs on only 6% of all hourly observations. Days * Weather data were extracted from the station logbooks. Obser- vations were made at four-hour intervals. Sil Thunderstorms are relatively uncommon, averaging 0.06 to 0.10 are next commonest, followed by days with 0.11 to with 0.02 to 0.05 inches of rain are most common, days with 0.20 inches. six a year (Standen, ms.).- Wind direction at Kure Atoll by semi-monthly Table C-l. periods, 1961-68 (expressed as percentages). 1-15 Januar 1961 1962 1963 1964 1965 1966 1967 1968 Direction OOO+F+0MNNAN OCHA GG Qae Wn oa Meo Ste} Qe \O 1 MNO OV COM MO O + WN GG) 8) EN CO Sp i= 6A) GAO) CSP i fal Gal ol DAINAMMA INO FANN FO eh eh de A leh pie ie es sel Une) ee rem. se. WOMNANNWO AMO AAO sz An il tI WAMNDWOMO AM+OO =O! @ NE HONGO) NOES UG = ad vA HA\OO NO OFT +t N OT + ~eO ADM AN ANAWMA AV ode ode OOOO AtHTeENMONO OVC VOAMNMNO AD AN O i GY GA) cl AAOrMY+A YOO AMOAO CF 20% er ier e! Pep Ser eer ery sel oe 16-31 Zi ©) ©) CS) NQY Iginy Oya et Cer em ee) ee ers fe, aleink en Fein yerce tere. 8 NOAAMMNMNAOAN rl r= GQ tah oe WACO AWN MYM INNA A Qiica @ Se anim ON es Se r tl AQ) ml NOE TE COs NIE CON) ON GO 1 ON A+O MIND + OF os qo qo IQ eeSP (SS SQN) a2 No) [ @a)esr (ovine oie 205.) (el ert ney, sersee te nen at ler, fe m+ SCO af Sp ar CO) ta} GANS CO) AOWONVNOWMNErHADODOHO (o-b ei eee en 8 ade Oe 10) OL Oh et te. — Kerr, <0, MN Qeer BY WY 69) si Uys GS oN ed OOWVOONM+OrOO 4H CVOOOMNMNAN AMO 4 oo ee Wan Ins MDNtONDAOMEN On rele (os ac8)” CHa, (es ae ele [er 8) Ue OM+F MME wnt O +t AM odoin (Tw StF Were (lineal Pees reel Hvar | er Jom Heuer Pome | OOOCOCCOON OWMmodonstroOomMmu OONAHAUNNMNMM™M Ow one 0 8 0 E & Db 0-0 mM tt tAdAdb dtd Add A Ae al Ga Ie UNS ea A OMO NDA AAAYA mM 1-15 Februa Ops OES Orla S NO Se ONO) tO -0\0 O So + 1c NAN OvO iN th ONXO + ra —t © Why ONO SN oo eo Qo OC © ON\O ON = \O = \O a) 31-60 61-90 91-120 0-30 32 (continued ) Table C-l. 1-15 Februa 1961 1962 1963 1964 1965 1966 1967 1968 Direction Ot M+ AON O O+ wo dONO vA a Ot OD AHON Ot SS Orrin N ode PIO ON tre CANO WIWONMAN+ MY ONOWNOAANEN NORE rt St UNO 1 oe oat Ee YQUTON CG) Rp ke) ee eh Kel ale, On) ek. fem, Je AOMAr OMS Cl al Ge rl tl OM =n © in OC © oNNoOANCO Qt a OmMmM+t + CO OC OO ee er ee Oar ) ia a LA OA Teh 121-150 151-180 181-210 211-240 241-270 271-300 301-330 5315359 a 16- Oooo oN Fst AO OAO SCDSOCOOHK A+ HAWNO HoNMe4 doeVUNOO LANDA O MOD OO JO OR OSORIO IO a CH er) AON NN OW + N 11ND \O it fal aN LON S OEOTO O25) CAIONNO © HW A++tOOCAHONO moO 4 WAMNMEA AHO WOM NOOrYANOWMNWOMAO aa re dN AOOCDMDHAWARWAA St Om: “OS FOeR0) BO COn OF Sa oie tee b= GQNO =f on) NS SS) nr~rn NO OMADLEWMWANEOO sa SST Cox) NST Try NG CaS) done UVN MO NNO GD NON O ele ietcicrey | tem ten* rer’ mer fers Lier jek Jem qo N qo Ore OC! OO" S' @' On ey MS) lear hp SO OONAANA mm OQNOEON REIS tet (iq) US 0 Ue teat) tal teal Gel are aS NCO ESAT oe OND ADAAAANN mms Mareh 1-15 OMOLMAN+OON 4 bh 4 WO SQ) ett ea ~eeN 4 AN OVO MH\O\O + +t CO NW N COLOR rl ANN et O'CON A oma qo SONMNANAWO ADA NAM Ol ae Ol ORO), Ol Ten, <0, (210) 5:0! “ie se) © 6 oa ies Noh(ee) Ent ae oa CONES 2a NO PN ON ©) ESN OO ROL NO Olas 6) FO; | .08> 20;( 0) 0; 0 OO NM M+ NAO WO CO iN HAN fFS-OHOON rE MOANA ODF Lele ane), C82 “ese cree (e750) > e INDO MM AWN + NON i tah tl COP CO1CO salt iestO rte) OHe ole Sees et) Ke ey ee) —-.0%e- ley = en ee.8 Say Oo Bed NO) teal AO os 19) NN OO ANON N AN AO®d ORR ets re). .eh- lel seae, oF) cet aye ey “ce te BSNS) CONG) IS SS a eas os qo Hon S2O7 OO. OO OFON DAV NCO ate [SS on) oN COON AANAUANMMNM @Q@XoO Ov | Ut FoR PF tb 4 mM bt badd Ata eee LS Ges) eal fees ISS ENS) ea Ny a WOMO NA TAANN mY 16-31 a 0 4 AMON \O Nin —t + \O ta ~ + NO Owe CU COTA 9.0 a ral AMESE \O & ON Lea) {09 i) GA May b= \O OM re 1) UN XO te OV == \O NCO KN M4 0-30 31-60 61-90 (continued ) Table C-l1. March 16-31 OCIA EOG 2 NaIGoSMEIOG hs LOO) a lOGG) OG aloe Direction QROTOJOS OOOO Che te wt tak QS) NAOn A Ast WNM +MOMNMNMNO AO il tl OOIMAO MINT M Os Oy tC Or ae. Pee 10: ey 9-8. mt tJ OC eas NE} ty ONES odode MmWnNAANAT MON 4 re fa MIANNA TAHOMA. OO NNHAHHAOW WM +tMnNnMNAMHA AN OO Ou, 8 aOR OL mS) SO. Saw MUMOINA MAO FH hes N 4 OOOArRNVNAAN Ora cer Kee Oey 107% Tey es ie: Fa AS areal OM rat ret) MO WN OC INO CO N \O HA~rANN Ee MN eb Nd qo April 1-15 AANOMAMrHAHHAOCONMNM HAHAANEMOHHAHOONM®M =P Ga) ta) tf] AMMMOOHODOON SO AMMMMNOCHODOOANO UNNN ~eMMat Or ATtOHTLEN O MMs AOO+tArTON - re oe OWVNArRRKHAHAWMNOCOO M+ SHOW Hla © OM & MAAN WAMMNHADDODOOVOAN KaMAooDOOOONN —t —+t ONt tO AWMWMA OL HPSS) Wa) SANOstal tae eS UN Nd ~KrryeerooNWwonoo Mo Cy ie ess tO FUOs SOO Se Ome emis a men) WOn00rFONOHOOMWN HooHNH 4 NWOMr~VUst HANAN O OO oaepaetHVNse be MAONN GA) i ode OOo © © © © OW Sy ENED ag yp SSS OONHnAHAYA mM) OW) ONS] 0 ob UU 0 0 BOO my! bt tAtdhAd Aad A AA Ae tH ANNO nASsT Ee OM OMO NAA AANUNANNMM®M 16-30 OFrOMNAUNNONNOOO WAUWVIANANSANDOSO N= Mt OODAMEAAHA GRY MNES) I GA) NO) ty} GUI mde COMrONnNMNHAO HH OV+t ©. sey je} tera re. Pie: 1 ere 6) =e Se se: ADOMANNAOHAHLKEM® fa om 4 Co) [SSCo) Sp (©) C0) CO) i) © [See Sr O N= ACO LO O DWINM YM imal i N NMDMNDODOODDVOCOVOO Col Kono Ko Ro Ronononone) a+ MOH Co)! OCo) Tal=p © Gay lt> var © SH OMS © AN a= SO mans) IVMA\O O'\O ANOCODOCO Oi" ez in Xe: eye werecer 1:0" Se stge, er ice> ike OMNHAN+\OFOCOOO +t AN xc trnmMmrnNtOmHA Ono eo) Oe Tiel. fe ape Ker. 0 res) 10) Ne Gwie ste UNED [Re lo 9) SUC) G9 aI CO) ode ra GLOVOVTOLOFOEO VON OWMOoOndTErO MW OONAANAUANMNMM™M @NOtONet ah ote Sa ee Nl’ tata AAA ed PHA ANNO AsO OM OMA NDAAAHANAMM 34 (continued ) Tabplete=15. 1-15 Ma 1OGT BAlG62 AAO6s) MplOO O65 gil O66 a LOG amllOG. Direction NNAMOANANANA AH ANANODNANHHSA rt NOE@ ECON = ©, Or) O-@rr- CY Sy Ng NO al SY es eee GAY) GASP =O) tal CaGayP Cy-= pt CAIN Gajsr Gaia) Garoa) ors r + M4 ONHHODDOOHOLK DA +tOOHODOOOHOUON MM 4 +tNMNNNNONHDAO HO ANMAOND Aint oe ap GUNS ON) aq) le) Qu Xe) =f Qii ess OPQ gal QO NS ANN Nd imal PORES? QS) QUOC) Z=7 NO)-| reo) Er IN SS) Wie) urea; sap (NET) WIVWUNDOOOON+TONO MYM Os iat ate Sel SE Or tO Eta QEO@ZOZOLO.OLOPON (Sy WeCO) pa) =r SSS) na yn DONA ANNNMMM NG) OU Ue 0 eo Gay UU 0 eb cal ca) Galt eh cal fAoagmAN NO ASF Om @ © ONS StS NN wai (OQ) WS pei VAN AWS) [S(O (SS) OP ex er ce) ee) Cuvee, Vel @ ~e-" e@ CS) SY ENCOUN GO) IIS NGO alia AN 4 QUA Seah) Lal QO) oO (Oo) ALCODOANs HACCO Vad A Wey ON (a) tal Gal tal ear teal Se Ga) i) O) a OmN SS eae S| CO) ISS) tas ral ON Alte! tol talteY Oy) Tate) NOPON EO) ' QUEQEOP Srlaca Ol rt mo 4 (coyesp CO) ©) ta) sr ESO) QI eSp I te) AMP-NOVUMNOHANHO dann QCo) 1S) (YOO) Q1Oy@) (2) (OS) NA+ AOODODCCO ‘Oo NA OS Ss lS Mint ini Nie ©;©@ TAA SS Sit “Oro. NOES NOM QI@ Srarerd red Oy oe eee) a ey 65) se) Cre be Je) <6) te COO) SO CIO) Se) ON (SO) UVC On Ear [SS18) Caja OONHHAUANNMMN™M OROTO) Has le ET ae it Coa ee i a OT DT OT ee OO ee | eter QUIN COURSES Og OMIA NDAHAHAANAANMNM June 1-15 OV ON ON ONO \1O F-1INTH O O'™O Che 0) SO Sie Oe Cee rt) SOS Oe ONE Otel OO StS Si ONDOKRNDAATHTAM CNDOONEAHOtUM Qt lit WN Dal iss Gri) Wey onwWoyeap (QN fear OO) Orr Ors Oe oe) are) tery) Ke), Srein xe CONOR ee ae CORN NEE ONQIRONON ISH ONIN Ol QO on Of Ve a 6-6 8) Hie. el aie) ce- ep ie: ON eo eee eh CON C0) Stabe) NAN A (09) PSN a7 [SSCo) Gals est ONE yor eo se; Ne, 2.0) ne. se ie. eles ey Tee” eo). 7:6: ~KEMANMANM+AANA (Utah its! tl Qt Wh CO) NOS gt OO COPS UN One Sb NOR MNT Ora: rr ood qo CO) Sony) tI Wot te) tS tea) ta Mit Ares ets Ona Qi ot ODNDMMNNMNANMNAKKA A OtMNANMNNANMNMHIH MAN dH @QIOA@FO@VO. Orv Oron OAINCOsrt =F SO Fanta OON A ANNAMNNMNM™M CNS) (ON Hee fe Ue Te te 0 mM! § tad AAR Te AAA eH tae At ANNO Ase Le OM OMA NDAAAAHANNMNM SD) (continued) Table C-l1. June 16-30 1961 1962 1963 1964 1965 1966 1967 1968 Direction OO rON + NO ATOM eo} repel = Le Sle) Ne 0) =e Nr we eh | Hel ce G7 MANS) SU ar CO) otal Sr INO) at) el cl ©) OO) GH! © GANO! © cI} NO) QO ON) ES GONE SF tI NOES aN OO eels HNO War oOo QM © OL sie) Seg lel= fe ser Ser ie: ier ee eh Se OODMDAKANMHANOHO HaAmM4 WO OIN ON + FCO MOCO CY \O AO ND OwWUiwW0 oO NENMOMWOAACOO NINA NOMMNHAAOOO dAaMNN fo UN INO) Wy NUD) CE CC) OOO sr © ro) AIS 6A NO OOO © dae Coa OO MOO FAtrOMMAM Oh OM Sag Os yO) eu Ne) 0 ey lev “ens en ie. OMFtOOhAWMNSH HN oa) ta ml G9) Ase Sse OnrlOOoO One Haman+t+tOAHOOCOFH ot 4 OOO OO © © © Oimo Att oOmin OONAANUNNAMNM™®M OW) OY! 0 0 UU U0 0 0 0D @o) U 0 0 cebrelce) cel tal eel cal (HAAN NOH OOM OMA AAAANANMM® July 1-15 GA Qiao ro OOO Oo e@)- ies § (Orkee es | 10) fe: = fe) = Jen" Key ce. fer e GAG Wis} Gri ao oo @Ee —t in RAT ANE AA HOH M IGN ON EINOES IIo nic aa a) Qe ~rMOrF-MOWO HADVOONA OcR Choe ECs Sa hore Soe ait Sm) aet amie Sore Laue av WO) SoU OC Ory O CO © tele) HANAN 4 WIMWOMrNMDMNMNMNOODODOOO UN+nt+tadndoococodo Ua XO GHC) 1 OOOO © © © HHNDOHOSOSOOOO WN \O 0) Sp CO) tr] QW) © [= W) Sp Sp ra) © Wt PAVOON++ 40 doc ANNERFOYWONAHOO emia) ele vet se le: le; 0") Yew ie. ele ce OD OW COMNAHAHOO aN Mm OU MINDA NAM INO AHO HN OD) Fey ears I MGUY EO) fe) al ©) NN Oooo ©o © © O CO) Wey CO) ra) SP [SO GO) UD OONAANANMM™ QROLON Seti ne ett 0 ee Pe OT ae Oe Oe TF oD | ta) tt cal QU) Weyeo) Ge) Sr [SS © EA) OMO NAA AAHAUANNMM®™ 16-31 00) O-Fl H OC OO OOo © © OADNCCOOCCCSO A+ UNE ONS tn)! CO, OO OF O@ r-| MNOHAOMOAHHAOCOM ANN 4A ONO ONAHAHAUNODOAODWM Oe Wiles ce NOP see Fe 580 Aer ale) ierane, DINODHANNONMNOMYM Qt mi CHL=NC) CO OOo OO OO a Oh SO Ore Oe ae Ce Cue yy pen et) \O WN OMMmMmANOCODCOOO HOMNN+OOCOO0OO UG) NEINIMNAODOOOO0O MmWUNEEMUNODODOOCO oe) NaeNUMNN AOOOHO SIDI EG NO CV OOS ©) oe) N QO ORO OEOLOTOR@ INAMWMNMMNOWOODOOCO Qa A ODOOCOO ON Sy eS al Ay Ao OONHANYA mas OW) Oe 0 0 fo 0 0 0 0D oy) 0 0 0 el ele A a Se ce ProA HANNO HASTE OM ONO DAAHUNNMM® 36 (continued ) TableniC=ir August 1-15 1961 1962 1963 1964 1965 1966 1967 1968 Direction ODODMDONDHADHOOO ONE MN Mh ArAOOO No9 Tat Cy QW Sp OOS? mle Wor) © COIN OlO =a Or sO CA):tent fl GU oa (alee [SS pC) Tae) (SO) OOS) HI AOO MOOS SO OS GUENe) Fl tal tl OTD O OOOO ma) © rl tal tal C0) Cr ©) GOH (OC) E) FAROO IGA retire NOES NOLO) CORT OOO, OS) @ Oe Oi SOF SO OO SOF a See OO QD NOICOE at NEO OE OiOr©@ =o (QU i! ~-NOAWMWODOODOOOOLH ANNNDBDOOCOCOMme]A praia rN ici Ore) © ©) OO) © Handdddddddc QUES OnyOSY ONi=) hq) il Ore; Cre) © Ql Weyico) Cn tal (SS) LON OVS) (@) ©) Nat N SOLO ZOFOLOTO ON ONNO St © MIA OONAANANNMMM®M @NO ONS ie te ei (op ee Bn cs BT Ts Oe Oo oe | Oil Gal ted (No) a ear i (C) Ga) © MAO NOAA ANA AMM 16-31 NFAWMNE MANOA MAO Cieeee 20a 0 ue) Wee 1e)) Keys 5 6) Ne” le ar (Sey Gove Wain Syne) fot (NY ph cl Tt NOl ERO TA OzOl@r@(Or@ ei,- 10) em, 0) ie, es ley . @ se” fe. Je se fol teal (NES EO} TO) TO) ONO) (SO) MNO mal Wey ONS) Way Cee) OS) (SS) eh Ster Rael vier. oe Ve sie; tien 20" Vel ce Je (QS PC): [> [SQ (S) (fo ON(S) {S) QS it +StUOANAANMIMNIMN OO ey Pie ee. hm 10/1". 0% Ons 0!) (0) es 18° ‘76: MM MN MO+bhRHHAOO QU satel Gad VQNestGg) t= ONO @! aiggianict rt tdtVACOHNANAA Pleep 16] CPLOMION=e CN. NEO OrN! @ uN CORMNORINAH IQUI@* OQ Ort ANN =e Ql Gajes) Coon =-a7 Wye) (©) Or a Ore OP OY.) FOr Or, 0 oO ee 10) + Siete 18) Xs ONO UN CONS UN AU gO OO NAA NO) tral No) (CG) Co)Esr =p Ge ©) OS) GALS) Co) QU] =piCoN ine) SO) () S) mm 4 COLO TOLO}).@EO7ON (SY W\CO) ta) Sp Ie CO) ayn OONHAANNANMM™Y @QNOFON Se ie ate em). Do Lb Teel teal al fs toy el foe te US fell tren! Ta) 148] Wasco) ae (SS) Ga) Sigal ONT TSE Geanian September 1-15 WAANANDOWVOMLE «A Sari 200" @ @FO1O*WANol Ht oOoNN INDADOARNADDDOOVOO0OO Spree Co) = Qe © 2) &) © 0000 Nan OMim OORT IN Unie onic Orel CORA TNC) CON Oa ialitat Gata! AMIN AAAAwWNOWMN +tNA0DDODOMNM+t+HAn N dad HA) OMIONTC OO (OY KO) (QU XO) IAS) (2) +ON+tOOCOOCHOOOSO aot +tONWOOMDMNONOO ~-ONDMADHAHANDHOO N Qar| PS Wap esp WAY ONC) (CF Gaited Ca) ©) INAOADA VANOCOHONO ANA MORARANKEEMONYA Ee ee iat Oe ali! tl toh XO) OLOL@ 1 Or@ OO. On Or'WNO Ast O Min OONHTHAUANNMMM®D GINS one) ah a A ee te oe el en ee ee ee | tee ANNO Aste Om OMIA AAAAHNNUNAMNM Si (continued ) INL Galle September 16-30 IQS IGS62 1963 IGS eS NOES ikea Alea) Direction OO NO A+\0NNOCOOO Cie Ome nen (en dale: Seu en, ek ers fe Lene VUNCO) © 1 eSP ny NI WM OO Oe © NAN il mises GO ONC! NO coo Oo eb peiee Kee te) 16r ee Oh en ner ceiw. jer el) ae Ey Bay Gal SED UNM ON GAT O © rl tl fe) hm Uu0 N+ \O MO M+ \O CO WA Mihesk tal GNC CA-ap Ose ~m 4 =p Un rl ON) © ON-Se Way GQ Se ON ee cen ek) el 0W wi lee Nex cen ied) sie? 4 Tet ve NMOM~MAOWMWAN NAN WL aN NMARPANMOOMNAMNNA oP Pel, ese rien Welw re: erey ear cel, “ler ey 726) NOE ONNOA NO OLN Ste HANAN A ONC AYP ONO OOOO CLOROE= S| Ou @) GIOEO TORO Amy Qi) CO) Sp QO WO Sp OVE WO DMAOMAMNOWMWONNA AO N49 SROEOROROTOROVON OMmodnHIterOMWMD OONHANAUANMMNM™M OW Ore! 0 0 UU 0 0 0 Mt tt tdAbd Aa Aa (HAHA VN NO HST OOM OMO DOA AANUNUNAMNM® Oetober 1-15 QUGOROTOVOTOsOIOOAOTOS Ove neti ies O ee yee IT ere el Rheky Yelgremeanen rie Min OLOLOEO TOTO OLer@ ee AN i= No) (0) No) Sp fr) @) NO) 2a7 @a)'©) GA) NO) (ONE e919) fel (SO) =r aig) GUI Xoo) (GM NA 4A OYA OVO AW Coo o © © Orel len) 40 xe er) cel Sete Nelle Se; 8 C6) Qi@ irs! QV CLO OG OOO HANM4 Q QQ =r HO Oe © oO OF) OFHNEOODOOOKN Nat NU YN Uy SU ON BS ©) Ga! GA) oa! cal ce OF Olen men iceh i Nereis ae) mer 5 eh re: pce! aAaANUANMMNNANA 4 QQ el el OV FAO Ott ANOO OT ie. Oo Cee rel eu ey en ei fe) ep ie NO) CF Sy, SND) 69) 6) GQ IOS) NA 4d ONDANMANAOODOOOO ey oere Ter Gel ce rie. 10n se: els lem rey ie ©) Co) Ql Ga) GQ IN Ll Se) Oo) OOS HANNAH KMstOo0NOrnKMmMeH O5. ist (0) = cen ever ey ier) es ie, en xe) > ie Keto OONOrK MMH HOA 4 SGVOTOTOTORO] ©@On Sy eS Ay ah A OOS OCOONHAAN (oa oe Oe) ONOVOWA SS a a a Mt taba Aaa ae PHaAH ANNO COM OMA NDNAAANAUNMM® o ~ \O qo NMNNODOAHAWMNMHAO MNMtTtOTANN AON 4 NO roa OOWM+AO00OrRNANAHO QQ ON=t = X00 fT A 4 1 © Qua) GN COs. @9) tal ar (esr cal cal Sp aq) oe cel ee iere 36) Jeet e elie,» f-er es emer ee) at \Ot+\O0ONOMOMEAA LEW Nod Hd OTOR OURO EOF@EOTO Me IN© A) [SS NO) Ga) QS) O] OO) Ors! [= AON A QUE O NOEs eee OROTOT © Stele OOm a NOL Otor@© oa o@) INO MINONOHHOCOOO oD Kents, Or Oia ee) els Once: len ie late NOMVWNAONOOCO ANNA S+tONMNONNANM+A M (Ole te eZ (0) — F050) = 16) 1 ein Nicer \ve) 5 he) ie Set Un © 9 ON NI = UN CO UN aaa) @NC0) 09) cl Qe) OT Or ey) 507) ers et Uren Rexx ewer) 18m 07). terre DO'OWMNINDHAOHOHA 4 (a ee) Qe @aQ©oO Oo © © OX (S) WeyCo) Ta) =r (SoS) Gai OONAANAUNAMNMY Ow Nel 0 0 0 U 0 a ob 0 mM! tt bid AaAd A AA ed Ee NCO STATE OM NOnAAAHAY mY 38 (continued) Table C-lL. November 1-15 LOGIN 4 NOCL O63 LOG 1965s OCG OG loo Direction Qi Way Ql Cy tale=p (SO) =p Co) tS Ue) QC) pales O)Se SS NGS) rr rt aI) Gal NO) COS) OQ) (OC) GUO) CO) a) URS Gal Wy SS CO) OS) TOYO GIS) ml QQ cS) ol NOINN ON NNMNINAN BE ay SSO Bama CAS vieues SP iit Coy QU] spe) QI Woy @) tal rare Se SE Te NGS SEINE UOC) Ga tee iG 09) QA) tal ce! OGY TONCO=ON-= SO) @) OO) iO CS] WNiCo) RS Copesp ny CPCS) Ie) reCURQisrt +DOMMNMNMNHA NDA NANA ~UN AO DN NO ANNM dH il tal ral @XOEt COE ONO ON OlON © ONO MMO +O-H OOD rl cl Uy oA) S) QW [SSCOKS) tal) 1) A) ta! ONDrAOAMWODHOAN HH Qt! tal el GS) (GS) GY ©) CY OO) OANCO al = ©. FANS OONMNAHAANNANMNMM @NOTON ES th alte mt t tad dA A A AHA ed ere re AGNCO ret ©) 1a) OMA NAAAANANANMMD 16-30 ONGO" ONS = COS Orr Gain (69) {S60 NO) Ea [Res e= PAW ay eal = 0.0) au o PS) 0.9),S) 69) (S= eit NOES Patan OL Or MO eee) 10), e310! Ore pe. ee ken le NOVAS GSS ING) ep SSI Ta er AS HN NICONGATOMIGCOD IGN =O = @).COiCOrs| Cer Ora ie SO ren ee a eee Cee at Yona ee N rH rl c! NOHO) Nig == PQ E= Qu @) See (Qu) SOMNANAHAOHOQD uN NA NO =F NON © Fw iA Ho OENey Oise Teese) =O) eh sel 58° -fe5 Xe: OINMNMNNONDAOMNA VOW fk foal tel NE TI C0):CO)G) [PO | OVO) Ca)\-Sr Oa)-=r Ci ceeme er le) eb ese ve. 0°" ie fe, .-0-. OMDAMNOAWODOMA MN mA qo rb Ay NS Ste) Teal ISS [SS rah CO) SS) P| (an) QENG) Talc) Noy Wop ita! @) CY ©&) fo OO O@) NOICOSI— ONO NOES Qari Or@ Wy SSN) 1) WN We aee (Nira! pe @) oe ee) @) @) 12) CY GSO) (On OUNCO So O@) GAIN OONAAHNANANMMNM ©) Ney onc Oe) 1 we Or Dee OM! tt AAA AA Ae [ira teal eal RN eo) teal ae [Cl Ga) O Mo Nn AANAATMs December 1-15 tet GOL OMar tO O=GaisiGniG CO NOR@ Sree srt Ov@ seer ete (9) A ON MW MNDOWONNANMOWMH WN Rt eer tes) tele ten SCS) 19) ANA A SS 3 Teal Mp WN To@): |S—a(6 Oe eS) CLO L a Oia ene. eu em fe Ser tel sni6; ie N rh ol al OM I= MO DW\O0 10 OOO O Ot COk Ewe U8. 461 Fel 0p eet, Buel ss erp oc 6 Wigy ONCo) Qi Gn) Galsricsr Ost esr fl Gal fal Gal ol Way We NOOO) Spo |S IS Sn ay Wevese ie FS ae SP COCO COE omy Site 20 aad i} fal NON OD ir! bk Or ONCOL O70 09 OO 105 mers (0S Mowplie) enue. ve: te! ce WO'0 NA MNOAO MIND MAN qo qo N ANMM+tMNMOYrOHAML ey Revd eo ey ferme: et se fe Se. 2 ell ale OMNM+t NMOCO INA MO os) teal Geol Teal tel Tl WNOrres QE NOS NNW) COP @ Cy Oe el Ch Ot Oe eee) oa = HN i SY) (continued) Table C-l. December 16-31 HOGI G62 e963) O64) l965" 1966 | 1967 119068 Direction MFA AO Rt WMO INA N | 4, VU MO ht NO ND MM oo eH N+ O AN WOO N\0 CO CO NINO ANWAOAODHNOCO NUNA NMMAMNMNOKN MMA NAA WMatINAdHOANMMKM® Hoag WMWoOMMNAMNMO CO ESO MN TN ONO) er NYG O + oon OOO ONWMNANMNVNEST AY COO OMNO WH st OO FA HANNA ONA AA AO OWMO Or NNAAN A ORS MA AN oN 4 +toAiMAAANMMNOO MM SERS) GA) CH) UNNI AG) WpveONO OHoN MANO HM] NOMA FAH Co) QW Say ©)-sp [S>-SP NO) pI Wo) GQ Gy HNN 0-30 31-60 61-90 91-120 121-150 151-180 181-210 211-240 241-270 271-300 3 Ol-33,0 SLOSS) LO Average semi-monthly wind speed in Beaufort Scale (numbers in parentheses are range) at Kure Atoll, 1961-68.* Table C-2. 1962 Nun 0.0) fo! em © WN d — MW moO tO Qe) aoe) Qt ON Nd 1968 8-12 knots, 4 = 13-18 knots, 5 = 19-24 39-46 knots 3 4-7 knots, knots, 6 = 25-31 knots, 7 = 32-38 knots, 8 knot, 1 = l=3eknots,. 2 Q= *Beaufort Scale (continued) Table C-2. August September October November December Jul 1962 OA) UA NO uN NO N a — rl UOT}ITpedxy LJaxseuel, zequON “HM °4T “STIOUL Touuostad *TIoO1V erny go sfaamns TeotsoToTq role Tz-61T Azenuelr QZ yoreW Q-€ 19q0490 6 Kew G aun Z2-9l Ttady ge wore T96T O96T 6S6T QS6T LS6T S61 CGI6T aAadng Jo 34ed °T-Sd eTqeL 50 ¢ (9961) Ssutqqoy SMiSN “suTQqoYyY *S JeTpueyD Q-9 ySnsny qzemaysg “V [ned (9961) sutqqoYy suTqqoy *§ seTpueyp >SMISN q-g Ateniqat gZo96T suey pue Ustad TpemeH Saptspoom ptaed BTUsosT Te) fo AQTSTaAtug ‘S‘Taurem *| preyoty owey pue YUSTta Tremey “‘TayTeM “TI pleuoYy TremeyH jo A4Ts -JaAtuy ‘yasno4tA *f UTYITeW BTUsOSTTeD fo A4tsTaatug fTaBsuTsSM WeITTIM eTqumTop yst4tag jo AqTs -ZaAtun ‘Apzeapn *A°d SOTXTW BTULOJT[TeO Jo AXYISTAA -Tun ‘ptodoeT zayzeqs “Vy TTemey jo AXTISTAA -Ttuq ‘xneamnowey *H saTrzeuo TTemMeH fo A4IsTaatuq ‘*soeqysar °O parempy euozyazy jo Aqtsaaa =-tuq ‘*‘°ae *‘taqy—Tng °qd aBato0ay "(4961) touzem pue Apreapn *(196T) ApreapN SHOTd *(196T) xnarnoweT *(€961) TesuTsn pue 1284 [Ng -Tpodxy aspt[ooD °*S prloreH HI-ZT tequeqydeg T96T aAING Jo 3a1eq SqydtazosnueW pue SsuOTzJeOTTGQnd 4JUeITNSSY Toauuos iad (penutyuco) ‘*T-Sd eTqeL 5) AL STMeyT sowuer “ZL Zaqny °“N sousTMBT G Arenuere 1,96T *Zoded stuL >d9d0d 09% O€ Aaquaoaq Q 1990490 *Zaded stuL Tauucsied ggqod snotzep Ca. Joe sree UasTaSTOYH °Y uewroNl *teded styy ‘(°sw) sutqqoy Smisn ‘sutqqoy °S tJaTpueyo OT-9 Arentge, uosdwouL °9O XeW STMeT sower ‘I, *zeded sTuL *dSd0d «= QT-9T Aaenuer 996T STMaT sower °°] QT taquaced *zeded stuL >dSd0d 09 ZI Aaequaaon *zoded styy ‘(°sw ‘Q96T) Z94TM *(9961) epnsi *(196T) eT2829S “(g96T) SUuBTIBqON pue Katats *(g96T) eeN ‘(696T *L96T) teTdey ‘(*sm) 6T ysnsny S96T Ze°0eTA *(Q96T) PreMpoomM pue ddeTD ‘(g96T) uosaowy Teuuoszad qSdOd snotzeA 04 ET Zaqueides €96T KaTqats °0 pera SUeTICION °“M J1AQoy ‘zeded styl xx? dSdod OT-g ACW sioa1eM uyoLr sutqqoy °S JaTpueyO "(996T) suTqqoy ‘Qmasn L-€ Aaenaqeg €O6T sqydtzosnuew pue suoTyeottTqng JueITNSaYy Teuuostad aAING Jo a7ed (panuTyuoo) “T-Sd eTAeL be * loaded *zaded *roded zoded STUL STUL *zaded sty} ‘(*sw) sutqqoy Ssydtazosnuew pue suotieoTTaqnd jue I TNSeYy werIsoig AaAING [TeoTSoTo}T_ ueasc0Q OT ftoed xx DOTAIOS SJTIPTIM pue USTH saqzeqs peytuUN x Tauuosazad gsqod snotzep ddeto °gq raes0y eThd *T Jyraqoy *dSd0d prieMpoom *M TNed Jowp~ng TayTem *dSdod ueuyoeyH setsnog °9 *>dSd0d STIIeH *“L ueA suTqqoy *g Je Tpueyo “SMaSn Tauuossadg Og eunr 696T 04 9¢ AeW TE€-9ge voreN QI6T 6 Atnr 04 + Lew O€-9¢2 yore 21-6 Aszensqeq 196T asAadng Jo a1eq (penutyuoo) ‘T-Sd eTaeL tn ssorzeqTe pe ,oos *zaded styy ‘(*sw) sutqqoy -yoeTd paepueg *(*su) sutqqoy Tetqray SUOT *(°su) sutqqoy -eATESqo eTOUssIO *(°su) uofuay *(qoQ6T) eoTY [eltray (“sw) 9 *(°su) uofuaey ‘(qoQ6T) a0TY Tetsay y) “(*su) uofuay ‘(qQo96T) a°oTY Tetley *(*sm) uofuay *(qoQ06T) eoTY Telrzavy 9 *(*su) uofuey *(6S6T) 2e2TY pue uokusy Tetzrey "(*su) uofuay ‘(6S61) e°0TY pue uofuay [elroy *(°su) uofuay ‘(6561 ‘9S6T) e°TY pues uofuay [etqley (G5) s0TyY pue uokuaey ‘(°sw) *T@ 49 YOTIPLY Tetzlay "4STC TBARN U9hT quepueuMoD O04 SauwaeyH SSN Teolsjo Sutpuew suot4 -wop jo Jtodey saatyory TeuotyeN “Gy DY -BATaSqo atoUssJO Sqydiaiosnuewm pue suotieoTTaqng 1Uey[NSoy aAING JO eamnideNn 2dTATaS SJTIPTIM DUS UST sazeqs peqtuN x SMaSN ‘sutqqoy °S tJaTpueyo SMISN ‘suTqqoy *S rteTpueyo Smaisn ‘sutqqoyd °*S tJaTpueyo SMiSo SMASN SMISN SMASN. SMISN SMISN. SMISN. ¥SMISN Swereal Tih BOP Cala Sowden ssn Tauuos iad 21 Atenuee €96T G2 UaLeN = O96T 9-92 ZJaqueydas 6G6T Qg aunt co Kew qT Ttady gS6T ST JIqusovoq qt AeW et Azeniged 1S6T Iz iequeced 6 zequaseqd 9S6T QT taqueydeg QT6T aAINeG Jo a3eq *TTOVV eany go skaAINs [ROTSOTOTG TOUTIN °g-Sq aTqez "(S96T) t8USTA "SUOTYBATASGO oTVeUIEISAS ON *SUOTJeATASGO oOTZeWAaYSAS ON °(996T) paempoom pue ddeto *SUOTIeATaSGO oTJeWAISAS. ON *(S961T) t9USTa sqydtzosnuewm pue suotyeottqng JueITNSaY 54 Asarmns ATosang Koaamns ATosang Xaarns AzTosing Xaains ATosang aAINg JO 9i1nIeN Ayatoog uoqnpny TremeH “pxzQO TaeyoTW *M a4nytysul qusweseueN OSTIPTIM ‘uosTetzqey “N etl Ayatoog uoqnpny TeuoTIeN ‘S‘JUaptserd ‘zaystayyong *M T4129 STOUTTTI ureyynog jo AYTSsaaa -Ttun ‘ioeusta *[ Aaarey suUeTIBTOW “M VIAaqoy ‘Ie SuosSZawYy uOTUTg “VY *dSd0d KaTqts *O par sUBTISION °M ZI9QOY ‘ie ‘uosdewy uotuTg “VY >dSd0d TI ‘Z42TM °O WETTLIM aUeTIeAOW °M ZAAQoYy :d9dod SsTOUTTTI uzreyynos jo AXTSsraA -Tun ‘wweTy y29q0y Touuosiad QT Azenuer +96T IL Iaqueseq O€ aun OT yoren og Arenigayq lz Atenuel €96T aAING fo a1ed (panutjuo.) ‘*g-Sd eTqeL owe) pue usTy TTemeH “TayTeM °T preuoy DD dSd0d ‘Suo0T y1eqoy SMISN ‘LaTpTzy auesng TremeyH jo AXTstaatun *(996T) AaTspireeg faaIns Airosang ‘faTspresg °M uyor HI taquaqydasg yted -- STees sanqdeg aITT eas SATTAY souwer OT-Qg tequeydag TremeH Jo Aqjtszaatugn *zeded stu, Aaains .ueqTd ‘xnounoueyt *H saeTzeu9 OT-ET YSnsny wat qoid ewey pue usty utaf, AZoog ssnostqd TTemeH Saptspoom ptaeq 6-/, Ttady dsdod djey = suey pue ysta tremey -- ‘sTees oanideg ‘IaYTeM °T preuoy Og-ST yoreyN SMASN. ‘sutqqoy °§ ZJaTpuey9 GT-cT yore SMISN “suTqqoy °S rteTpueyD AaeN "9° “SpaeuoeT ssoy AACN -- Aaarins Liosang SSoieMOZ 17. Somaeus) QT Atenuer +96T SJdTiosnuem pue suoTZeotTand 7UeITNSey dAING FO 3sind eN Tauuos dag eAING JO 34eq (penutjuod) *2-Sq aTaqeL 56 ‘zaded stu “GQ96T ‘TT Ttady -TE youre *TTemey SOFTTPTIM pue usta Joj Adejyazoag queqystssy ‘yaoder dtay Sqydtizosnuew pue suoT IeoOTTGng JUeITNHSaYy aAqIng Jo aanien ouey pues YSTq TTeMey AaAIns T[eag ‘IOYTeM “TT preuoy AYTSTIATUQ 34849 sesuey SWUeTYyY Yraqoy STOUTTTI utesuynog jo AXTSsSTIATUN Aavains ArosImM) ‘Taustd *[ Aaarey awey pue usTW TTeMey Seyesoy ‘dq Ts9uTT SMASN STeeas yuNnoD Suokuay *M [Ley awey pure ust Tremey ‘a0TY UOSsTAaN faamms Larosang ‘I9YTEM °T preuoy SMASN ssoryeqre pueg ‘suTqqoy *g JaTpueud SMISN. ‘suTqqoy °S tJaTpueyp SMISN “‘TeTpTzy suesng IOTIazZULT ayy jo AIG 1ATI9G JuUeISTSSV Xaarins AtTosang ‘uteg ‘y AaTueys Tsuuositadg (penutquod ) €T-OT [tady ¢€ Areniqeaq -T¢€ Arcenuer gz yoreN 4 Ttady QT-ST yoreM Z ttady aAING jo 696T 996T 996T S96T 27% ‘g-Sd eTAel Table BS-3. Total man-days spent at Kure Atoll by POBSP personnel. Personnel Days Whinllleitenmnls Open Weistec zee 316 Robert R. Fleet 267 Paul W. Woodward 220 Vernon M. Kleen 150 Dennis L. Stadel 149 David A. Bratley 106 Cameron B. Kepler 103 Thomas F. Dana 85 T. James Lewis 82 Paul G. DuMont 81 James P. Ludwig 80 Alan H. Anderson 70 George R. Wislocki 7O Robert L. Brownell 68 Robert A. Sundell Dil Roger B. Clapp 56 Robert L. DeLong 56 Ralph W. Schreiber 54 Norman N. Heryford 50 Robert S. Standen Ke) Kenneth A. Amerman yh Warren B. King Sil Richard L. Maze 36 Walter Bulmer 11 Robert di. seydle 9 Lawrence N. Huber 6 Charles A. Ely 4 C. Douglas Hackman y Robert W. McFarlane 2 Fred C. Sibley 2 Max C. Thompson 2 A. Binion Amerson, Jr. ale Total Man-Days Ze 478-373 DT 58 Subtotal Table BS-4. Man-days spent at Kure Atoll by POBSP personnel (listed by years). Personnel Dates Days 1963 James P. Ludwig September 13-December 1 80 William O. Wirtz II February 20; September 20- November 27 69 Kenneth A. Amerman October 30-December 12 yh Roger B. Clapp December 5-31 7 Warren B. King December 26-31 6 Robert W. McFarlane February 20; March 10; May 8-10; June 30 2 Fred C. Sibley March 10; May 8-10 2 A. Binion Amerson, Jr. March 10; June 30 al Subtotal 2a 1964 Robert R. Fleet January 26-June 7; October 11-December 23 209 William 0. Wirtz II February 23-May 4; July 23- November 25 197 Dennis R. Stadel July 12-October 4; December 10-31 107 Paul G. DuMont August 6-October 25 81 George S. Wislocki May l-July 9 TO) Robert A. Sundell June 7-August 2 Th Cameron B. Kepler October 26-November 29; December 23-31 yy Warren B. King January 1-31 Syl Roger B. Clapp January 1-26 26 Charles A. Ely November 8-12 ae Subtotal 826 1965 David A. Bratley May 6-August 19 106 Alan H. Anderson February 15-April 25 70 Cameron B. Kepler January 1-February 28 59 Robert R. Fleet March 4-April 30 58 William 0. Wirtz II March 28-May 16 50 Norman N. Heryford April 22-June 10 50 Robert S. Standen January 24-April 2 Ke) Dennis R. Stadel January 1-February 11 he T. James Lewis November 12-December 16 285 Table BS-4. (continued) Personnel Dates Days 1966 Paul W. Woodward April i () Aew AT ree-Ttady 04eT ATae yoem 4sT-AeH eunf Syaem g 4SeT-TTIdy-PTW ounf YoomM 4ST-T AEN Rew AT 1e8-yoOTeN- PIN sUN(-PTU-Yorey 9} BT Kew 078T-Ttady-PtIN Kew o.et-Azenuer 03e7 eune-[Tady syeem g 4YSITA Ken Aqne AT see-auNL 34eT OT Azenagqeq-Arenuer 318 T ZI Jaquesaq-1equaeaoN 948] IJOqUeAON 32427 sutAe,T sda yeod °69-96T ‘TToVy eAiny 7@ SutTAeT B3e yeed pue ButfeT ssa jo esuey 8 Kem Tt-ttady 9g ACW Yoeom 4ST ATnL Yoom 9ST-ouner 3 SYyoomM go 4SeT LeW Trady Trady yoom 1S8]T syeem g 4seT Ge Gz aunr-9T Ttady ZJoqueides ATzee-TtTady 3427 eunr ayet-€ Tpady eune a4yeT-Go [Tady aune ATAe98-Te Arenige, ounr o1eT-Arenuer 94eT Qg ysnsny-g Areniged qsn3ny ATszee-G Arenuer qsnsny ptw-Azeniqoq 97 T aun e4eT-[Tady 03eT ATNf-pTw-g oune Azceniged-ptw-)7T Arenuer Taqueced-ptw-)[ TequisAoN Taquedaq-pTu-+T TequeAon aun oun P= pT UW Syoom Zg 4SITq -AeW yoom 1SeT aun (:)£eW-PTN syoom Z 4SRT Ge ULaL 2FTUM AppoN umMorg ular, payoeq-Aery uray, A.oo0g PIF Gez,estIq Yeosy Aqoog peo s-pey Aqoog uMoIg kqoog peoejs-anTd prtqotdotL, pettey-pew JOLSMIV9UG SewySsTIYO Joyvemieaug poeTte.-aspom Terzed utuog ssoayeqrty uesfeyT ssorzeqTy pe yoos-yoe la oune Setoedg °C-98 STQeL urtaL 3>TUM Appon uMotg uda, peyoegq-KLery (peanut 7U0d ) Setoeds ‘g-0d STdeL 76 Presumably the timing of these cycles reflects the periods for each species when food was most abundant for egg production and raising of young (see Lack, 1966, for full discussion). It is not surprising that, at Kure Atoll, with well-defined seasonal variation in air temperature, sea surface temperature, day length, et cetera, birds bred in well-defined cycles; their food supply probably varied regularly and predictably throughout the year. During POBSP studies only 4 Red-tailed Tropicbird, 2 Blue-faced Booby, and 8 Brown Booby nests were found outside the species' normal breeding periods. The majority of these nests failed, indicating that environmental condi- tions at these times were unfavorable. It has been demonstrated on numerous occasions that lenghtening days stimulate breeding in temperate zone species (Rowan, 1929; Blanchard, 1941; Wolfson, 1952). At Kure the same stimulus is probably responsible for the initiation of breeding in spring and summer breeders. Shortening days may stimulate the albatross. Although the causes for these variations are unknown, it is probable that they are related to an actual scarcity or to the unavailability of food to the birds near the atoll. High winds, low temperatures, rain, and local storms are the most likely factors influencing food abundance and availability. Factors Affecting Nesting Success Details of nesting success are recorded in the appropriate Species Accounts. The following discussion summarizes those factors that af- fected success. Not all of them were equally important, and accurate quantitative data are lacking that showed the relative contribution of each, although rat predation and winter storms were obviously the most important. Polynesian Rats By far the most important factor affecting nesting success at Kure for most species was Polynesian Rat, Rattus exulans, predation. This mammalian predator has been present since at least 1870 when they were abundant (Read, 1912). Although unproven, it is probable that Polynesians accidentally introduced them long before Europeans discovered Kure. Thus, rats and birds have long been associated on Green Island, yet ap- parently no species has been extirpated by their predation. The first record of rat predation was Robbins' (1966) observation that they ate Brown Noddy eggs that had been left uncovered for only a few minutes. Since 1963 an additional 9 species (Table NS-1) have been recorded as victims of rat predation. Laysan Albatross, Bonin Petrels, Red-tailed Tropicbirds, Sooty Terns, and Gray-backed Terns have suffered most heavily from this predation. Adult and young birds that were attacked typically had open wounds, sometimes 1 to 2 inches in diameter, between the scapulae, slightly Ka anterior to the uropygial gland, or slightly posterior to the legs. Kepler (1967) describes the method of attack on Laysan Albatross in detail. Typical rat-damaged eggs were hollow, with a large opening aig One end Predation was not equally intense in all years. In early 1964 only a few Laysan Albatross and Red-tailed Tropicbirds were preyed upon. By the time albatross began breeding again in the fall of 1964, and through the summer of 1965, predation was extremely heavy. Black- footed Albatross, Red-tailed Tropicbirds, Sooty Terns, and Brown Noddies were also victims of rat predation. The peak of predatory activity was reached in the spring and summer of 1966 when large numbers of birds were destroyed, the area of predation increased, and more species were recorded as being preyed upon. Table NS-l. Avian species preyed upon by Polynesian rats on Green Island, Kure Atoll, 1963-69. Stage Preyed Upon Species Eggs Young Adults Black-footed Albatross x Laysan Albatross xX x BOnwnwee creel: x 5% Wedge-tailed Shearwater x ¥ Red-tailed Tropicbird x x Great Frigatebird ne Sooty Tern Ke x Gray-backed Tern x oe Brown Noddy x x White Tern b One year later only young Sooty Terns were noted being eaten. Predation became serious again in 1968 when rats destroyed most eggs and young of Sooty Terns, and all young and eggs of Gray-backed Terns. However, no albatross were known to have been lost. From mid-December 1968 through February 1969 several Laysan Albatross were destroyed. By June 1969 rats were noted preying heavily on eggs and young of Red- tailed Lropiebimds. Details of predation on each species are discussed below: Black-footed Albatross: One nestling was eaten in early June 1965. In late April and early May 1966 at least seven nestlings in the northwest beach-Scaevola ecotone were destroyed. Nestlings on the open sand near the north point escaped predation. Laysan Albatross: During the latter part of the 1963-64 breeding season at least 12 adults and a few nestling Laysans were eaten. In 78 the 1964-65 season over 50 adults and several nestlings were destroyed. The damage in these seasons was confined to the central plain, mainly the south antenna field. Laysan Albatross suffered most heavily from predation in the 1965-66 breeding season. From 8 to 10 February 1966 Chandler §S. Robbins found 57 adults that had been eaten by rats. Another two rat-destroyed adults were found in late April. Robbins estimated that there were 310 nests, mainly with nestlings, northeast and east of the barracks, primarily in the north (90) and south (110) antenna fields. By mid-April only four remained: one in the north antenna field, one in the south antenna field, one along the northwest beach, and one adjacent to the central plain. Only the latter bird fledged; the others were eaten by rats during mid-May. Robbins banded 400 nestlings in mid-March. At least 46 of these birds were eaten by rats. An additional 14 nestlings were destroyed in May and June. Predation was heaviest in the central plain, along the northwest beach, along the road to the pier, and behind the fuel tanks. No predation was noted in 1967 or during the 1967-68 breeding season. From mid-December 1968 through mid-January 1969 at least 32 adult Laysans were destroyed by rats. By February few adults were being molested by rats, but over 30 nestlings were killed and eaten, primarily along the runway, in the central roost, and along the southwest beach. Bonin Petrel: Rats probably caused the complete breeding failure of this species from 1964 to 1968. Rats were commonly seen running in and out of burrows and the remains of eggs were found in these burrows. To test this hypothesis, three study areas were established in 1969: one control area and two areas that were poisoned with Warfarin. No young Bonin Petrels were raised in the control area, but several fledged in the other two areas. Although other young fledged from areas that were not poisoned, it was probable that the poisoning had reduced the rat population in these areas as they were close to the study areas. One young Bonin Petrel, which was collected, had rat damage on the back. Wedge-tailed Shearwater: Several egg shells broken in typical rat fashion were found. Rats were also suspected of eating chicks. Red-tailed Tropicbird: In all years eggs and young of Red-tailed Tropicbirds were heavily preyed upon. Fleet (ms.) found that in 1964 53.6 percent of all eggs laid in a study area and 88.9 percent of nestlings hatched were lost to predation. Comparative figures for 1965 were 64.9 percent and 100 percent, respectively. 1 Great Frigatebird: On 16 May 1966 an adult male Great Frigatebird with a typical rat wound on its back was captured. Another adult male and 2 adult females with openings in the back were found on 10 June 1966. All of these birds were roosting. There was no evidence of pre- dation on eggs or young. sooty Tern: In 1965 rats were seen carrying Sooty Tern eggs from the colony when the incubating birds were disturbed. They straddled the eggs and punctured the end before carrying them away. From 1966 to 1968 both eggs and young were destroyed by rats. In 1968 they destroyed all the eggs and young. Gray-backed Tern: Two nestling Gray-backed Terns with openings in the back were found in May 1968. Eggs opened in typical rat fashion were also found. In 1968 the rats evidently destroyed all the eggs and young. Brown Noddy: Both eggs and young were eaten. Nests on the open beaches generally escaped predation. In 1965 there was a 27.9 percent nestling loss in a two-week period. Only 8 of the 147 chicks lost were found, and all had wounds in the body cavity. White Tern: A White Tern nestling with its whole back ripped open was found in June 1968. This bird may have been eaten by rats. Discussion; Two distributional patterns of predation were noted. In those species, such as Brown Noddies and Red-tailed Tropicbirds, where only eggs or young were eaten, predation was widespread, occurring almost everywhere the species wes present. The exception was individuals breeding on the open beaches who appeared to be immune to predation; individuals breeding in the densest vegetation appeared to be most susceptible. In the case of Laysan Albatross, however, where adults were eaten, the area of predation was localized. In 1964-65 it was confined to the central plain, mainly the south antenna field. In spring 1966, preda- tion was again extensive there, and had spread to areas north of the central plain, mainly along the northwest beach, where Black-footed Albatross were also eaten. By May it had progressed to the area by the barracks and behind the fuel tanks. Before the young fledged in July, a few along the runway were eaten. During the 1968-69 breeding season predation was noted along the runway, in the central roost, and along the southwest beach. The localized nature of albatross predation suggests that only a small group of rats, which had learned to attack these large birds, was involved... —Lossiply, they had overcome an initial fear of size), The fact that Fleet (ms.) found that tropicbird chicks more than 17 days old were not preyed upon supports this theory. Also, the progression of predation in 1966 suggests that a group was moving around the island attacking albatrosses. 80 As stated previously, predation reached a peak in 1966, when rats were extremely numerous. Starting in the fall of that year the Coast Guard began intensive poisoning around the island. By spring 1967 very few rats were noted, possibly as a result of the poisoning program or possibly as a naturally occurring population crash. Almost no predation was noted that year. In 1968 and 1969 rats were again numerous and predation wis heavy. Thus, there was an-apparent correlation with the number of rats present and the amount of predation. Not all species were equally susceptible to predation. Species such as Red-footed Boobies or Sooty Terns, whose breeding adults were either aggressive towards intruders or flew away quickly when disturbed, were predation-free in the adult stage, while others such as Blue- faced Boobies, whose adults vigorously defended their young, were free from nestling predation. Apparently rats could not break through the shells of albatross and booby eggs, so these were not eaten. The following model of predation is presented: Rats prey on eggs and young of many species every year, with the intensity of predation directly proportional to the size of the rat population. Another important factor is probably the amount of plant matter, which com- poses the bulk of the rats' diet (Wirtz, ms.). In years when plant food is scarce predation is heavier. During periods of peak rat abundance larger birds are preyed upon. The localized nature of this predation suggests that certain individuals learn to feed on larger birds and then move around the island. During the 1966-67 and 1967-68 alba- tross breeding cycles predation on albatrosses was reduced or eliminated, suggesting that most of the individuals that had learned to eat alba- trosses had died. It seems likely that there must be this type of predation cycle (i.e., intensive one year, reduced the next) on these birds or otherwise they would have become extinct at Kure Atoll. Storms Winter storms, with accompanying winds and high water, accounted for much of the loss of albatross nests. In 1964 from 16 to 22 December the island was battered by winds up to 65 knots and received several inches of rain. Water came up over much of the western beach and ca. 400 yards of the ends of the island were washed away. During this storm 12 of 25 Black-footed Albatross study nests and at least 54 Laysan Albatross nests along the exposed lagoon beach were destroyed. On 13 and 22 December 1968 storms with heavy winds hit the atoll. Several nests (exact number unknown) of both albatross species were destroyed by these storms. Some incubating adults were almost entirely buried by the drifting sands. Although POBSP observers were not present during the complete albatross breeding cycle each season, late spring observations suggested that wind-blown high water was responsible for at least some nest loss 81 every year, especially along the eastern beach where water usually reached the Scaevola. In May 1968 the high-tide line along the lagoon beach was within 5 feet of the Scaevola and 3 to 4 feet of the vege- tation along the edge showed evidence of extreme sand scouring. High Tides On 8-9 January 1969 water caused by high tides washed to the edge of the runway and into Scaevola along the northeast and southeast beaches. Several (exact number unknown) Black-footed Albatross nests were washed away. In early September 1968 high tides washed away 5 Brown Noddy eggs dice? nOrch pou. bh these sides ‘hadwoccurnned earlier in the year when most noddy nests contained eggs, almost all the nests would have been destroyed as the waves washed over the entire breeding area. Rain Heavy spring and summer rains were not known to have caused directly any nest loss, but an observation in May 1967 suggested that rain could indirectly be responsible in Red-footed Booby nest loss. After several days of heavy rains many Red-foot nests became water- logged and were bending over. Possibly nestlings and eggs fall out if the nests become too heavy. Undoubtedly long exposure of young birds to cold and rain could cause death, but this was probably a minor mortality factor. Human Disturbance Human disturbance affected nesting success indirectly, generally by causing the eggs or nestlings to be exposed to rain or cold. Some species, such as the Red-footed Booby, deserted their nests if handled or flushed repeatedly. Great Frigatebird Predation Although Great Frigatebirds are known to prey on other seabirds (Schreiber and Ashmole, 1970), none was seen to do so at Kure. However, when flushed from their nests, adult frigatebirds would occasionally re- turn to the nest and peck a hole in the egg, or pick the nestling up, fly away, and drop it to the ground. This action probably occurs rarely in an undisturbed situation. Great Frigatebirds also destroyed Red-footed Booby nests when humans flushed the boobies from them. seals Seals occasionally destroyed nests when they hauled out. For example, on 26 May 1965 a seal crushed 18 Sooty Tern eggs in a study plot near the northeast beach. 82 Dogs Dogs occasionally destroyed nestling Brown Noddies and probably nestling Christmas Shearwaters. otarvation No mass starvation, such as occurred on Ascension Island (Ashmole, 1963), was noted. However, every year a few emaciated Laysan Albatross nestlings were found. Some young Brown Boobies also apparently starved to death. Miscellaneous Factors Nesting success was also affected by infertile eggs, nest desertion, and adults attacking the young in such species as the Sooty Tern. Movement Through the recapture of previously banded birds it has become well established that seabirds of several species travel between various islands in the central Pacific Ocean. At Kure Atoll individuals of 13 species banded on other central Pacific islands or atolls were captured (Table M-1). In addition, an American Golden Plover and five Ruddy Turn- stones banded in the Pribilof Islands, Alaska, were collected at Kure. Four hundred and sixteen individuals of fifteen species from Kure were recorded in the central Pacific (Table M-2). Figure M-1 shows the loca- tion of these islands. Details of inter-island movement are recorded in the individual Species Accounts. Rigorous mathematical analysis of movement is difficult due to the following: (1) only a few individuals of some species were captured at Kure and/or banded on other islands, (2) some species were not banded on other islands, and (3) the banding and recapturing effort was inconsistent with respect to year and seasons--important parameters to consider in any analysis. Therefore, no detailed analysis of inter- island movement is attempted in this paper. Discussion of the movement of each species in the Hawaiian area will be published elsewhere. Although the POBSP banded thousands of seabirds in the Line and Phoenix Islands, there was only one definite movement recorded between these areas and Kure Atoll. It appears that in the central Pacific movement to and from Kure Atoll was restricted to the Northwestern Hawaiian Islands, Wake Atoll, and Johnston Atoll. Besides the inter-island movement in the central Pacific, forty-one individuals of seven species were recovered at sea or on islands outside the central Pacific. Figure M-2 shows the general location of these movements, which helps reveal the post-breeding dispersal patterns of several species. How much actual movement there was between Kure and 83 "eSIOA-S0TA IO pamyAdeosar useq sAey SpITq popueq-smMyYy ateym OTJToeg Terques uT STTOIe TO spueTST JO UOTZeOOT “T-W SmNsTT GNV1MOH - NOLVY NOLSNHOF~* IIWMVH a’ Iwnvy VOHIN®e YayDIN, SIVOHS 31V9 Yd HON3Yd” SJTDVNNId YaNdsVOe NVSAV1" ‘ DISNVISI 433Y S3WY3H PUO TaVad, AVMGIW, Toiw* aun» 84 ‘OT FTOe Terques ayy SpTsqno TO vas 4e pataAodear pue TTOIY emmy 4e pepueq spitq Jo suoTyeooyT *2-W amstTz VINVWSVL “aNVi1vaz aN A SI DIGvWwuaN” a+ | 4e403 VINVaeLsny oe WOOD vpndl= ;, Py ; s3aing “SI. ~ yowvs svsanouww! silnviaxo1 sOovdvivd si xINJOHa- + ia SONVISI @ ilolw vouruyor | APPON uMmoig Dx PsiqayoBisy 40915 @ . Aqoog uMo1g ¥. Paiqridos Pal!oi-P2y CO ssoajyogjy uosho7 SSO4JDQ|Y P2400j-420/g EY uJ VUSV1V 85 OgS°E — 1 Init Ge cg € i GE LT 12 LOT Ogu STCVOL te = = i = = = € O U € OT AppoN yore ta og O O O © = 0 O O O O rs AppoN umorg GLS O i HT O - - 0) q @) 2 SS udtay, ky009 rT O - O O - - 0 O T 0 9 JAAOCTd UspToH) ueolreuy 89 O O t = = I OT G € on € paitqez,estaiq yeory 06 0 OT an 2g 1 ©) SI 9 9 S 92 Aqoog peyoos-pey 9 1 O O O = O O O O + T fqoog umoig €T O 0 © 0 - © t © G € t kqoog peoej-antTd rf O ©) I - = O O 0 © ©) t patqotdot], poTtey-peu 4 = 0 O O g © c @) O ©) 0 TeyvemTesyg peTtey-sspem Zl - - - - = - T 0 € € S Teryed utuog S9S = = - - - 0 O c G re 9ES SsorpBqTy ueskey QST = - - - - O rs 0) O eg CET SSOLYeqTY peyoos-yoeTd Sotoedg Ghee GlO°k GBS G6 6LE GET 64 (SeTTMm TeoTyneu) soueqsTd ae | a) = wm oO A 4 © a EA mtd = (@) Oo © ct O o 0 Ph: th iy Fs: OM Oo hs ct = wy © jsr Sy G a @ fg |” Kk © on a) ar oO PS € o Oo 5 ory Hb: BR = = © Hn Hs o na ) i) O o n is ned ct ley fey 5 iS |” fd Q ct O oO i) ie) (o) 5 oO wr 90). 1S) ne is? es [=e @ Qu ct bh tT bs @) (e) 0a La) = L} |= "69-6661 ‘TTOVV eany ye pemmgjdeoer pue oTgToed Terjued ey} UT SpueTST TeyzO UO pepueg SprTd “T-W oTOeL ott + HE Se 9 OT 42 90€ co = O AL O O O AL € O O O O a O oC LAT T 9 0 c 0 S €€T aT O O O O O O T il = = = O O O al ale O O O O O O aL deat O O 9 iL € aL: t 88 cg 9c Sl T 8 t HE Cc 1 O O O O i O €T ©) oc T iE € T S il O O T O O O O t = O O O O T O € = = O O O € O. HOT = = 1c al: O o OOT TE = = O O ali 9 td 916 BER 669 US 99¢ Set + STBLOL TTO1V TIOVV S[Teoysg eye ueskelT TYsueTSTI Jooy sowleH T[To1y ayemM uo JSsUyOr -STAq youaery pue [zeog AemptIW SOTTW TeoTyNeU STe1OL AppoN Yoetd AppoN umMorg uzey, Ayoog suojsuany, Appny MeTIND peystyy-seTIsTag J2AOTd uap[oy ueotszouy PATQezesT IZ 7eetH Aqoog pe jooj-pey Aqoog umorg Xqoog peoej-antgd patqotdorz], peTteq-pey JOVEMIVSUG PeTTeI-espomM Teq,eq utuog ssormzeqrty ueskeyT SsOorZEATY pe yoos- Yoel Sotoodg aoue std Ne) eo) "OTJTOed TeijUs. 9|9y. UT SUOCTIeOOT Tayo 4e poanqydeoar pue [TO1V emmy 4e pepueq spxtTg °Z-W eTdeL 7 the Marshall Islands is unknown due to the small number of birds banded there and the limited recapturing effort in that area. It may well be that the Marshallsare part of the normal range for Kure birds such as Red-footed Boobies and Great Frigatebirds. Recovery records show a generally southerly or westerly movement from Kure outside the central Pacific, but this may simply be the result of the greater probability of recapturing birds in the western Pacific because of the greater number of islands in that area compared to the eastern Pacific. In summary, seabirds from Kure Atoll moved commonly to other islands in the Northwestern Hawaiian Chain, to Wake Atoll, Johnston Atoll, to a lesser extent to the Marshall Islands, and less frequently throughout the Pacific Basin--to California, to Japan, to Indonesia, and to the North Pacific. Species Accounts In the following accounts the observations refer only to Green Island. Sand Island observations are summarized later. The reader is referred to Figure K-l for the locations of the various areas discussed in the text. BLACK-FOOTED ALBATROSS Diomedea nigripes Status Abundant winter-spring breeder; 200 to 350 pairs annually. Present from late October to third week of July. Breeding begins in mid-November and continues until the last young depart in July. Populations Black-footed Albatross were first recorded from Kure Atoll by George H. Read (1912) of the U.S.S. Saginaw. He reported that "the main source of food will be the seal and brown albatrosses. Both seem plentiful..." and that the albatrosses were nesting at the extreme westward point. Lt. Commander Montgomery Sicard, Captain of the Saginaw, reported (in the Annual Report of the Secretary of the Navy on Operations of the Department for the year 1871) in a letter to Rear Admiral John A. Winslow: "As regards food, I commenced by sending out parties to kill seal and birds, but after about a month I found that, owing to the rapid diminuation of the seal, I was obliged to cut the allowance down, and killed one seal and twenty birds per day for the whole crew." Therefore, ca. 600 albatross were killed in December 1870 ana an unknown number, probably in the hundreds, in November 1870, indicating a Black-footed Albatross population of the same order of magnitude as was found in the early part of this century (Table BFA-1) and during POBSP studies (Table BFA-2), but considerably higher than Kenyon and Rice's 1957 and 1958 counts. 88 Table BFA-1. Kure Atoll. Population Previous records of Black-footed Albatross on Green Island, Date of Survey Estimate Breeding Status, Remarks, References 1870- October 29- 71 January 4 1915 1918 1923 1956 1957 1958 USS) 1960 1961 1962 1963 March 28 September 15 April 17-22 December 9 December 21 June 5 May 9 October 3-8 March 28 January 19-21 September 12-14 February 2-4 August 6-8 February 3-7 2 1,500 ca. 600 alos "Brown Albatrosses"” eaten by crew of U.S.S. Saginaw. Found nesting at extreme point to the westward (Read, 1912: 32, 68). Nesting in small colonies along the north- ern shore, western end. Few along the southern shore (Munter, 1915: 137). Offshore (R.G. 45, Nat. Archives, Report of Commanding Officer U.S.S. Hermes to Commandant 14th Naval District). 300 nesting pairs. Well-grown downy young in little colonies on open sandspits or scattered along beaches (Wetmore, ms.). Aerial count of 2,200 of both albatross species (Aldrich ety all... ms. )). Group of 64 counted from the air (Kenyon and Rice, 1958: 189). Estimated 50 young: count of 14 nestlings along south beach and 28 along north beach. 10 unemployed birds seen (Kenyon and Rice, 1958: 188-9). Estimated 50 young (Rice and Kenyon, 1962: 367). (Robbins, 1966: 53). 95 nesting pairs; 280 adults on island at one time (Robbins, 1966: 53). 160 nesting pairs; 200 adults on island at one time (Robbins, 1966: 53). (Udvardy, 1961). 65 nesting pairs; 100 adults on island at one time (Robbins, 1966: 53). (Robbins, 1966: 53). 200 nesting pairs; 235 adults on island at any one time (Robbins, 1966: 53). 89 Table BFA-2. POBSP semi-monthly estimates of Black-footed Albatross on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 ' January 1-15 - 650 700 - 500 - 700 16-31 - 650 700 * - - 700 February 1-15 - * 700 B25) DO: = (HO 16-28 * 650 600 ~ - - 1, 000 March 1-15 - 550 600 - - - 820 16-31 - 550 600 * * 400 780 April 1-15 - 275 400 - - - 675 16-30 - 275 300 300 - - 400 May 1-15 100 275 300 225 4.33 - 400 16-31 - 275 250 200 402 396 380 June 1-15 oe 200 235 182 ok 392 310 16-30 - 48 150 28 * 48 * July 1-15 - 4 6 4 * 2 - 16-31 - * 2 6) = 0 - August 1-15 - © O O - O - 16-31 - O - euuen© - O - September 1-15 - : 0 - © - O - 16-30 O O = © - ) - October eI) 0 0 = - - © ~ 16-31 * alae = a 2 % e November 1-15 32) 275 - - - * - 16-30 475 500 550 - = * - December LI) 570 700 550 - - * - 16-31 650 700 - 500 - * - *Birds present, number unknown 90 Whether or not the population increased from 1957 to 1963 is difficult to determine without early season estimates. POBSP data clearly demonstrate the fallacy of comparing estimates made late in the breeding season. For example, in May 1966 104 nestlings were counted; one year later 200 were present. It could easily be con- cluded that the breeding population had doubled in size, but in reality it was approximately the same both years as determined by December nest estimates. The differences in number of nestlings re- sulted from varying rates of nest loss each year. How much nest loss there was in 1957 is unknown but it probably was considerable since a tidal wave hit the atoll on 9 March 1957 (Rice, 1959: 19), two months prior to the Kure census. Breeding Black-footed Albatross were easily enumerated because of the relatively small numbers involved and the conspicuousness of their nests. In three of the six breeding seasons that POBSP personnel worked on the island, accurate nest counts were made early in the season; accurate nestling counts were made late in all seasons. These data indicated a yearly breeding population of 235 to 335 pairs. In contrast to estimates of breeders, estimates of non-breeders were inadequate. Generally they were based on a few counts and on the individual observer's judgment. From 26 December 1964 to 27 February 1965, and from February to June 1969, however, counts of non-breeders were made each week (Tables BFA-3-4). In 1969 most breeding Black-foots were streamered to facilitate recognition. These data showed that non- breeding Black-footed Albatross composed O to 55 percent of the total population using the island during a week. The only other population data of importance were the number of Black-footed Albatross captured during an entire breeding season (Table BFA-5). In 1963-64 almost four times as many birds were caught as the highest nest count. Assuming that all breeding birds were handled, the non-breeding population at least equaled the breeding population in size. At the same time it was estimated that only 335 individuals, including 235 breeders, were present at any one time. Obviously there was a large daily turnover. Evidently many of these birds were from Midway Atoll (see Banding and Movements), 49 miles to the southeast, and some were subadults returning to the atoll for the first time. The banding history of the 309 non-breeders captured in 1968-69 indicates the general structure of this population. Thirty-nine of these birds (12.6 percent) had bred in previous years at Kure, 106 (34.3 percent) were raised at Kure but had not bred yet, 12 (3.9 percent) had been banded on other islands, 21 (6.8 percent) had been banded at Kure but were not found breeding there, and 131 (42.4 percent) were unbanded. These latter birds were most likely raised on other islands. Therefore, the majority (53.1 percent) of non-breeding Black-footed Albatross was not of Kure origin. Table BFA-3. Counts of non-breeding Black-footed Albatross, Green Island, Kure Atoll, 1964-65. Number of Percent of Date of Count Non-breeders Total Population* December 26 1.55) 29.0 Sub 236 Sone Samuel 165 3052 16 182 36.4 23 123 B.S 30 233 np) Ik February 6 BSL DD oS 13 110 29.3 20 124 Ril 6S) ei N53} BIS ol * Total population includes the number of non-breeders and twice the number of active nests. Table BFA-4. Counts of non-breeding Black-footed Albatross, Green Island, Kure Atoll, 1969. Number of Percent of Date of Count Non-breeders Total Population* February 24 170 34.6 March 3-5 150 33.4 10-11 125 29.4 17-18 80 OO 24-25 80 2 © all 250 WS 5 April 11-14 110 26.8 20-21 80 21 @ 29-30 60 W6o 7 May 5-6 50 WS 12 10 2 2 20 3 L560 26 O 0.0 June 2-3 10 353 9 iL O43 16 @) 0.0 *Total population includes the number of non-breeders and twice the number of active nests. 92 Table BFA-5. Number of Black-footea Albatross handled during several breeding seasons on Green Island, Kure Atoll. Breeding Season Breeding Birds Non-breeding Birdst Total 1963 -64 216 TT6 992 1964-65 305 Treye) oo | edt 1965 -66 314 208 522 1966-67 216 4g 265 1968-69 538 382° 920 Many of these individuals were probably breeding but were not found on nests. 2 Almost all definitely not breeding. Annual Cycle Black-footed Albatross arrived on 24 October 1963, 25 October 1964, and 29 October 1968. The September 1918 record is unique. The population increased slowly at first and it was not until mid- November that large numbers of albatross were present. By mid- December the population peak, which continued through February (1965) and March (1964, 1969), was reached. Non-breeding birds were common from at least late December through March. In April they began to decrease rapidly until they were almost all gone by mid-May. After February, breeding Black-foots returned to the island only briefly to feed their young, so most albatross seen on the island at this time were non-breeders. Limited data show that 5-year old Black-footed Albatross were present from at least December through April, 4-year old birds from at least February to late May, 3-year old birds from at least February to mid-May, and 2-year old birds from early March to early May. Some 5-year Black-footed Albatross bred (three records in 1969). Black-footed Albatross bred on a well-defined annual cycle that showed almost no yearly variation. Figure BFA-l shows the 1964-65 cycle when the most accurate data were collected. The first egg was laid on 14 November in 1963, 1964, 1965, and 1968. Egg laying continued through mid-December, with a peak the last two weeks of November. From 64 to 70 days (X=67, n=11) after laying, the eggs hatched; the first egg hatched on 19 January in 1964, 14 January in 1965, 17 January in 1966, and 18 January in 1969. Hatching continued through the third week of February, with a peak the last two weeks of January. 3 One of the pair remained with the nestling 15 to 29 days (x=22.1, n=35) after hatching. Afterwards both adults returned periodically to feed their young. The fledging period for young Black-footed Albatross was from 137 to 170 days (x=151.2, n=13); the first young fledged in mid-June and the last one by: 18 July in 1964, 20 July in 1965, mid-July in 1966, and 18 July in 1968. Most young left the atoll in late June or early July. These young did not return to Kure for at least one and one-half years after fledging. 300 2 200 S eggs 5 = 100 =} =a N D | F M A M j j 1964 1965 Figure BFA-l. Breeding cycle of Black-footed Albatross on Green Island, Kure Atoll, 1964-65. Nesting Success Table BFA-6 summarizes Black-footed Albatross productivity for 1963 to 1969. Major factors in nest loss were winter storms, especially in 1964-65 and 1968-69, which washed away or buried nests with blowing sands, and rat predation in 1966. Hatching success was calculated in 1963-64 from 50 nests as 66 percent, in 1964-65 from 30 nests as 36.7 percent, and in 1968-69 from 331 nests as 57.7 percent. Thirty-three percent of the loss in 1964-65 was due to a storm which washed away 10 of the nests. Eighteen (36 percent of eggs laid and 54.5 percent of eggs that hatched) of the young fledged in 1963-64, and 145 (43.8 percent of eggs laid and 75.9 percent of eggs that hatched) fledged in 1968-69. In the incomplete study of 1964-65, nine young (36 percent of eggs laid and 81.8 percent of eggs that hatched) remained on 30 April. Crude rates of survival to maturity of nestling Black-footed Albatross were obtained (Table BFA-7). Although inexact the figures for the 4- and 5-year old birds are probably accurate in a general way as they will be breeding in a year or two and the ones that are still alive should be on the island. The lower rates for the 1966 and 1967 cohorts reflect the greater tendency of 2- and 3-year old albatross to remain away from the island. (Ge) een (OO) (0°0) 6d 6a Onc 0°OOT =< = => == = = +OT 99- S96T (Ge) eee) (6°0) (0°) eene ote Gaia ote 0°OOT -- == -- -- == Men G9-496T (O°CE) HO?) 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"69-€96T ‘TTOVV aany ‘pueTsI usery uo ssozyeqry peqooj-yoeTa Jo AYTATZONporg *9-WAE STAPL 95 *pomideos spxitq jo aeseqyueored ou} ST oINSTJ puoodss 9y4 pue SATTS Useq SsAeY OF UMOUY SPITq jo ssequeored |yy SqUeseadaz aaNSTyJ SATA x 0° OOT == ae =e = = ae a5 ae a6 SHI 69-896T (0°0) 0°0 0° OOT = == mi =e == ao 25 -- +02 89-L96T (Geer). (Or) GT Goi 0° O0T ao == == == == =- == 002 L9-996T S-B9a GePuviol AS-IS CS-Goor GOaCer We-ESer SG=gsior ASUS USHOSal OS-oIs1 QEIDENEEL CIE IES) pomydeossy uoseag Sutpssrg BUT pseIg (penutyuo0o) =" )-WH ETAL 96 Ecology Black-footed Albatross bred mainly around the island's perimeter on the open beaches and the beach-Scaevola ecotone. A few pairs also bred at the east end of the runway in the sand-Eragrostis association, along the east side of the runway, and in the open area at the southwest end of the runway. They laid their single egg in a simple depression in the sand to which occasional bits of vegetation had been added. The main concentration of nests was along the southwest beach and near the north point (Fig. BFA-2). In 1967, 61.5 percent of the nests were along the lagoon beach. This tendency for nests to be more common along the western beach than the eastern one was noted in all years by the POBSP. Non-breeding Black-footed Albatross roosted in the same areas as breeding birds. Before fledging the young albatross congregated on the beaches, generally near the water. Banding and Movements From 1960 through 1969 1,614 adult Black-footed Albatross were banded at Kure Atoll. Recapture rates of these birds from one breeding season to the next were high (Tables BFA-8-9) and probably approached the survival rate. As would be expected, however, Black-foots banded as breeders were recaptured more frequently than birds banded as non-breeders. Thirty-one adult Black-footed Albatross were recaptured on other islands: 24 at Midway, 6 at Pearl and Hermes Reef, and 1 on Lisianski. One of the Midway birds bred there. Ad adult banded at Kure on 16 Nov- ember 1963 was recaptured on Southeast Island, Pearl and Hermes Reef, on 13 March 1964 and then was found breeding at Kure on 5 December 1968. Eight hundred and seventy-nine nestling Black-footed Albatross were banded. The recapture of these birds has been summarized previously. Recovery of Kure-banded Black-footed Albatross at sea showed a general northeasterly post-breeding dispersal from Kure (Table BFA-10). One hundred and fifty-eight Black-footed Albatross banded on other islands were recaptured at Kure. The majority (133) was from Midway Atoll. Most of these birds had been banded by Dale W. Rice, Chandler §S. Robbins, and Eugene Kridler of the United States Fish and Wildlife Service, and Harvey I. Fisher of the University of Southern Illinois. One hundred and twenty-four were nestlings at the time of banding and 22 bred at Kure. Twenty-three of these Black-foots were banded at Pearl and Hermes Reef (10 as nestlings). Nine bred at Kure, but not at Pearl and Hermes. In addition, two nestlings banded at French Frigate Shoals, one in June 1963 and the other in June 1966, were captured. The 1963 banded Black- foot bred at Kure. NM 6 PUCTST Usemy U LO6T SPN STTOVV ean ) uO SssomzeqTY pe yooj-yoeTg Sutpsesizq jo souepunqe pues eee ‘ I q Td ‘e-Vad SAnsT OROOm (1°99) 1°99 (9°61) 9° 6h (€°6t) €°6t 98 89- L96T g9-L96T (0°89) Se 0°OOT -- (erm) Cresip) 4 GoSe S66 O°OOT (Gi) Stee) Se) O° HE 4° Sh 4°2S L9-996T 99-S96T S9-H96T 49-€96T O*OOT pornydecay uoseeg Sut peerg *¥69-096T ‘(Sesequeorted se pesseidxe) erley} peunydeoes pue [TOV eany 4e Sstoepeerq-uou se papureq ssoryzeqTy peJooj-YyoeTd FInpe fo soyes amnydeoey (1°99) 1°99 O°O0OT == (este)! 5 (2 4S) (exes) T'9S I Ah 6°28 (g°92) (€°92) (2°99) +°6S 6°S9 0°62, CE-iG)=e (COL)) = 6) 22) 9°89 6222: €°€Q (ie) = (Gee). 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SZocS FINPV €Q6T TOQUeACN + C96T Arenue Ll T.6€,60T NiO€.8E FINPV 496T YOLEN +T L96T ATRL ST T,OToLHT NOS TH x}TNPe BUT pese1g €96T Azenuer JT c96T AINE M.Otochl N,Ov.04 SUT TISON O96T YOLEN QZ puno,y usyM puno,wy o7904M | sutpueg jo poepueg a1eq owt. 4e easy “OTJTOeg [e14U90 91 apTsqno pateAooar pue [TTOVV eINy 4e pepueq ssozzyeqrTy peZooj-yoeTg “Ol-VAd STaqeL *pamades spatq jo zaqunu ou} ST SINSTJ puoodses ay. pue SATTe Udeq aAeY OF UMOUY SpAitq Jo seseyueorted ayy. sqUuosomdet sINsTJ YSATA x O° OOT -- -- -- -- -- TET 69-996T (O° Ss) (OO) 0° Sg 0° SZ 0° OOT -- -- -- 0g 1,9-996T 9-896T 99-L96T 19-996T 99-S96T S9-496T +49-€96T “u poepueg uosees perangdeossey uoseeg Sutpeerg BUT pooarg (panutjuco) *6-VWid eTqeL 100 LAYSAN ALBATROSS Diomedea immutabilis Status Abundant winter-spring breeder; 800 to 1,600 pairs annually. Present from early November until mid-August. Most abundant December- March. Breeding begins the third week of November and continues until mid-August. Populations All population estimates of Laysan Albatross prior to 1961 (Table LA-1) are considerably lower than POBSP estimates (Table LA-2), indicat- ing an increase since 1923. Undoubtedly this increase resulted at least in part from the bulldozing of trails in 1959 and the construction of the LORAN station which increased the potential breeding habitat. Presently at least 50 percent of the population breeds in areas that were inaccessible to them prior to 1960. The main source of the new breeders probably is Midway Atoll, 49 miles to the southeast. The Midway population was subjected to heavy human disturbance in the late 1950's and early 1960's and it is likely that this was also a factor in the population increase at Kure. In three of the six breeding seasons that POBSP personnel worked on the island, accurate nest counts were made during the early part of the season; accurate nestling counts were obtained in all years. These data indicated an annual breeding population of 1,600 to 3,200 Laysan Albatross. As with most species studied at Kure, studies of non-breeding Laysan Albatross were inadequate. Only two worthwhile studies of this type were conducted. In February and March 1965 large numbers of albatross were handled and painted according to their location on the island: north beaches, central roost, central plain, and the area south of the runway. It was hoped that subsequent ratios of painted to unpainted birds would yield information on the total number of Laysan Albatross using the island. On six days during late February painted and unpainted birds were counted in the central roost. Using the following formula, Number painted observed = Total number painted, Number unpainted and painted observed Total population the following "estimated populations" were obtained: 101 Number Number % of Pop- Painted Unpainted ulation Estimated Date Observed Observed Painted Population February 17 86 154 35 ld 2,480 23 169 261 39.3 2,028 ok 102 61 62.6 2, 220 25 87 ho 68.6 2, 240 27 54 23 Oval 2,550 28 106 61 6B 5) Ae On 28 February the total population using the island was calcu- lated to be 6,120. Since 3,279 adults were painted during February and an average 30 percent of those handled each time were unbanded, it is not unreasonable to assume this estimate was fairly accurate. At the conclusion of the study 4, O51 Laysan Albatross had been handled. During this time the maximum number of breeders was 1,070. Even if it is assumed that all breeders were handled, it still means that 73.6 percent of the total population using the island at least once were non-breeders. From late January to mid-June 1969 an attempt was made each week to capture as many non-breeding Laysan Albatross as possible. Due to the large number of birds handled, two nights were necessary to complete this work. Breeding birds had been streamered previously to facilitate the separation of breeding from non-breeding albatross. Weekly counts (Table LA-3) showed that on the average 713 non-breeding Laysan Albatross roosted on the island each night during this study. At their peak of abundance these non-breeders composed 42-58 percent of the total popula- tion using the island. By late June 5,197 non-breeding Laysan Albatross had been handled. Since only 2,000 albatross were breeding, non- breeding Laysan Albatross composed at least 72 percent of the population that used Kure Atoll during the 1968-69 breeding season. The composition of the non-breeding Laysan Albatross handled was as follows: 486 (9.4 percent) had bred previously at Kure, 943 (18.2 percent) were raised at Kure but had not bred yet, 727 (14.0 percent) had been banded at Kure as adults but had not been found breeding there, 339 (6.4 percent) were banded on other islands, and 2,702 (52.0 percent) were unbanded. Therefore, at least 58.4 percent of the non-breeding Laysans were probably of non-Kure origin and 27.6 percent were of definite Kure origin. Annual Cycle After an absence of almost two and one-half months, Laysan Albatross returned to the atoll in early November (earliest records: 4 November 1963, 5 November 1964, and 4 November 1968). The September 1918 record OZ Table LA-l. Previous records of Laysan Albatross on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 300 Scattered over island, main colony near middle of island (Munter, 1915: 137). 1918 September 15 2 Offshore (R.G. 45, Nat. Archives, Report of Commanding Officer U.S.S. Hermes to Commandant 14th Naval District). 1923 April 17-22 LOO Young present; ca. 50 pairs nesting on sand beaches at edge of bushes, either alone or with colonies of Black-footed Albatross (Wetmore, ms.). 1956 December 9 ? Aerial count of 2,200 of both albatross species (Aldrich, et al., ms.). 1957 June 5 805 270 nestlings counted; 75-100 adults present (Kenyon and Rice, 1958: 189). 1958 May 9 600+ Count of 300 chicks (Rice and Kenyon, 1962: 367). 1959 October 3-8 9 (Robbins, 1966: 53). 1960 March 28 hoot 75 nesting pairs; 425 adults on island at one time (Robbins, 1966: 53). 1961 January 19-21 1, 100+ 550 nesting pairs; 700 adults on island at one time (Robbins, 1966: 53). September 12-14 0 (Udvardy and Warner, 1964). 1962 February 2-4 2, 160+ 1,080 nesting pairs (Robbins, 1966: 53). August 6-8 hO+ 20 nests (Robbins, 1966: 53). 1963 February 3-7 2, 900+ 1,450 nests (Robbins, 1966: 53). 103 Table LA-2. POBSP semi-monthly estimates of Laysan Albatross on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 1-15 - 3,600 4,000 - 1,450 - 5, 000 16-31 - 4,900 4,000 - - - 5,000 February 1-15 - * 4,000 2, 000 1, 000+ - 5,000 16-28 * 45500 6,120 - - ~ 6, 000 March 1-15 - 35250 6,500 - - - 5,000 16-31 - 3,250 7, 000 - * * 5,000 April 1-15 - 3,200 4,500 - - - 3,500 16-30 - 3, 000 1,500 1,300 - - 3,000 May 1-15 1, 000 3, 000 1,100 IL MOO 2,458 - 1,500 16-31 - 3, 000 950 100 2,281 1,400 iL, ZOO June 1-15 - 3, 000 950 990 2, 209 1,220 1, 000 16-30 - 1,250 950 955 2,133 1, 200 * July 2-15 - 400 200 100 * 300 - 16-31 ~ 50 TN) 14 - XQ) - August 1-15 - 14 y 2 - 4 - 16-31 - 0 - 0 - 2 - September 1-15 - O - ©) - 0) - 16-30 O O - ©) - O - October 1-15 O ©) - - - 0 - 16-31 O ©) - - - ©) - November ss 1-15 12S; 200 - = = a 16-30 900 1,000 3, 000 - - * - December 1-15 3,450 4,000 4500 - - * - 16-31 3,600 4,000 - 1,450 - * - * Birds present, number unknown, 104 Table LA-3. Counts of non-breeding Laysan Albatross, Green Island, Kure Atoll, 1969. Number of Percent of Date of Count Non-breeders Total Population* February 24-25 950 43.8 March 3&5 1, 000 Wh 4 10-11 W250 Ome 17-18 800 42.6 24-25 850 ToT) Syibeal 1,400 B53) April 11 & 14 1,250 55 4 20-21 1, 000 51.3 29-30 850 Whe. May 5-6 865 hres? ee ee 330 25.8 20-21 415 30.4 26 110 TOs June 2-3 250 ele 9 70 T8O 16 25 2.6 * Includes twice the number of nests and the number of non-breeders. is unique. They were not numerous until late in the month and not until December was the population peak reached. By mid-April the population began to decrease, mainly due to the departure of non- breeders; by mid-June adults were rarely seen. By late August all Laysan Albatross had left the atoll. Non-breeding Laysan Albatross were present from at least late December through June. They were most abundant, composing as much as 58 percent of the total population using the island, from at least January to mid-April. Limited data showed the following periods of occurrence for various Laysan Albatross age classes: 4-year olds and 5-year olds-- at least late January through June, 3-year olds--at least February to mid-May, and 2-year olds--from mid-March to early May. The first three age classes were most common in March and April, while the latter group was never common. IOS) Laysan Albatross bred on a well-defined annual cycle, from mid-November to mid-August, which showed no significant yearly variation. Figure LA-l1 shows the cycle for the 1964-65 breeding season when the most accurate data were collected. 800 2 600 as eggs iS 400 g . = 200 N D J F M A M j j A 1964 1965 Figure IA-l. Breeding cycle of Laysan Albatross on Green Island, Kure Atoll, 1964-65. The first egg was laid on 21 November in 1963, 20 November in 1964, 17 November in 1965, and 19 November in 1968. Eggs were laid until late December, with a peak in late November and early December. The eggs hatched 63 to 74 days (x=66.9, n=15) after laying; the first eges hatched on 25 January in 1964, 26 January in 1965, and 22 January in 1969. Peak hatching occurred the first two weeks of February. By the end of that month all eggs had hatched. HorMlO) vo ell idayis "(x—15.4/7.. n=e3)) atver hatching abeleast one of the pair remained with the chick. Afterwards they returned only briefly to feed it. The fledging period for Laysan Albatross was 137 to 163 days (x=150.5, n=8), thus, young albatross began to leave the atoll during the first week of July and all were gone by late August. Most birds fledged the first two weeks of July. Some of these young Laysan Albatross returned to the atoll for the first time ca. two years after hatching. Nesting Success Table LA-4+ summarizes Laysan Albatross productivity from 1963 through 1969. The smaller number of young that fledged in 1965, and especially 1966, resulted from rat predation. In the latter year only one young fledged from the area northeast and east of the barracks. Sandstorms, which buried many nests, caused much nest loss in 1968-69. Hatching success was determined for 25 nests in 1963-64 as 44 percent, for 25 nests in 1964-65 as 60 percent, and for 864 nests 106 in 1968-69 as 61.7 percent. Fledging successes for the respective years were 24 percent, 20 percent, and 48.3 percent of all eggs that were laid, and 54.6 percent, 33.3 percent, and 78.2 percent of eggs that hatched. Table LA-5 summarizes the recapture of nestling Laysan Albatross in subsequent years. The low rates for the 1966, 1967, and 1968 co- horts reflect the tendency of younger Laysans to remain away from the atoll rather than low survival rates. Ecology Laysan Albatross bred in most of the open areas on the island, avoiding only the area around the barracks and, generally, the exposed open beaches where the similar Black-footed Albatross bred in largest numbers. They were also found in fairly dense Scaevola thickets and under Tournefortia where there was enough open area underneath to permit the adults to waik from the nest to an open area. The greatest number of nests was present in the central plain, along the runway, behind the fuel tanks and along the lagoon beach-Scaevola ecotone. Very few nests were found along the east beach. Figure LA-2 shows the general location of the breeding areas in May 1967; the sites were similar to those of previous years. The single egg was laid on the ground. As incubation progressed the adults built a mound of vegetation around their bodies, thus form- ing a substantial nest. The range of the non-breeders coincided with that of the breeders. Nests often became the focal point for roosting aggregation. Towards the end of the breeding season young Laysan Albatross began to form groups along the beaches where they made their first attempts to fly, often ending up in the lagoon and swimming back to shore. Table LA-4+. Productivity of Laysan Albatross on Green Island, Kure Atoll, 1963-69. Maximum Approximate Percent Maximum Nestling Number of Fledged from Breeding Egg Count Count or, ~ Young Maximum Number Season or Estimate Estimate Fledged of Nests 1963 -64 1,600 1,600 1, 000 62.5 1964-65 B41 475 470 OW yate. 1965-66 700 80ot 75 52.6 1966-67 650 1,050 1, 000 -- 1967-68 -- 701 600 -- 1968-69 1, 000 625 465 46.5 1 Also 100 eggs (February 7-10), C.S. Robbins estimate. 107 °196T ASW STTOVV eany ‘pueTSI user) uo ssozveqty ueskey Sutpeszq Jo uoTANqT14STC "Z-WI eamnsTa 108 *peanjdes spztq jo aseqyueozed 9y3 ST aINSTJ puodas 9y4 pue SATTB Useq aAeY OF UMOUY SpITq Jo aBequoeozed oy4 squeserdar aInSTJ YSATAx 0°00T cy =e = a = =e 5 or =F LO 69-896T (0°0) 0°O 0°O0T = zs = == at =5 ss a TOL 89-L96T (E°e) (0°0) E"S Soe Q°O0T 2s sa = =a “9 oe == 56 L9-996T (Ue) — -(@°O)) (0°0) T' t2 T*t2 Tt 0° OOT = =e, a = as ae LS9 99- S96T (E-9i)) (0) (4°0) (0°0) T' On T'9t T'9t T' On 0° OOT =2 == == ae == elt S9-496T (Q°gG) — (VES) (0"¢) (T"0) (0°0) Q°as 9° €S 9° €S 6°€S 6'€S 0°O0T == -- == =< Qe6 9-€96T (1°9) (0°0) (0°0) (0°0) (0°0) (0°0) (0°0) L°9 IPS) L°9 MSE) LAO Mi? ® L°9 0°OOT =< =n GT Z9-T96T Gi ec) ee (ORO) (2°T) (9°2) (G2) (9°2) (0°0) (0°0) tt ec 4° €S 9° €2 c° 92 €° Ge G°QE G°ge G°Qe 0°OOT =- ATS T9-096T (9°S) (0°0) (0°0) (9°S) (26m) (ert) 120) (0°0) (0°0) 9°S 9°S 9°S 9°S Guage EES eree (SoKcis €°€€ O°OOLT QT 09-6S6T 99-S96T S9-196T +9-£96T €9-c96T Z9-T96T T9-O96T 09-656T “UU pepueg uoseas peinjdeosay uoseeg SuTposrg Sut posrg Ga 890m 19> 2061 89> 99er *x69-666T ‘(seBequeoded se posseidxe) aztey4 poringdeoer pue [TTO1V ony 1e sBut{T4yseu se pepueq ssoreqrty ueskeT Jo soyzer amydeoay °C-VI eTqel 109 Banding and Movements From 1960 through 1969, 11,711 adult Laysan Albatross were banded at Kure Atoll. Tables LA-6 and 7 summarize the recapture rates of these birds in subsequent years at the atoll. These recapture rates are probably near the survival rates as large numbers of Laysan Albatross were handled, especially during the 1963-64, 1964-65, and 1968-69 breeding seasons (Table LA-8). As would be expected, a larger percentage of breeders than non-breeders were usually recaptured. One hundred and four of these adult Laysan Albatross were captured on other islands: 100 at Midway Atoll, 2 at Pearl and Hermes Reef, 1 at Laysan, and 1 at French Frigate Shoals. Thirteen of the Midway birds and the Laysan bird were recaptured later at Kure. From 1960 through 1969, 4,475 nestling Laysan Albatross were banded. The recapture of these birds has been discussed previously. One banded on 15 May 1964 was found dead on Eastern Island, Midway Atoll, in December 1967. The recovery of Kure-banded Laysan Albatross at sea (Table LA-9) indicates a basic northeasterly post-breeding dispersal from the atoll. One Laysan Albatross banded as an adult by POBSP personnel and 27 banded as nestlings on other islands were recaptured at Kure. The adult was from Midway Atoll and the nestlings were from the following locations: Midway Atoll (3), Pearl and Hermes Reef (21), Lisianski (1), and Laysan (2). None was found breeding. In addition, 537 Laysan Albatross banded by non-POBSP personnel, mainly Harvey I. Fisher, Dale W. Rice, and Chandler S. Robbins, were captured at Kure. The majority (532) had been banded at Midway Atoll; 54 as adults and 478 as nestlings. Eleven of the adults and 55 of the nestlings were found breeding at Kure. The other Laysans were banded as nestlings: 1 at Pearl and Hermes Reef and 4 on Lisianski. 110 *pomnydes spitq jo eseqjueozed oy Sst dINSTI puodas 9y} pue SATTS Udseq SACU OF UMOUY SpAtTq jo asequaeorzed ayy sqyuasordezt ainSTty YSITA x 0°OOT am aS o4 = = == a = qLe 69-896T Gace Gace Gace O°OOT -- -- -- -- -- QT 99- S96T (Oi72) = (0 0) (€°g) (0° 22) O° ld Ow (Sea ES 6° Lt 0°OOT -- == ais a Qt G9- 796T (Sete) 2 (60) (9°9) (6,2) “(e"2e) Gad 0° Se 8° 9d ots eee) 0° OOT -- -- -- OS 9-€96T Giga) (60) Cig) (ee) SSS) = (ea) lige G° l2 6°92 9° On Pale) 9° HQ 0°OOT == ae OFF €9-296T (yee) (1°0) (9°9) (E266) (S06) SCeHO2) Cr: Ec) + °c? eee G° 92 T°0S 6°09 9° €Q 4° SQ ©°OOT -- 256 Z9-T96T Caen) = (Ort) (eS) (9762) & (S79c)e (9°29) (261) aCe 42) eked i oO L206 H°6€ f° 6h G° SL Gay, 9°19 0°OOT 96€ T9-096T OECSSIE 1692296 = WiIG=9 9099S S96 S9= TOC TOSe9One sc9[e 90 Lc9a=L96 L9=096T2 =U PepuRe UOBBOS poungdeoey uoseag Sutpesrg SUT paar g °¥69-O096T ‘(Se8equeoted se passerdxa) o10y} pernqdeoes pue TTOVV euny ye scepserq se pepueq ssozyzeqty ueskel[ 4Inpe fo seyer eanqdeooy °9-VI eTqeL ILE 94% ST ocNSTJ puoodses oy} pue SATTSe Useq oveYy 9-896T 99-L96T 0° O0T (6°2t) 2° €9 (€°TE) Gos (€° TE) 07 UMOUY O*OOT = (€°9) 0°0S poanjdeosy uoseeg suTpoeorg Spatq fo (€°9) 0°0S 99-G96T S9-496T 49-€96T €9-296T <9-T96T *poanjdeo spitq jo esdequaored aVeyuooted ay} sqyuosorder aamsTtjJ YSITA x Aovh=@ OT * x69-6S56T ‘(sesequeoted se pesseidxe) ezeyy pornqdeoes pue [TOV amny 7e@ Sdepserq-uou se pepueq ssouyzeqry ueskeT 4[npe jo sezer omnj,deoay popueg uosees BUT p9o1g 69-9896T 99- L96T 1,9-996T 99- S96T S9- 4961 n9- £96T 09-6S6T *2-VI eTIeL qe Table LA-8. Number of adult Laysan Albatross handled during several breeding seasons at Kure Atoll. Breeding Season Breeding Non-breeding Totals 1959-60 0) 16 16 1960-61 396 O 396 1961-62 1, 040 13 1053 1962-63 1, 030 an TOs 1963 -64 TSS: 3052 4,670 1964-65 359 3, 787 4,146 1965 -66 129) 1,929 3, 220 1966 -67 342 294. 636 1967-68 0 168 168 1968-69 I Sully 5,197 6,514 Table LA-9. Laysan Albatross banded at Kure Atoll and recovered at sea. Age at time Date Banded of Banding Where Found When Found 20 January 1961 Breeding adult 32°00'N 173°00'E March 1961 3 February 1962* Breeding adult 27°45'N 130°10'E 26 April 1965 8 February 1962** Breeding adult 42°00'N 144°00'E 2 August 1967 5 February 1963*** Breeding adult 42°30'N 169°20'E 10 September 1965 28 December 1963 Breeding adult 36°12'N 141°25'E 24 January 1964 26 January 1964 Adult 39°35'N 146°38'E 5 September 1964 14 March 1965 Adult Whe es 'N 173°O02'E 31 October 1965 26 March 1965 Adult Marshall Islands: 20 May 1965 Mejit Atoll Q February 1965 Breeding adult 42°30'N 169°20'E 10 September 1965 Q9 December 1963 Adult 31°02'N 131°33'E 8 March 1967 30 March 1967 Adult 42°5O'N 160°X)'E 17 August 1967 21 January 1964 Breeding adult 42°11'N 155°18'W 25 July 1968 9 December 1963**** Breeding adult 21°50'N 159°20’W 10 March 1969 1 May 1961 Nestling 4O°50'N 150°00'E 1 November 1962 29 July 1964 Nestling 432°19'N 155°27'E 27 July 1965 27 June 1967 Nestling 41°49'N 150°06'E 17 October 1967 15 May 1964 Nestling 36°O9'N 143°46'E 10 December 1966 * Recaptured at Kure 16 November 1963. and 30 November 1965 (breeding). KEKE 1968 (breeding). Recaptured at Kure 15 January 1964 (breeding). Recaptured at Kure 14 December 1965 (breeding) and 4 December Recaptured at Kure 27 December 1963 (breeding), 20 November 1964 TIS) BLACK-FOOTED x LAYSAN ALBATROSS Diomedea nigripes x immutabilis A hybrid albatross was collected on 26 January 1964. This individual, a female, resembled a Black-footed Albatross in plumage and bill size, but had a lighter-colored head than usual, a whitish abdomen, and a horn-colored bill similar to that of the Laysan. NORTHERN FULMAR Fulmarus glacialis rodgersii In 1964 four dead Fulmars were collected: one each on 16 January, 26 February, 28 March, and 30 March. These latter two birds were fleating in the surf off the west point. Only the January specimen, a dark-phase individual, was prepared as a study skin. The color phase of the others was not recorded. Fulmars have been also recorded in the Northwestern Hawaiian Islands from Midway Atoll and French Frigate Shoals (Clapp and Wood- ward, 1968). BONIN PETREL Pterodroma h. hypoleuca Status Abundant winter-spring breeder; maximum estimate 2,500. Present from early August to June. One July record. Peak populations present from September or October through mid-March. Breeds from late January to late June. Populations Munter (Table BP-1) probably overlooked Bonin Petrels during his brief diurnal stay, while Wetmore gave no numerical estimate so it is impossible to determine whether or not the Bonin Petrel popula- tion has changed numerically since the early part of this century. Robbins' October 1959 estimate, made prior to the alteration of the island, is in general agreement with recent POBSP ones (Table BP-2), suggesting that the LORAN station construction had little effect on the population size. Recent POBSP estimates indicated a maximum of 2,000 to 2,500 individuals using the island during a two week period. These estimates, however, are considered unreliable and the total population may be considerably higher. For example, during the 1963-64 breeding season 1,620 Bonin Petrels, about 900 less than the highest estimate, were banded. The comparable figures for the next season were 1,005 and ca. 1,000, respectively. It is unlikely that 64 percent of the total population using the island in 1963-64, and 50 percent in 1964-65, were banded; these maximum estimates were probably too small. 114 Table BP-1. Previous records of Bonin Petrels on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 -- (Munter, 1915). 1918 September 15 Zi Offshore (R.G. 45, Nat. Archives, Report of Commanding Officer U.S.S. Hermes to Commandant 14th Naval District). 1923 April 17-22 ? None found nesting (Wetmore, ms.). 1957 June 5 -- (Kenyon and Rice, 1958). 1959 October 3-8 1, 000 (Robbins, 1966: 53). 1960 March 28 -- (Robbins, 1966: 53). 1961 January 19-21 500 (Robbins, 1966: 53). September 12-14 2 I resting (Udvardy and Warner, 1964: 2). 1962 February 2-4 200 (n 1966: 53). August 6-8 il (Robbit. [Su S)\e 1963 February 3-7 10 (Robbins, Even more illustrative of the difficulty in estiwating the size of Bonin Petrel populations was a roosting study conducted from 11 February to 14 April 1969 (details to be published elsewhere). On alternate nights at ca. 2100 hours as many Bonin Petrels as possible found roosting or bur- rowing in either. the north antenna field or the open area just south of the barracks were captured. Unbanded individuals were banded and the band numbers of previously banded ones were recorded. During this study 1,747 different Bonin Petrels were handled; yet the maximum number handled on any single night was only 213. Obviously there was a considerable amount of turnover. Usually at least 90 percent of the birds captured on any night had not been handled the previous night nor on any other given night, and at least 40 percent had not been handled previously during the study period. Since only 451 Bonins were captured more than once, the Kure population was highly transitory. Based on an average daily total turn- over rate of 50 percent, and a daily population of 100 birds from September through April, the total number of Bonin Petrels using Kure at least once would be 12,000. Figure BP-1 shows the number of Bonin Petrels captured each night during this study. This is a better indication of the size of the population than the semi-monthly estimates. INS Table BP-2. POBSP semi-monthly estimates of Bonin Petrels on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 1-15 - 225 LOO = 2, 000 - 4OO 16-31 - 225 1,500 - = - 800 February 1-15 - * 1,500 2,500 2,500 - 2, 000 16-28 - 2,500 1,500 - - - 2,500 March 1-15 - 2,500 2, 000 - - - 2,500 16-31 - 750 2,000 - * 100 1, 000 April 1-15 ~ 750 200 - - - 500 16-30 - 750 100 150 - - 250 May 1-15 * 300 25 20 100 - 200 16-31 - O 1 il 10 1 200 June 1-15 - O O O 1 0 100 16-30 - O O O 0 0 - July 1-15 - 0 ©) O 0) ©) - 16-31 - O O 0 - il - August ae - O ho 50 = aL = 16-31 - 50 - 200 - 500 ~ September 1-15 - 500 - 750 - 1, 000 ~ 16-30 * 500 - 1, 000 - 1, 000 - October IES) * 500 - = = x 4 16-31 * 1,000 - - = * - November 1-15 750 1, 000 - - - * = 16-30 400 1, 000 700 - - * - December 1-15 400 1,500 1,000 - 4 * 2. 16-31 500 ISCO ~ 2,000 - * - * Birds present, number unknown. 116 April March February 10 90 fo} io-) 70 0 0 40 30 20 10 140 130 126 100 spsig jO Jaqunyy two study areas, 11 February-14 April Number of Bonin Petrels captured each night in 1969, on Green Island, Kure Atoll. Figure BP-1. ILL Since the maximum nest count was 500, most Bonin Petrels using the island were non-breeders, probably birds from other islands (see Banding and Movements). This would account, in part, for the rela- tively stable population size despite the almost complete breeding failure of this species. Annual Cycle Bonin Petrels returned to the atoll in early August (earliest records: 6 August 1962, 10 August 1965, and 8 August 1968). At first only a few individuals were present but by September or October they were abundant. Peak populations were present through mid-March. The size of the population then decreased rapidly and by May few petrels roosted on the island. Usually all birds had departed by late May (latest records: 24 May 1965, 25 May 1966, 4 June 1967, and 27 May 1968). In 1969, when young Bonins fledged, adults were present until at least mid-June). The July 1968 record is unique. Although courting and burrowing began shortly after the adults returned, egg laying did not begin until late January (23 January 1964 and 1965 and ca. 17 January 1969). Apparently most eggs were laid in late January and early February. In 1969 a female with an egg in the oviduct was captured on 19 February. This was in accordance with the termination of egg laying for other years. Eggs were known to have hatched only in 1969, when the first ones hatched in early March and the last ones in early April. Most eggs hatched from ca. 1€ to 20 March. The first chicks fledged in early June and the last ones the third week of June. [It is not known how many years will elapse before these young Bonin Petrels return to the engollice Nesting Success From 1964 through 1968 no young Bonin Petrels were known to have fledged from the atoll. In 1968, however, one young may havebeen suc- eessfully reared, as on 17 July an adult was captured that regurgitated stomach oil when handled, a sign in most procellariiformes that young are present. A careful study of burrows in the area revealed no young. The only success data were for 1969 when three study areas, one in the open area just south of the barracks, one in the north antenna field, and one in the south antenna field, were established in February. The former area was unprotected; the latter two areas were poisoned with Warfarin to protect them against Polynesian rats. In the area near the barracks 20 nests were marked in mid-February. By 25 February they had all disappeared due to rat predation. The eggs had been broken and the shells were often found outside the burrow. 118 Four eggs of the 9 (44.4 percent) marked in the south antenna field hatched. Two of these young (50 percent) fledged. In the north antenna field 24 of 40 (60 percent) marked eggs hatched and 10 of these chicks fledged (25 percent of all eggs and 41.7 percent of eggs that hatched). Rats destroyed a few of the nests and others were probably lost at the initial disturbance, but the cause of loss later in the year was unknown. An estimated 80 young Bonin Petrels, all from the central plain, fledged in 1969. Most of these birds were outside the protected areas, so they were subjected to possible rat predation. Whether the poisoning program was sufficient to eliminate these mammals from the central plain, or whether the population was naturally reduced, is unknown, but the data from the study areas strongly suggest that Polynesian rats were the major factor in the breeding failure of this species at Kure Atoll. Ecology Bonin Petrels occurred mainly in the area just south of the barracks, the small antenna field by the tennis court, and at the northern edges of the north and south antenna fields. The major breeding habitat was the sand-Eragrostis association. Here they dug burrows from one to six feet long, generally under a grass clump. Some birds, possibly young ones, dug shallow trenches, but these were not utilized for breeding. The single egg was laid near the far end of the burrow. Other burrows were found under Scaevola near the grassy areas, in the Tribulus-covered central plain, and in Open areas of the central roost. Not all of the sand-Eragrostis area was used by this species. In 1969 the majority of young Bonin Petrels fledged from burrows along the edges of Scaevola at the northern end of the south antenna field. Banding and Movements Robbins banded 48 adult Bonin Petrels and the POBSP banded 6,050. The low recapture rates of these birds (Table BP-3) agree with the previous discussion of the large amount of turnover in this species. Eighty nestling Bonin Petrels were banded in 1969. Since Bonin Petrels burrow and when they are roosting on land their legs are in almost constant contact with the ground, bands are subjected to heavy abrasion. After three years or so the bands become so thin they fall off. Thus recaptures after three years are Lower than would be expected. Three Kure-banded Bonin Petrels were recaptured at Pearl and Hermes Reef: an adult banded on 10 March 1965 was recaptured on Seal 119 *poanydeo spatq jo esequaored 5Y4 ST oINSTJ puodes oy} pues SATTe Useq aAeY OF UMOUY SpaTq Fo aSequeoszed ayy Sjueserdet eansTy 4SaTA x 0°O0T -- -- -- -- “= -- -- -- -- 966 ‘T 69-9961 (Che SP) (0°0) Lee: AEE 0°O00T ae 25 Jo ae 22 oo ae TH6 L9-996T (rds) (O20) (4° 22) 9°22 Caged a et 0° OOT == -- -- -- -- << T6E 99- S96T (ae2/) (0°0) (g°2) (6°2) Bed. Med, 9° €T O° Gir 0° O0T =~ -- -- -- -- Goo ST S9-196T (6°S) (9°0) (9°) (9°2) (7g) 6°S 6°S G6 67 tE eG 0° O0T -- -- -~ -- 0g9 ST 49-€96T odd Z°de Z’dcd Goce cece 0°O0T .-- -- 6 Z9-T96T 0°O 0°O O°O 0°O 0°0O O°O Ono 0°0 0°O 0°O0T -- al T9-096T ©°O O°O 0°O 0°O 0°o 0°O 0°OOT BE 09-6S6T 99-S96T S9-796T 9-€96T £9-296T 29-T96T popueg uoseag peingdecsy uoseeg SuTpee7rg BUT poeorg 9-896T g89-L96T L9-996T "x69-6661 ‘(Sesequeored se pessoidxe ) aZoy} peanjdeoez pue TIo1VV ezny 4@ syIMpe se poepueq sTezjed uTUog Fo soyer oanqgdeosy *¢-dd STAeL 120 Island 8 days later and then captured at Kure on 23 November 1965; another adult banded on 23 November 1964 was recaptured on Seal Island on 19 March 1965; and an adult banded on 21 October 1964 was captured on Southeast Island on 16 March 1965. Twelve non-breeding Bonin Petrels banded on other islands were recaptured at Kure (Table BP-4). Table BP-4+. Adult Bonin Petrels banded on other islands and recaptured at Kure Atoll. Place of Banding Date of Banding Date of Recapture Midway Atoll 29 March 1960 3 October 1964 Midway Atoll 5 February 1962 30 November 1964 Midway Atoll 6 February 1962 8 February 1966 Midway Atoll 7 December 1963 9 September 1966 Midway Atoll 17 December 1963 15 September 1966 Pearl and Hermes Reef 16 March 1965 10 September 1966 Pearl and Hermes Reef 7 March 1963 11 February 1966 Pearl and Hermes Reef 1 March 1963 8 January 1965 Lisianski 12 March 1965 25 February 1969 Lisianski 12 March 1965 13 March 1969 Lisianski 13 March 1965 23 April 1969 French Frigate Shoals 12 March 1967 27 August 1968 KERMADEC PETREL Pterodroma neglecta On the evening of 20 April 1923 Wetmore collected a male, dark- phase Kermadec Petrel as it flew over the central plain. Another bird, possibly this species or the similar Herald Petrel (Pterodroma armonjoniana heraldica), was seen flying over Green Island on 31 July 1964. Wirtz described this individual as follows: "size and build of a tropicbird, lighter brown on back than a Wedge-tailed Shearwater, very sharp brown to white demarcation line on breast like Brown Boobies, rest of underparts very white, white underwings." Kermadee Petrels occur commonly at sea in the central Pacific. MURPHY'S PETREL Pterodroma ultima Ludwig collected a female Murphy's Petrel on 7 October 1963 as it flew over the central plain. Murphy's Petrels have also been recorded in the Northwestern Hawaiian Islands at French Frigate Shoals (Clapp and Woodward, 1968). 121 BULWER'S PETREL Bulweria bulwerii On 11 September 1964 DuMont saw a small all-dark bird with a wedge-shaped tail and a light band across the upper wing surface gliding over Scaevola. DuMont acknowledged that this individual may have been a young Brown Noddy rather than a Bulwer's Petrel; hence, the occurrence of this species at Kure is considered hypo- thetical. WEDGE-TATLED SHEARWATER Puffinus pacificus Status Abundant summer-fall breeder; 500-1,000 pairs annually. Present from March to early December with peak populations from April through October. Breeding begins in June and continues through early December. Populations Estimating the size of the Wedge-tailed Shearwater population was difficult, not only because of their fossorial and nocturnal habits, but also because they bred under the dense Scaevola. Most estimates were based on the observer's judgment or on banding data, rather than on any consistent sampling. However, in 1967 and 1968 a method was utilized that allowed comparison of the population size on a weekly and yearly basis. Once a week, after dark, Wedge-tailed Shearwaters were counted in the central plain and the major breeding area south of the barracks. It was assumed that this number represented about 10 percent of the total population. This estimate took into ac- count the larger area where the species occurred, the presence of undetected birds in burrows, and daily turnover. During 1964, 2,481 adults were banded. If this species was Similar to the other procellariiformes on the island, the total popu- lation using the island each year was at least twice this figure. A Lincoln Index calculation also suggested that the population size was at this level. In 1965, 173 of the 404 Wedge-tails handled had been banded in 1964 as birds in adult plumage, thus: 173 = 2,481 where x = total population. Lok X Therefore, x = 5,794. Thus POBSP observations indicated a total yearly population of 5,000-6,000 individuals using the island. An estimated 1,000 to 2,000 Wedge-tails bred each year. Data are inadequate to determine how many of the non-breeding birds using the island were Wedge-tails that had bred previously at Kure or were subadults raised at Kure that had not bred yet. The majority of the Kure population was light-phased. Only one adult of 137 (0.73 percent) handled in 1967 was a dark-phase morph. 22 Comparable figures for 1968 and 1969 were 0.69 percent and 0.43 percent, respectively. Only 19 dark-phase birds (out of 4,319) were banded by the POBSP. Comparison of earlier observations (Table WIS-1) with POBSP estimates (Table WIS-2) is of limited value, but the available data tend to indicate no significant change in the size of the population during this century. The major breeding area was always in the central plain, an area that was not greatly altered by the construc- tion of the LORAN station. Whether more Wedge-tails breed under Scaevola now than in 1923 is not known. Table WIS-l. Previous records of Wedge-tailed Shearwaters on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 -- (Munter, 1915). 1923 April 17-22 300 Calling and burrowing (Wetmore, ms.). 1957 June.5 common Numerous burrows (Kenyon and Rice, 1958: 189). 1959 October 3-8 5,000 Young (Robbins, 1966: 53). 1960 March 28 common (Robbins, 1966: 53). 1961 January 19-21 ) (Robbins, 1966: 53). September 12-14 2 Hundreds of burrows (Udvardy and Warner, 1964: 2). 1962 February 2-4 O (Robbins, 1966: 53). August 6-8 1,000 (Robbins, 1966: 53). 1963 Februarv 3-7 © (Robbins, 1966: 53). Annual Cycle Wedge-tailed Shearwaters were first seen flying over the island on 2 March 1964, 19 March 1965, and 17 March 1969. The population in- creased in size rapidly and by the end of March hundreds were present. Apparently the largest populations were present from April through October. The 1967 and 1968 counts (Fig. WIS-1) suggested that Wedge- tails occurred on the island most commonly prior to egg laying and 23} Table WIS-2. POBSP semi-monthly estimates of Wedge-tailed Shearwaters on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 1-15 - O O - - - ©) 16-31 - O O - - - O February 1-15 = O O O O ~ O 16-28 O O ) - - - O March 1-15 - 300 ©) - = a 0 16-31 - L750 300 - * 25 1,000 April 1-15 - 3,500 1,000 - - - 5,000 16-30 - 3,500 1, 000 750 ~ - 5,000 May 1-15 * 3,500 1,500 1,000 3,300 - 5, 000 16-31 - 2,500 2,000 1 (50 3,960 3,990 5, 000 June 1-15 - 2,500 2,000 OO 25350 1, 260 5, 000 16-30 - 2,500 2, 000 LSD CR OKO 1; V4.0 * July 1-15 - 2, 000 2, 000 1,800 * 4080 - 16-31 - 2, 000 2, 000 2, 000 - 6, 230 - August 1-15 - 2,000 2,000 2,500 - 3,790 - 16-31 - 2, 000 = 2,000 = 5,410 - September 1-15 - 2, 000 - 2,000 = 2,440 = 16-30 * 2, 000 - 2,000 - 3,530 - October a1 * 3, 000 - - - * a 16-31 * 3,000 = a = x 2 November 1-15 750 3, 000 - - - * - 16-30 1D 50 800 = = 4 ie December 1-15 * O 1 - - * - 16-31 O ©) - ©) - O - * Birds present, number unknown. 124 SUIYI}EH °99-L96T ‘TTOVV emmy ‘pueTST User TO uTeTd Terqus. 949 UT SZoTeMITeSYS peTTe7.-espem fo squnod Aine auny Kew le vz Zt ol € 9% 6l rl S 67 zz Sl Buike] 333 ‘T-SIM Omnst aT ool 002 00e 00) 4 00s 009 002 008 006 Jaquny SPp4lg 40 125 hatching, and least commonly during the incubation period. By November the population began to decrease and all Wedge-tails had left by the middle of December. Although courting and burrowing began shortly after the adults returned to the island, egg laying did not begin until June (11 June 1965, 17 June 1966, 8 June 1967, 18 June 1968, and 12 June 1969). Evidently peak egg laying occurred the last two weeks of June and in early July. Some eggs were probably laid as late as the third week of July. No yearly differences in the timing of peak egg laying were evident. The first nestlings were found on 31 July 1964, 30 July 1965, early August 1966, and 9 August 1968. Hatching continued at least through August. Young Wedge-tails began to fly 99 to 111 days (X=103, n=10) after hatching. Thus fledging began in late October and was completed by early December. Young Wedge-tailed Shearwaters did not return to the atoll until they were ca. 2 years old. Nesting Success In 1964 50 nests with one egg each were studied to determine hatching and fledging success. Forty-seven eggs (94 percent) hatched and 40 nestlings fledged (80 percent of all eggs laid and 85.1 percent of the eggs that hatched). Seven nestlings died before fledging. One was killed by an adult during defense of its nest, one was aban- doned, and the remaining five were suspected victims of rat predation. In some years hatching success may be considerably lower. For example, in August 1968 only 11 of 178 (6.2 percent) active Wedge- tailed Shearwater burrows checked in the central plain contained eggs or chicks. Whether all these burrows once had eggs which were destroyed, or whether only a small number of the burrows were utilized by this species, was not determined, but it suggests that nest loss may be heavy. Broken eggs with typical rat openings at the ends were commonly found in the central plain. An estimated 1,000 young in 1964 and 400 in 1965 fledged. Ecology Wedge-tailed Shearwaters bred mainly in the central plain and in the grassy area just south of the barracks. Here they dug burrows, generally at the base of Bragrostis or other vegetation. Some burrows were also found under Scaevola. Other Wedge-tails laid their egg on top of the ground under Scaevola, where little or no direct sunlight penetrated. Generally the egg was placed in a shallow trench rather than on flat ground. Sixty nests of this type were found from 24 to 28 June 1967, indicating that they were not uncommon. They were found wherever there was a con- tinuous growth of Scaevola but appeared to be more common in the north roost. Bae Bae 126 When they were not at sea, in burrows, or flying over the island, Wedge-tails roosted in the same areas where they bred and along the beach-Scaevola ecotone. Banding and Movements Robbins banded 235 adult Wedge-tailed Shearwaters and the POBSP banded 4,319. The percentage of these birds recaptured at Kure was relatively small (Table WIS-3). In some years (e.g., 1967 and 1968) little effort was expended in capturing this species, resulting in lower recapture rates, but even in years when large numbers were handled (e.g., 1964, 1965, 1969), they were low, suggesting a high rate of turnover. An adult Wedge-tailed Shearwater banded the night of 12-13 March 1964 was recaptured the next night on Southeast Island, Pearl and Hermes Reef, 135 nautical miles to the southeast. Three hundred and seven nestling Wedge-tailed Shearwaters were banded (4 in 1959, 5 in 1963, 237 in 1964, 57 in 1965 and 4 in 1966). Three of the 1964 cohorts were recaptured in subsequent years at Kure: 1 on 23 July 1966, 1 on 5 September 1966, and 1 on 31 July 1968. Four Wedge-tails banded as adults on other islands were captured at Kure. One banded at Kilauea Point, Kauai, on 17 August 1964 was recaptured on 23 September 1964 and another one banded there on 26 March 1964 was captured on 5 September 1966. ‘Two banded at French Frigate Shoals, one on 8 August 1964 and one on 19 June 1966, were recaptured on 19 November 1965 and 29 August 1966 respectively. SOOTY SHEARWATER Puffinus griseus The remains of eight Sooty Shearwaters were found on Green Island: 1 on 18 May 1965, 1 on 23 April 1966, 2 on 4 May 1966, 2 on 28 May 1966, 1 on 26 May 1968, and 1 on 29 May 1969. Five of these birds washed up on the lagoon beach and three on the ocean beach. On 19 May 1966 Woodward captured a Sooty Shearwater with ex- tremely worn flight feathers as it swam in the lagoon. Sooty Shearwaters are regular migrants through the central PAaciiic CHRISTMAS SHEARWATER Puffinus nativitatis Status Uncommon spring-summer breeder; ca. 25 pairs annually. Present from March through October. Breeds from late April until October. 127 *pornjydeo spitq jo esequaorzed ayy ST oINSTJ puooses oy} pue SATTe Useq aAey OF UMOUY Spatq jo asequeor1ed ayy sqzUuosoTdet aInSTy YSATA x <2 ae 2s == =e oe == -- -- -- -- TOL 696T (6°L) 6a O°OOT ss == -- -- -- 5° oe == 7 oe: JLT —- Q96T (S°9) (9°2) ar 6 0700 == -- -- -- -- ae oe =5 got —- 4.96T oe Er) eS) ©)? J, 6°93 COTE 0°0OT -- -- -- -- -- -- -- OSt 996T (orm) (orn) (SO) = (eS) Ho €°g 0° O0T 33 a =o oe ac = 902 S96T (S°O) <(e2O) — Cert) Creep = (@ou) S°0 IO CoS NX G6 O°COE -- -- -- -- -- TSn°S 96T (O°O) (O°O) —rn)— Gies) (erin) (O°O)) 0°0 0°0 Oba i) Sot OP Ait 0°OOT == a == -- 96T €96T BE) (Eo). Ceroe})) (20) Said G2 e° ee Ere OZOOM -- -- -- QL, Z96T (O20) (20) — (©) (GRO) (0) (OO). (OO) (OPO) 0°O O°O 0°0 9°0 9°0 9°0 9°0 9°0 OOO LST 6S6T 96T 896T L961 9961 S96T H96T CO6T ZT T96T O96T G6T popueg pounqydeossy re3az TCoR °x69-656T ‘(sesequeoted se pesserdxe) etey} pornqdeoet pue [TO1V einy 1e sartnpe se papueq sro 1eMTeeyg poTtey-eSpem Jo soqyer aanqydeooy °C-SLM eTqeL 128 Populations Robbins banded four adult Christmas Shearwaters and estimated the population as 125 in October 1959 (Table CS-1). This represented the first known record for the atoll. Since there was little habitat change from 1923 to 1959 it is probable that Wetmore overlooked this species in 1923. This is not surprising, due to the small population and secretive nature of Christmas Shearwaters. POBSP estimates are given in Table CS-e. Fairly accurate population data were collected for three years. In 1967 four nests and 15 additional nest sites were located, indicat- ing a breeding population of at least 38. One year later, between late May and mid-August, 48 different Christmas Shearwaters, two nests, and at least 23 potential nest sites were found. In 1969 12 hests, 2 nest sites, and 52 adults were located. Thus, if this species was similar to other procellariiformes on the island, the total population was at least twice the number of potential breeding birds, or 76 in 1967 and 100 in 1968 and 1969. The Lincoln Index can be used for an approximation of the total population using the island. Nine of the 42 adults banded in 1968 were recaptured in 1969. An additional 39 adults were banded and 4 banded in 1967 were recaptured. Thus, _9=42 where x = total population. Therefore, x = 238. This figure is not unreasonable as almost 25 percent of the estimated population was handled in both 1968 and 1969 and a con- siderable number of adults were unbanded in 1969. In summary, the total Christmas Shearwater population using the island was 100 to 200, with ca. 25 pairs breeding annually. Table CS-l. Previous records of Christmas Shearwaters on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 -- (Munter, 1915). 1923 April 17-22 O (Wetmore, ms.). 1957 June 5 -- (Kenyon and Rice, 1958). 1959 October 3-8 IPS 4 adults banded (Robbins, 1966: 53). 1960 March 28 -- (Robbins, 1966: 53). 129 Table CS-1. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1961 January 19-21 0 (Robbins, 1966: 53). September 12-14 2 (Udvardy, 1961: 46). 1962 February 2-4 @) (Robbins, 1966: 53). August 6-8 ©) (Robbins, 1966: 53). 1963 February 3-7 ) (Robbins, 1966: 53). Annual Cycle Christmas Shearwaters were first seen on 11 March 1964, 15 March 1965, and 11 March 1969 after an absence of almost four and one-half months. By the end of the month most of the population was present. The population peak was maintained through at least mid-September. The population decreased as the young fledged and by late October all Christmas Shearwaters had left the atoll. A breeding cycle was constructed from data from the 30 known nests (2Bine963— Siciny 19645. Se ine lOG5pes mm lOG6n 4h inlo67, 3) in 1968, and 12 in 1969). Data were inadequate to detect yearly variations. Egg laying occurred from late April (earliest record: 26 April 1964) through at least May, hatching from mid-June to mid-July, and fledging from late September to late October (latest nestling records: 25 October 1964 and 28 October 1968). The limited data suggested an egg-laying peak in early or mid-May. Ecology Figure CS-1 shows the approximate location of Christmas Shearwater nests found on the island. The majority (59.3 percent) was located under scaevola south of the barracks, suggesting that this was their center of abundance, or at least that nests were found most easily there. The single egg was placed on leaf-covered ground, under dense Scaevola, Scaevola-Ipomoea tangles, or Boerhavia-covered Scaevola. Here little or no direct sunlight reached the incubating adult. In 1967 four nests were under Scaevola, an average of eight feet from an open area. In 1968 the abundance center of Christmas Shearwaters shifted from the area south of the barracks to a path just north of the transmitter building. The cause of this shift was unknown but may have been cor- related with the increasing abundance of Verbesina, and consequent 130 Table CS-2. January 1-15 16-31 February T=15 16-28 March 1-15 16-31 April 1-15 16-30 May Weis) 16-31 June 1-15 16-30 July 1-15 16-31 August 1-15 16-31 September 1-15 16-30 October 1-15 16-31 November 1-15 16-30 December 1-15 16-31 * Birds present, number unknown. 1965 1966 1967 100 = 100 715 100 75 100 75 100 715 100 * 50 - 50 - 25 = * Ss O = 1968 POBSP semi-monthly estimates of Christmas Shearwaters on Green Island, Kure Atoll, 1963-69. 1964 196 ILsyal “TTOVW Sany “puelTsT user) uO STet1EegG WI0IS AQOOS pue SsTE_QemTesUS SseMASTIYO SutTpserq JO UOTANGATIYASTC J244aq W4O}S Ajoos & JOJOMIDAYS SOWISIIYD @ °T-SO aansta eye decrease in available roosting sites, where the birds were formerly abundant. By 1969, however, they were again common south of the barracks. Christmas Shearwaters roosted at the edge of Scaevola in the interior, along the beach, and in open areas along the runway. Banding and Movements One hundred and sixty-four adult Christmas Shearwaters were banded. Table CS-3 summarizes the recapture of these birds at the atoll in subsequent years. These figures suggest a high rate of turn- over on the island, especially for 1967 through 1969 when a large percentage of the population was handled. The lack of any Christmas Shearwaters banded prior to 1967 captured in 1969 is probably the result of band loss. In this species as in the preceding one, bands wear away after three years or so. Five nestling Christmas Shearwaters were banded (2 in 1963, 2 in 1964, and 1 in 1968). None was recaptured. LEACH'S STORM PETREL Oceanodroma leucorhoa On 10 May 1964 Wislocki found a Leach's Storm Petrel washed up on the beach. Another dead one was found on 21 December 1964 by Stadel. Leach's Storm Petrels occur commonly at sea in the central Pacific. SOOTY STORM PETREL Oceanodroma tristrami Status Uncertain; probably rare breeder. Few individuals present from October through mid-May, but no eggs or nestlings found. Populations The first recorded Sooty Storm Petrel for the atoll was an adult collected on 1 January 1964. Since that time POBSP observers found them irregularly in small numbers (Table SSP-1). POBSP data indicated a yearly population of perhaps 20 individuals. Earlier observers probably overlooked this species. Annual Cycle Sooty Storm Petrels arrived in late October or early November (earliest records: 31 October 1968 and 2 November 1964) and departed by mid-May (latest record: 14 May 1967). At least a few individuals were found in every month between these two dates. 133 ST dINSTJ puodss 93yd pue SAT TS 0°OOT -- -- -- (4° Te) 4°12 0°OOT -- -- (OG) > (OO) 0° S2 0° S2 0°O0T -- (OO) (©) (9°ez) 0°O O°O 9°92 0°O00T (©°O) (OPO) (aes) (hed) 0°0 0°0 ESD t° ST (O°) i. (@7@) (AG) (hg) 0°0 0°0 L°S Me G (©°O) (O°@) ~~ (©°@) ~ (O°O)) 0°0 0°O 0°0 0°0 (©°O) (©°O) (©°O) (O°o) 0°0 OLO 0°0 OKO (O°@) ~~ (O20) (O°O) (©°O) 0°0 0°O 0°0 0°O poingdecoez pue [TOVV eany 4e sqyTnpe se papueq sieyemreous seuystTazyO jo soqyer sanqdeooy useq sAey OF UMOUY SpziTq Jo S96T ©" OO ad =i) (0°0) 0°O O° OOT ee (O°@)) (0°0) On@ 0°O O° OOT (O°O) (O°©) ~~ (@°O) 0°O 0°O 0°O €96T pounjzdeosy rzeez ROOT ‘pomnqdeo spitq jo e8equeorzed oy4 °x69-6661 ‘(Se8equeozed se pesserzdxse) szeyy aseyueoied 94 sqUesemdat aInSTF YSITA x -- -- == 6€ 696T ats = = on 896T =— =e == ot LO6T ot mG 2 val 996T -- -- -- €T G96T ao = == 3 HO6T -- -- -- €96T -- -- -- iT c96T (©°O)) —- (O°) 0°O 0°O 0°OOT + 6S6T T96T O96T SOT pepueg Teak "€-SO eTqeL 134 Although no eggs or young were found, the evidence indicated that a few pairs bred. Burrowing was noted on 1 January 1964, 2 April 1964, and 5 November 1964, while storm petrels calling from burrows were heard in late December 1966, early January 1967, and 14 May 1967. In addition, 5 of the 7 (71.5 percent) collected individuals had enlarged gonads. Ecology Sooty Storm Petrels had the most restricted range of all species occurring regularly at the atoll (Fig. CS-1). The vegetation within the range was mainly Eragrostis, Boerhavia, and Solanum at the edge of Scaevola. At least 12 of the 20 adults banded were near Eragrostis. In January and May 1967 this species was heard calling from Bonin Petrel burrows about four feet long, which were located under Scaevola ca. 4 feet from the edge of the grassy area south of the barracks. On 5 November 1964 a Sooty Storm Petrel was found digging under Solanum. Banding and Movements The POBSP banded 20 Sooty Storm Petrels (11 during the 1963-64 breeding season, 6 in the 1964-65 season, 1 the following season, and 2 in 1969). None has been recaptured. RED-TAILED TROPICBIRD Phaethon rubricauda Status Abundant late winter-summer breeder; ca. 1,000 pairs annually. Recorded in every month but essentially absent from December through mid-February. Most abundant from April to August. Although nests have been found in every month, the main breeding season extends from March through November. Populations Despite their conspicuousness and abundance, Red-tailed Tropicbirds were difficult to enumerate, due mainly to the difficulty of locating all their nests under Scaevola. POBSP estimates (Table RTTB-1) are based on an extrapolation from a study area at the west end of the run- way for the number of nests present, and on the maximum number of tropicbirds counted in flight over the whole island at 1400 hours (these birds were non-breeders). In June 1967 to test the validity of the estimates approximately 60 percent of the island, mainly the northern half and sections of the southern end, was surveyed for tropicbird nests. At this time 658 nests were counted. A projection for the remainder of the island gave a total of 1,000 to 1,100 nests at the height of the breeding season; this agreed with estimates made in 1964 and 1965. There was some apparent yearly fluctuation in the gize of the breed- ing population although its exact magnitude was unknown. From 1965 135 Table SSP-1. POBSP semi-monthly estimates of Sooty Storm Petrels on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 115 - 2 O - 20 - O 16-31 - 10 @) - - - O February 1-15 - a O ) aL - O 16-28 - O 0 - - - ile March 1-15 - aL O - - - il 16-31 - O 0 * 0) O O April de) 5 3 0 = - - O 16-30 - @) © aL 0 - O May 1-15 - O O O 10 - O 16-31 - O O @) O O O June 1-15 - 0 O O @) O O 16-30 - O O O O 0 O July 1-15 - O @) O O 0) - 16-31 - O 0 O - O = August 1-15 - ©) @) O - O = 16-31 - O - @) - O = September ILI) = ) - 0 - 0) un 16-30 0) O - 0) - O = October 1-15 O @) = & = 0 o 16-31 O ) = s # By 2 November 1-15 ©) 4 - = 2 10 zs 16-30 O aL yy x 5 s December LoS) O il i - - O - 16-31 O ) = 20 - O = * Birds present, number unknown. 136 Table RTTB-1. POBSP semi-monthly estimates of Red-tailed Tropicbirds on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January ALN) - 5) Ti - 5 o/s 5 16-31 - 8 6 - - - 6 February 1-15 - 60 30 20 - wi 16-28 * 225 300 - - - M@) March 1-15 - 500 200 - - ~ 200 16-31 - 1, 000 400 - 450 IW) 1, 200 April 1-15 - 1,100 900 - - ~ 1,200 16-30 - 15 50 1,000 750 - - 1, 200 May 1-15 2, 000 1, © 1,800 1, 000 2, 000 - 1, 200 16-31 - ISS 50 2, 200 1,400 2, 000 1, 244 1, 200 June 1-15 - 1,350 2,200 2,500 2, 000 1, 295 1,000 16-30 - IL 35O 2, 200 2,200 2, 000 IL 25 * July 1-15 - 15: OO 1,600 2,200 2, 000 1, 095 - 16-31 - 1,500 1,400 2, 200 - 1,095 - August 1-15 - 1,200 1, 200 2,000 ~ * - 16-31 - 1, 000 - 2, 000 - * - September 1-15 ~ 800 - 1,000 - * ~ 16-30 * 500 - 350 = * w October 1-15 * 900 - - - * - 16-31 * 500 - - - * = November Eas) (© 150 . : 3 s = 16-30 8 a 18 - - * - December 1-15 i! 10 5 ~ z * Z 16-31 8 7 a 5 us % i *Birds present, number unknown. ILS) through 1968 the southwestern-most clump of Scaevola was checked regu- larly for tropicbird nests. In 1965 there were 97 nests; in 1966, 119; in 1967, 118; and in 1968, 109. These data suggest some minor variations, but no significant changes. Previous estimates (Table RTTB-2) suggest that Red-tailed Tropicbirds were always common at Kure, but that the population probably increased after the construction of the LORAN station which increased the available ecotonal regions that this species favored. Table RTTB-2. Previous records of Red-tailed Tropicbirds on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 100 Eggs (Munter, 1915: 137). O25. April li(=22 200 Fresh eggs to downy young (Wetmore, 7S 5 ))g 1957 June 5 1, 000 Nesting (Kenyon and Rice, 1958: 189). 1959 Octover 5-8 50 Nesting (Robbins, 1966: 53). 1960 March 28 100 Nesting (Robbins, 1966: 53). 1961 January 19-21 © (Robbins, 1966: 53). September 12-14 ? Downy young to almost fledged young (Udvardy and Warner, 1964). 1962 February 2-4 25 (Robbins, 1966: 53). August 6-8 250 25 nests (Robbins, 1966: 53). 1963 February 3-7 25 l-nest (Robbins, 1966: 53). Annual Cycle At least a few Red-tailed Tropicbirds were recorded in every month. However, they were essentially absent from December through mid-February. Figure RTTB-1 shows the average number of tropicbirds counted flying over the island at 1400 hours during POBSP studies. Although most of these birds were non-breeders, this Figure best indicates the annual population cycle at Kure. The bimodal peak in summer may have been caused by inadequate data, by an influx of birds that had not bred previously, by individuals that lost nests earlier, or by a combination of these factors. 138 350 300 Pp) pe) =) on fan) So — on fa) Number of Birds 100 90 J F M A M J J A S 0 N D Figure RTTB-l. Annual population cycle of Red-tailed Tropicbirds flying over Green Island, Kure Atoll, 1964-69 (based on averages of counts made at 1400 hours). Although Red-tailed Tropicbird nests were found in every month, the main breeding season extended from March through November. POBSP data indicated that breeding outside this period was rare. Due to the high rate of nest loss and renesting (e.g., in 1964, 46 of 62 [74.3 percent] of first breeding attempts in a study plot failed, and 54.3 percent of these pairs renested), it was difficult to reconstruct accurately a breeding cycle for this species; thus the 1967 and 1968 cycles, and to a lesser degree the 1966 cycle, were less accurately known than the 1964 and 1965 ones which were observed from initiation. Figure RTTB-2 shows the estimated number of nests present during the 1964 to 1966 cycles. The available data (Table RTTB-3) indicate that there was some yearly variation in the timing of peak egg laying, perhaps as much as two weeks. Figure RITB-3 shows the number of eggs laid each semi- monthly period for four years in a study area at the southwest end of the runway. Peak egg laying occurred earlier in 1964 than in 1965 and extended later in 1966 than in 1965. 1S) 1964 400 300 200 eggs 100 1965 800 600 400 200 1966 1000 eggs 500 Tio Me Tenia ho eos = Oo Ne D Figure RTTB-2. Breeding cycles of Red-tailed Tropicbirds on Green Island, Kure Atoll, 1964-66. Ge g-AeW-ptw 4seeT 4V Kew ATQeqorg sune AT Tea 4SB9T 7e-YoTeW ~prw 4seeT 4V 96T "69-196T ‘TTOVV eany 140 TaquaAoN -ATne 34e7T oun a4eT pue Kew 3127 ysn3ny ysnoiryy 1SeeT 42 -Keyq AT Ie9 ¥SB9T 4V aune pue AeW o1e7 Atnre -ptu 4seeT 4e -YOTeW a1eT SBT 4V ‘pue[sI uaety uo aTofd Butpaezq partqotdoay, POTTe4-pey oud UT £-9UN ?-PTN Ce 3-AeW AT Ia 4SP9T 4V oune -PpTw 4seoeT 72 09 AoW sune ATIe9 qSeeT 72 -yorey, ATree 4SB9T 4V 4-3Sn -Sny AT1ea 4SB9T 4V ATnL -ouner 34e7 ZTaquey -dag ATIe09 1SeOT 12 -Trady 94eT 4ySeeaT 4V sun c= AeW ysn3ny ~ptu 4seoT 4e -yoren ATIe9 7SeeT 4V Taq -waoeq ATIe9e -ATNP-ptW oun ysnsny -PpTwW yseeT 42 -[tady AT seq ounL- pTu-AeW asn3sny AT are9 qseeT 7e-Are —niqeaq 32487 TOQqUeAON 33eT -ysnsny AT req ysnany Aqree pure aune AT Aze9e -LeW 374e7T ysn3sny 34eT YSBeT 7e-TTady-PtW aunt ut auo ATepuo -oes e@ f{[tady JO S¥aom OMY FSATH ATNL -pTtw 4seeT 4e -yorewW ATIeq SuUTSpeTA SUT YO eH weed BUFO eH Sut sey S3q yeod Sutkey 33q 990T 96T HOOT spotzed zoey potdied °C-GLLE eT qeL 30 20 10 1964 1965 1967 J F Ialerbege)) ISUMIMS IS We NO DATA NO DATA oer eS On ON OD Number of Red-tailed Tropicbird nests in which first egg was laid each semi-monthly period in study area at west end of runway on Green Island, Kure Atoll, 1964-67. 142 Normally the first eggs of the new cycle were laid in late February or early March (earliest records: 20 February 1965 and 5 March 1964). Some eggs, usually from renestings, were laid as late as August. Com- bined data for the study area showed a peak of egg laying in May and June. By early April some of the eggs began hatching, 40 to 46 days (x=42.4, n=110) after oviposition. The last eggs hatched in early Sep- tember. The period of peak hatching varied yearly from late May to early August. Combined data from the study area showed a peak from mid-June to early August. Four POBSP records fall outside the limits outlined above: (1) an egg found on 2 November 1964, (2) a nestling of unknown age reported in early March 1964, (3) a nestling which fledged in early April 1965, indicating that the egg was laid in late November and hatched by mid-January, and (4) a nestling less than 10 days old found on 17 January 1966, indicating the egg was laid in late November. Young tropicbirds began to fly 78 to 105 days (x=85.4, n=38) after hatching. Thus fledging began in mid-June and continued through Novem- ber. After fledging the young tropicbirds flew out to sea. The earliest record the POBSP has of a young tropicbird returning to Kure is three years; it was banded as a chick in October 1963 and found breeding in September 1966. If Red-tailed Tropicbirds are similar to other seabirds, however, young return to the island at least one year prior to first breeding. Also, tropicbirds with heavy speckling on the dorsum, indicating that they were subadults, were seen in November 1964. Unpublished POBSP data indicate that this species attains adult plumage about two years after hatching. Thus Red-tailed Tropicbirds probably first return to the atoll when less than two years old. Nesting Success Table RTTB-4 summarizes Red-tailed Tropicbird productivity at Kure Atoll. Detailed succsss data were collected in a study area of ca. 0.58 hectares at the southwest end of the runway from 1964 through 1966. The results of the first two years are being reported by Fleet (1972) and are briefly summarized in Table RTTB-5. In 1966, 119 pairs bred in the study area. Of these, 30 (25.2 percent) fledged young on the first attempt and 2 (1.7 percent) on the second. Fourteen pairs (11.8 percent) renested after the first nest failed. Palynesian rats were the major cause’ of nest loss in all years. 143 Table RTTB-4. Productivity of Red-tailed Tropicbirds on Green Island, Kure Atoll, 1964-68. Maximum Maximum Maximum Estimated Breeding Nest Egg Nestling Number of Season Estimatet Estimate Estimate Young Fledged 1964 700 350 500 450 1965 1,,000 825 335 350 1966 IL I5O 1, 000 500 320 1967 1,305 1230 200° 2 1968 94.0 750 400 300 1 Maximum estimate for any semi-monthly period. 2 Minimum figure. Table RTTB-5. Results of Red-tailed Tropicbird breeding in a study area on Green Island, Kure Atoll, 1964-65. 1964 1965 First nesting: Number of pairs 62 97 Number of eggs lost 21 34 Percent of eggs hatched 66.1 64.9 Number of young lost 25 43 Percent of young lost 61.0 69.4 Number of young fledged 16 19 Percent of survival (yg fledged/eggs laid x 100) 25.8 19.6 Number of nest results unknown - il Percent of nest results unknown - A @ Renesting: Number of pairs renesting 25 12 Percent of pairs renesting (renesting/ failures x 100) G8 IS 36 Number of eggs lost tt 3 Percent of eggs hatched 76.0 Mma Number of young lost 2 5) Percent of young lost LOSS 100 Number of young fledged ale/ O Percent of survival 68.0 OO Number of nest results unknown - mt Percent of nest results unknown - Beos 144 Table RTTB-5. (continued) 1964 1965 Total young fledged 33 19 Total success (percent of total pairs) Doe 19.6 Total success (percent of total nestings) Svan) on eeedlapglt * Fleet (1972). Ecology Perhaps more than any other species, Red-tailed Tropicbirds were associated with the air space over the island. Their conspicuous aerial displays and vocalization were in marked contrast to their relatively secretive breeding habits. During their peak of diurnal activity at midday they were most abundant at the west end of the runway and over the southwest quarter of the island, less abundant but still common along the northwest beach, and scarce over the rest of the island. Presumably this abundance pattern reflected the distribution of nests. Red-tailed Tropicbirds bred mainly under Scaevola, and secondarily under Tournefortia. Due to their awkwardness on the ground, their breed- ing sites were restricted to areas where they could crawl under the vegetation to find an open area from which to take off. Consequently, most nests were within 10 feet of the beach=-Scaevola ecotone or around the edges of isolated patches of vegetation. The adults dug a scrape in the soft sand, shaping it with their bodies. Here a single egg was laid. The distance between nests was determined by the availability of nest sites and the distance the incubat- ing bird could reach with its bill. In 1968 detailed measurements were made of tropicbird nests in Scaevola clumps near the north point and along the lagoon beach. The former area measured ca. 864 inches by 720 inches -and contained 2 nests, or 1 per 51,480 square inches. These nests averaged 66.4 inches from their nearest neighbors and 76.6 inches from the edge of the vegetation. Comparable figures for the latter area were 1 per 22,252 square inches, 62.1 inches, and 110.8 inches. Non-breeding tropicbirds were found either flying over the island or roosting under Scaevola in potential breeding areas. Red-tailed Tropicbirds were frequently harassed by Great Frigatebirds until they disgorged their food. Banding and Movements Robbins banded 25 adult Red-tailed Tropicbirds and the POBSP banded an additional 1,973 adults. Recapture rates (Tables RTTB-6 and 7) of 145 ST aanStj puooses ay} pue SATTe Useq eABY OF UMOUY Spatq JO 0°OOT -- -- (1°9) 19 0°O00T -- (O°O) — (©°O)) 0°0 0°0 0°O0T (g°h) (O20) ~~ (O°S) Go Ub Ca LOU (S°G) —(©°O).- ~ (©°O) GS G°S GS (em) —(©©) (SO) cma 8° t 9°S (0°) (O'@) (S72) 0°0 0°O Ga? O°OOT °x69-656T ‘°( ‘pomngdeo spatq jo esequeored ayy aseyueorzed ay} squesertder ornsTy 4SATH x — = -- -- -- -- oe +6 696T we oe — -- -- -- -- GT Q96T ae — zt = -- -- -- Tal LO6T == == — -- -- -- -- LEZ? 996T 0°OOL -- -- -- -- ae == EG S96T (9°T2) Ory - O00 = == =F = 5 H06 967 Chew) (O° Gg) O's 9° Ge ©°OOT -- -- -- -- O2ZT C96T (o°@) (O°@) — (O°O) an Gt Sy oraor -- == ce 2961 (C26) (S° SS) (O°) (WO) (OO) (@*O) OFO eo SS Cece C° ee Sao cece O°OOT € 6S6T G96T +96T pornydeoay re98z OG popuegd Teak gosequeorted se pessetdxe) e7euy pomnydeoaz pue TI01W eany 7e Sdepesig-uou se papueq spitqoTdoz], peTTeq-pod FINpe Jo se jes ernqydeosy = 9-GLI STAPL *pounqdes spitq jo eSequeorzed ay} ST eanStz puooes oay4 pue oATTS Useq aAeY OF UMOUY SPATQ Jo ase qusozed 9y4 sqyuesormdexr amMSTI Isai » 0°OOT == -- rains -- -- 6LT 696T (0°0) 0°0 0°OOT =e == =- -- Ik R96T (29) = (O20) TE) TES) O° OOT -- -- -- Ec L96T (Seb) (0x0) > (62H) G°QT G°QT Ge?) 0° OOT -- -- T9 996T (Za) 5 (O20) (nO) = 109561) Mek eh Diet), 2° On 0°O0T == 6€ S96T 0°0 O'h O't O't O22 a Tte = 0200 92T 96T 96T 896T LO6T 996T S96T HO6T pepueg pomqydeosay reaz Teak °x69-196T ‘(Sede quaozed S@ pesseidxe) eteyy, paingdeoaz pue [Toy eany 1% saepaerq se pepueq SspatqotdorL peTte}-pey yimpe fo sejyer eanqgdeooy *)-qLIY eTqeL 146 Way these birds were low due in part to the small number of tropicbirds handled in 1967 and 1968 (Table RTTB-8). An adult banded on 2 April 1964 flew aboard a ship at 4°40'N, 154°20'W on 30 November 1967. Eight hundred and five nestling Red-tailed Tropicbirds were banded at Kure: 6 in 1959, 31 in 1963, 346 in 1964, 103 in 1965, 83: in 1966, 189 in 1967, 41 in 1968, and 6 in 1969. One of the 1963 cohorts, 5 of the 1964 cohorts, and 1 of the 1965 cohorts were re- captured in 1969. All but one were breeding. The 1963 banded Red- tailed Tropicbird also bred in 1966. A nestling Red-tailed Tropicbird banded on 14 September 1964 was found breeding on Tern Island, French Frigate Shoals, on 8 June 1969. Another two of the 1964 cohorts were recovered at sea: 1 on 13 Febru- ary 1967 at 31°39'N, 123°22'W and the other on 28 April 1970 at 38°30'N, 178°30'W. These recoveries and the one adult record indicate an easterly post-breeding dispersal of tropicbirds from the atoll. Three Red-tailed Tropicbirds banded on other islands were re- captured at Kure: an adult banded at Midway Atoll on 29 July 1962 was found breeding on 21 September 1966, another adult banded at the same location on 25 June 1963 was found breeding on 20 July 1966, and an orange-streamered adult from Johnston Atoll was seen, but not caught on 8 August 1968. Table RTTB-8. Number of Red-tailed Tropicbirds handled on Green Island, Kure Atoll, 1963-69. 1963 1964, 1965 1966 1967 1968 1969 Breeding adults 0 140 184 225 Ful Te Ou7 Non-breeding adults 120 922 213 293 15 15 116 Totals OR OCe 397 518 66 22 363 WHITE-TAILED TROPICBIRD Phaethon lepturus Kepler observed a White-tailed Tropicbird as it flew over the southeast beach on 6 February 1965. This species breeds in the Main Hawaiian Islands (AOU, 1957) and has been reported breeding on Midway Atoll (Bailey, 1956). 148 BLUE-FACED BOOBY Sula dactylatra Status Common winter-summer breeder; ca. 54 pairs annually. Maximum yearly population ca. 160. Present all year. Breeds from late January through November. Populations Evidently the Blue-faced Booby population size decreased after the construction of the LORAN station as estimates made prior to 1959 (Table BFB-1) were larger than recent POBSP ones (Table BFB-2). To what degree it decreased is difficult to determine, however, since neither Munter nor Wetmore counted adults or nests. Kenyon and Rice's count of 80 nests in June 1957 was 10 higher than the maximum POBSP one. Undoubtedly the construction of the station and consequent dis- turbance of the breeding area played a significant role in this decrease. Since POBSP studies began in 1963 the breeding population has fluctuated from a maximum of 140 in 1964 to a minimum of 78 in 1966. Enough data are available to examine some of the possible causes of these fluctuations, especially the 35 percent decrease from 1965 to 1966. One hundred and forty-five adult Blue-faced Boobies which bred in 1964 and/or 1965 had been banded prior to 1964. In 1966, 28 of these birds (19.3 percent) disappeared from the population, presumably as a result of mortality. Twenty-five adults that bred in 1965 did not breed in 1966, 16 individuals reversed this pattern, and only 4 bird bred for the first time. Overall there was a net loss of 33 individual Blue- faced Boobies. Based on this, the major factor in this decline evidently was excessive adult mortality. By comparing individual breeding records for 1966 and 1967, however, a different picture develops. In 1967, 35 individuals bred that did not breed in 1966, 4 birds reversed this pattern, 10 young birds bred for the first time, and only 1 bird was known to have disappeared; thus there was an overall gain of 4O Blue-faced Boobies, which increased the size of the breeding popula- tion almost to the 1965 level. Since the breeding population returned to the 1965 level so rapidly in 1967 (from 39 pairs to 56 pairs), due mainly to individuals breeding that had bred prior to 1966, this sug- gests that other factors besides adult mortality caused the decrease in 1966. One striking feature of the 1966 breeding season was its lateness compared with other years (see Annual Cycle section). Perhaps in that year a segment of the population (those that bred in 1967 but not in 1966) reached the proper hormone level for breeding in January and February, but extrinsic environmental conditions at that time were 149 Table BFB-1. Previous records of Blue-faced Boobies on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 200 Nesting near center of island (Munter, UCN g IST ))o 1923 April 17-22 ca. 400 Common in open space at northern end of island; nesting; some had 2 eggs, others young 2 weeks old (Wetmore, ms.). 1957 June 5 170 80 nesting pairs; few nests contained 2 eggs; majority had newly hatched to nearly fledged young. Few birds in im- mature plumage (Kenyon and Rice, 1958: 189). 1959 October 3-8 85 Adults (Robbins, 1966: 53). 1960 March 28 110 55 nesting pairs (Robbins, 1966: 53). 1961 January 19-21 80 No nesting; adult count (Robbins, 1966: 53))c September 12-14 numerous Young from downy nestlings to almost fledged young (Udvardy and Warner, HOGI. *2))): 1962 February 2-4 80 3 nesting pairs (Robbins, 1966: 53). August 6-8 100 22 nesting pairs (Robbins, 1966: 53). 1963 February 3-7 56 3 nesting pairs (Robbins, 1966; 53). unfavorable; consequently those birds did not lay eggs. When conditions again became favorable these birds had entered a refractory period and were unable to produce the necessary reproductive hormones; thus fewer birds than normal bred. Apparently this decline in 1966 was caused in part by high adult mortality, and possibly in part by the internal physiology of the birds combined with as yet undetermined extrinsic environmental conditions. Since almost all Blue-faced Boobies were banded, many of them as nestlings, population studies of this species were more detailed than those of any other species on the island. It is, therefore, worthwhile to summarize the preliminary findings here. 150 Table BFB-2. POBSP semi-monthly estimates of Blue-faced Boobies on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 1-15 - W417 is2 - 80 - 120 16-31 - 140 125 * - - 120 February 1-15 - * 140 gO 110 - 120 16-28 * 165 140 - - - 125 March 1-15 - 150 150 - - - 120 16-31 - 150 140 - 140 120 130 April 1-15 - 145 124 - - - 130 16-30 - 45 127 125 - - 130 May 1-15 100 145 129 116 129 - 130 16-31 - VAS 125 p12 135 114 135 June 1-15 - V45 132 129 118 108 Th5 16-30 - 145 130 125 veh 12 ca July 1-15 - 150 130 125 - 119 - 16-31 - 150 138 125 - 120 - August 1-15 - 150 140 125 - 112 - 16-31 - 150 - 125 - 104 - September 1-15 - 150 - 125 - 110 - 16-30 * 150 - 125 - 116 - October 1-15 * 168 - - - * - 16-31 * 160 - - - * - November 1-15 275 156 = - - * - 16-30 225 168 60 - - * - December 1-15 190 173 50 - - * - 16-31 41 17 ~ 80 - x 2 * Birds present, number unknown. 15 i All available data indicated that all population growth in this species resulted from the recruitment of Blue-faced Boobies raised on Green Island. For the first two years of life these individuals wan- dered extensively and only occasionally roosted at Kure. By their third year, however, those Blue-faced Boobies that would eventually breed on the island roosted there regularly and began to pair off. Some even bred at this age, but the majority did not begin to breed until the fourth year. Limited data suggested that no individuals bred for the first time after their fifth year of life. On the average, 13.1 percent of the adults breeding one year dis- appeared the next, presumably as a result of mortality. Balancing this loss was an average annual recruitment rate of me a WETCES Mice! Orig percent of which were banded as nestlings. Every year an attempt was made to capture all breeding Blue-faced Boobies to determine the structure of the population. Tables BFB-3 and 4 summarize these data. It is evident that the Kure population was a healthy one with younger birds gradually replacing older adults which presumably died. In 1968, 25 percent of the breeding population was composed of known-age individuals. From 1967 through 1969 the size of the breeding population remained fairly stable although its structure changed. By comparing breeding records of individual Blue-faced Boobies we can determine the actual changes in these populations each year. From 1967 to 1968 there was a decrease of 2 which resulted from the following: 21 individuals present in 1967 bred in 1968 (17 for the first time), 2 birds reappeared, 1 adults that bred in 1967 were unaccounted for, and 11 bred in 1967 but not in 1968 although they were still present. Another 84 birds bred in both years. Comparable statistics for 1969 were: 17 present in 1968 bred in 1969 (9 for the first time), 12 adults that bred in 1968 were unaccounted for, and 9 that bred in 1968 were still present but not breeding; there was, therefore, a loss of 4 individuals. Another 86 birds bred in both years. Using the preceding data it is possible to predict the size of the 1970 breeding Blue-faced Booby population. In 1969 there were 103 breeding Blue-faced Boobies. An estimated thirteen of these birds will die before the 1970 breeding season and about 10 will be alive but not breed--a loss of 23 adults. To balance these anticipated losses there were, in 1969, 15 adult Blue-faced Boobies banded as nestlings that had not bred and 12 adults that had bred previously but not in 1969; thus in 1970, if all these birds breed, the breeding population could be 107. Based on 1968 and 1969 data, however, only 9 of the younger birds will breed, so the estimated population will be 101, or a slight decrease from 1969. The main reason for the recent slight yearly decreases was the low survival rate of the 1965 cohort (see Nesting Success). Also, relatively 52 Table BFB-3. Structure of the breeding Blue-faced Booby population on Green Island, Kure Atoll, 1964-69. Number nesting in: Banded as adults in: 1964 1965 1966 1967 1968 1969 1959 i 5) 2 3 2 2 1960 3 3 1 1 wl 0 1961 52 42 ek 34 25 22 1962 48 36 22 28 25 22 1963 19 14 ial 20 20 19 1964 0 3 2 4 4 4 1965 - - 1 iL 1 O 1966 - - O 2 2 i 1967 - - - O O 1 Banded as young in: (Nestl) ISB) il 1 it 1 il 1 (Imm ) 1962 1s 1 4 10* 7% Bx (Imm) 1963 0 0 0 3 9 7 (Nest1) 1964 - O O 2 10 14 (Nest1) 1965 - - O © 0 2 Not recorded: IAL 6 10 3 i 0 Totals 142 111 78 112 108 103 Table BFB-4+. Structure of the breeding Blue-faced Booby population on Green Island, Kure Atoll, 1964-69 (expressed as percentages). Percent of the total population: Banded as adults in: 1964 1965 1966 1967 1968 1969 1959 Tae) 15 2uG ean an) 1559 1960 Bre AL Ont 3 0.9 0.9 0.0 1961 36m 3720 BOR, 30.4 Bevel Le? 1962 BBR Beek 28.2 25.0 23.1 Piles 1963 13.4 1226 1d: 17.9 ORD: 18.4 1964 0.0 ena 2.6 3.6 Ban 3.9 1965 - © Li 0.9 0.9 0.0 1966 - - O TS) 1.9 KO 1967 - - - 0.0 OZ SO Banded as young in: (Nest1) _1959 Ony 0.9 Iba) 0.9 0.9 5 © (Imm ) 1962 0.7% 0.9 Bal 8.9% Glebe FE stokes (Imm ) 1963 0 O O een 8.3 6.8 (Nest1) 1964 - ) O 1.8 9.3 1326 (Nest1) 1965 - - ©) O20 OO 1.9 Not recorded: Teeth By lt 1228 2 0.9 OZ) * | banded as subadult ——— 153 few Blue-faced Boobies were raised in 1966, and apparently large numbers of them did not survive to maturity. The next few years the size of the breeding Blue-faced Booby population should either remain stable or decrease slightly. High survival rates of the 1967 to 1969 future cohorts will be necessary to increase the size of this population. Besides the decrease in the size of the breeding population there was a decrease in the total number of Kure-based Blue-faced Boobies using the island (Table BFB-5). Excluding the subadults, many of which were probably from other islands, the island population decreased from 145 in 1967, to 140 in 1968, and to 133 in 1969. Subadult Blue-faced Boobies formed an insignificant portion of the total population. Estimates included 4 from April to October 1966, 7 in May and June 1967, 3 in June 1968, 10 in July 1968, and 2 in February, 1 in April, 9 in May, and 16 in June 1969. There was an apparent in- crease of subadults in 1968 and 1969 when many, probably from other islands, were found roosting on the beaches. Most subadults were individuals that had been raised at Kure in the previous two breeding seasons. They roosted for one or two days in the breeding area and then disappeared. Occasionally a Blue-faced Booby banded on another island was found roosting on the beach but only thrice, on 5 and 12 June 1968 and 21 April 1969, was a bird banded on another island found roosting on the central plain. On the night of 16 July 1968, 6 Blue-faced Boobies were found roost- ing at the north point. This was the first time that any group which could be considered a club was found during POBSP studies. Smaller groups, composed mainly of subadults banded on other islands, were seen on 18, 2, and 25 July 1968. Since Green Island, in contrast to other islands, sel- dom had such roosting groups, its subadult population size was smaller than that generally found on other islands. Table BFB-5. Number of Blue-faced Boobies handled each year on Green Island, Kure Atoll, 1964-69. 1964 1965 1966* 1967 1968 1969 Breeding adults 131 105 68 108 Oy, 103 Non-breeding adults 29 31 el Bi 33 30 Subadults 6 @) 3 5 7 20 Interisland © O IL 2 8 al Totals 16 136 93 152 55 15 * No attempt made to catch all birds. 154 Annual Cycle Blue-faced Boobies were permanent residents on Green Island. POBSP data indicated that they were more common during the breeding season than in the fall and early winter. After most young had fledged few adults roosted during the day in the central plain, but they returned to roost: there at night. Whether any adults left the atoll Torsexvendeds periods wasenotdererminedr Subadult Blue-faced Boobies were present irregularly from May to February. There were a few records for the other months. It appeared that they were most common during the summer. The Blue-faced Booby usually bred from late January through November, with a peak of egg laying in February and March, a peak hatching period in late March and April, and a peak fledging period in late July and August. Breeding in some years, however, deviated from this general cycle. For example, in 1966 egg laying did not begin until late March, and in 1965 and 1966 there were no apparent peaks of egg laying. Table BFB-6 compares yearly breeding cycles, and Figure BFB-1l shows the number of nests present each semi-monthly period. The first eggs of each breeding season were laid on: 5 January 1964, 20 January 1965, ca. 16 March 1966, ca. 15 February 1967, ca. 29 January 1968, and ca. 21 January 1969; and the last eggs on: 29 June 1964, 13 July 1965, 5 June 1966, 1 July 1967, and 17 June 1968. Two POBSP records fall outside the above limits. On 20 October 1963 an egg was laid which disappeared by early December. On 12 November 1963 a young booby ca. three weeks old was found; presumably the egg had been laid on cated September and hatched ca. 22 October. This chick was not reported again. Peak periods of egg laying varied considerably from year to year; Figure BFB-2 shows data for the north antenna field. When the data for all years were combined there was a major peak in February and March and lesser ones in April and late May (Fig. BFB-3). Much of the egg laying in May and June was the result of renesting. On Kure, Blue-faced Boobies laid clutches of 1 or 2 eggs (Table BFB-7), with an average clutch size of 1.88. In 1966 the average dif- fered significantly from most other years due to the lateness of the breeding cycle or to an inadequate food supply. Renesting pairs had an average clutch of 1.67 egg (n=24). From 2 to 12 days (x=5.3, n=99) elapsed between the laying of the first and second eggs. Incubation began with the laying of the first egg. Eggs hatched after an incubation period of about six weeks (Table BFB-8). The first eggs took on the average of one day longer to hatch than second eggs in the clutch. From 2 to 9 days (x=4.7, n=50) elapsed between the hatching of two eggs in the same clutch. 155 1Taqo -4090 97eT 4S8eT ye-xATne AT Teg KewW SY¥9eM OM YSeT aun a4eT 4SeeT qe-YorLey Z Trady pue Arenige,s ‘ATenuee 3787 Kew 22eT qS8eeT 7e-Are -nuee Tg °eo 6961 996T LET "69-196T ‘TIO7V etny ‘pueTsI usery uo eTofo Sutpeezq Lqoog paoej-anTd ey} UT spotsed zoCeW TEquoAON YooM pac-Aqne LT (Ata) Trady-ptu -YorTey-PTW Ayne J2 °es -Yorem TT “ed (eunr AT re9a-KLey 24eT) yore TE-ST ‘Are -niqged ST-T eune /T[-Are -nuer 6g °ed I2q401900 94°T -xTNf 94eT Lew pue [Tady oune Og 4SeeT 38 -YoreW gg ° ed Trady-ptw - Yo en- PTW Aqne AT1e9e qseeT 7e-Are —nigey AT req Taqueaon-ptu -ysnsny 31e7 Aqtne AT ite pue Lew Aqn¢L QT-Ttady 2 Key ayeT ‘Ttady ayeT SyoreN a4eLT PTeL sade qSOM °9uON eune G -yoren AT rea FSBeT 4V JOquavaon “PTW 4SeeT 7e-ATNL-PTW Ajne At ree pue Acw AINE GT 4SeeT 7'e-YOTeN GT yoren a4eT pue Arenz -qo7 prelL sada SOW °ouUOoN asnsny AT1e98 -AZenuer Og TJOQqUeACN 34°eT -ATNL 3a4eT Ttady-pTw ~YoTeN- PTW ATNE 33eT -AZTenIGeA TZ yore prw -AICNIGS,T sune 62-Are -nuer ¢ °@O poyeptodsza,ul x SUTSpeTA BSUTYOV eH yead BUTYL eH sutfeT S3y yeod sutkey S3y potied °Q-Gdd eTqQel 156 1964 50 eggs 25 1965 NO DATA 1966 NO DATA 1967 NO DATA 1968 30 20 eggs 10 1969 NO DATA Figure BFB-1. Breeding cycles of Blue-faced Boobies on Green Island, Kure Atoll, 1964-69. 1966 1967 Number of Nests O 38 423 15.4 0 167 83 Wd) J F Figure BFB-2. 23.6 364 18.2 3.6 0 0 65 32 IS 3.6 18 18 #18 #+%18 36 O te) 125 150 105 150 0 200 0 200 0 0 0 0 43 217 87 261 130 0O 0 0 35.5 226 129 0 97 32 65 NO DATA tay es) OS 0 0 38 #O NO DATA 28 WW. 27.8 139 56 28 O M A M J J A S 0 N D Number of Blue-faced Booby nests in which first egg was laid each semi-monthly period in north antenna field, Green Island, Kure Atol1,1964-69 ( figures above bars indicate the percentage of total nestings that year). 158 60 50 a 40 i = Sa 20 Number of Nests J F M A M J J A S 0 N Figure BFB-3. Number of Blue-faced Booby nests in which first eggs were laid each semi-monthly period on Green Island, Kure Atoll, 1964-69 (combined data; white bars indicate re-nesting). Table BFB-7., Clutch size of Blue-faced Boobies on Green Island, ‘Kure Atoll, 1964-69. year 1-Egg 2-Egg 3-Egg x n 1964 4 53 10 93 4 1965 5 34 0 1S 39 1966 6 8 ©) IOS 14 1967 2 3 0 1.60 5 1968 3 20 0 1.86 23 1969 Th Tie) a 1.94 54 Totals oh 167 iL 1.88 192 159 Table BFB-8. Incubation periods for first and second eggs of Blue-faced Boobies on Green Island, Kure Atoll, 1964-69. Year First Egg Second Egg Range x n Range x n 1964 4O-48 days 43.9 Sif 4o-45 days 4.5 en 1965 40-47 days 43,5 22 41-47 days 42.9 20 1966 43-46 days 43.9 i 41-44 days NP) 5 4 1968 4o-4h days 43.0 4 40-43 days Lea, 3 1969 4o-45 days 43.6 27 41-45 days 42.9 22 x Lo-48 days 43.6 97 4O-L47 days 42.7 "3 The first eggs hatched on 21 February 1964, 15 March 1965, 27 April 1966, ca. 28 March 1967, 11 March 1968, and 2 March 1969, and the last eggs hatched on 31 July 1964, 15 July 1965, 16 July 1966, at least 20 June 1967, and 24 July 1968. Peak hatching periods occurred 6 weeks after the egg laying peaks. Combined data (Fig. BFB-4) show a peak in late March and April. 95 45 35 25 Number of Nests J F M A M J J A S 0 N D Figure BFB-4. Number of Blue-faced Booby nests in which first eggs hatched each semi-monthly period on Green Island, Kure Atoll, 1964-69 (combined data; white bars indicate re-nesting). 160 In 44 of 102 two-egg clutches (43.1 percent) laid from 1964 through 1966 both eggs hatched but only one young, usually the first to hatch, fledged. In 34 of the clutches (33.3 percent) one egg hatched and in the remaining 24 clutches no eggs hatched. Where both eggs hatched and the first chick was the one to survive, the second chick lived from 1 to 15 days (X=3.7, n=38) after hatching. The guard stage in this species was 29 to 84 days (%=56.9, n=65). Young boobies began flying 109 to 151 days (x=123, n=44) after hatch- ing; hence fledging began in early July and continued through November with a peak in late July and August. These young returned to the nest site one to two months after fledging and then left the atoll; there- fore, by mid-February none of the preceding breeding season's young remained on the island. Usually by early May a few of these birds again roosted on the island for short periods. Nesting Success Table BFB-9 summarizes Blue-faced Booby productivity from 1964 through 1969. In 1965 14.6 percent of the young were produced by re- nesting pairs. Comparable figures for other years were 5.9 percent in 1964, O percent in 1966, and 22.5 percent in 1968. Five young in 1965 and one in 1966 were produced from second eggs that were given to pairs that lost eggs or young. In all casés the donor pair's first egg hatched. Sufficient data were collected from 1964 through 1966 to determine the success of the two clutch sizes (Table BFB-10).and the relative contribution of first and second egg to productivity (Table BFB-11). These data showed that two-egg clutches were more successful than one- egg clutches and that second eggs contributed significantly to overall productivity, suggesting that two-egg clutches have adaptive significance at Kure. Kepler (1969) discusses this adaptation in detail. Detailed nesting success data were gathered in the north antenna field colony during 1964 and 1965, and to a lesser extent from 1966 through 1968 (Table BFB-12). It is obvious from this table that in all phases of reproduction, Blue-faced Boobies were successful breeders, especially considering the fact that although they may lay two eggs, a pair can only raise one young. Rates of survival to sexual maturity of young Blue-faced Boobies (three years of age) were also high (Table BFB-13). Ecology Blue-faced Boobies bred in two distinct colonies, one in the north and one in the south antenna fields (Fig. BFB-5). The majority, 81.5 percent in 1964, 72.8 percent in 1965, 64.0 percent in 1966, 67.9 per- cent in 1967, 64.9 percent in 1968, and 64.8 percent in 1969, occurred Kure Atoll, 1964-69. Table BFB-9. Number Year of Nests 1964 70 1965 60 1966 39 1967 56 1968 5M, Hab 1969 ‘ Produced by 6 re-nesters and 5 foster chicks. Number of Nests % of Nests Fledging Young Fledging Young 20, (ebo5 41 68.3 27° 69.2 39 69.6 312 57 4 4? 86 3 7 produced by re-nesters. 4 4 not included in total. Number remaining on 20° June, including 6 foster chicks. Table BFB-LO. Productivity of Blue-faced Boobies on Green Island, Number of Young Banded 50 38 27 35 Sul 21 = 1 foster chick. Success of first nesting attempts for two clutch sizes of the Blue-faced Booby on Green Island, Kure Atoll, 1964-66. Year 1l-Egg Clutch 2-Egg Clutch 1964. Success 2 (50.0%) 32 (65.3%) Failure 2 (50.0%) 17 (34.7%) 1965 Success O ( 0.0%) 25 (61.0%) Failure 6 (100.0%) 16 (39.0%) 1966 Success 5 (41.6%) 11 (84.6%) Failure 7 (58.4%) 2 (15.4%) Totals Success Tt (32.28%) 68 (66.0%) Failure 15 (68.2%) 35 (34.0%) Table BFB-1ll. Contribution of first and second eggs to Blue-faced 161 Booby productivity on Green Island, Kure Atoll, 1964-66. Number of Young Produced Year lst Egg end Egg 1964 29 (85.3%) 5 (14.7%) 1965 17 (68.0%) 8 (32.0%) 1966 14 (100.0%) O (0.0%) Totals 60 (82.3%) 1S} GLY 5) 1°69 jaan B° HE 1°99 Ete 0°09 €°0g 6° €€ ie re a €°06 4° tt 2°64 O° 49 Gea cert 9° LS Gly €° 4S 2, Se CO o°€9 L°€S EAS 6°09 S4SON SUT YO} eH oq peyo eH [WOT peSpeT4 Sunok jo % S33q JO % 162 "69-H96T *TTOFV eany ‘sqdweqye SuTyseu 43ITJ uo peseq pleats BuuUaejue YYIOU ey4 UT SeTqoog peoevj-entg jo AyTATZoONpord *€2 oL isk OT 61 9E pespeTa sunoxX jo Zaqunh St 0g ES fe) GE L9 peyoyeH prey s33y jo s33q jo ZTaquny TIqunyl ‘om’ Og UO BSUTUTeWAT TaquMN x Sy) 6S £9 ge ql, OLT HE GE ge S¢ Of LS SYSeN Jo TaqunN 696T 896T L96T 996T S96T 4961 *Z1-ddd eTqed 163 UMOUY SpaTq Jo esequedzed 9343 JUSeSeTdet SaInsTy ISITHW (9°62) 9°62 (9° ST) 9° ST (0° ge) 0° 9¢ (Giga) H° EE (Eo eT) Q° ST (0°g¢) O°9E (2° €2) oc? (1° §2) Sor *poingded spaitq fo esequeorzed 34} ST OINSTJ puodas ayy pue aATTe Useq sABY OF O*OOT (GO) iesoll (0°9) One (SO) Z°O€ O°OOr -- -- -- -- =< (0°2) O° th 0°O0OT -- -- -- -- (Or) = (07) O° SE 9°6E 0°OOT ae ao 2 (are Cerwe) (CS? Ue) + HE 9°On €°9¢ 0°O00T -- -- (©O°O) (©°O) —(©°@) (Oro) (€°9) (€°9) E°9 €°9 €°9 €°9 e°9 e°9 G96T 496T €96T Z96T T96T O96T pomydeooy reaz *x69-6961 ‘(seBequsoied se passerdxa) TLOVV euny SpueTsST usery uo payoyey sqzoyo%e Aqoog peoej-enTg snotszea JO soyer TeATAING *pospeTyT oaey 04 UMOUY 979M SPAITQ 98944 TIV x Lig 0S eh As OT 996T S96T 96T €96T ZI6T 6S6T 4. Z0YOO “eT-d4d OTdeL 164 ‘TTOVV emmy ‘pueTsI use IQ uo geTqoog uMorg pue pedej-enTg Sutpeezq jo u oTqnqt 44st Aqoog umoig a Aqoog parn}-anig : °C- Gad emsta 165 in the former area. In 1964 one pair bred in the open area just south of the barracks, where there used to be a few nests prior to the construction of the Coast Guard station (Robbins, personal com- munication). Nests were either clustered or solitary with the average distance from the closest neighbor 23 feet in 1964 and 25 feet in 1965 (Kepler, 1969). The range was from 5.5 feet to 89 feet. The size of the pair's territory was ca. 240 square yards, but only 60 square yards were actually utilized (Kepler, op. cit.). Unlike Brown Boobies, Blue-faced Boobies bred in the open area of the central plain rather than along its edges. For the most part the breeding areas of these congeners did not overlap, although they some- times bred as close as five feet to each other. The nest was a simple scrape on the ground made by the male with iiosmcail. the eggs were laid in a slight depression in the center of the scrape. Young Blue-faces eventually wandered from the scrape and soon Tribulus grew over the area. In 1967 a female, breeding for the first time, laid her egg on top of Tribulus. The spreading of Verbesina in the central plain has changed the distribution of nests since 1964, especially in the south antenna field. In 1968 only an area of ca. 100 x 25 yards that had no Verbesina remained in this area. Most pairs bred there, but a few bred among the plants where there was enough open area. Previously nests had been widely distributed in this field. The area just north of the transmitter building was also overgrown with Verbesina and only two pairs bred there in 1968 and 1969, although several pairs had bred there previously. As yet the plants have not invaded the major breeding area in the north antenna field. Unmated birds and pairs that were not breeding generally roosted among the breeders. In 1968, however, many non-breeders roosted in the same general area of the north antenna field, with no breeders among them. There were breeders near the periphery of this group. Eyen these birds maintained territories. Subadult Blue-faced Boobies, mainly Kure-raised birds, also oc- casionally roosted among the breeders, generally near their natal site. They also roosted on the beaches, generally at the north point. Banding and Movements Two hundred and twenty-two adult Blue-faced Boobies were banded at Kure Atoll. Table BFB-14 briefly summarizes the recapture of these birds at the atoll. Details of banding and recapture will be pubdished separately. ST oInSTJ puooses ay} pue sATTe O° OOT == (0°00T) (0°00T) OLOOw = = OHO (0°0S) (0*00T) 0°0S 0°O0OT (O2O)) {COR@One) 0°O O° OOT (Hott) tr) tt Ho th (EES) (EZ) SS EG (EZ) *CeeSi) CONS 8° St (O°9€) (O° th) O°9€ O° th (CORO): S(O OI) O2O1L O-O1 (Ore) (Org) 0°8 0°8 0° OOT (0°O0T) 0°OOT (0°OOT) 0°OOT (4° tr) ah (Grows) 0109 (6°25) G°2S (erg) E° es (0°OT) O°OT 0° OOT (0°O0OT) OL OLOR (9° SS) 1°99 (2° 8%) 0°09 CESS) Ges (L°2) EIEG (0°OT) OnO¢g (0°9T) O°9T useq O° OOT (9° SS) g° LL (6°29) Cot (Ser) €°9Od, (2°29) €° G9 (0°0€) OOS (0°02) Ot *poinides spatq jo eseqjueorzed ayy SD oS = os =a on ‘S -—=— = — — — — A —_— = —— —— —— — —— ro — — = — —=— —— = E 0°OOT == == == == = 6 Eh) 2S + 07001 vee = = == Ge (S°16) (S°T6) c° 6 6° 46 0° OOT -- == == 6S (Se27))e= (e186) (= 19) VEMOVE €°S9 0°99g O0°OOT Se oS cL (OfOE)_ (O°0t) <(O" On)» - (Or 01) O°0€ O'Or 0°0S 0°0S 0°OOT == OT (O42) \(O7gg) (0242) @lo°et) (Co-0) Ore O°Ze Orge O"9€ O°9€ O°OOT G2 aAaey O4 UMOUY Spatq jo esequaozed 9yq squesertdez samn3Sty 4SATA x 696T L96T 996T G96T 496T COQ6T Z96T T96T O96T 6S6T pepued 166 S96T pornqgdeooy ze3ex d19y} pornjydeoser pue [ToVV euny 1e pepueq seTqoog pooej-oenTg 4itTnpe jo seyer sanqdeoay HOOT °x69-6661 ‘(seSeqjueoi1ed se passeidxa) LEK “HT-dld eTael 167 Five of these adult Blue-faced Boobies were captured on other islands: 4 on Eastern Island, Midway Atoll and 1 on Laysan. Three of the birds found at Midway were breeding at the time of recapture. None had bred at Kure, nor did they return to Kure. Robbins banded one subadult Blue-faced Booby in August 1962 and the POBSP banded 14 (2 in 1967, 2 in 1968, and 10 in 1969). Only the one banded in 1962 was recaptured. It bred at the atoll in 1964, 1967, 1968 and 1969. Recaptures of nestling Blue-faced Boobies reaching sexual maturity have been summarized previously. In addition, 35 nestlings were banded in 1967, 31 in 1968, and 21 in 1969. Four of the 1967 cohorts were re- captured at Kure in 1968 and four in 1969, including one captured the previous year. In 1969 6 of the 1968 cahorts were captured. Young Blue-faced Boobies raised at Kure traveled more extensively than the adults. Eight were found on other islands; 1 on Eastern Island, Midway Atoll, 1 on Southeast Island, Pearl and Hermes Reef, 3 on Lisianski, 1 at French Frigate Shoals, and 2 on Sand Island, Johnston Atoll. Two of these birds, one from Pearl and Hermes and one from Lisianski, returned to Kure. The Blue-face traveling to Midway was especially interesting. It was banded on 6 August 1962, recaptured at Kure on 26 October 1964 and 5 December 1965, and found breeding at Midway on 22 August 1966. This is one of the few records of a Blue- faced Booby hatched on one island breeding on another island. Relatively little movement of Blue-faced Boobies from other islands to Kure Atoll was recorded: 1 from Eastern Island, Midway Atoll, 3 from Pearl and Hermes Reef, 5 from Lisianski, and 4 from French Frigate Shoals. This is not surprising since the atoll generally lacked permanent aggre- gations of non-breeders in which most interisland birds roost on other islands in the Northwest Hawaiian chain. All but two of these birds were banded as nestlings. BROWN BOOBY Sula leucogaster Status Common spring-summer breeder; ca. 40 pairs annually. Maximum year- ly population ca. 100. Present all year but most numerous during breeding season. Breeds mainly from April through October. Populations Available data (Tables BB-1 and 2) indicated that Brown Boobies were about as numerous during the 1960's as earlier in the century. Munter's estimate was larger than any POBSP estimate, but since he did not count the birds it is difficult to determine the validity of his 168 Table BB-1. Previous records of Brown Boobies on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 ca. 200 Eggs present (Munter, 1915: 137). 1923 April 17-22 100 Eggs present (Wetmore, ms.). 1957 yume, 70 27 nests counted; out of 13 examined, 2 had 1 egg, 5 had 2 eggs, 1 had 2 newly hatched chicks, and 5 had 1 chick, the largest of which were half-grown and covered with down. Few birds in imma- ture plumage (Kenyon and Rice, 1958: 189-190). 1959 October 3-8 50 Nesting (Robbins, 1966: 53). 1960 March 28 20 Nesting (Robbins, 1966: 53). 1961 January 19-21 1 Not nesting (Robbins, 1966: 53). September 12-14 2 (Udvardy, 1961: 46). 1962 February 2-4 5 Not nesting (Robbins, 1966: 53). August 6-8 LO 19 nests (Robbins, 1966: 53). 1963 February 3-7 25 2 nests (Robbins, 1966: 53). estimate. The only other previous numerical data were Kenyon and Rice's count of 27 nests in early July 1957. POBSP counts at the same time of year were similar, indicating little change in a decade. Since this species bred mainly along the edges of the central plain, its breeding habitat was not greatly altered by the construction of the LORAN station. During POBSP studies an attempt was made each year to handle all Brown Boobies at the atoll. These data (Table BB-3) indicated a total yearly population of ca. 100 individuals, with no more than 84. breeding in any one year. Although the size of the breeding population remained relatively stable at ca. 4O pairs during POBSP studies, its structure, determined from the recapture of previously banded birds, changed (Tables BB-4 and 5). Like the Blue-faced Booby, the Brown Booby population maintained its stability through an influx of young birds and a decrease of older adults, a sign of a healthy population. From 1967 through 1969 there was an average annual adult mortality rate of 18.7 percent and an average recruitment rate of 1/7. percent. 169 Table BB-2. POBSP semi-monthly estimates of Brown Boobies on Green Tsland, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 196 January 1-15 - 25 75 - 85 - 100 16-31 - 115) fe * - - 100 February 1-15 - * 75 2 8 - 100 16-28 * 25 1D - - - 60 March Loi) = 25) 50 - = = 50 16-31 - 35 50 - 25 30 60 April 1-15 = 35 46 = = 2 75 16-30 - 63 45 30 - - 75 May 1-15 15 63 hg Te) 43 - 80 16-31 - 87 50 yy 62 48 90 June 1-15 - 87 70 65 66 4g 100 16-30 - 87 80 75 74 46 * July 1-15 - (5 100 715 - 62 - 16-31 - 95 100 90 - 69 - August 1-15 - 95 90 100 - 64. - 16-31 - 95 = (2 - 62 = September 1-15 - 80 =- vals - 64 - 16-30 * 65 - 70 = 75 és October 1-15 * 60 - = S % zs 16-31 * 15 - - = * ss November 1-15 150 64 = 2 = * aa 16-30 150 64 4s = = * 3 December 1-15 88 55 4O - = * : 16-31 65 50 - 85 : x 2 * Birds present, number unknown. 170 Table BB-3. Number of Brown Boobies handled each year on Green Island, Kure Atoll, 1964-69. 1964 1965 1966 1967 1968 1969%* Breeding adults 52 52 70 69 68 60 Non-breeding adults 25 35 13 alia i 2X0) Tk Subadults 5 il 5 4 6 3 From other islands O O 3 @ O O Totals 82 88 gl 8 9 i * No attempt was made to catch all non-breeding birds. Table BB-4+. Structure of the breeding Brown Booby population on Green Island, Kure Atoll, 1966-69. Number breeding in: Banded as adults in: 1966 1967 1968 1969 1959 2 2 ib ik 1962 12 11 rl 6 1963 15 15 13 10 1964 23 20 16 14. 1965 le 8 10 6 1966 il 5 5 4 1967 - 4 4 3% 1968 - - 4 4 Banded as young in: 1957 1** 1** 1** 1** 1959 dh O 1 O 1962 dl al ale 3} 1963 ib Z 4 3) 1964 @) 0 3 4 1965 0 O O¥x* ib Not recorded: 12 ff 8 14 Totals 82 "i 80 MG * One from Pearl and Hermes Reef. ** From Midway Atoll. **X* One from Howland Island. IW Table BB-5. Structure of the breeding Brown Booby population on Green Island, Kure Atoll, 1966-69 (expressed as percentages). Percent of total population breeding in: Banded as adults in: 1966 1967 1968 1969 1959 2.4 2.6 Iks8} iL 4/8 1962 14.6 Wh 55 8.8 Soak 1963 US 53 1G) oH 16.8 13.45 1964 28.0 26).3 20.0 18.9 1965 O65 10.5 ILLS Gr 1966 A5) 6.6 6.3 5 4 1967 - 553) 550 41% 1968 - - 50 54 Banded as young in: 1957 1.O%* 1.3** hig eee 1.4%** 1959 2 Ono Tes 0.0 1962 1,2 1.63 These) OAL 1963 Lok 2.6 50 ML 1964 0.0 O20 Boe) 54 1965 0.0 0.0 2. 5*** Lott Not recorded: 14.6 9.2 ORO 18.9 * One from Pearl and Hermes Reef. *x From Midway Atoll. *** One from Howland Island. Unlike the case of the Blue-faced Booby, some of the Brown Booby population growth was the result of an influx of birds from other islands. Also, there was no indication that young Brown Boobies began roosting in the central plain one year prior to breeding for the first time. The number of non-breeding birds present one year does not indicate how many newapceedens therci wills bemthemMextuyecar: its Thus dmpossibille: sto pre= dict the size of future breeding populations on the island. Subadult Brown Boobies formed an insignificant proportion of the total Brown Booby population. During the year a few were present every month. For example, in May and June MOGI. 1 to 3 were found each census night, whike in 1968, from late May until early August, 1 to 3 were present on 3 of 10 census nights, and only 6 different individuals were captured during the period. Generally, these birds had been raised on Green Island in previous years. (2 Annual Cycle Although Brown Boobies roosting in the central plain were easily enumerated, many individuals, especially during the non-breeding season, did not roost there. Therefore, counts included only a portion of the population and were less accurate than the Blue-faced Booby censuses. Consequently the Brown Booby annual population cycle was not accurately determined. However, POBSP data suggested the following cycle. Brown Boobies were present all year, with maximum populations during the breeding season. After breeding most adults left the central plain and roosted in Scaevola, on the reef, or on the beaches. At this time they were relatively inconspicuous. Some adults may actually have left the atoll. As breeding progressed, more and more birds roosted in the central plain. Brown Boobies bred in all months, but the main breeding season was from April through October. Figure BB-l shows the number of nests with eggs and the number with young for each semi-monthly period when POBSP personnel were present. Unlike Blue-faced Boobies, Brown Boobies showed little yearly variation in the breeding cycle (Fig. BB-2 and Table BB-6). Only in 1968, when breeding began in February and no egg laying peak oc- curred, was any significant variation noted. Eggs were laid in all months except December, but most egg laying occurred from April through July. Earliest egg dates for each breeding season were 1 April 1964, late March 1965, 14 April 1966, ca. 19 March 1967, ca. 8 February 1968, and ca. 10 April 1969. Combined data show a peak period of egg laying in May and June (Fig. BB-3). Brown Boobies laid a clutch of one to three eggs, but the usual clutch was two (Table BB-7). Three to nine days (x=5.2; n=85) elapsed between the laying of the first and second eggs. Since incubation began before the second egg was laid, hatching was asynchronous. On Kure the interval between hatching of eggs in the same clutch averaged 4.6 days (range 1-9 days, n=52), slightly less than the average interval between egg laying. Incubation periods for the first and second eggs laid did not differ significantly (Table BB-8). No third egg hatched. In the study years the first egg hatched on: 18 May 1964, early May 1965, 25 May 1966, ca. 30 April 1967, 21 March 1968, and 20 May 1969. Some eggs hatched in January, October, and December, but no young fledged from the three nests observed. Combined data show that the peak period of hatching occurred in June and July (Fig. BB-4). Although both eggs may hatch, only one of the young usually survived. In 1964, for example, in 18 of 23 (78.1 percent) two-egg clutches, both eggs hatched, but in only one case, where the eggs hatched one day apart, did both young fledge. Only one other nest produced two young: in 1965, Lys 1964 35 25 15 =bes 1965 NO DATA 1966 NO DATA 1967 25 15 NO DATA 1968 25 15 young 5 eggs 1969 J F M A M J J A S 0 N D Figure BB-1. Breeding cycles of Brown Boobies on Green Island, Kure Atoll, 1964-69. 174 Nests —_ te) O70) (0) 10:0) 20.0) 25015 25:085'010!0)) 2152/5 0) 0) 0) 0) 40) vOy 0) 9.310), 20'2.453'08 16.9) 11910) 2 310 20 NO DATA ODN2:9 35:9 00:1 92718 OleOs 3) a rdirG.2 ° oO °o oO °o oa on of Number 10 0 oO 0 0 QO 10.3. 10:3) 3:5 22:7 241 13:8 10:3 NO DATA 10 0 36 O 36 10.7 36 7.1 107 10.7 143 143 36 36 36 10.7 J F M A M J J A S 0 N D Figure BB-2. Number of Brown Booby nests in which first egg was laid each semi-monthly period in north antenna field, Green Island, Kure Atoll, 1964-68( figures above bars indicate the percentage of total nestings that year). LS Ge Ge g-AeW 24 eT Ke -ptu yseeT 4e -[tady 94e7 aunt AT Tee 4seoT 4e -Trady AT sreq 696T "69-796T “TIOVV eany SpueTsI weer) uo aTofo 6961 Arenuelr OT-ATNL 6T qysnsny-pTw -ounf-prtu peyoyey saso qSOM ° oUON 840490 6 -Yorey TZ °ed oune pue Lew UT pTeL Sado SOW ° dUON qsnsny Qog-Are -niqed g *8o 890 ZJIqueaAoN-ptw -xfTnf 24eT ATnf-ptu -ounP-PTtW ysnsny AjTre98 yseaT 42 -TTady o€ °Bo eK ouUne a4eT 3SBeT 42 -yorey 6T *eO LO6L ZJequaedsaq ayetT-ATnr 92 AqTar 13qo71900 2, “eo-Ken Sz Kew syoom OM} 4SeT qsnsny YZ -THady 41 9967 z3q -wooeq AT1e9 -ysnsny 9 ATNL-ptu -oUnf-PTW ZJequeides 6 “eo-Lew AT Ieq Kew Aqne 62 -YoreW 9427 S96T Sutpeerzq Aqoog umMorg 9y4 po yerTodze4ul x TJOQqUaAOCN 33°eT -ySnNsnY 0427 ATNL-suNnL asn3sny eyeT-ALeW QT Kew 248T -T tady-PtW ASTHAWO TE -[tady [T °eo HOOT ur spotzed soley SUTSpOTA Sut yor eH yeed BUT Yo eH sutf£eT S3qy yeod Sutkel 334 potzed “Q-da eTASL 176 65 99 45 30 Number of Nests 25 15 J F M A M J J A S 0 N Figure BB-3. Number of Brown Booby nests in which first eggs were laid each semi-monthly period on Green Island, Kure Atoll, 1964-68 (combined data; white bars indicate re-nesting). Table BB-/. Clutch size of Brown Boobies on Green Island, Kure Atoll, 1964-68. | Year 1 Egg 2 Eggs 3 Eggs xX n 1964. 2 eo 3 2.03 31 1966 in 22 3 1.97 29 1967 O 16 2 Daa 18 1968 1 14 O 1.93 15 Total 7 (7.5%) 78 (83.9%) 8 (8.6%)- 2.01 93 IT: Table BB-3. Incubation periods for first and second eggs of Brown Boobies on Green Island, Kure Atoll, 1964-68. Year First Egg Second Egg Range 5 n Range x n 1964 hO-46 days 42.3 23 ho-44 days 42.2 22 1965 41-45 days 4.1 10 No data O 1966 39-48 days 42.6 18 4Wi-bh days 42.4 12 1967 ho-45 days 42.3 3 hi-42 days 41.5 2 1968 42-43 days 42.5 8 Ho-44 days 41.9 8 x 39-48 days 42.4 62 Ho-44 days 42.2 yy 40 2 30 azo E = 10 J F M A M J J A S 0 N D Figure BB-4. Number of Brown Booby nests in which first eggs hatched each semi-monthly period on Green Island, Kure Atoll, 1964-68 (combined data; white bars indicate re-nesting). one egg hatched on 9 July and the other one on 12 July; both nestlings were still alive and healthy on 15 August. With these exceptions, the life span of the second chick on Kure ranged from one to seven days (eh oe oe). Much of the egg laying in the latter part of the season was the result of re-nesting. Table BB-9 summarizes re-nesting data for 1964, 1966, and 1968. 178 Table BB-9. Re-nestiag of Brown Boobies on Green Island, Kure Atoll. One Egg Two kggs 1964 1966 1968 Total 1964 1966 1968 Total Number of nests lost 6 4 5 15 4 3 im 73) 10 Number of re-nestings 3 2 2 / O it 3 4 % of re-- nestings SOsO2 SOOM eLORO 46.7 O B85 Sut LOOKO 40.0 Three Eggs Nestling 1964 1966 1968 Total 1964 1966 1968 Total Number of nests lost ae 2 O 3 2 O 2 4 Number of re-nestings O a O 2 O O 2 } % of re-- nestings O 100.0 0 66.7 ) 0 100.0 50.0 Combined 1964 1966 1968 _Total Number of nests lost 13} 9 10 32 Number of re-nestings 3 5 i IB) % of re-- nestings eee RS OO 46.9 Young Brown Boobies fledged 85 to 103 days (x=94.7; n=30) after hatching. Thus, in five seasons of observation fledging began as early as late July and continued into December, with a peak usually occurring in September-October. Fledglings returned to their nest sites for one or two months after fledging, but all those that survived eventually left the atoll. POBSP data indicate that most Brown Boobies were absent from the atoll for their first two years of life and returned sometime during their third or fourth year. Some three-year-old birds bred. ILS) Nesting Success Tables BB-10 to 12 present Brown Booby productivity data for 1964 to 1968. The most detailed data were collected in 1964 and 1966 in the north antenna field when all nests were checked daily. As can be seen, Brown Boobies were relatively successful breeders and ca. 30 young were produced annually. Enough data were collected in 1964 and 1966 to determine the relative contribution of first and second eggs to productivity (Table BB-13), the relative contribution of re-nesting pairs (Table BB-14), and the nesting success of the various clutch sizes (Table BB-15). These data indicate that two-egg clutches were adaptive on Kure as they produced more young than any other clutch size, and the second egg laid contributed significantly to overall productivity. Although the nesting success of Brown Boobies was high, survival to sexual maturity (three years), or at least the rate of return to the atoll, was low (Table BB-16). This may be related to the long distances young Brown Boobies traveled after fledging (see Banding and Movements). Their destinations were often inhabited areas where they were probably sub- jected to heavy human predation. Ecology Brown Boobies bred in small groups or, rarely, in individual pairs on the central plain and adjacent grassy areas. Most nests were found in the north antenna field (43 in 1964, 34 in 1965, 31 in 1966, and 29 in 1967, 1968 and 1969), rather than in the south antenna field (5 in IGS, Ii Way MSHS MO Ga MSS, Cydia ISHS S) aia MC AS4 eral Cy ha MC Se) a ane) the former area nests were placed along the edge of Scaevola at the western side of the field, resulting in a rather linear distribution extending ca. 100 yards in a northeast to southwest direction (Fig. BB-5). In the latter area nests were confined to an area within 50 feet of an isolated Scaevola clump at the southern end of the field. Occasional nests were present on Tribulus-covered dunes just north of the north an- tenna field. Only rarely was a Brown Booby nest nearer than 5 feet toa Blue-faced Booby nest. The nests, generally substantial structures of Scaevola twigs and leaves with some Solanum, Tribulus, Verbesina, Eragrostis, and Boerhavia, were built on the ground. Some nests contained very little vegetation, with the eggs almost on the ground, while others were 3 inches deep. A sample of 10 nests had an average outer diameter of 12.7 inches, an average inner diameter of 7.1 inches, and an average depth of 1.4 inches. The central portion was depressed from the weight of the incubating bird. After the eggs hatched, most of the nest was destroyed by the movements of nestlings and adults. Tribulus and Scaevola surrounded many of the nests. 180 *“ATne AT Tee UT Sututewer rTequmN *popnTout you mog oc i "2TQe4 STU} UT UOTSNTOUT JOoZ aqyenbepeuy e4yep 696T x G° 19 I @) Js g Q96T 0°08 t 0 at 15 L96T 0°0g g 0 Qg OT 996T 0°O0OT wl 0 TEE TL G96T 0° OOT S O S S 496T ssooong poeonpordg 9dw9837V 10wW311V SYSON jo Teoh qUus018g ZTaquny puoosg uo Sunoz YSITWq uo Bunoz ZTaqunyt Teq10L SUTSpeTW SYSeNn SUTSpeTA SIsSeNn jo Zaquny jo zZaquny "¥89- 196T ‘TTOVV eany *pueTs]I usery “‘pTety euuezUe Y4NOS 9344 UT SeTqoog umMorg Jo AITATYONpOIg “TT-dd eTaqeL ‘tne ATIee UT BSutTutTewer Tequny *papnTout Jou udsAeTY S uf *aInSTJ umMWTUIW ¢ ‘4S9uU owes WOTT peSpeTy BSunofk omy, 3 ‘papntTout ou 3auC : "aTqey STYI UT UOTSNTOUT ZO aVenbepeut e yep 696T x 4°65 €°19 SHE 1° 9S 6T ctf 9S AS Q96T 0°OOT O° SL 9° St 6° #9 ack cte cle 48t L96T tad, WOE O'Or O°SS 42 Ee 09 Tle 996T Trek USGL: 6° St 6°9S ALS ce 69S Tee G96T Omer! O° +S O'gE 4° OL ald OS ale, EOS +96T 4S9N uTYoY eH a peyoyeH pespeTd peyoyeH PTeT sqseN Jo eax >woTW pespeTW Sunox Jo % SS3q SunoxX jo ss3q jo SS3q jo Zaqunn JO % Taqunn Taqunn Taquny "¥99-+96T ‘sq7dueq7e SUT}SOU JSAITJ UO paseq PLTeTJ euUusZUe YIAOU 944 UT SaeTqoog uMoug fo AZTATJONPOTG ‘“OT-dd eTqAeL 181 Table BB-l2. Productivity of Brown Boobies on Green Island, Kure Atoll, 1964-68*. Number Number of Nests % of Nests Number of Year of Nests Fledging Young Fledging Young Young Banded 1964 kot 21 Th 5 29 1965 aut 3h cd 23 1966 41 34 82.9 34 1967 232 22° 95.5 17 1968 hO 29 TAS 29 * 1969 data inadequate for inclusion in this table. 1 one not included. e Fifteen not included. 3 Number remaining Wal Senolke awllyy7g Table BB-13. Contribution of first and second Brown Booby eggs to productivity, Green Island, Kure Atoll. Year lst egg end egg 1964 21 (80.8%) 5 (19.2%) 1966 20 (86.9%) 3) (ssc) Totals ki (83.7%) 8 (16.3%) Table BB-l14. Contribution of re-nesting Brown Boobies to productivity, Green Island, Kure Atoll. Year First attempt second attempt 1964 26 (96.3%) 1 (Soa) 1966 23 (92.0%) 2 (8.0%) Totals ko (9h 2%) 3 (5.8%) Table BB-15. Success of first nesting attempts for various clutches of Brown Boobies on Green Island, Kure Atoll. Year l-Egg Clutch e-Egg Clutch 3-Egg Clutch 1964 Success 1 (25.0%) 23 (82.1%) 1 (33.3%) Failure 37 (75.0%) Bless 5)) 2 (66.7%) 1966 Success I (83.03%) 20 (87.0%) 2 (50.0%) Failure (66.7%) 8 Br On) 2 (50.0%) Totals Success 2 (28.6%) 43 (84.3%) 3 (42.9%) Failure 5 (71.4%) 8 (15.7%) 4 (57.1%) 182 umouy Spaitq jo (0°0) (0°0) OO 0°O (at). Grea) qt en (C2 Sp eal(es At) O° eT 9° 12 (S26) (202) eer GeGg C2292) Gl) aoe dy? SIit (Or0)e (9°41) O20 On *‘poinqydeo spitq jo esequeorzed 9y4 ST oanSTyJ puooes esejusoied 04} sqUuesomdar oin3Ty ysitgq (9°98) 8°g 0°OOT == 25 =r (het) (OLED) t° + One O° OOT == Sie (GeO (G6) (Gac) 9° l2 GW 9° l2 0°OOT -- (O20) (O20c)- (O10) = = (O60) G29 9° 9¢ 9° 9g OOS O°OOT C290) 3935)" 1G ip) 9T (0-0) GAA CuGe 9° Lg GaSe ESE (oA) (Sp) 2 (OO) = (Or) = (COe)) On O° 4 O°h 9° 4 7h pespeTy saey oF 0*°OOT -- (Ox@) = (OKO) Ot Sai 90} pue SATT@ Usaeq aaey 04 UMOUY eZTeM SPAITQ 9Sout ITTV x aha a HE 996T -- -- c2 G96T -- -- 62 496T -- -- els CO6T -- -- QT Z96T (0°0) o'r 0°OOT co 6S6T 4TOYOO TIOVV sany SpueTs] useerty wory syroyoo Aqoog umMourg snotzeA Jo saqyer oingdeoay pomnqydeosy zeaz °x69-6S561T ‘(sesequeored se passerdxa) “QOT-dd ETAL 183 Non-incubating Brown Boobies roosted either on the ground next to the incubating birds or in the Scaevola near the nests. While guarding the younger nestlings, the adults roosted on the ground, but many of the older young roosted on Scaevola next to their parents. Some breeding adults roosted at night outside the central plain in such areas as the north tower, or in Scaevola along the northwest beach. Non-breeding Brown Boobies generally roosted outside the central plain on the north tower, at the north point, in Scaevola along the northwest beach or, more rarely, in the north roost, along the south- west beach, or on coral blocks near the southeast beach. During the day Brown Boobies were commonly seen standing on the reef. Banding and Movements One hundred and forty-eight adult Brown Boobies were banded at Kure Atoll. Table BB=-1/ briefly summarizes the recapture of these binds. 9 Details of) this study will be published Tater. One aduilt female Brown Booby banded on 8 November 1964 was recaptured at Pearl and Hermes Reef on 18 March 1965 and 21 March 1967. One subadult banded in 1962, one banded in 1966, and two in 1968 were not recaptured. The recapture of nestling Brown Boobies reaching sexual maturity has been previously summarized. In addition, 17 nestlings of the 1967 cohort were banded and 29 of the 1968 cohort. Four of the 1967 cohort were captured in 1968 and 2 of the 1968 cohort were recaptured the Me xipmmyie ans. Young Brown Boobies wander extensively. An immature banded on 4 October 1963 was captured at Wake Atoll on 19 June 1966 and later at Kure on 2 January 1967. Four other Brown Boobies banded as nestlings were recovered in the Pacific Basin (Table BB-18). These data suggest a southerly dispersal of young Brown Boobies after fledging. Three Brown Boobies banded on other islands bred at Kure: a nestling banded on Eastern Island, Midway Atoll, in 1957 bred at Kure each year from 1966 to 1969; a breeding adult male banded on Southeast Island, Pearl and Hermes Reef, on 28 August 1967 also bred at Kure in 1969, and an adult female banded on Howland Island as an immature on 1 February 1965 was found roosting on Green Island on 15 June 1966 and 8 July 1967, and breeding in 1968. This is one of the few recorded movements of a bird from central equatorial Pacific islands to the Northwest Hawaiian Islands. Another three Brown Boobies (two banded as nestlings and one as an adult male) were recaptured at Kure, but not found breeding. 184 aInSTJ puoodss ayy pue SATT® Uaeq aAey OF UMOUY (2°22) Cace (6°24) 62h (Ze Ty) hth (1°24) Tht (Eee) Los (4° 9) uc She (eon) Gok (Ee EI) TAT O*OOT (Geo TKS) es 21S (€°9S) €°9S (6°2S) 9°9S (1° S¢) 6°SH (6°2t) Q° eG (So Lz) One (EAL) Eo EE O° OOT (€°9S) L°99 (Tit) Lg (4°2S) Q° 4S (LG) T° LS (6°0%) 6°On O°OOT (Ar) G°9L, GieG) Ong (€° 49) €° +9 (0°0S) 0°0S (3° ed) Gee’ O° OOT (9° +S) (6) (4° 9) Eng (1°6S) 9° £9 Glestat) aoe S96T O°OOT -- (205) SO ee OMOOM (2°89) (9°) Dike ds Line (ards) (@°O)) O° Gar Gude HOOT €96T peanqdesey reaz "x69-6561T ‘(sae8equeored se passeidxa) areyy peangdeoaz pue [TTO1V eany 1e papueq setqoog umMoug YInpe Jo seer sanqgdeooy Spaitq jo *pomydes spaitq jo aseqyueozed ayy St aseqjusoted ayy squosezder aanSTy YSATA x 6 Q96T i; L96T al 996T Msi S96T on H96T ge €96T ce Z96T 6 6S6T pepueg Teak “JT-dd eT aed 185 Table BB-18. Brown Boobies banded as nestlings at Kure Atoll and recovered at various locations. Distance Date (nauti- How Banded Where Found When Found cal miles Obtained 4 October 1963 Marshall Islands: 97 Novas, WSS «aL sXers) Trapped Majuro Atoll 22 July 1966 Indonesia: Central 21 March 1968 3,564 Found af- Moluccas ter storm 30 August 1966 Ellice Islands: 3 March 1967 2,187 Found dead Funafuti Atoll Ly Seley USES) Ellice Islands: 9 May 1970 2,187 Shot Funafuti Atoll RED-FOOTED BOOBY Sula sula Status Abundant winter-summer breeder; 175 to 430 pairs annually. Present all year with peak numbers from May through October. Breeding generally begins in March and continues through November. Populations Both Munter's and Wetmore's estimates (Table RFB-1) were larger than any POBSP estimate (Table RFB-2). At the time of their visits Scaevola covered considerably more area than it does today, consequently, there was more available breeding habitat for this species. Whether the number of Red-foots has decreased, however, is questionable, for today large areas of Scaevola are not utilized by breeding birds, suggesting that breeding Red-foots used no more area in 1915 and 1923 than they do today. Perhaps these observers were over-enthusiastic in their estimates, a real possibility -since they made no counts. Kenyon and Rice's estimate was based on a nest count during the peak of the breeding season and was on the same order of magnitude as recent POBSP estimates, again suggesting that Red-foots are as abundant today as earlier in this century. POBSP estimates were based mainly on counts of roosting birds. This method was accurate only in a general way because it was difficult, if not impossible, to see from the radar reflector and north tower all Red- foots roosting in the Scaevola. A simple example will illustrate. On 18 May 1967, 106 Red-foots, including breeding birds, were counted at 186 Table RFB-l. Previous records of Red-footed Boobies on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 2,000 Eggs (Munter, 1915: 137). 1923 Aprid) 17<22 2, 000 Few eggs; most pairs nest building or selecting nest sites (Wetmore, ms.). 1951/7, pune 5 Gaye 100 Estimates 240 nests based on count of 218 (Kenyon and Rice, 1958: 190). 1959 October 3-8 300 (Robbins, 1966: 53). 1960 March 28 600 Nesting (Robbins, 1966: 53). 1961 January 19-21 ) (Robbins, 1966: 53). September 12-14 2 Nesting (Udvardy and Warner, 1964: 2). 1962 February 2-4 15 (Robbins, 1966: 53). August 6-8: 500 2h0 nesting pairs (Robbins, 1966: 53). 1963 February 3-7 2h0 (Robbins, 1966: 53). sunset in the central roost. The next day at sunset, 95 individuals were seen in the north roost. At this time there were ca. 147 nests in the central roost and ca. 282 in the north roost, so the roost counts did not even equal the number of incubating or brooding adults. Far more accurate, especially for comparison, than roost counts were the nest counts of 1965 through 1969, and the 1964 estimate. These were fairly consistent at ca. 200 pairs, but in 1967 there was at least a 147 percent increase from 1966 and a 74 percent increase from 1965, the previous maximum count. What caused this increase is unknown, but a few possibilities, such as an influx of birds from other islands or a large number of young Red-foots nesting for the first time, will be examined with the aid of banding and recapture data. Breeding Red-foots were sampled three times, on 27 June, 2 July, and 4 July 1967, in as random a way as possible. One hundred and twenty- five individuals were handled. Since the maximum nest count was 429, these samples represented ca. 13.8 percent of the breeding population. Seventy-nine of the 125 (63.2 percent) Red-foots handled were already banded. Table RFB-3 summarizes the history of these birds. By using the derived percentages we can determine the number of each class in the 187 Table RFB-2. POBSP semi-monthly estimates of Red-footed Boobies on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 __~_ 1966 1967 1968 1969 January Los = LD 700 = ISK = (oe 16-31 - ney 400 * - - 800 February 1-15 - 400 400 225 380 - 600 16-28 * S15 600 - - - 700 March 1-15 - 550 600 - - - 800 16-31 - 550 500 ~ * 450 800 April 1-15 - 550 350 - - - 800 16-30 - 550 350 400 - - 800 May 1-15 1,000 550 400 500 OMS - 1, 000 16-31 - 550 OO 500 LOG 1,000 1, 000 June 1-15 - 550 550 400 La OVs 1, 000 1, 000 16-30 - 550 550 700 LO 1, 000 * July 1-15 - 600 550 500 IL, OS) 1, 000 - 16-31 - 700 600 700 - 1, 000 - August 1-15 - 700 550 750 - 1, 000 - 16-31 - 700 - 700 - 1, 000 - September 1-15 - 550 - 550 - 1, 000 - 16-30 * 700 - 600 - 750 - October 1-15 x 700 - - - * - 16-31 * 700 - - - 500 - November 1-15 400 800 - - - 500 - 16-30 400 800 350 - ~ 500 - December 1-15 400 800 300 - - 500 - 16-31 350 646 - IB - x _ * Birds present, number unknown. 188 Table RFB-3. Number and percentage of total number of previously banded Red-footed Boobies captured in samples of breeding birds on Green Island, Kure Atoll, 1967. Number Captured:* Year of Banding Adult Subadult Immature _. Local 1962 rly (s38)) 0 0 3t1B (3 9g + 1-38) 1963 10 (12.7%) © 4 (5.1%) © 196) Be (256) oe) CU ee hy ele Raa) 0 1965 4 (5.1%) O ©) 0 1966 26 (32.9%) 3 (3.8%) 0 0 * Age at banding. A = Banded on Wake Atoll. B= Banded on Midway Atoll. total population. For example, since there were 542 (0.63 x 858) banded Red-foots in the population and 1.3 percent of these were banded in August 1962, it then follows that 6 (542 x 0.013) individuals’ in this class were present. Table RFB-4 gives the calculated totals for each class. Table RFB-4. Structure of breeding Red-footed Booby population on Green Island, Kure Atoll, 1967. Unbanded Red-footed Boobies: 316 Banded Red-footed Boobies: 542 in the following age classes: Age when banded: Vearsor Banding Adults Subadults __Immatures Locals 1962 6 0 0 21+6B 1963 69 O 28 0 1964 14, +6A 6 28 0 1965 28 O O 0) 1966 179 | al O ) Totals .06tOA 27 56 21 6B A = Banded on Wake Atoll. B = Banded on Midway Atoll. 189 With these data we can examine three possible reasons for the 1967 increase: (1) an influx of Red-foots from other islands; (2) large numbers of young birds breeding for the first time; and (3) many of the Red-foots banded as adults in 1966 breeding for the first time. Based on banding data these would result in the follow- ing increases, respectively: 6.7 percent, 24.7 percent, and 51.7 percent, by themselves not enough to account for the total increase. Even if all the unbanded individuals were added to each class, re- sulting in increases of 97 percent, 115.6 percent, and 142.6 percent, they.do) not account for the increase. it is allso unlikely that all unbanded birds belonged to only one group. When considered together, however, they may provide a sufficient explanation. It is worth men- tioning at this point that roost counts made in May and June 1967 were at the same level as those made during the same period in 1966. Perhaps there was actually no appreciable change in the population size, but rather that more Red-foots bred in 1967 because of favorable environ- mental conditions which at the moment are unknown. In this respect it should be noted that this increase occurred after the year when the smallest number of Red-foots recorded during POBSP studies bred, and that the population in 1968 was only slightly above the 1965 level. The 31.3 percent increase from 1968 to 1969 followed a similar pattern, i.e., an increase after a decrease and a later breeding season. This correlation between size of the breeding population and the timing of the breeding cycle has been discussed previously. The breeding popu- lation was not sampled for banded birds in 1969 to determine other possible causes for the increase. Unlike the other sulids on the island, subadult Red-footed Boobies formed a significant portion of the total Red-foot population. For example, in late May 1966 this group composed an estimated 20 percent of the population. These birds were present all year, but data were in- sufficient to determine accurately population fluctuations through an entire season. Available estimates include: O in late February 1964 and late March 1968, 3 in late March 1965, 1 in early January 1967, 100 in late May 1966, 200 in May and June 1967, 60 in May 1968, 100 in June 1968, and 150 to 200 in July 1968. These data show that subadult Red-foots occurred more commonly in summer than in winter. Beginning in April 1966 an attempt was made each month that POBSP observers were on the island to catch as many subadult Red-foots as possible. It was soon discovered that the plumage of these birds varied considerably. Each individual captured was placed in one of three categories -- subadult brown, subadult white-brown, or subadult white-- based on its plumage. (Details of this study will be published later). Although the exact chronology is unknown, the available data based on 51 birds banded as nestlings indicate that it takes three years to acquire adult plumage and that subadult browns are one year old, sub- adult white-browns are two years old, and subadult whites are three 190 years old. This knowledge enables one to comment on the age structure of the subadult population. Table RFB-5 summarizes the age structure of the subadult population for those birds that were handled. Table RFB-5. Age structure of subadult Red-footed Booby population on Green Island, Kure Atoll, in May-July 1967 and 1968 (expressed as percentages). Age 1967 1968 One year 19.7% 50.9% Two years 55 23% 37 .0% Three years 25 .0% 12.1% Number handled 76 108 The relative increase of one-year-old birds in 1968 probably re- sulted from the high productivity of the breeding population in 1967 compared with 1966. Although population studies of young Red-foots are still in the embryonic stage, it may be worthwhile at least to summarize the avail- able data. Tables RFB-6 to 8 list by year the number of subadults that were caught, how many were previously banded, and the history of these banded birds. These data show or suggest the following: (1) sub- adults were present from April to October; (2) Red-foots from other islands were present from May to October; (3) birds from the 1966 cohort were present in June, July 1967; and May, June, July 1968; (4) birds from the 1967 cohort were present in May, June, July 1968; (5) a relatively large percentage of the previously banded birds had been banded on other islands (60 percent in 1966, 40 percent in 1967, and 23.6 percent in 1968); (6) the 1966 Kure cohort was apparently more common in 1968 than in 1967. This last statement requires some ex- planation. In 1967, 15 percent (6 of 40) of the banded Red-foots handled had been banded as nestlings in 1966; in 1968, 41.2 percent of the 51 previously banded birds were in this group, suggesting that more had returned to the island since 1967. Without data from a complete year it is difficult to attach much significance to these observations. Annual Cycle Red-footed Boobies occurred throughout the year with peak popula- tions present from May through October, the period of peak breeding activity. The smallest populations were recorded in January, February, and December. Whether or not all members of the breeding population were present every month was not determined. Red-footed Boobies bred on a well-defined annual cycle which usually began in March and ended in November. Like their congeners, Sula dactylatra, Red-foots showed some yearly variation (Table RFB-9). 191 Table RFB-6. Summary of subadult Red-footed Boobies handled on Green Island, Kure Atoll, April-October 1966. April May June July August September October Previously banded O Wh 13 tl 5 3) e Unbanded 8 BS) thet BAL el 9 iL Repeats 0 @) O 14 i rh O Totals Cea ei ee soe (SLOT Te a ae Distribution of returns: April May June July August September October Banded on Kure as: Imm. 1963 O 0 1 1 O O O Local 1964 0 O 2 © 0 © O Imm. 1964 O 6 3 3 0 iL i Subadult 1965 O IL a © O 0 O Banded on: Midway Atoll O 4 2 O 2 al 1 Laysan O O O O a 1 O French Frigate Shoals O O 1 O O O O Mokapu Point, Oahu 0 O al 1 ©) 0 O Sand Island, John- ston Atoll 0 3 O 2 2 ©) ©) Wake Atoll O O 2 O 0 0 O For example, the peak period of egg laying was almost two months later in 1966 than in 1964. Figure RFB-1 shows the number of nests with eggs and the number with young for each semi-monthly period POBSP personnel were present. Egg laying was known to begin as early as late January, but usually did not stant until early March. |)The peak period of egg Laying varied yearly from mid-March to early June. Generally there was only one peak, but in 1968 there were two, slightly over two months apart (Fig. RFB-2). Only one egg was laid per mest. Eggs were laid as llate as late June. Eggs hatched approximately six weeks after being laid. Therefore, hatching began as early as mid-March and continued until late August. O amy XO) T is? © € @. 0 O Ie 0 c eS _© O be «@ o Oe eik O LE O € G ea o OQ. © O ul O O Ee O Ore yak is ue © Aine eune Lew 192 sumnjezr Jo samyonz1s THSUCTSTT - KemMptW :uUO pepueg 996T “eqns H96T ‘ull. :Se ainy uo popueg oyeM TYSUeTSTT ues ket AemptW :uO popueg 996T * Bans 996T “wu Q996T “TIS8N $88 aInNy uo poaepueg Teney :uO popueg 996T * wut 996T “13S58N :se aIny uo papueg ‘TTo1y eany “SpueTSsI usery uo paTpuey yInpeqns QL 0° Sg 61 €°GS oh 2°61 GT uotyeTndo JO % peTpuey zJaqunu [e107 “pepuequn - xxf *(suamnjet) pepueq ATSnoTtAedg - xd Te 101 g lie 20 t t O xxM (PTO zeek €) + Ih OF = 4 o4TUM cL 92 S GI 2 «ef (PTO 2ze0f g) 6 se ee Cae coed UMOTG=-94 TUM € IE a 2 9 © xn (plo 1284 7) TE G 1 ¥O uMOLg TNL eine few >poTpuey Jequnn SStTO "1961 ATne-KeW Satqoog pe .oos-pey yfnpeqns jo Arewums */-d4d eTqeL 193 “pepuequyl) - xxi] *(suanjez) papueq ATSnoTAetg - xg T © © eyesTIq@ yousry O O t TYSUCTST'T T O O SowzoH pue [Ledd 0 O--¢ TLOvV AeCMpTW SOT TeIOL suo papueg 9 U 9 T © © SGSk “Tsai 8Se einy uo pepueg iaacal eT € i C swigl (gto aes C) S O € ¥d 34TUM O O a TTOFV oxeM 1 OO eyestAq youserq O © THSUBTSTT :uO popueg ©) © € L96T “eqns oo Qg OT Gt g Ww CSor clase 382 eIny uo pepueg OWS On 9 G € xx (pTo azeef g) OT € I xd@ UMOTG-39 TUM ali O O uo VSUuYyOr-pueg € © © TYSUCTSTT :uo pepueg aE Q Cr t OS G ASE ASN 25S 6°0S SS 9g g GQ wei) (iSO aeeVS Tr) Aqne sune Lew aIny uo pepueg 9 9 G ie uMOTg suinjaz Jo aanjons49 uotzeTNdo peTpuey Te oune eI sseTto 4[Inpeqns ZJaqunu [eo], ;peTpuey Zoquny Jo % "996T ATnr-ABW *‘TTO1V emmy “pueTs] useert5 uo peTpuey setqoog pe,ooj-pey atnpeqns jo Azewuns °Q-d4y eTqeL 194 Taquadsaq é g-ATnL 04eT £-ATNL- PIN g-4snsny Taquadad-=% -AqTnp *e0 SUTSpeTA ATne syaem Key 2 ysert ‘xen ATneL Suto eH SY9eM So 4YSeI = sYyeem g 4SITT eune ATTeq syoom J YSaATq ANE Yyoom 4ST Kew yeog oune ysnsny ayeT YSeeT 38 asnsny Zz °@o i} 4sn3ny eT -pTW 4SPOeT euNne 34eT -YoIeN- PTW -T Tad y-PTW -Ttady-pIw -Kewq AT Ie qe-KeW- PTW -Keq AT IER BUT Yo eH sune Trady-ptw syeem J 4SATT Key Trad y-ptw But key -YoleW- Pin ‘yoreW o4eyt [Ttady a7e 7] Yoom 4SeT Kew Yeom pare -Yorey, 972 T S3yq yeod aun jo pus 4seeT Kew AT ree 7e-YoL ey qseeT 3e-Are Seo jie CED aun sune AT area -ptw $Aze Kew-ptu- £12 —nuer 3487 -yorey AT Ley -yorey, AT AeY -Yyorew o4eT -nuer 34eT -nigqaq 34e7 Sutkeyl 33 696T Q96T L96T 996T S96T +96T potted °69-+96T ‘TTo1y emmy ‘pueTsI ueery uo aToko Sutpseeriq Aqoog pe,ooj-pey eyy ut spotszed zofeMm °6-da eTqeL oD 1965 150 30 NO DATA 1966 200 100 eggs YOUN 1967 400 eggs young 200 NO DATA NO DATA 1968 300 200 100 1969 400 200 NO DATA J F M A M J J A S 0 N D Figure RFB-1. Breeding cycles of Red-footed Boobies on Green Island, Kure Atoll, 1965-69. 196 100 80 60 40 Number of Nests 20 Pe. J F M A M J J A S 0 N D Figure RFB-2. Number of Red-footed Booby nests in which first egg was laid each semi-monthly period on Green Island, Kure Atoll, 1968. Hatching peaks varied from May to July. Not enough numerical data were available to determine the peak periods of egg laying and hatching for all years combined. Young Red-foots began to fledge in mid-July. Fledging continued into December. Approximately 13 weeks elapsed between hatching and fledging. These birds returned to roost in the nesting areas for one or two months after fledging. Most, if not all, eventually left the island. In 1969 all of the 1968 cohort had left by mid-February. However, by at least May some were again roosting on the island. Nesting Success Table RFB-10 summarizes Red-footed Booby productivity for 1964 through 1969. The only detailed success data were collected in 1968 in the central roost for three different groups of nests (Table RFB-11). These figures compare favorably with data for other sulids at Kure. 197 Table RFB-10. Productivity of Red-footed Boobies on Green Island, Kure Atoll, 1964-69. Maximum Maximum Maximum Estimate Approximate Nest Count Egg Count Nestling Count of Young Percent Year or Estimate or Estimate Or Estimate Fledged Fledged** 1964 200 - 200 200 - 1965 2h. 189* 200* 200 82.9 1966 174 1770 150 IBIS TS 1967 42x 395* 3ehx 300 66.9 1968 297* 121 256% 225 US olf 1969 390 350 2D ORK = = * Count. EK Percent of young fledged from highest nest count or estimate. **xx Number remaining on 20 June. Table RFB-ll. Nesting success of three groups of Red-footed Booby nests in the central roost, 29 May-10 August 1968. Group I: Nests that contained eggs on 29 May n= 2/ 70.4% hatched 70.4% young remaining on 10 August from eggs laid 100.0% young remaining on 10 August from eggs hatched Group II: Nests that contained young on 29 May iq ee ine 100.0% hatched 34.9% fledged by 10 August 58.1% young remaining on 10 August 93.0% probable fledging success Group III: Nests started after 29 May ia = BS 65.4% hatched 61.5% young remaining on 10 August from eggs laid g4.1% young remaining on 10 August from eggs hatched All Groups: Nests with eggs 67.9% hatched 66.0% of all eggs laid will probably fledge young 97.2% of all hatched eggs will probably fledge young 198 Table RFB-ll. (continued) All nests 82.3% hatched 78.1% of all eggs laid will probably fledge young 94.9% of all hatched eggs will probably fledge young Ecology Red-footed Boobies bred mainly in Scaevola, and also in the northernmost Tournefortia along the northeast beach. Here they built bulky nests of Solanum vines and some Ipomoea and Scaevola. Occasional sprigs of green Boerhavia were added during the incubation period. Red-foots gathered these materials in open areas, mainly the central plain, by pulling vigorously at the live plants until the vine broke. Almost all, if not all, of the birds used only one or two areas from which to gather vegetation during one breeding season. After a while little green vegetation was left in these areas. As far as could be determined, only adult males gathered nesting materials. A few sub- adults were also seen pulling at the vines. Nests were built by placing the vines on a supporting platform of branches, which were sometimes on the same bush and at other times formed by interlocking branches of different bushes. Generally, nests were near the top of Scaevola bushes, but some were well below the level of sur- rounding vegetation. It appeared that some Red-foots broke off Scaevola leaves and twigs surrounding the nests. The height of nests from the ground ranged from two to at least seven feet, depending on the height of the vegetation. In 1967, 10 nests averaged 44.4 inches from the ground. These same nests had an average outer diameter of 13.2 inches, an average inner diameter of 5.8 inches, and an average depth of 3.1 inches. Usually Red-foots bred in small groups among other Red-foots or among Great Frigatebirds. The distance between nests was determined by the availability of nest sites and the range of the incubating bird's bill, nests being no closer than a bird could reach. For the most part, nests were found in the center of vegetated areas rather than along the edges. The main breeding areas were in the north and central roosts. Maximum nest counts for the former area from 1965 through 1969 were 163, 122,262, o1166, fand#i20, and! fontitheslattersarea 76.525), U7 eae and 270. In 1965 and 1966 a few pairs bred in Scaevola southwest of the runway. Figure RFB-3 shows the general location of breeding areas in 1967 and 1968. As can be seen, many areas were not utilized although they appeared suitable, at least to humans. Prior to the construction of the LORAN station, Red-footed Boobies bred in the Scaevola east of the south antenna field (Robbins, personal communication). 199 "(seere Sut4soor OsTe Siem Sseore Sutpeetq ) 9g9-L96T ‘TTOU emmy ‘pUeTST UueerTy) UO severe SuUTpSezq pue But ysoor ALqoog pe,ooj-pey fo uoTINqTs74STC Buljsooy se sipah yjog = "€-gay omsta 200 Non-breeding Red-footed Boobies roosted in the same general areas as the breeders. However, many also roosted along the beach-Scaevola ecotone. Sometimes these birds intermingled with the breeding birds, while at other times they formed groups by themselves. It was noted several times that when a nest was built in an area where Red-foots were previously absent, it soon became a focal point for roosting birds. Very few Red-foots roosted along the south beach, east beach, or on man-made structures. The main roosting areas are shown in Figure RFB-3. Although breeding Red-foots landed on the ground while gathering nesting materials, none roosted there overnight. All breeding Red-footed Boobies at Kure were light-phase: all white with black flight feathers. Banding and Movements One thousand and eleven adult Red-footed Boobies were banded at Kure Atoll. Tables RFB-12=-13 summarize the recapture of these birds at the atoll. Even though large numbers of Red-foots were usually handled each year (Table RFB-14), these recapture rates are low in com- parison with the preceding two species where most individuals on the island were captured each year. Undoubtedly if all or most Red-foots had been handled each breeding season, the recapture rates would have been at least as high as the Brown Booby and possibly as high as the Blue-faced Booby. Thirty-six of these adults were recaptured on other islands: 24 on Eastern Island, Midway Atoll, 3 at Pearl and Hermes Reef, 1 on Lisianski, 1 on Laysan, 6 on Sand Island, Johnston Atoll, and 1 on Wilkes Island, Wake Atoll. Interestingly, 10 of these Red-foots were recaptured again. at Kure (4 from Midway, 2 from Pearl and Hermes, 3 from Johnston, and 1 from Wake), suggesting a type of migratory movement. Three of the birds captured at Midway Atoll bred there. Four hundred and twenty-five subadult and 607 nestling Red-footed Boobies were banded at Kure Atoll. Tables RFB-15-16 summarize the re- capture of these birds. As would be expected, fewer young Red-foots were recaptured than adults. However, more young traveled to other islands. Twenty-nine Red-footed Boobies banded as nestlings were re- captured at the following locations: 3 at Midway Atoll, 2 on Lisianski, 9 at French Frigate Shoals, and 15 at Johnston Atoll. Three of these birds were recaptured later at Kure (1 from Midway, 1 from French Frigate, and 1 from Johnston). The French Frigate Shoals Red-foot bred at Kure. Twenty Red-footed Boobies banded as subadults were recaptured at the following locations: 6 at Midway Atoll, 1 at Pearl and Hermes Reef, 5 on Lisianski, 4 at French Frigate Shoals, and 4 at Johnston Atoll. Eight of these birds returned to Kure (4 from Midway Atoll, 1 from Lisianski, 2 from French Frigate, and 1 from Johnston). One Midway bird bred at Kure. The subadult Red-foot recaptured at Pearl and Hermes Reef was banded at Kure on 14 October 1963, recaptured on North Island 20ik on 20 August 1964, and found dead at Taka Atoll, Marshall Islands, later that fall. No attempt to analyze this movement was made because of the great variation in the amount of effort expended in recapturing Red- _foots on the various islands. Table RFB-12. Recapture rates of adult Red-footed Boobies banded as breeders at Kure Atoll and recaptured there (expressed as percentages), 1964-69%. Mean Year Recaptured Banded fal. 1964 1965 1966 1967 1968 196 1964 32 100.0 432.8 40.6 BS 5 T 21.9 TLS) af) Q235)) — (28) C25) (Qa). (CUS) 1966 9 -- -- 100.0 55) 0 33} 03) 35}038 (ADSI) > (SSeS) > (858) 1967 58 -- -- -- OOKO Hu 8 22.4 (29.3) (22.4) 1968 3 -- -- -- -- 100.0 33 53 (33.3) 1969 8 -- -- -- -- -- 100.0 Pehinch Ligure represents the percentage of birds known to have been alive and the second figure is the percentage of birds captured. Enough Red-footed Boobies banded on other islands by the POBSP were recaptured at Kure from 1964 through June 1969, however, to permit a fairly detailed analysis of the nature of this movement. Table RFB-17 summarizes these data. Only Red-foots banded through 1968 are included in the totals. In addition, 4 Red-foots banded at Midway Atoll (2 as adults and 2 as nestlings) by non-POBSP personnel were captured at Kure. To eliminate the differences between the number of birds banded on the various islands, the number of recaptures is expressed as a percentage of the total number of birds banded on a particular island... It is assumed that enough Red-foots of all ages were handled over a sufficiently long period tO minimize any bias) resulting from recapture techniques. The final figures are a movement index of Red-foots from other islands to Kure. These data show or suggest the following: (1) Only Red-foots from the Hawaiian Islands, Wake, and Johnston traveled to Kure; (2) a greater percentage of young Red-foots traveled than adults; (3) movement indices for the Hawaiian chain decreased with increasing distance from Kure; (4) considerably more Red-foots from Wake and Johnston moved to Kure than would be expected if interisland movement were only a function of distance, suggesting that there may be a "migratory" movement between these areas. Too few data are available to determine yearly and seasonal differences. 202 *poanqded spitq jo e8equeorzed 3yy ST oINSTJ puocodsas 9} pue SATTS Useq aAeY OF UMOUY Spatq Jo aeSsequeorted 9y. 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One Ee awls SOE Bsunok se pepueq 9% oIny 1’ perngdeoez OZ© Jn" ©) Org S°0 GO 0°O Ore) €a¢g sqy—Tnpe se pepueq % € OT GT CT 9 9 S oe amy 1e peunjdeoaz requmnu Te OL S CT cL OT G 9 Z QL eany ye pemydeosaa 3unok 3@ pepueq zrequnyl O rT € € tt O € 4 eany 72 pornjdeoet sz[npe se pepueq s7equnN pepueq Jequnu Te OL €0L °T Oge 949 Q9T ‘T 962 66T ST 6ST pepueq xsunok tequnyl ttre ty €ST LT9 409 499 OST HLT pepueq sq—npe zequmit SoeTtu “yneu) soueqstd "S] UeTTe® TLOYV TLOvV uods speoug oye ueske] Tysuel jooy sewzeH T[TOIV 7WO.LT -Mey UTeN oyeM -uyof -STIqg youerg S Sito pue [Taieeqd AeMPTW *69-H96T “TIOVW eany 0 spueTsT seyyO uO dgdod aux Aq pepueq satqoog peqooj-pey Jo JuewercW “LI-ddd OTAPL 206 GREAT FRIGATEBIRD Fregata minor Status Abundant late winter-summer breeder; 120 to 400 pairs annually. Present all year with peak numbers present from May through mid-July. Breeding begins in late February and continues to mid-December. Populations Apparently the Great Frigatebird population did not decrease as a result of the construction of the LORAN station; earlier estimates (Table GF-1) were no higher than POBSP estimates (Table GF-2). Although Scaevola covered considerably more area in 1915 and 1923 than in the 1960's, Munter and Wetmore's observations indicated that this species bred in only a small portion of the available habitat. Even today Great Frigatebirds do not roost or breed in all the available Scaevola. Table GF-1. Previous records of Great Frigatebirds on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 1, 000 1 colony near center of island: ‘eges (Munter, 1915: 137). 1923 April 17-22 200 100 pairs nesting in Scaevola near central plain (Wetmore, ms.). 1957" dune 5 325 100 nests estimated; roosting in Scaevola; 3 concentrations between central plain and southeast beach (Kenyon and Rice, 1958: 190). 1959 October 3-8 275 Nesting (Robbins, 1966: 53). 1960 March 28 800 Nesting (Robbins, 1966: 53). 1961 January 19-21 Me) None nesting (Robbins, 1966: 53). September 12-14 2 Nesting (Udvardy and Warner, 1964: 2). 1962 February 2-4 50 None nesting (Robbins, 1966: 53). August 6-8 250 115 nests (Robbins, 1966: 53). 1963 February 3-7 100 Not nesting (Robbins, 1966: 53). 207 Table GF-2. POBSP semi-monthly estimates of Great Frigatebirds on Green Island, Kure Atoll, 1964-69. 1963 1964 1965 1966 1967 1968 196 January 1-15 = 250 100 - 15 - 350 16-31 - 250 200 - - - ‘LOO February 1-15 - * 300 160 70 - 250 16-28 * 650 300 - - - 250 March 1-15 - 750 700 - - - 300 16-31 - 150 700 - * 450 500 April oS) = 750 550 = = - 500 16-30 - (50 550 600 = - 600 May 1-15 900 750 550 700 1,500 - 800 16-31 - 750 550 700 1500 1, 000 850 June 1-15 - 750 1,000 700 1,500 1, 000 1,500 16-30 - 750 800 600 1. 500 1, 000 * July 1-15 - 500 750 300 1,500 600 16-31 - 1L5 KOO (50 250 - 600 August 1-15 - 1, 000 700 225 - 500 16-31 - 950 - 300 - 500 September 1-15 - 1 O5O - 120 - 300 16-30 * 1 OO - 325 - 300 October 1-15 * 1,100 - - - 500 16-31 * OO - - - 500 November 1-15 400 1,300 - - - 300 16-30 400 700 300 - - 300 December 1-15 250 700 450 - - 100 16-31 700 197 - 15 2 * * Birds present, number unknown. 208 The most accurate population data were nest counts made in 1965, 1967, 1968, and 1969 which revealed minimum breeding populations of 532, 512, 240, and 380, respectively. The 1967 and 1968 counts were conservative since they were made in May when a large number of nests probably had already been destroyed. These data were insufficient to determine if there was any significant change in the size of the breed- ing population during POBSP studies. If Great Frigatebirds breed only every other year as some authors (e.g., Nelson, 1967; Schreiber and Ashmole, 1970) suggest, then the total Kure breeding population was 1,000 to 1,100 individuals. Generally POBSP estimates were based on roost counts and therefore were accurate only in a general way. The limitations of these counts were discussed in the Red-footed Booby account. Enough banding data were collected in 1966 and 1967, however, to analyze the structure of, and to characterize, the Kure Great Frigatebird population. Tables GF-3 and 4 summarize the data. It was assumed that enough birds were handled to eliminate any bias resulting from the recapture technique; therefore, these samples are representative. It is evident that approximately equal numbers of adults and sub- adults were present, that only a small percentage of the population was banded, and that many of the banded birds were from other islands. These facts suggest a high turnover rate. For example, in 1966 only 12.2 percent of the adults and 7.3 percent of the subadults banded previously at Kure were captured. In 1967 only 5.1 percent of the adults and 2.8 percent of the subadults banded in 1966 were found. Since 318 frigatebirds were banded prior to 1966 and 561 in 1966, the total number of birds using the island during a year must be in the thousands. A simple Lincoln Index calculation shows this. Prior to 1967, 879 Great Frigatebirds were banded at Kure. In 1967, 429 frigatebirds were handled; 49 had been banded previously at Kure. Therefore: 879 = 49 where x = total island population. x 29 Therefore, x = 7,700. Although this figure is probably too high, it indicates that the Kure population numbers in the thousands. If this is true, then most of the frigatebirds using the island are non-breeders. The relatively large numbers of interisland birds (56 or 34.4 percent of all banded birds captured) also showed the transitory nature of the population. An interesting and unexplained feature of the subadult population was the preponderance of females. In 1967, 151 subadults were sexed by bill measurements (co’ less than 110 mm., 9? greater than 112 mm, POBSP, unpublished). Of these, 49 (32.5 percent) were males and 102 (67.5 percent) were females. Further work will be necessary to determine the significance of this observation. 209 Table GF-3. Summary of Great Frigatebirds handled on Green Island, Kure Atoll, April-October 1966. Adults Subadults Total Total number handled Tl 264 Su Number banded 255 228 483 Number previously banded on Kure 15) 5) 30 Number previously banded on other islands iG 21 28 % banded Ted) 135 O77 % interisland (figure in parentheses is percent of banded birds) D5 (Bias) SeOnGers)) 5 2 (C833) Table GF-4+. Summary of Great Frigatebirds handled on Green Island, Kure Atoll, 5 May-8 July 1967. Adults Subadults Total Total number handled 213 216 eg Number banded 178 182 360 - Number previously banded on Kure 30 19 Tie) Number previously banded on other islands 5 105) 20 % banded 16.4 15 NG gal % interisland (figure in parentheses is percent of banded birds) 23 (58) 658) (Naa) Leen (2OR0)) Annual Cycle Great Frigatebirds were present throughout the year with peak populations present from May through at least mid-July. POBSP data suggest that there was a general exodus from the atoll during the winter. The first indication of a new Great Frigatebird breeding cycle was the reddening and enlargement of the male's gular sac, usually in November. By late December or January males began sitting in Scaevola, inflating their gular sacs, and calling to females overhead. Courtship activity was intense from at least late January through April, and some males with inflated gular sacs were seen into late May. Great Frigatebirds bred on a fairly rigid schedule, although there were some yearly differences in the timing of peak egg laying (Table GF-5). The most accurate data were obtained in 1965 and from 1967 to 210 Ce Ce ,-ALew AT Teg Ttaidy-pTw -Yoren-PTW Kew Aj ie9e YseeT 72 -yorey AT I1eq 6961 °69-196T STTOFV eany ‘pueTsI useerH uo eTokdD SuTpeertq prtqeyestag{ JvertH eyy Ut spotzed sz0lew Taqueadaq AT ITea-13q -weidag 34e7 Sun f=pTwW -LeW 97eT Aqtne AqTree -Ttady 07 eT Tyady Yyoom pare -YoleN 9327 Kew AT Ie -yorey AT ITegq 89OT Taqueceg AT 1e9a-13q -we4deg-ptw Ley SYO0M g 4SeT Atne AqTze9 -TTady-ptw Trady SYoomM g 4SITW Ken -yorey AT req L96T ZJaqueydag Ge Trady qseoT 4V Kew AT TST Kew - 3 9961 raqua -09d-pTu-7193q -weqydeg a4e7T oun Syoom Z 4YSITy A[ne-ptw -iTpady 04eT Trady Syoom Z 4SeT ounr AT rTe9e-Are —niqay 34eT S96T Taequesaq -ZJaqueydeg oune IO Kew AT qQeqorg ATRL -Trady Ay szeq Trady IO YoLeW AT Qeqord sATne -Areniged TZ HOOT SUTSpa TA SUTYo4. eH yBed BUTYO?. eH Ssutkey B3y yeod Sutke] 33q potted *G-d) eTaeL (ab 1969. Figure GF-1 summarizes the 1965, 1968, and 1969 data, while Table GF-6 compares the 1967 and 1968 nest counts. &ggs were first laid in late February or early March, and a few were laid at least as late as early June. The peak period of egg laying was usually in April. Eggs began hatching in early April and some hatched as late as mid-July. The peak period of hatching was in late May or June. Young frigatebirds began to fly as early as September and the last ones fledged in December. At least some of these young birds remained in the colony until the following June. Table GF-6. Comparison of 1967 and 1968 Great Frigatebird nest counts on Green Island, Kure Atoll. 1967 1968 Number % with Number % with Date of nests eggs of nests eggs May 9 202 % 1 256 GH 6O ey 239 5O>2 29 120 W7 5 June 6 ILO, 36.4 14 91 IRS 19 185 2B} odl Nesting Success Table GF-7 summarizes Great Frigatebird productivity from 1964 to 1969. The exact causes for the almost complete nesting failure in 1966 are unknown, but it may have been rat predation. At least six adult Great Frigatebirds with typical rat wounds on the back were found that year. However, no rats were actually seen attacking adults or nestlings. Although no detailed breeding studies were conducted, it was evident that a considerable number of nests were destroyed by unknown causes, most likely by the breeding adults. For example, from mid-May to mid- June 1967 ca. 23.8 percent of all nests disappeared. Kceology The main Great Frigatebird breeding area was in the north roost within 50 yards of the northernmost Tournefortia along the northeast beach. In 1964 large numbers bred in the central roost, but during the next two years none bred there; in 1967 only three pairs, in 1968 only six pairs, and in 1969, 26 pairs were found in this area. Figure GF-2 shows the general breeding areas in 1967 and 1968. In the latter year at least eight pairs bred in the Scaevola ca. 10 yards east of the 212 1965 300 200 100 1968 100 oC 1969 200 100 young NO DATA J F M A M J J A S 0 N D Figure GF-1. Breeding cycles of Great Frigatebirds on Green Island, Kure Atoll, 1965, 1968, 1969. °99-L96T ‘TTO1V emmy “pueTS] usexp uo svete SuTpeerq PITGSVeSSTIT yeorH JO uwotTynqt.z4sTd L4G "2-9 eIMsTT 214 Table GF-7. Great Frigatebird productivity on Green Island, Kure Atoll, 1964-69. Maximum Nest Count Approximate Number Percent Wear or Estimate of Young Fledged Fledged 1964 hoot 200 +5080 1965 266 250° 9h .0 1966 200 ale On5 1967 256 1857, 72.44 1968 120 Te 64.3 1969 190 752 is i Probably too large. 2 Number remaining in mid-August. 3 4 Number remaining in mid-June. Number remaining in early August. 2 Number remaining in late June; 75 eggs had not yet hatched. north tower, an area that had not been utilized previously. Prior to the construction of the LORAN station, this species bred along the east edge of the south antenna field (Robbins, personal communication). Usually Great Frigatebirds bred in small groups with other frigate- birds or among Red-footed Boobies. The distance between nests was determined by the availability of nest sites and by the range of the incubating bird's bill, nests being just out of a neighbor's reach. For the most part, nests were placed in the center of vegetated areas rather than along the edges. Great Frigatebirds bred mainly in Scaevola, and also in Tournefortia. Here they built bulky nests of Solanum vines with the addition of some Ipomoea and Boerhavia. Males gathered these materials while flying over open areas, mainly the central plain. Nests were built by placing the vines on a supporting platform of branches; sometimes these branches were on the same bush and at other times were on interlocking branches of different bushes. Unlike Red-footed Boobies, frigatebirds always placed nests near the top of the vegetation where there was enough area overhead for the adults, with their large wingspan, to take off and land easily. The height of the nests from the:ground ranged from three to at least eight feet, depending on the height of the vegetation. In 1967, 10 nests were an average of 50.1 inches from the ground and had an average diameter of 12 inches. Only a slight depression was formed by adults incubating the single egg. The main roosting areas for Great Frigatebirds were in the Casuarina west of the power plant, along the northwest beach from the north tower to the north point (including the north tower), along the northeast beach north of the central plain, among breeding Great Frigatebirds in the north ZAG) roost, among breeding Red-footed Boobies in the central roost, and along the southwest beach south of the easuarinas. Occasionally they roosted along the southeast beach and on the LORAN guywires. They usually roosted in small groups with other Great Frigatebirds or with Red-footed Boobies. Scaevola was the dominant roosting site, followed by Casuarina. and Tournefortia. Roost sites varied considerably from day to day. For example, one night large numbers roosted along the northwest beach while the next night almost none was present there. Active nests became the focal point for much of the nocturnal roosting and these sites were subject to less daily variation. Banding and Movements Seven hundred and three adult Great Frigatebirds were banded at Kure Atoll. Compared with the Red-footed Booby, relatively few of these birds were recaptured at the atoll (Table GF-8), again suggesting the highly transitory nature of this species. Four Great Frigatebirds banded as adult females were captured on other islands in the central Pacific: 2 on Eastern Island, Midway Atoll, 1 on Lisianski, and 1 on East Island, French Frigate Shoals. None was recaptured again at Kure. Seven hundred and seventy-one subadult Great Frigatebirds were also banded. Although relatively large numbers of frigatebirds were handled, especially in 1966 and 1967 (Table GF-9), few subadults were recaptured (Table GF-10). However, over twice as many birds banded as subadults were recaptured on other islands than adults: 1 on Southeast Island, Pearl and Hermes Reef, 2 on Lisianski, 1 on Laysan, and 7 at French Frigate Shoals, including 1 that returned to Kure. Robbins banded 95 nestling Great Frigatebirds (48 in 1959, 10 in 1961, and 37 in 1962). Three of the 1959 cohort were recaptured (1 in MOGSE eam) 1066. ‘and in 1967) and2 of) the 1962)\cohort) (2 in) 11966) and 1 again in 1968) were also recaptured. The POBSP banded 170 nestlings (29 in 1965, 113 in 1967, and 28 in 1968). Three of the 1965 cohort, 8 of the 1967 cohort, and 6 of the 1968 cohort were recaptured the year after banding. Two of these Great Frigate- bird nestlings banded in 1965 were found at Midway Atoll in early 1966. Six Great Frigatebirds banded at Kure were recovered outside the central Pacific (Table GF-11). These data show that after breeding some adult frigatebirds traveled to the western Pacific. Movement of previously banded frigatebirds to Kure was analyzed in the same manner as movement of Red-footed Boobies (Table GF-12). In addition, 2 Great Frigatebirds banded at Midway Atoll (1 as a nestling and 1 as adult male) by non-POBSP personnel were found breeding at Kure Atoll. It is evident that there was decreasing movement, on a percentage 216 *poainqdeo spitq fo esequaozed 94 ST oINSTJ puodss 9y. pue SATTS Useq sAeY OF UMOUY SPITq fo sBequedzsed 9y4 sqyuesoudedr oINSTI YSITA x O0°OOT -- -- -- -- -- -- -- -- -- -- Got 696T (9° t) gh O°COE Ae == =m ae + == == oe oe 49 896T (aes) ei) qr? S°9 0°O0OT == a == as =" =e a = 69T LO6T Csi) ee leg) = (LS) Diet, ey; Goalae 0°OOT a =< == = = == == G€2 996T (OP) (O°O) = (OZ (eeu) O'H Ont Oe O°oT 0°OOT -- -- -- -- -- -- G2 S96T (a) (Ss ae aE 9) (Sec) (Or) Les Ee Oe Gua 9° QT 0°OOT -- -- -- -- -- 49 +96T (G25) ene) = (9200) a(920c)| = (Ged) = (O20) 6°S Whe ail G° 92 3° Qe 2° ge Ze 0°OOT -- -- -- -- HE €96T (©°@) ~ (©°O)) -— (O2O)) - (O"O) = (O20) — (OPO) 1 {0°O) 0°0 0°0 OO 0°0 0°0 0°0 Ox0 OZO0R -- = a2 S Z96T O °6) 0°O (26) 0°OOT Z 6S6T 9OT 896T L96T 996T S96T HOOT €96T c9OT T96T O96T poanydeosay re8z Teoz "469-6661 ‘(Sedequeored se pesserdxa) a19y} pomnjdeoer pue TTO1W emmy 1e saTnNpe se pepueq spitgqezeStaq@ yeery go saqexr oanqdeooy °Q-d) eTqeL Abi basis, with increasing distance from the atoll, and that young birds traveled more frequently than adults. The relatively large volume of movement from Pearl and Hermes Reef probably resulted from an insuf- ficient supply of roosting sites at that atoll. All Great Frigatebirds banded as nestlings on other islands captured at Kure Atoll were taken at least two years after banding, indicating they remained in the breed- ing colonies for at least a year after hatching. Table GF-9. Number of Great Frigatebirds handled each year at Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 Breeding adults © @) ) 6 22 il 37 Non-breeding adults 34 64 A. 27 se ie One Subadults 56 70 Gil Bi) BG 62 69 Totals CO 138 107. +582 20) a3 207 LESSER FRIGATEBIRD Fregata ariel On 27 May 1967 Woodward saw an adult female Lesser Frigatebird flying over the north roost and later roosting in Scaevola among Red- footed Boobies and Great Frigatebirds. The bird was collected later in the day. Although the brood patch was bare, there was no indication that this bird was breeding. Another adult female was captured the night of 25 June 1968 as it roosted with five Great Frigatebirds at the edge of Scaevola along the northwest beach. The next day it was banded, photographed, and released. Lesser Frigatebirds occur abundantly in the central Pacific south of Johnston Atoll. They breed on many islands of the Phoenix and Line Groups, and, to an unknown, extent, in the Tuamotu and Marquesas groups (Sibley and Clapp, 1967). There are the first records for the Northwestern Hawaiian Islands. BLACK-CROWNED NIGHT HERON Nycticorax nycticorax (hoact1li? ) Woodward found the remains of an adult Black-crowned Night Heron under Scaevola in the northeast Sooty Tern colony on 3 June 1966. The race N. n. hoactli breeds in the Main Hawaiian Islands and has wandered previously to Midway Atoll (Clapp and Woodward, 1968). *pomngdeos spitq jo eSequsorzed ayy ST OINS}TJ puooses 3yy pue sATTe Useq sAeY OF UMOUY Spatq fo asequeozed ayy sqUuesetdat aanSTy YSITA x O0°OOT -- -- -- -- -- -- -- -- €9 696T (0°0) Or© 0°OOT -- -- -- -- -- -- -- ot 996T (9°0) (ee) 9°0 MOT 0°OOT -- -- -- -- -- -- ZT L96T (PE) (EE) (9°€) Wee ive Q°9 0°O0OT -- -- -- -- ~- 92 996T (0°0) (9°2) (9°2) (9°2) 0°O 9°2 G9 Om) 0°O0OT -- -- -- -- Mey G96T (S°T) (9°S) (4° 17) (1°8) (GS) Cal er) LONE Q°OT €°0¢ 0°OOT -- -- -- 69 496T (0°0) (0°0O) r=) (PO. (StS) (9°T) 0°O O20 Tee) OWE tale Tole 0°OOT == a0 9S €96T 0°O 0°OOT I T96T 96T Q96T 196T 996T S96T 96T €96T Z96T T96T pepueg poaungdesay rea T@oR °x69-T96T ‘*(SoSequeorted se passerdxe) azeyy peunydeoor pue [TOVV euny 4e@ satnpeqns se pepueq sprtqe.,estiyq yeoery fo soqyer oangdeosy *OT-d) eTqeL 218 219 Table GF-ll. Great Frigatebirds banded at Kure Atoll and recovered at various locations. Age and sex at Time of Date Banded Banding Where Found When Found 7 October 1959 Nestling Philippines: Luzon: 10 November 1960 Camarines Sur: Lagonoy Gulf: Sagnay 11 August 1964 Adult 2 Philippines: Mindanao: 10 March 1965 Matanao: Barrio Banghal 9 August 1964 Adult 2 Philippines: Negros: 8 March 1966 Bacolod 10 June 1966* Adult 2 Philippines: Samar: 12 December 1967 Bobon 25 June 1966 Subadult Marshall Islands: Uge- 14 February 1967 lang Atoll 14 October 1963** Adult 2 Marshall Islands: Eni-- 18 March 1969 wetok Atoll: James Island * Recaptured at Kure on 2 July 1967. ** Recaptured at Kure on 21 July 1966, 3 July 1967, and 30 May 1968. EMPEROR GOOSE Philacte canagica Kleen caught a female Emperor Goose on 15 December 1968 as it sat along the beach near the east end of the runway. No Emperor Geese have been recorded previously from the Northwestern Hawaiian Islands, but their occurrence has been well-documented in the Main Hawaiian Islands on four previous occasions (Clapp, et al., 1969). EUROPEAN WIDGEON Mareca penelope An emaciated second winter female European Widgeon was captured by Coast Guard personnel on LO November 1964 near the east end of the run- way. Lewis saw another individual on 12 November 1965 feeding near the huUci Ans eie ComMlecledmiuwGwOndays maven. kui proved, GO! ibe autirsit winter female. These records and one for Midway Atoll are the only ones for the Hawaiian Islands (Clapp and Woodward, 1968). 220 *sa—Tnpeqns pue sounjzeuut ‘sSuT[4seu soepnTour x c°O G°0 S°O Omr EO 6° Tou aIny 1e@ pommydeoesz pepueq spitq [Te Jo % ZO eal 6°0 eon 9°0 4°9 0°O amy 78 pornydeoer Sunof se papueq % 0°O G0) ils ©) 10) 8°0 teed, 1S einy 1e@ pernqdeoes syinpe se papueq % any 4e pounjydeosxz raqumu [Te .oOL I € 6 € i G2 0 ainy ye peinydeoar Sunofk se pepueq Taqunyl O IE i 2 2 Dit i ony 7.2 perngdeoer sqqTnpe se pepueq Jaquny poepueq Jequmu Teo], O6t €9e 250 ‘T Te? 6LT 662 C pepueq xZ3uno0k requmyl T THS 090 ‘°T €Qz? one O€2 jae pepueq syr[npe sequny SOTTW °yneu) douUeIsSTCG eOUIN TTOVV uo1s S—Teoys oe ueskey itTysuet Jooy souzoy TIO1VV SWOT -uyore -STiq youerzg -STT pue [Tieog eMPIW *69-+96T ‘TTOVV eany 04 SpueTST Zeu1IO UO qgGod 9ui Aq papueq spatqer eStIWq¥ 1eeIH Jo JUeWAAOW “ZT-dD eTaeL 221 PINTAIL Anas acuta Robbins photographed a Pintail 3 to 8 October 1959 and found a carcass on 3 February 1962 (Robbins, 1966: 53). ‘These were the first known records for the atoll. Subsequent POBSP records include: a female collected on 1 October 1963 along the road by the barracks; a male shot on 7 November 1964; two males collected on 17 March 1965 and 19 March 1965; a male collected on 22 September 1966 in the north antenna field, and a male collected on 26 November 1968. The March 1965 Pintails, both in breeding plumage, were first seen with a female in a flood pond along the runway on 17 March. This species breeds across most of the northern half of the Northern Hemisphere (AOU, 1957) and winters regularly in the main Hawaiian Islands (Medeiros, 1958). TUFTED DUCK Aythya fuligula On 29 October 1963 Wirtz caught an immature female Tufted Duck near the barracks. Two other ducks believed to be this species were also seen, one on 30 October 1963 and one on 18 September 1964. Tufted Ducks have also been recorded in the Northwestern Hawaiian Islands from Midway Atoll (Clapp and Woodward, 1968). BUFFLEHEAD Bucephala albeola DeLong saw an adult female or immature Bufflehead on 12 November 1968 by the barracks. He saw it several times the following week, but was unable to collect it. Buffleheads are casual visitors to the Hawaiian Islands (AOU, 1957). Table D-l summarizes sightings of unidentified ducks made at Kure Atoll. Table D-1. Summary of unidentified ducks seen on Green Island, Kure Atoll, 1963-68. Date Seen Number Remarks 1963 September 30 IL -- October 1 aL In lagoon October 6 i -- October 7 AL Landed in antenna field. 222 Table D-l. (continued) Date Seen Number Remarks | 1963 October 18 iL -- October 28 1 -- October 29 il = October 30 ill -- November 12 i In puddle near fuel tanks. 1964 September 24 4 Flying over lagoon. November 5 1 Landed on east beach. 1968 August 29 il Near north point. August 31 al! -- September 28 i Along northwest beach. PEREGRINE FALCON Falco peregrinus (pealei?) Standen shot a female Peregrine Falcon on 7 March 1965 as it flew over Scaevola near the radar reflector. Dr. Clayton M. White identified the specimen as F. RD. pealei on the basis of the extremely heavily marked flanks and thighs; in every other regard the specimen matches the/darker examples of F. p. japonensis (Clapp and Woodward, 1968). This specimen and sight records from Midway Atoll and Lisianski are the only records of Peregrines in the Northwestern Hawaiian Islands (Clapp and Woodward, 1968). DOTTEREL Eudromias morinellus An immature female Dotterel was caught on 9 September 1964 in a mist net erected on the station grounds. This bird had been seen flying with a flock of American Golden Plovers for several days before it was collected. This is the only record for the Hawaiian Islands (Clapp and Woodward, 1968). 223 AMERICAN GOLDEN PLOVER Pluvialis dominica Status Common spring and fall migrant; common winter resident; uncommon summer resident. Populations ranged from 3 to 250 individuals. At least a few present in every month. Populations Only one pre-POBSP estimate (TableAGP-1) was larger than POBSP estimates (TableaGP-2) of American Golden Plovers. POBSP estimates indicated plover populations ranging from 3 to 250 individuals. The 1965 and 1969 estimates were similar, but smaller than the 1964 estimate. It is difficult to determine if these differences are significant since there are many variables (e.g., time of day, tide conditions, weather conditions) that could affect the number of birds counted on any given day. If the population did decrease, it probably was the result of conditions on the breeding grounds rather than condi- tions at Kure where no significant habitat changes occurred from 1964 to 1969. Annual Cycle Figure AGP-1 presents graphically the annual population cycle based on semi-monthly estimates and observations of Golden Plovers at the atoll. At least a few were present every month but they were common only from late August to mid-May. This species apparently migrated from March through June, and mid-August through at least October. Ecology Golden Plovers occurred commonly in the open areas, both in the interior and along the beaches. TableAGP-3 lists by season the per- centage and location of plovers seen on shorebird counts from 1963 to 1965. Plovers were more common in the interior than along the beaches. During the winter a few Golden Plovers established territories near the barracks. The main seasonal distributional changes were a greater abundance on the beaches and along the runway in the fall, and a decrease in abun- dance in the central plain during the summer. Golden Plovers commonly occurred in pure flocks or with Ruddy Turn- stones, and occasionally with Wandering Tattlers or Sanderlings. Un- fortunately, numerical data on species association were not obtained. 22k Table AGP-1. Previous records of American Golden Plovers on Green Island, Kure Atoll. Population Date of Survey Estimate Remarks and References 1915 March 28 ) (Munter, 1915). 1923 April 17-22 fairly Found in central plain with turnstone common and on the beaches (Wetmore, mg.). 1957 June 5 2 Along north shore (Kenyon and Rice, 1958: 190). 1958 May 9 2 On beaches (Rice, pers. corr.). 1959 October 3-8 25 (Robbins, 1966: 53). 1960 March 28 25 (Robbins, 1966: 53). 1961 January 19-21 50 (Robbins, 1966: 53). September 12-14 14 On runway and beach (Udvardy and Warner, 1964; 2). 1962 February 2-4 500 (Robbins, 1966: 53). August 6-8 Dp) (Robbins, 1966: 53). 1963 February 3-7 20 (Robbins, 1966: 53). Banding and Movements The POBSP banded 88 American Golden Plovers and Robbins banded 13 (Table AGP-4). Two of these individuals were recovered on the island at a later date; one banded on 12 November 1963 was collected on 30 April 1967, and one banded on 20 October 1964 was captured on 2 March 1965. In addition, one banded on 31 October 1963 was found dead on Sand Island, Midway Atoll, on 1 March 1966. An adult Golden Plover banded at Zapadni on St. George in the Pribilof Islands on 8 August 1966 was collected on Green Island 26 days later. Another banded on Lisianski by Eugene Kridler on 11 March 1964 was mist netted at Kure on 1 September 1964. 225 Table AGP-2. POBSP semi-monthly estimates of American Golden Plovers on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 1-15 - 5 all - hO - 60 16-31 - © ye = = = 50 February 1-15 - * 56 70 * - 4O 16-28 * 87 46 - = s 35 March 1-15 - 65 50 - - - 35 16-31 - 137 38 - 12 - TK) April 1-15 - 87 yy - - = ho 16-30 - 87 Dale 46 - - 35) May 1-15 LO 87 4S 1B 20 - 25 16-31 - 6 20 12 12 - 10 June 1-15 - 37 20 12 7 10 16-30 - Ta Zi el 14 il * July 1-15 - 30 20 15 = 2 zi 16-31 - 13 20 14 - 3 - August 1-15 - 33 30 13 - 20 - ‘16-31 - (2 - 57 - 20 = September 1-15 - 100 - ho - LO - 16-30 * 121 ~ 67 - 55) - October 1-15 * 136 - - - 50 - 16-31 * 87 - - - 50 - November 1-15 250 95 = = = 50 - 16-30 (9 i 65 - - 50 - December 1-15 63 85 65 - - 50 - 16-31 70 80 - ho ~ 55 a * Birds present, number unknown. 226 Territoriality ee eee | Breeding plumage a: ji Migration —_ 100 Number of Birds Jan. Feb. Mar. Apr May June July Aug. Sept. Oct. Nov. Dec. Figure AGP-1. Annual cycle of American Golden Plovers on Green Island, Kure Atoll ( based on semi-monthly estimates). 227 Table APG-3. Percentage of American Golden Plovers recorded by season in different areas on Green Island, Kure Atoll, during 1963-65 shorebird counts. Area Winter Spring Summer Fall Ocean beach MteS 53 ILO sll 4.8 Lagoon beach 9.1 9.3 28.4 23/40 North point 0.4 Lo ® Os2 O.7 West point 44 6.9 Toll Boh Runway 9.6 1Lg® 10.6 4e.1 Central plain MSA WS) 1 2.4 I2E5) Rest of interior 46.5 Mor oS 46.9 14.8 Table AGP-+. American Golden Plovers banded at Kure Atoll. Year Banded Adults Immatures Unknown Totals 1959 4 8 O 12 1962 O O ald a 1963 O 0 Se Bi 1964 O 0 43 43 1965 0 0 4 4 1966 O O 3} 3 1967 O 0 a iL Totals y 8 89 101 BLACK-BELLIED PLOVER Squatarola squatarola Single Black-bellied Plovers were seen on 19 and 23 October 1963 and 20 May 1964. The latter was at the east end of the runway. Black-bellied Plovers, which are found infrequently in the North- western Hawaiian Islands, have also been recorded from Midway Atoll, Lisianski, and Laysan (Clapp and Woodward, 1968). RUDDY TURNSTONE Arenaria interpres Status Common migrant and winter resident, uncommon summer resident. Populations ranged from 2 to 150 birds. Most abundant in spring and fall. Least common in summer. 228 Populations Comparison of pre-POBSP observations (Table RT-1) and POBSP estimates (Table RT-2) reveal no significant differences in the size of the Ruddy Turnstone population, nor were significant differences observed during the course of POBSP studies. These data indicate population on the island at any one time of 70 to 150 during periods of peak abundance, and 2 to 20 during periods of minimal abundance. It was not determined how many turnstones used the island during the year, although it was probably considerably more than the maximum estimate . Table RT-l. Previous records of Ruddy Turnstones on Green Island, Kure Atoll. Population Date of Survey Estimate Remarks and References 1915 March 28 100 Small flocks along the shore (Munter, LOTS asia) 1923 ~=April dif=22 common Found in central plain and under dense Scaevola (Wetmore, ms.). 1957 June 5 3 Along north beach (Kenyon and Rice, 1958: 190). 1958 May 9 if (Rice, pers. corr.). 1959 October 3-8 50 (Robbins, 1966: 53). 1960 March 28 50 (Robbins, 1966: 53). 1961 January 19-21 150 (Robbins, 1966: 53). September 12-14 35-40 Groups on beach (Udvardy and Warner, 1964: 2). 1962 February 2-4 125 (Robbins, 1966: 53). August 6-8 20 (Robbins, 1966: 53). 1963 February 3-7 27 (Robbins, 1966: 53). Annual Cycle Figure RT-l presents graphically the annual population cycle, based On semi-monthly estimates and observations, of Ruddy Turnstones at the atoll. At least a few turnstones were present every month but they were 229 Table RT-2. POBSP semi-monthly estimates of Ruddy Turnstones on Green Island, Kure Atoll, 1963-69. 1963 1964 1965. 1966 1967 1968 196 January 1-15 - ey 125 - 50 - 150 16-31 - 715 79 - - - 80 February 1-15 - * 83 50 * - 60 16-28 * 112 107 - - - © March 1-15 - 87 105 - - - 100 16-31 - 87 90 - 39 36 120 April La) = 87 66 - - ~ 76 16-30 - 87 7 SAL - - 80 May 1-15 10 TK@) 25 26 43 - 50 16-31 - 4 25 27 21 12 20 June 1-15 - Tal, 30 2 20 14 4 16-30 - 13 30 29 16 10 * July 1-15 - 15 30 14 - 16 - 16-31 - 21 20 12 - 36 ~— August 1-15 - 76 oO Dale - 60 - 16-31 - 85 - 120 - 50 - September 1-15 - 86 - ie = 90 ~ 16-30 * 92 - 56 - 90 = October 1-15 * ra - - - * = 16-31 * 86 - - - * = November 1-15 150 145 - = = ¥ zs 16-30 15 100 80 = = x i December 1-15 50 105 80 - = * = 16-31 80 106 80 50 - 100 - * Birds present, number unknown. 230 Breeding plunage ES area! Migration ie ae ol (tg Rn | 100 Number of Birds & on [op) ~S [oe) wo (=) oS (<=>) (=) (=>) S w [=n] ipo) © 10 Jan. feb. Mar Apr May June July Aug. Sept. Oct. Nov. Dec. Figure RT-1. Annual cycle of Ruddy Turnstones on Green Island, Kure Atoll ( based on semi-monthly estimates). 25. common only from August through April. Although exact data were lacking, this species apparently migrated from mid-April through June, and August through October. Ecology Ruddy Turnstones occurred commonly in open areas, both in the in- terior and along the beaches. Table RTI-3 lists by season and location the percentage of turnstones seen on shorebird counts from 1963 to USO 6 Turnstones were about equally abundant in the interior and along the beaches. The main seasonal changes in distribution were a greater abundance of birds on the beaches and along the runway in fall, and a decrease in the numbers on the central plain during the summer, probably the result of thespresence or absence, respectively, of migrants. Ruddy Turnstones commonly occurred in pure flocks or with American Golden Plovers, and irregularly with Wandering Tattlers or Sanderlings. Unfortunately, numerical data on species association were not obtained. Ruddy Turnstones occasionally were found under, or on top of, dense Scaevola. In 1966 Woodward noted turnstones feeding on oatmeal that was being used as rat bait. Table RT-3. Percentage of Ruddy Turnstones recorded by season in different areas on Green Island, Kure Atoll, during 1963-65 shorebird counts. Area Winter Spring Summer Fall Ocean beach 15) 5 AS 5h 16 5) ILL GS) Lagoon beach 16.0 19.2 27.9 203 North point 336i 2.6 1.9 4.3 West point WES 52 5.6 1OpiL Runway 9.2 8.6 9.7 22.9 Central plain 16.5 17/ oth He STL 16.4 Rest of interior 25.1 29.6 ayn 3 wh 2 Banding and Movements The POBSP banded 24 Ruddy Turnstones and Robbins banded 7 (Table RT-4.) . One of these birds banded on 31 October 1963 was shot at the atoll on 29 August 1966, and another banded in November 1964 and re- captured on St. George, Pribilof Islands, Alaska, in August, 1967, was collected at Kure on 9 January 1969. In addition, one banded on 31 October 1963 was recaptured at Midway Atoll on 1 March 1966. Four adult Ruddy Turnstones banded on St. George Island, Pribilof Islands, Alaska, in August 1966 were collected at Kure later that year. 232 Table RT-4+. Ruddy Turnstones banded at Kure Atoll. Year Banded Number 1959 i 1963 4 1964 WN 1965 i 1966 al 1967 1 Total ail PINTAIL SNIPE Capella stenura On 13 January 1964 King collected a male Pintail Snipe in the central plain. This is the only record for the Northwestern Hawaiian Islands (Clapp and Woodward, 1968). COMMON SNIPE Capella gallinago delicata Schreiber collected an immature male Common Snipe on 25 September 1966 in the south antenna field. A snipe, probably this bird, had been seen daily for the preceding five days in the same area. This is the only record for the Northwestern Hawaiian Islands (Clapp and Woodward, 1968). BRISTLE-THIGHED CURLEW Numenius tahitiensis Status Uncommon migrant, summer resident and winter resident. Populations ranged from O to 12. Recorded in all months, but may sometimes be absent in summer. Most common in fall. Populations Early observers recorded Bristle-thighed Curlews infrequently and in small numbers (Table BIC-1). POBSP estimates (Table BTC-2) were Similar to earlier estimates, indicating total populations ranging from QO to 12, and wintering populations of 1 to 5. Annual Cycle Bristle-thighed Curlews were usually present all year with peak numbers from September through at least February (Fig. BTC-1). In some years (e.g., 1968) they were absent during the summer. 233 Table BTC-1l. Previous records of Bristle-thighed Curlews on Green Island, Kure Atoll. Population Date of Survey Estimate Remarks and References 1915 March 28 O (Munter, 1915). 1923 April 17-22 several (Wetmore, ms.). 1957 June 5 O (Kenyon and Rice, 1958). 1959 October 3-8 6 (Robbins, 1966: 53). 1960 March 28 1 (Robbins, 1966: 53). 1961 January 19-21 6 (Robbins, 1966: 53). September 12-14 carme2 Loose groups on the beach; also fed on the open and low grassy areas of the island interior (Udvardy and Warner, 1964: 2). 1962 February 2-4 10 (Robbins, 1966: 53). August 6-8 i (Robbins, 1966: 53). 1963 February 3-7 1 (Robbins, 1966: 53). Limited data suggest migration periods from late July through at least October, and in April and May. Many of the curlews seen in the fall were present for only short periods. In September 1964, 9 Bristle- thighed Curlews were banded and tagged with orange streamers. By mid- October only 3 tagged birds were still present. Ecology Unlike the other shorebirds, Bristle-thighed Curlews occurred more frequently in the interior of the island than on the beaches. Only 13.8 percent of the curlews recorded on the 1963 to 1965 shorebird counts were seen on the beaches, while 48.6 percent were seen in the central plain, 15.7 percent on the runway, and 21.9 percent in other areas of the in- terior. This species usually occurred alone or in small intraspecific groups. On three occasions, November 1963, November 1964, and December 1965, curlews were seen eating Polynesian rats. 234 Table BTC-2. POBSP semi-monthly estimates of Bristle-thighed Curlews on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 1-15 - 3 4 - 1 ; - a 16-31 - 3 n - - - i February 1-15 - * 6 9 ib - il 16-28 - 2 6 - - - 3 March 1-15 ~ 2 4 - - - 2 16-31 - 2 3 - 2 - 1 April 1-15 - 2 4 - - - 2 16-30 - 2 1 2 - - 0 May 1-15 - 2 2 2 ils - O 16-31 - O ili al O O ie June 1-15 - O 1 ay O O 16-30 - O if ib O 4 - July 1-15 - i ak i - O - 16-31 - 2 2 2 - O - August 1-15 - 2 2 iL - O - 16-31 - 2 - ie - 3 = September 1-15 - 5 - 6 - 2 = 16-30 * 12 - 2 - ih - October 1-15 * 4 - = = 0 a 16-31 * 5 = = = O = November 1-15 4 TF - = = il s 16-30 4 3 3 - - 1 - December 115 4 4 al - - I = 16-30) 3 D - 1 - aL - *Birds present, number unknown. 25)5) : 10 Number of Birds on J F M A M J J A S 0 N D Figure BTC-1l. Annual cycle of Bristle-thighed Curlews on Green Island, Kure Atoll (based on semi-monthly estimates; bars indicate migration periods. Banding and Movements Eleven Bristle-thighed Curlews were banded by the POBSP (9 in September 1963, 1 in October 1963, and 1 in January 1965) and 2 by Robbins in October 1959. An orange-streamered bird was seen at Midway Atoll on 7 December 1964 and 25 July 1965. ‘These may have been dif- herent birds. WOOD SANDPIPER Tringa glareola Bratley collected a Wood Sandpiper of unknown sex in the north antenna field on the evening of 22 May 1965. -On 15 May 1969 Kleen shot a male as it flew over Scaevola near the radar reflector. It had been seen earlier feeding on an albatross carcass along the southwest beach. Wood Sandpipers have also been recorded in the Northwestern Hawaiian Islands at Midway Atoll (Clapp and Woodward, 1968). WANDERING TATTLER Heteroscelus incanum Status Uncommon migrant, winter and summer resident. Populations ranging from 0 to 14. Recorded in all months, but sometimes absent in summer. Most common in fall and winter. 236 Populations Wetmore in 1923 was the first biologist to record Wandering Tattlers on the atoll (Table WTa1). Since that time the species has been recorded in small numbers by most visitors to the island. Recent POBSP estimates (Table WTa-2) indicated populations of fewer than 15 Wandering Tattlers at any one time. These estimates did not vary significantly from the earlier ones. Table WTa-1. Previous records of Wandering Tattlers on Green Island, Kure Atoll. Population Date of Survey Estimate Remarks and References 1915 March 28 ) (Munter, 1915). 1923 April 17-22 zig 2 (Wetmore, ms.). 1957 June 5 aL One emaciated individual caught along north beach (Kenyon and Rice, 1958: 190). 1959 October 3-8 4 (Robbins, 1966: 53). 1960 March 28 @) (Robbins, 1966: 53). 1961 January 19-21 4 (Robbins, 1966: 53). September 12-14 8-10 On beach (Udvardy and Warner, 1964: 2). 1962 February 2-4 315) (Robbins, 1966: 53). August 6-8 1 (Robbins, 1966: 53). 1963 February 3-7 10 (Robbins, 1966: 53). Annual Cycle Wandering Tattlers were recorded in all months, but in 1965 and 1968 they were absent in June or July. They were most common from mid- September through February of the succeeding year (Fig. WTa-1l.) Limited data suggested two periods of migration: one from late March to mid-June and another from mid-August through November. Ecology Tattlers were found mainly along the beaches and on the reef. The majority (57 percent) of those seen on the 1963 to 1965 shorebird counts 25h Table WIa-2. POBSP semi-monthly estimates of Wandering Tattlers on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 196 January 1-15 - 4 ital - 2 - 5) 16-31 - 4 14 - - - 2 February 1-15 - * iL i iL - 2 16-28 = 3 10 : - 2 2 Warch 1-15 = 3 t = - - a 16-31 - 4 2 = 2 = 2 April. 1-15 - 2 6 - - - 2 16-30 - 2 © 5 - - 2 May 1-15 5 2 8 vt 6 - 2 16-31 - 2 2 3 3 1 5 June 1-15 - 4 2 3 3 6 3 16-30 - iL ) 2 ) 4 x July 1-15 - 5 © iL - O - 16-31 ~ 0) O 3 - ) = August 1-15 - 3 if iL - 0 - 16-31 = 4 - 2 - 2 - September 1-15 - 2 - iL - 3 - 16-30 * 4 = 7 Mm nf October 1-15 us 9 = = = 5) - 16-31 * 6 - - - 5 ~ November 1-15 * ule - - - 5 S 16-30 8 3 2 - - 5 = December 1-15 8 5 = - 5 = 16-31 4 iL - - 5 - *Birds present, number unknown. 238 — ipo) ine) on (=) on — (=) Number of Birds Figure WTa-l. Annual cycle of Wandering Tattlers on Green Island, Kure Atoll (based on semi-monthly estimates; bars indicate migration periods). was along the rocky ocean beach. About 16 percent were seen along the lagoon beach, 15.3 percent at the west point, 7.2 percent at the north point, 3.1 percent along the runway, and 1.4 percent in the interior. Usually Wandering Tattlers occurred singly or in twos, but occa- Sionally they were found in small intraspecific or interspecific (with American Golden Plovers and Ruddy Turnstones) flocks. Banding and Movements One Wandering Tattler was banded in October 1959 and one in May 1969. No Tattlers banded on other islands were recorded at Kure. LESSER YELLOWLEGS Totanus flavipes On 30 August 1964 DuMont captured an immature female Lesser Yellow- legs in the central roost Sooty Tern colony. The bird was lying on the ground and was very emaciated. Other yellowleg observations of ques- tionable validity were made on 28 September, and 9, 10, and 18 October 1963. Lesser Yellowlegs have been recorded from Midway Atoll, Laysan, Maui, and Oahu in the Hawaiian Islands (Clapp and Woodward, 1968). 239 SHARP-TATLED SANDPIPER Erolia acuminata Only two Sharp-tailed Sandpipers were collected: one on / October 1963 and one on 29 October 1963. In addition, four were banded: one on 30 O&tober 1963, one on 31 October 1963, one on 2 November 1963, and one on 21 November 1963. Most of these birds were found in the open areas near the barracks. Sharp-tailed Sandpipers Have also been recorded in the Northwestern Hawaiian Islands from Midway Atoll, Pearl and Hermes Reef, and Laysan (Clapp and Woodward, 1968). PECTORAL SANDPIPER Erolia melanotos Six Pectoral Sandpipers were collected: one on 14 October 1963, three on 25 September 1964, and two on 21 September 1966. In 1964 they were found along the runway with Ruddy Turnstones and in 1966 they were found on the tennis court. Pectoral Sandpipers are regular visitors to the Hawaiian Islands and have also been recorded from Hawaii, Oahu, Midway Atoll (Clapp and Woodward, 1968) and Laysan (Ely and Clapp, ms.). SHARP-TAILED or PECTORAL SANDPIPER Erolia acuminata or melandtos Due sco, the aditticulity of yseparating, these two) species in the duiellid; only collected individuals were included in the respective species ac- counts. Other unidentified Erolia were recorded in late October 1963 (20), early November 1963 (3), 29 December 1963 (2), 14 January 1964 (2), 28 January 1964 (1), 5 February 1964 (1), 24 September 1964 (1), 26 Sep- tember 1964 (1), 19 November 1965 (1), and 14 August 1966 (1). DUNLIN Erolia alpina sakhalina Lewis collected a female Dunlin on 15 November 1965 in the area just south of the barracks. On 28 May 1968 Woodward saw another Dunlin in full breeding plumage at the east end of the runway with two Ruddy Turn- stones and three American Golden Plovers. Dunlins have also been recorded in the Northwestern Hawaiian Islands from Midway Atoll, Pearl and Hermes Reef, and Laysan (Clapp and Woodward, 1968). LONG-BILLED DOWLTCHER Limnodromus scolopaceus On 3 October 1963 an immature male Long-billed Dowitcher was col- lected by Ludwig. Other dowitchers were noted on 4, 6, and 24 October 1968. 240 This is the only definite record of this species in the North- western Hawaiian Islands (Clapp and Woodward, 1968). WESTERN SANDPIPER Ereunetes mauri Huber collected a male Western Sandpiper on 30 December 1966. This bird was seen on both the lagoon and ocean beaches. It was always seen alone although Sanderlings were nearby. This is the only record for the Northwestern Hawaiian Islands (Clapp and Woodward, 1968). BAR-TAILED GODWIT Limosa lapponica On 9 May 1966 Woodward saw two Bar-tailed Godwits roosting along the lagoon beach. One of these, a male molting into nuptial plumage, was subsequently collected. Bar-tailed Godwits have also been recorded in the Northwestern Hawaiian Islands from Midway Atoll, Lisianski and Laysan (Clapp and Woodward, 1968). RUFF Philomachus pugnax On 11 December 1963 Clapp saw a Ruff feeding with American Golden Plovers and Ruddy Turnstones in a rainwater puddle at the south end of the runway. A male, collected on the beach later in the day, had heavy fat deposits and was in winter plumage. This is the only record for the Northwestern Hawaiian Islands (Clapp and Woodward, 1968). SANDERLING Crocethia alba Status Uncommon spring and fall migrant, uncommon winter resident, and rare summer resident. Populations ranged from 0 to 13. Present from at least mid-August to mid-May of the succeeding year; in some years may also be found in the intervening period. Populations Sanderlings were first recorded from Kure Atoll by Wetmore in 1923 (Table Sand-1). Other early observers recorded this species infrequently and in small numbers. Recent POBSP estimates (Table Sand-2) were similar to these earlier estimates. 241 POBSP observations indicated that no more than 13 Sanderlings were present at any one time. Table Sand-1. Previous records of Sanderlings on Green Island, Kure Atoll. Population Date of Survey Estimate Remarks and References 1915 March 28 0 (Munter, 1915). 23 April a7 22 4 (Wetmore, ms.). 1957 June 5 ©) (Kenyon and Rice, 1958). 1958 May 9 i, (Rice, pers. corr.). 1959 October 3-8 iL (Robbins, 1966: 53). 1960 March 28 0) (Robbins, 1966: 53). 1961 January 19-21 2 (Robbins, 1966: 53). September 12-14 0 (Udvardy and Warner, 1964; 2). (1962 February 2-4 2 (Robbins, 1966: 53). August 6-8 O (Robbins, 1966; 53). 1963 February 3-7 6 (Robbins, 1966: 53). Annual Cycle Sanderlings were present from at least mid-August to at least mid- May of the succeeding year, with peak populations from September through November (Fig. Sand-1). Occasionally. they occurred in late May, June, July, and early August. Limited data suggested two migration periods: one from March to mid-May and one from mid-Augustthrough November. The lack of a spring peak similar to the fall peak may indicate that spring migrants followed a different route than the fall migrants. Ecology Sanderlings occurred mainly on the sandy beaches, usually alone or in small intraspecific groups. They were infrequently seen along the runway with flocks of American Golden Plovers and Ruddy Turnstones. The majority of Sanderlings (62.1 percent) seen on the shorebird counts of 1963 to 1965 was found along the lagoon beach. Other areas ake Table Sand-2. POBSP semi-monthly estimates of Sanderlings on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 1-15 - 4 3 7 - ial 16-31 - 4 3 - - - 11 February 1-15 - * 1 ik 4 - 9 16-28 5 mM 2 : Z 6 March Hie 1.5 a 5) 3 = - - 8 16-31 - 3 3 - 4 April 1-15 - af 2 = = = 9 16-30 - 1 © 3 : : é May 1-15 - 1 3} O O - i 16-31 - 0 3 2 © 2 June 1-15 = O AL O O alt O 16-30 - O O O O aL O July 1-15 - 0 O 4 - il - 16-31 - O O O - al - August 1-15 - O ile - * - 16-31 ~ 2 - 2 - iL - September 1-15 - dial - 3 - 2 - 16-30 1 13 - 2 - ile - October 1-15 * 12 - - - x - 16-31 * fe) - - - * - November 1-15 * W - - - * - 16-30 8 6 5 - rd * i December 1-15 8 2 O - - x a 16-31 5 2 - 6 - WO - 43 — (Sa) — Oo Number of Birds J F M A M J J A S 0 N D Figure Sand-1. Annual cycle of Sanderlings on Green Island, Kure Atoll (based on semi-monthly estimates; bars indicate migration periods). where they occurred included the east or ocean beach (14.3 percent), the north point (14.3 percent), and the west point (9.3 percent). No seasonal differences in this distribution were noted. a Banding and Movements No Sanderlings were banded by the POBSP nor were any found which had been previously banded. RED PHALAROPE Phalaropus fulicarius Fleet found a male winter-plumaged Red Phalarope with an injured wing on the beach near the east end of the runway on 13 March 1964. Another Red Phalarope in winter plumage was found dead in the north antenna field on 16 April 1969. Red Phalaropes occur uncommonly at sea in the central Pacific and have been recorded in the Northwestern Hawaiian Islands from Pearl and Hermes Reef and Laysan (Clapp and Woodward, 1968). RING-BILLED GULL Larus delawarensis A female Ring-billed Gull in first nuptial plumage was found dead on 22 February 1963 by Coast Guard personnel and obtained by Sibley. Ring-billed Gulls have been recorded from Pearl and Hermes Reef, Molokai, and Maui in the Hawaiian Islands (Clapp and Woodward, 1968). HERRING GULL Larus argentatus vegae Sibley collected three Herring Gulls in first winter plumage near the west point on 10 March 1963. Other specimens were collected on oll 10 November 1964, 20 January 1965, 8 March 1965, 1 April 1965, and 8 January 1969. Two of the latter were in first winter plumage; one was in furst nuptial plumage; and two were in second winter plumage. On 2 July 1968 Woodward found a well-bleached Herring Gull skull on the north point. Herring Gulls are regular visitors to the Hawaiian Islands and have also been recorded from Midway Atoll, Pearl and Hermes Reef, Lisianski, Laysan, and Oahu (Clapp and Woodward, 1968). SLATY-BACKED GULL Larus schistisagus Fleet and Standen collected an adult male Slaty-backed Gull in winter plumage on 9 March 1965. This is the only record for the North- western Hawaiian Islands (Clapp and Woodward, 1968). Robbins (1966) reported two "black-backed" gulls, possibly this species, 3-7 February 1963. GLAUCOUS-WINGED GULL Larus glaucescens On 10 March 1963 Sibley collected a Glaucous-winged Gull in first nuptial plumage near the west point. Prior to this the only records for the atoll were a mummy found on 18 April 1923 by Wetmore, and two seen 3 to 7 February 1963 by Robbins. From 24 December 1964 until 9 March 1965 Glaucous-winged Gulls were commonly seen on the beaches of Green Island, mainly near the dump at the west point. An estimated 6 were present in late December, 7 in early January, 4 in late January, 2 in mid-February, and 3 in late February and early March. Five Glaucous-wings were collected as follows: 24 December 1964, 1 February 1965, 1 March 1965, 7 March 1965, and 9 March 1965. These specimens were all immature birds; 3 were in first nuptial plumage, 1 was molting into second winter plumage, and 1 was in second winter plumage. Two other individuals were banded in late December 1964. When captured, one of these gulls regurgitated a rat, while the other one re- gurgitated rat fur and crab chelipeds. Three additional Glaucous-winged Gulls were collected: two on 31 December 1966 and one on 28 November 1968. One of the gulls collected in December was seen flying over and standing in the central plain. One of the Decemver birds was in first nuptial plumage; only the wing of the other was found and it also appeared to be in nuptial plumage. The third bird was in second winter plumage. Glaucous-winged Gulls have also been recorded in the Northwestern Hawaiian Islands from Midway Atoll, Pearl-and Hermes Reef, Lisianski, Laysan, French Frigate Shoals, and Necker (Clapp and Woodward, 1968). 245 GLAUCOUS GULL Larus hyperboreus On 3 January 1965 Kepler saw a Glaucous Gull on the beach. One or two individuals were seen during the rest of the month and one, a male in first winter plumage, was collected on 1/7 January. This species occurred most frequently at the west point early and late in the day. One Glaucous Gull fed in the lagoon or on dead fish that washed up on the beach. Wetmore found a beach-dried mummy on 18 April 1923. This species has also been collected at Midway Atoll, Laysan, Kauai, Lanai, and Maui in the Hawaiian Islands (Clapp and Woodward, 1968). BLACK-LEGGED KITTIWAKE Rissa tridactyla On 24 December 1964 two adult and one immature Black-legged Kitti- wakes were seen by Stadel and Kepler. One of the adults was seen for the last time two days later near the north point, and the immature, which was injured, was collected on 30 December. Hackman saw another adult kittiwake flying along the edge of the reef on 28 March 1967. The next day he found the mummified remains of another adult along the edge of the runway. Black-legged Kittiwakes have also been recorded in the Northwestern Hawaiian Islands from Pearl and Hermes Reef and Laysan (Clapp and Wood- ward, 1968). BLACK TERN Chlidonias niger On 20 June 1967 Woodward saw a Black Tern in breeding plumage flying north along the east beach. This constitutes the first record for the Northern Hawaiian Islands. They have been recorded on Oahu, and Maui in the Main Hawaiian Islands (Clapp and Pyle, 1968). ARCTIC TERN Sterna paradisea On 29 July 1968 Woodward found the mummified remains of an adult Arctic Tern along the southern edge of the runway. This was the first record of this species for the Northwestern Hawaiian Islands. Arctic Terns are occasionally seen at sea in the central Pacific. 246 SOOTY TERN Sterna fuscata Status Abundant spring - summer breeder; 2,000 to 10,350 pairs annually. Present from late January or February through, early November. Peak numbers from May through July. Breeding begins in late April or early May and continues through August. Populations Sooty Terns were recorded infrequently by early observers and prior to 1963 apparently did not breed on the island (Table ST-1). In that year an estimated 500 pairs nested at the northeast end of the run- way. Since that time they have bred every year and the population appears to be increasing (Table ST-2). Table SI=1. » Previous) records) of Sooty Terns on ‘Green Island; Kure Aycolale: Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 O (Munter, 1915). 1923 April 17-22 ©) (Wetmore, ms.). O57 siune! 5 3 Flying over (Kenyon and Rice, 1958:, 190). 1958 May 9 29 Flying over (Rice, pers. corr.). 1959 October 3-8 14 (Robbins, 1966: 53). 1960 March 28 O (Robbins, 1966: 53). 1961 January 19-21 0 (Robbins, 1966: 53). September 12-14 2 One found dead or injured (Udvardy and Warner, 1964: 3). 1962 February 2-4 0 (Robbins, 1966: 53). August 6-8 50 No nests found (Robbins, 1966: 53). 1963 February 3-9 ) (Robbins, 1966: 53). 247 Table ST-2. POBSP Semi-monthly estimates of Sooty Terns on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 1-15 - 0) 0) - O - iL 16-31 - 0 5 & a a 0 February 1-15 - O 5 ©) ©) - ©) 16-28 O * 30 - - - ho March 1-15 - * 100 - - - 2,000 16-31 - * 1, 000 - 750 150 2,000 April 1-15 = * 1,000 - - - 2, 000 16-30 - 4, 000 3, 000 5, 000 - - 5, 000 May 1-15 50 4, 000 6,060. IO;C00 “17,000 = 25000 16-31 - 4,000 3,000 io,000 17,000 15.000 25,000 June 1-15 - 4, 000 3,000 16,009 il7,000 i15,000 — 25,000 16-30 * 4,000 SOOO 15,000 /,000 5,00 * July 1-15 - 6, 000 6,000 16,000 . 17,000 - 15,000 - 16-31 - 6, 000 55000 15,000 - 5, 000 = August 1-15 - 6, 000 51000) OOOO - 2, 000 - 16-31 - 3, 000 ~ 1,000 - * - September 1-15 - 200 - - = x i 16-30 * x = = = * = October 1-15 * * - = = % 5 16-31 * * = = = * = November 1-15 * * - - - 0 - 16-30 O O O - - ©) - December 1-15 fe) 0 ©) - - 0 - 16-31 O ©) - iL - ©) - *Birds present, number unknown. 248 Available data indicate that Kure was colonized at least in part by Sooty Terns from Midway Atoll, 49 miles to the southeast. Thirty- six individuals banded on Midway from 1949 to 1962 were found breeding on Green Island 1964 to 1969. During POBSP studies large numbers of Sooties banded at Midway after 1963 were also found breeding (see Banding and Movements section). Undoubtedly the construction of the LORAN station created suitable habitat that was lacking éarlier. Another factor of possible importance was the purposeful harassment of the Midway colony, especially in 1957 and 1958, to prevent Sooties from breeding near the runway on Sand Island. Perhaps many birds from Midway moved to Kure after being harassed but did not breed until enough adults were present and there was suitable habitat. From 1966 to 1968 fairly accurate estimates of breeding populations were made by counting and marking all eggs found during the height of the breeding season. This method was believed to be at least 90 percent ac- curate. These counts indicated a population of 14,986 in 1966; 16,530 in 1967; and 5,700 in 1968. Table ST-3 breaks these counts down by colonys Table ST-3. Populations of Sooty Terns breeding in the various colonies on Green Island, Kure Atoll, 1966-69. Colony 1966 1967 1968 1969 Central roost 4,090 4704 1, 226 O North roost 1, 284 7, 746 O O Northeast colony 9,612 4,100 ©) O South antenna field O ) Aah 20,700 Except for 1968, these counts were considerably higher than the earlier estimates of 4,000 in 1964 and 7,000 in 1965. .Whether the size of the population actually increased or the higher counts were due to greater accuracy is not known. Assuming, however, that yearly egg and chick loss were fairly constant, it is unlikely that there was any significant change from 1964 to 1966, for in all three seasons an estimated 2,000 young fledged. In 1969 an estimated 20,700 individuals bred, but this estimate is not comparable to the more accurate counts. In 1967 and 1968 the structure of the breeding population was determined by walking through the colonies with a string of 100 bands, capturing the first definitely breeding bird seen, banding it if unbanded, recording the band number if it was banded, and then capturing the next closest breeding birds until 100 birds were banded. From these data the percentage of the breeding population that was banded and the history of banded individuals were determined. In both years at least 25 percent of the population was handled and all areas in the colonies were covered, thus making it as representative a sample as possible. Table ST-4 com- pares the 1967 and 1968 data. (In 1968 all Sooties banded in 1967 were considered an unbanded birds.) 249 Table ST-4+. Percentage of banded Sooty Terns in the breeding population on Green Island, Kure Atoll, 1967-68. Banded 1967 1968 Interisland 18.6 128 1964* PP 3 19.9 1965* 553 5 1966* 53.48 61.9 * All banded as adults. This analysis showed that the population structure was fairly consistent in both years and that the 65.6 percent decrease in 1968 was not the result of excessive adult mortality. Further support of this idea was provided by the rapid recovery in 1969 of the breeding population to the 1967 level. Unfortunately, the sampling of breeding birds in 1969 was insufficient to determine if this increase resulted from the influx of new birds; however, the structure of the population of banded birds remained relatively unchanged from the previous two years. Possibly a large segment of the 1967 population bred on other islands in 1968, probably Midway, or were present at Kure but did not breed. Sooty Terns which bred at Kure one year do breed on other islands as illustrated by two cases: one bird that bred on Kure in 1966 bred on Southeast Island, Pearl and Hermes Reef, in 1967; one that bred in 1967 at Kure bred in 1968 at Johnston Atoll. From the 1967 data it can be calculated that no more than 50 percent of the Sooty Terns that bred in 1966 bred again at Kure the next year. In 1967, 5,876 breeding birds (35.9 percent of the total breeding popu- lation) were handled. Of these, 1,751 (29.8 percent) had already been banded: 390 at Kure in 1964, 92 in 1965, 943 in 1966; and 326 on other islands. Of the 1966 birds, 669 were breeding when banded. Extrapolating for the total breeding population of 1967, there were 4,926 (.298 x 16,530) previously banded birds, 1,882 (.382 x 4,926) of which were breeding in 1966. In 1966, 4,400 breeding Sooty Terns were banded. Therefore, in 1967 only a maximum of 42.8 percent of these birds was breeding at Kure, suggesting a large amount of turnover, especially when we consider that the population increased slightly. Whether these adults were present but did not breed, whether they bred on other islands such as Midway, or whether Sooty Terns do not breed every year was not determined. The total number of Sooty Terns using the island at least once in 1967 was considerably larger than the maximum breeding population estimate of 20,700. With data collected from the breeding population samples, the total number of Sooty Terns using the island was calculated with the Lincoln Index as follows: 250 1967 1968 Number of adults banded on Kure prior to year of recapture 12,544 16,720 Number of previously banded Sooties found breeding on Kure 1,425 = OE Number of breeding birds captured By SO er, Percent of breeding populations that was banded at Kure PB 7 43.9 Total calculated population 48,809 38, 087 These figures are not unreasonable as over 16,000 adult Sooty Terns have been banded, yet only 50 percent of the breeding population in 1968 had been previously banded. Annual Cycle Sooty Terns usually returned to the atoll in February. The birds heard 8 to 10 January 1969 were probably stragglers rather than return- ing migrants. Returning migrants were first heard over the island at night on 23 February 1964, 29 January 1965, and 17 February 1969, but it was not until 28 February 1964, 27 February 1965, and 25 February 1969 that they were first seen. At first only a few were present, but they, increased in numbers until by the end of March hundreds were seen over the island during the day and considerably more heard at night. They were first seen on the ground in early March 1964, on 31 March 1965, and 27 March 1969. By the end of April thousands of adults were landing. During this early portion of the cycle the size of the population varied daily, depending in large part on weather conditions with few birds present on cool rainy days. Shortly before egg laying began, large flocks of Sooties swirled over the potential breeding areas, landed, and courted. Eggs were laid shortly thereafter, but were soon deserted. In 1966 Sooties began laying eggs in six areas, but deserted all nests within a day after egg laying began before finally settling in a permanent area. Usually egg laying began the first week of May, with a peak later that month; a few eggs were laid into late June. Peak periods of egg laying within the colony varied as much as three weeks between years (Table ST-5) and two weeks between the various subcolonies within one year (Table ST-6). Most eggs were laid within three weeks after the start of nesting, with a resultant sharply defined breeding cycle (Fig. ST-1). 2p Ce aun P=-PpTw euNe 34eT qseeT ye-LeW GZ Key ZT1-6 AeW 34eT -{jady Gg °eo 696T "69-1961T “‘TIOFV eany ‘pueTsSI ueerty uo aptoko Butpeeiq usa, AyooSs ey4 qysnsny ayeT-ysnsny + A[nNE yeem 4ST -ounf Yoom pre ATne Yoom prg-ounP OT eUNL YoeemM 4ST -AKeW yaom pre aune 34eT -Xew TT 9961 aunt? Té-T g-AEN Qe 4sn3sny yoom pre-ysnsny 6 aunt €g-2gT AqTne -ptu-oun? gT Kew So- ST “Kem Teroues e@ ut ATUO ayemMooe BYeCx 4Ssnsny 34eT -ATN 94eT asn3sny aunr Gt-+ (2) Aime ky 1e9e Aqnc -ptw-oune + ATnpe-sune keg 21-9 (3) APN O4eT aun P-pTw (3) Aen -Ttady 0€ ayeT-AeW Ss G96T *xh96T UL Spotized sz0feNW SUTSpaTA SuTYyo eH yeod SUT Yo VeYH sutheT say yeod SutkeT 33q potted °C-LS eTqIel ZO Table ST-6. Peak egg laying periods in various Sooty Tern colonies on Green Island, Kure Atoll, 1966-69. Colony 1966 1967 1968 1969 Central roost 15-22 May 2nd week May 3rd week May - North roost 15-17 May 3rd week May - ; - Northeast colony 20-25 May lst week May - = South antenna field = = Last week May- 9-12 May lst week June Adult Sooties were extremely vociferous during the egg laying period but once incubation began were only rarely heard. At this time most of the birds in the colony were nesting. Eggs hatched 26 to 31 days after laying (x=28.1; n=30). Usually hatching began in early June and continued until the third week of July, with a peak one month later than the egg laying peak. Adults were conspicuous during the hatching period, but within a week or so after most eggs had hatched, they were uncommon during the day. In 1968 when most nests failed, swirls, similar to pre-breeding swirls, were present during the first three weeks of July. Fledging began in late July and was completed by the end of August. Shortly after fledging young birds left the island. None returned for at least two years. When the chicks fledged, the adults essentially disappeared and were seen only infrequently from September until early November, usually flying overhead. Except for a single December and three early January records, Sooty Terns were not seen from mid-November until late January or February. Their destination after they leave the atoll is unknown. Nesting Success Table ST-7 summarizes productivity of Sooty Terns from 1964 through 1968 on Green Island, Kure Atoll. The low rate of success in 1968 was due mainly to rat predation on eggs and young. Hatching success was calculated for 80 eggs in 1965 as 53.5 percent, for 39 eggs in 1966 as 49.1 percent, and for 141 eggs in 1967 as 74.5 percent. The greater success in 1967 was due to the relative scarcity of Polynesian rats and the consequently reduced predation. Ecology Sooty Terns bred in several different areas on the island (Fig. ST-2 to 4). For convenience the colony areas were named central roost, north roost, south antenna field, and northeast colony (the area northeast of 253 1965 4,000 2,000 eggs | 1966 8,000- 4,000 eges, 1967 8,000 4,000 NO DATA 1968 3,000 2,000 eggs 1,000 1969 10,000 5,000 NO DATA J F M A M J J A S 0 N D Figure SI-l. Breeding cycles of Sooty Terns on Green Island, Kure Atoll, 1965-69. 254, Table ST-7. Productivity of Sooty Terns on Green Island, Kure Atoll, 1964-68. Maximum Nest Count Estimate Number Percent Year, or Estimate of Young Fledged Fledged 1964. 2, 000 2, 000 ; os 1965 3,500 2,100 60.0 1966 7,493* 2, 000 26.6 1967 8, 265* 2,000 a4 .2 1968 2,850* 50 16 (County. the runway and east of the south antenna field). In 1964 Sooties bred solely in the central roost after they were chased from the runway. From 1965 through 1968 the colony divided into smaller units that gen- erally did not breed in the same areas each year. Of the areas, the central roost was the only one utilized consistently from 1964 through 1968; it was deserted in 1969, possibly as a result of the spread of vegetation in that area. In 1969 Sooties bred only in the south antenna field. What factors control the selection of the breeding area is unknown, but Sooties swirled over several different areas before settling down. Since 1963 some, or all (1964), of the birds in the colony have attempted to nest along the runway in the sand-Eragrostis association but have been chased away because of the potential danger to aircraft. If left unmolested, it is likely that they would breed there year after year. At Kure, Sooty Terns bred in two major habitats--open areas of bare sand and scattered vegetation such as Eragrostis, Boerhavia, Tribulus, and Verbesina, and under Scaevola. Usually one egg was laid, but in some nests two eggs were found. In 1966 ten of 7,493 (0.13 percent) nests counted contained two eggs, while in 1968 eleven of 2,850 (0.39 percent) had two eggs. Whether the same female laid both eggs was unknown. The egg was usually placed on bare sand but in a few cases was laid on top of matted vegetation, or, more rarely, in the midst of Eragrostis clumps. There was no evidence that adults modified the surroundings prior to egg laying. Rather, a slight depression was formed as incubation progressed. Although Sooties bred in open areas, most eggs were laid at the edge of some vegetation. Egg densities were determined for six 100-square-foot plots in 1965, three in 1966, and six in 1967. In 1965 the number of eggs ranged from 4 to 22 (xX=13.3); in 1966 from 4 to 19 (x=13.0); and in 1967 from 11 to 38 (%=23.5). 255 "GQ-"96T “TTOVY emmy “pueTSI usez) uo severe SuTpseiq Ute] kyoog FO uoTINGT.4STC °2-LS emst a 256 "19-996T ‘TT°4V eany ‘pueTs—T usery uo severe SuTpeerq use, 900g Jo uoTANgGT.z4STC °€-I9 emst a | - "69-996T ‘“TTOV eMMy ‘pueTST userp uo severe SuTpeserq use], AJZOog JO uoTINATsAST "+H-LS east 258 Non-breeding Sooty Terns roosted on the beaches and in the breeding colonies, or flew over the colonies. Although many Sooties were found roosting on the beaches prior to egg laying, no case of egg laying in these areas was known. In 1966 ca. 1,000 Sooty Terns roosted on the beach adjacent to the northeast colony for several days in late May. Polynesian rats preyed heavily on Sooty Tern eggs and young. Yearly differences in hatching success were in large part attributable to the population size of these mammals. Generally, Sooties did not breed in the same areas as the other tern species. In 1967 the Gray-backed Tern colony was engulfed by Sooty Terns in late May, but no detrimental effects were noted. However, in. 1969 the Sooties apparently destroyed several Gray-back nests when the Sooties moved into the south antenna field. Banding and Movements The POBSP banded 18,534 adult Sooty Terns. Tables ST-8 and 9 sun- marize the recapture rates of these birds in subsequent years at Kure. As would be expected, breeding Sooties were recaptured more frequently than non-breeders. Although large numbers of adult Sooty Terns were usually handled each year (Table ST-10), these recapture figures are low, especially for the 1966 and 1967 breeders. Only in 1966 and 1967 were large numbers of breeders handled; in 1968 the breeding population was greatly reduced and in 1969 only a relatively few breeders were handled. Thus, the chanees of recapturing a 1966 or 1967 banded bird were reduced. Not all adult Sooty Terns banded at Kure were recaptured there. One hundred and forty-five of them were captured on other islands: 133 on Eastern Island, Midway Atoll; 5 on Southeast Island, Pearl and Hermes Reef; 2 on Laysan; 4 on Sand Island, Johnston Atoll; and 1 on Wilkes Island, Wake Atoll. All but four of these birds were banded as non- breeders. However, 105 were breeding on the island of recapture (100 at Midway Atoll, 4 at Pearl and Hermes, and 1 at Wake Atoll). In addition, the POBSP banded as nestlings 5,569 Sooty Terns (1,601 in 196//; 1,600) in 1965's). 1.600 “in 1966; 500 in 19673) and 63 im W966)F However, an undetermined number banded in 1965 were actually adults since three of those banded as nestlings in 1965 were found breeding the next year. By June 1969 20 (1.3 percent) of the 1964 cohort had been recap- tured at Kure (4 in 1966, 10 in 1968, and 6 in 1969). Eight of the 1965 cohort. had also been captured (2 in 1968 and 6 in 1969), but it is possible that these birds were adults when banded. None of these Sooty Terns was found breeding. Two Sooty Terns banded as nestlings were captured on Sand Island, Johnston Atoll: one banded on 24 July 1964 was found on 12 August 1967 and another banded on 26 July 1965 was captured on 30 August 1967. 259 Table ST-8. Recapture rates of adult Sooty Terns banded as breeders at Kure Atoll and recaptured there (expressed as percent- ages), 1965-69*. Year Recaptured Year Banded n. 1965 1966 1967 1968 1969 1965 145 100.0 47.6 26.2 10.3 505) (28.9) (16.6) (6.9) (5.05) 1966 4,400 -- 100.0 227 9.3 3.6 (15.2) (6.1) (@-6) 1967 hh 25) -- -- 100.0 Tole BE6 (4.3) (3.6) 1968 800 -- -- -- 100.0 259) (2-9) 1969 769 -- -- -- -- 100.0 Table ST-9. Recapture rates of adult Sooty Terns banded as non- breeders at Kure Atoll and recaptured there (expressed as percentages), 1964-69*. Mears Year Recaptured Banded filo. MO 1965 1966 1967 1968 196 MSGS OO | LOOL0 29.0 Pilosk i 5) 550 Dol. (2,9) (15.8) C@s6)) (3.2) (Aoi) 1965 468 -- 100.0 hoy 22.2 Gaal. 3.6 (2ASc%) (als 4) (4.7) (3.6) IG n/t -- -- 100.0 HLL 4.3 alge (8.0) (Zar) (Loi) 1967 Bil -- -- -- 100.0 20 2.0 (0.0) (2.0) 1968 173 -- -- -- -- 100.0 0.6 (0.6) 1969 72 -- -= “= -- -- 100.0 * First figure represents the percentage of birds known to have been alive and the second figure is the percentage of birds captured. 260 Table ST-10. Number of adult Sooty Terns handled each year at Green Island, Kure Atoll, 1964-69. 1964 1965 1966 1967 1968 1969 Breeding ©) 181 55022 5,890 1, LO WET Non-breeding 3, 808 546 4,001 99 ekg 133 Totals 3, 808 727 9,023 5,989 LISS) 1,410 Movement to Kure Atoll of Sooty Terns banded on other islands was considerable. In four years of detailed breeding population study, 501 adult and 3 nestling Sooty Terns from other islands were found breeding at Kure. The majority (495) had been banded at Midway Atoll (36, in- cluding the 3 nestlings, prior to 1963) but there were also 3 from Laysan, 5 from Sand Island, Johnston Atoll, and 1 from Wilkes Island, Wake Atoll. At least 315 of the Midway-banded Sooties were recorded as breeding at the time of banding. How commonly this interchange of breeding birds occurs is unknown. Besides the Sooty Terns found breeding, another 65 birds banded as adults and 6 banded as nestlings on other islands were also captured. Again the majority was from Midway Atoll (59), but there were also 2 from Southeast Island, Pearl and Hermes Reef, 1 from Laysan Island, and 9 from Sand Island, Johnston Atoll. The Sooty Terns banded as nestlings were from Midway (5) and Johnston Atoll (1). The lack of movement from French Frigate Shoals and the main Hawaiian Islands is surprising and at the moment unexplainable. Two recorded movements involving Kure and the central Pacific equatorial islands are probably in error. On 11 May 1965 an adult with band number 793-59416, presumably a Phoenix Island bird, was recaptured at Kure. By changing the 59 to 95 it becomes at Kure band. Another band number 813-90913, can be changed to a Kure band by changing the O to al. Since these band numbers can be converted so easily to anes used on Kure, and since neither was verified by a specimen or noted as unusual at the time of capture, it is best to consider them highly questionable. Although the number of Sooty Terns banded on other islands was large, when expressed as a percentage of the total number of birds banded, the amount of movement was relatively small. For example, 113,744 Sooty Terns were banded at Midway Atoll by the POBSP prior to 1967, yet only 517 (4.6 percent) were recaptured at Kure. On Laysan 117,311 Sooties were banded prior to 1968 and only 4 (0.003 percent) were found at Kure. 261 GRAY-BACKED TERN Sterna lunata Status Uncommon spring-summer breeder; 20 to 39 pairs annually. Present from February or March until mid-September. Breeding begins in early April and continues to at least mid-August. Populations POBSP Gray-backed Tern population estimates (Table GBT-1) were larger than previous estimates (Table GBT-2), which is not surprising Since these earlier trips were of short duration and only two, April 1923 and June 1957, occurred at the peak of the breeding cycle. However, the Gray-back population evidently changed little in size from 1923 when Wetmore, in late April, considered them "fairly common," a term applicable to recent POBSP April estimates. Even POBSP data showed large differences in population estimates over a relatively short period. For example, in early July 1964 the maximum estimate was 4, but in late July 13 young Gray-backs were found which had been overlooked the previous two months. POBSP estimates ranged from 50 to 100 Gray-backed Terns during the peak breeding periods. Most of these estimates were based on the number Cmpundsmscen) tne the breeding area andewenrenmuaererore, conservative, in 1967 and 1968 the most accurate data were obtained by finding a majority of the nests. Based on these nest counts, the breeding population in 1967 was 78, and in 1968 was 52. Since many nests were probably destroyed before observations began in 1968, the difference was probably not sig- nificant. Earlier estimates actually agreed fairly closely with the accurate data, if nest losses were considered. For example, in 1964, 15 young Gray-backs were found in July and August. Based on Sooty Tern data, there was at least a 50 percent nest loss, so these nestlings represented 30 eggs at the beginning of the season, suggesting little or no change from 1964 to 1968. Annual Cycle In February of March, after an absence of at least four months, Gray-backed Terns returned to the atoll (earliest records: 6 February 1969 and 11 March 1964). Whether they were absent in February 1964 and 1965 or were merely overlooked is unknown. At first Gray-backs were present Only at night, flying over the island or roosting on the beaches. By mid- March 1969, however, they were also present during the day. Evidently the population arrived en masse rather than building up slowly. Maximum numbers of Gray-backed Terns were present from late 262 Table GBI=1. January 1-15 16-31 February 1-15 16-28 March 1-15 16-31 April 1-15 16-30 May 1-15 16-31 June 1-15 16-30 July 1-15 16-31 August 1-15 16-31 September 1-15 16-30 October 1-15 16-31 November 1-15 16-30 December 1-15 16-31 *Birds present, number unknown. POBSP semi-monthly estimates of Gray-backed Terns on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 = 19 i - te 90 1S 90 (5 90 1 90 50 90 30 - 50 = ¥ co 2) = O = 1968 196 263 Table GBT-2. Previous records of Gray-backed Terns on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 12 1 egg at the foot of bunch grass on bare ground (Munter, 1915; 137). 1923 April 17-22 fairly None nesting (Wetmore, ms.). common 1957 June 5 8 Flying over (Kenyon and Rice, 1958: 190). 1958 May 9 1 Flying (Rice, pers. corr.). 1959 October 3-9 O (Robbins, 1966: 53). 1960 March 28 0 (Robbins, 1966: 53). 1961 January 19-21 0 (Robbins, 1966: 53). September 12-14 2 1 dead or injured bird (Udvardy and Warner, 1964: 3). 1962 February 2-4 O (Robbins. 1966s) 53). August 6-8 hO 20 nesting pairs (Robbins, 1966: 53). 1963 February 3-7 ©) (Robbins, 1966: 53). March to mid-August. Numbers decreased rapidly in late August and by mid-September Gray-backs had disappeared. In 1968, when no young fledged, the decrease occurred in mid-July. Pre-breeding behavior was not recorded in detail, but the 1966 data: were probably typical. During the last two weeks of April, Gray-backs were common throughout the day and night in open areas just north of the runway. At least one egg, which was later deserted, was laid under an Eragrostis clump. By early May they were not seen in these areas and, in fact, very few were seen at alll. In late May the breeding colony was found under Scaevola in the north roost. While most Gray-backs shifted areas, one pair started breeding along the runway and another one along the north- easupbeach, | chs) shitting onthe colony, from jche open areas) tothe north roost accounted, in part, for the difficulty in 1964 in locating the colony. Egg laying began one to two months after the birds returned. Munter found an egg on 28 March 1915, but there have been no recent egg dates 264 this early. The earliest POBSP egg records were 10 April 1965 and 9 April 1967 (with interpolated data). Egg laying continued through the third week of June. Sufficient data were collected only in 1965, 1967, and 1968 to determine peak egg laying periods. In the former two years egg laying began in early April. In 1967 the peak of egg laying occurred the last two weeks of April, while in 1965 it was not until the last two weeks of June, at least for eggs that hatched. This suggests that the eggs laid in April 1965 were destroyed and the latter egg peak was due to renest- ing. In 1968 the peak occurred the last week of April, slightly later than the 1967 peak. Combined data show an egg laying peak from mid-April to mid-May (Fig. GBT-1). 35 25 15 Number of Eggs J F M A M J J Figure GBT-1. Number of Gray-backed Tern eggs laid each semi- monthly period on Green Island, Kure Atoll, 1967-69. Hatching began in mid-May (earliest record: 9 May 1967) and con- tinued until the third week of July (latest record: 21 July 1968), with the peak about a month after the egg laying peak. Most eggs hatched by the end of June. The incubation period was unknown. Fledging began as early as late June (1967) and was completed by the third week of August. : Figure GBT-2 shows the number of eggs and young present for each semi-monthly period where data were available. The most accurate data were for 1967 and 1968. Nesting Success Table GBT-3 shows the productivity of Gray-backed Terns from 1964 through 1969. The complete destruction of eggs and young in 1968 was probably the result of rat predation. ° 265 1965 30 20 eggs 10 Oy ine 1966 20 eggs 10 ae 1967 1968 15 10 eggs 5 ZN 1969 NO DATA ee ee A ei a One One ONS Figure GBI-2. Breeding cycles of Gray-backed Terns on Green Island, Kure Atoll, 1965-69. 266 Table GBI-3. Productivity of Gray-backed Terns on Green Island, Kure Atoll, 1964-69. Maximum Maximum Maximum Nest Egg Nestling Approximate Count or Count or Count or Number Percent Number Year Estimate Estimate Estimate Fledged Fledged Banded 1964 25 O 25) 25 -- Ny 1965 25 25 9 7 28.0 9 1966 20 20 5 2 110) 50) O 1967 39 34 29 28 Wales) 28 1968 26 19 10 © “O50 9 1969 20 15 12 -- -- 2 In 1967, 24 of 34 marked eggs (70.6 percent) hatched. The comparable figure for 1968 was 6 of 19 (31.6 percent). No fledging success rates were determined, mainly due to the difficulty of finding the young Gray-backs under the Scaevola. Ecology Gray-backed Terns generally bred in small colonies under Scaevola in the north roost (Fig. GBT-3). Only in 1964 and 1965 was the main colony in the same general area. Solitary nests were found under Scaevola along the runway and northeast beach in 1966, and near the south antenna field in 1969. Munter (1915) found an egg under a clump of Eragrostis; only one nest (1966), which was later deserted, has been found recently in a similar situation. In 1967 there were 39 nests in ca. 4,365 square feet, or one nest per 112 square feet. The single egg was placed on gravel under Scaevola, gen- erally near the base of the bush. Ten nests averaged 3.75 inches from the trunk of a Scaevola bush, and were under Scaevola of an average height of 32.2 inches. Although there was much vegetation on the ground in the area, none was present in the nests. Non-breeding Gray-backed Terns roosted on the open beaches, mainly at the north point and other open areas in the north roost. Many Gray-backs flew over the central plain catching moths on the wing. This was the only seabird species that caught food on the island. Banding and Movements The POBSP banded 50 adult Gray-backed Terns on Green Island, Kure Atoll, and Robbins banded 2 in August 1962. Table GBT-4 summarizes the recapture of these birds. None was found on other islands, nor were any Gray-backed Terns banded on other islands found at Kure. Only one of the 63 nestlings banded was recaptured--one 1964 bird in 1969. 267 *peanqydeos spatq jo esequeozed ay. st amnSTy puooss ay} pue SATTS Useq eABY OF UMOUY Spatq Jo asequaedzed ayy squesetded eanSTJ iSsITaq x 0°O0T == = == == == == = c 696T (0°0) 0°0 0°OOT Ss == = == == == T 896T (0°0) (2°9T) 0°0 2°QT 0° OOT -- -- -- -- -- EI L961 (0°0) (0°0) (0° Se) 0°0 0°0 0° Sg 0°O0T == =< -- -- 4 996T (0°0) (6°S) GS2e2) = (ear) 0°0 6°S G° 2? 2] Ge 0°OOT -- -- -- Ae S96T (0°0) (2°9) (1°9) (1°9) (0°0) 0°O L°9 Caer 0°O¢d 0°02 0°OOT -- 2 CT H96T (0°0) (©) (0°0) (0°0) (0°O) (0°0) (0°0) 0°0 O20 ©°O 0°0 0°0 0°0S 0°0S 0°OOT c Aeron 996T L96T 996T GS96T H96T €O6T c96T pepueg poingdedsay eax Leak °¥69-296T ‘(SeSequeotad se pesserdxa) areyy peanqdeoes pue TTOVV einy 7e pepueq sural peyoeq-Aerp y[Npe fo sejyer singdeosy “+4-1d) eTqeL 268 "69-H96T “TTOVV SAMY ‘pueTsT user) uo suze peyoeq-fery Sutpserzq JO uotynqtszastq °¢-rEy sIN3T IT 6961 8961 S96L'VO6L 33% 6961 @ 8961 O 2496.0 9961 ¥ S96L 269 BROWN NODDY Anous stolidus Status Abundant spring-summer breeder; 400 to 800 pairs annually. Present from March through December. Usually absent in January and February, although there were two records for January. Breeding from late April through early November with a peak in June and July. Populations Comparison of pre-POBSP Brown Noddy records (Table BN-1) with POBSP estimates (Table BN-2) indicate that this species increased in numbers after the construction of the LORAN station, which was not sur- prising because much of the area (mainly along the runway) now utilized by Brown Noddies was covered with dense Scaevola prior to 1961. Brown Noddies were one of the easiest species to enumerate because of the conspicuousness of the birds and their nests. Recent POBSP esti- mates indicated a maximum population of 1,000 to 1,700 birds. Table BN-l. Previous records of Brown Noddies on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 O (Munter, 1915). 1923 April 17-22 ? (Wetmore, ms.). 1957 June 5 252 66 eggs found (Kenyon and Rice, 1958: 190). 1958 May 9 70 (Rice, pers. corr:). 1959 October 3-8 500 Nesting (Robbins, 1966: 53). 1960 March 28 ©) (Robbins, 1966: 53). 1961 January 19-21 0 (Robbins, 1966: 53). September 12-14 2 (Udvardy, 1961: 46). 1962 February 2-4 0 (Robbins, 1966: 53). August 6-8 200 75 nests (Robbins, 1966: 53). 1963 February 3-7 i (Robbins, 1966: 53). 270 Table BN-2. POBSP semi-monthly estimates of Brown Noddies on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 1-15 = O 3} - * = fe) 16-31 . ©) 150 - - - 0 February 1-15 - O O 0 O - 0 16-28 O O O - ~ - 0 March 1-15 - 0 2 - ~ - 0) 16-31 - 55 400 - ©) @) 200 April 1-15 - 250 400 - - - 400 16-30 - 750 500 500 - - 400 May 1-15 500 750 700 600 15650 - 400 16-31 - 750 900 700 1,650 500 400 June 1-15 - 750 OO 900 1,650 1, 000 450 16-30 * 750 3 50 900 1 6510 1,200 * July m5 - 600 900 900 INO 1,200 - 16-31 - 1,000 750 900 - 1, 200 - August 1-15 - 1, 000 750 800 ~ 1,200 - 16-31 - 1, 000 - 700 - 800 - September 1-15 - 800 ~ “L400 - 800 - 16-30 * 500 - 400 - 600 - October 1-15 * 400 - = a * Ye 16-31 * 250 - - - 300 - November 1-15 400 150 ~ - - 300 - 16-30 715 15 100 - - x E December 1-15 75 50 20 ~. = 4 a4 16-31 0 14 i * x 0 eS *Birds present, number unknown. ———— a Call In 1965, 1967, and 1968 population estimates were based on counts of adults and nests (Table BN-3). These data suggest that the total population changed only slightly from year to year. However, the number of noddies breeding each year varied--1,000 in 1964, 1,056 in 1965, 802 in 1966, 1,568 in 1967, and 978 in 1968. The reason for these fluctua- tions are unknown. No counts were made in 1969 so it is impossible to determine if the apparent decrease that year was real. Table BN-3. Comparison of Brown Noddy counts on Green Island, Kure Atoll, for 1965, 1967, and 1968. Date of Count 1965 1967 1968 March 14 2 -- -- 21 1 -- -- 27 il -- -- Mopesial | 8) 99 =a oe 10 IBS -- -- 18 365 -- -- 2h 336 -- -- May 1 428 -- -- 8 594 739 -- U5) DiS 624 fe == 655 627 -- 29 770 681 sabes Juae: 5 914 -- -- 6 -- 760 -- 8 a a 367 12 784 -- se 14 -- 858 552 22 -- -- 564 26 869 -- -- 29 << -- 3 soy 6 -- -- 719 10 Why -- -- 13 -- -- 614 == ae -- 645 ak 490 =e ee 29 a, -- 730 aie Annual Cycle Brown Noddies were generally absent from mid-December until the last two weeks in March (earliest records: 30 March 1964, 14 March 1965, and 19 March 1969). Following an initial appearance on the sand- spits west of Green Island (in 1969 they were first seen at the north end of the island), they invaded the southern beaches of Green Island, moving gradually northward and encircling the entire island; then they started flying over the interior. By mid-April they occupied all of their breeding areas. Peak Brown Noddy populations were present from May through August. Courtship was first observed on 9 April in 1964 and 6 April in 1965, but it was not until May in both years that eggs were laid. Al- though the first eggs were laid at approximately the same time each breeding season, yearly differences in breeding peaks existed (Table BN-+). For example, in 1967, 512 nests were present in late May, while one year later only 30 were found in that month. Figure BN-1 shows the number of eggs and young present each semi-monthly period for which there are data. Egg laying began as early as late April (1967) and some eggs were laid as late as late August. Most egg laying, however, was completed by mid-July. Usually there was an extended period of egg laying, with numerous eggs laid one to two months after the start of breeding, which resulted in a broad egg peak. However, in 1968 most eggs were laid during a three-week period. Figure BN-2 compares the number of first eggs laid each week at the north point in 1967 and 1968. The 1968 pattern may be explained in part by the lateness of the breeding cycle. POBSP data showed that peak periods of egg laying in this species occurred no later than early July. Therefore, if breeding was delayed, most noddies would reach breeding condition at the same time, with a resultant sharp egg laying peak as in 1968. Usually only one egg was laid in a nest, but in 1968 two of 89 nests (2.2 percent) at the north point contained two eggs. Neither nest pro- duced young. On Kure eggs hatched 34 to 40 days after laying (x=36.1, n=76). Eggs hatched from late May until late September. Hatching peaks occurred from the last two weeks of June through July and showed the same yearly varia- tions as the egg laying peaks. On the average the young birds began’to fly 46.5 days after hatching (n=66, range 40-56). Fledging, therefore, began in late July and continued until at least early November. Most young began flying in August and September. After they left at the end of the breeding season, young Brown Noddies did not return to Kure for at least two years. is I3Q0990 AT Ie38-ysnsny 3 Yaem SITS g-ATNP-pTw Aqne oune é YOoM SCT SY¥9OM CZ ISBT qsnsny j ayeT-amL ZZ g-Ken O€ Aqne Yyoem 4ST-oune 3 SY9eM Z YSeT Ke Laquay aun,” jo -deg AqTzee yazed yseeT 4e a-Kew 1, -Keyl QT =-[Tady 93eT L96T °69-196T *‘TLOVV emmy ‘pue[TSI Uusery uo aTofd SutTpaerq AppoN uUMorTgG ay} UT Spotted sole di, -isnsny AqTzee AqTne syaom co FS4ATE qysnsny ayeT-eunl OF aune syaom co ts84TF Aq[ne a yeT-Aen SL 99OT g-ATNL 94eT AINE 34eT -aunf 23eT g-9UNl PT aun p- ptu=- Aen YooM {SCT qsnsny A[Iea YSesT ye-KeW Q ZTIQUIAON=PTU -Aqn¢-pTw g-ATNE 94eT Iaqueydes gg-eunr JZ aun qsnany aqyeT-AeW ET SuTSpaTA SUuTUOV eH yeod SUTYOVeH Ssutfkey s3q yeod Sutkey 33 potted “H-N OTABL 274 1965 NO DATA 1966 400 200 1967 600 400 200 young NO DATA 1968 400 eggs 200 1969 100 eggs 50 NO DATA J i M A M J J A S 0 N D Figure BN-1L. Breeding cycles of Brown Noddies on Green Island, Kure Atoll, 1965-69. 2 "Q9-L96T “TTO9V ezny ‘pUeTST USserH uO eeze Lpnyas quTod y410U 4e Yyeom Yoee pteT ss#se AppoN umOTG FO Laqunyl °Z-Nq@ eammst7T ysn3any Ayng auny Kew Ol—y €-8% LZ-1Z OZ—-vl E1-L 9-OE 67-E% ZZ-91 GSI-6 8-2 I-92 SZ-6L S8l-Z tl-S y-8z ol =a Ss Og. = ® o OG a. nn Or 896l 0S Viva ON S]SON JO Jequiny 276 Nesting Success Detailed nesting success data for Brown Noddies were collected at the north point in 1967 and 1968. In the former year, 26 of 33 eggs (78.8 percent) that should have hatched by 20 June did so. Comparable figures for 1968 showed that 59 of 93 eggs (63.4 percent), or 59 of 89 nests (66.3 percent), hatched. Young fledged in 1968 from 38 of the 89 nests (42.7 percent), representing 64.4 percent of the young that hatched. Table BN-5 summarizes productivity of Brown Noddies from 1964 through 1968. Table BN-5. Productivity of Brown Noddies on Green Island, Kure Atoll, 1964-68. Maximum Nest Count Approximate Number Percent Year or Estimate of Young Fledged Fledged 1964 500 250 50.0 1965 528 260 49.2 1966 400 187 46.8 1967 784 450 Ni oh 1968 489 210 42.9 Ecology Brown Noddies bred in most major habitats on the island, from open beaches to dense Scaevola thickets. Figure BN-3 shows the general location of the 1967 breeding areas. This was a typical pattern for all years. Although they bred over a considerable portion of the island, they avoided several areas such as the central plain and northeast beach. Only 13.5 percent of the nests were located in the midst of Scaevola, The majority, 33.8 percent, was located along the lagoon beach at the beach-Scaevola ecotone. Lesser concentrations were found along the western edge of the runway (15.8 percent), the west end of the runway (10.2 percent), and the north point (13.4 percent). A similar distribu- tion pattern was noted in 1968. Nest construction of Brown Noddies was as variable as their nest sites. Four general types of nest were found, varying in location and the amount of vegetation. Most nests were on the ground--some with no vegetation added (Type I), some with a little vegetation (Type II), some with large amounts of vegetation (Type III), and some bulky nests were built in Scaevola (Type IV). In 1967 all nests found were classified according to the preceding types with the following results: Type I - 208 (30.4 percent), Type II - 410 (59.8 percent), Type III - 26 (3.8 percent), and Type IV - 41 (6.0 percent). AUT *J96T “‘TTOVV etny ‘pUBTSI USseZD UO SETPPON UMOTE suTpeetq FO UOTINGTIASTA "€-Nq omMsT aA 278 Scaevola, Eragrostis, Boerhavia, Ipomoea, and algae were the main types of vegetation present in the nests. Many nests also contained sea urchin shells, ropes, crayons, and/or other debris. During the day Brown Noddies commonly roosted on the beaches and on coral blocks in the water. At night they roosted on Scaevola, mainly along the lagoon beach. Some noddies were also found on man- made structures such as the radar reflector. In the summer of 1965 many young Brown Noddies were found with open wounds in the body cavity, just ahead of, or behind, the wings-- a typical sign of Polynesian rat predation. Rats were also observed eating noddy eggs. It is well known that downy Brown Noddies are polymorphic, varying mainly in the color of body down. Usually the color information given is the proportion of dark and light birds in the population. However, considerably more variation than has been recorded in the literature was found at Kure. Basically downy Brown Noddies vary in three characters: (1) amount of white on the head, (2) color of the abdomen, and (3) color of the rest of the body down. The following six classifications were used for the amount of white on the forehead: (1) none, (2) lore white, (3) forehead white, (4) one-half of head white (area from bill to just above the eyes), (5) top of head white (to posterior portion of skull), and (6) whole head white (extending down neck). The abdomen may be: (1) white, (2) the same color as the rest of the bird (dark), or (3) somewhat lighter than the rest of the body but not white (light). The rest of the body down may be (1) white, (2) medium gray, (3) deep gray, or (4) black. Some individuals may also have combinations of these colors such as white down with gray edgings, or medium gray down with white edgings. Using these classifications, the plumage of 265 downy Brown Noddies was described in 1968 (Table BN-6). The color of the body down is con- sidered to be the major category in this table. lLight-phase birds, especially ones with medium gray body down, predominated. However, when using all three characters, those with white body down and gray edgings were most abundant. Percentages of the various color phases found in 1968 were similar to those found in 1967. Two individuals were unique. One had white body down with gray edgings and a black line which curved forward along the side of the head to the mandible. The other had a black back and collar while the rest of the down was white with gray edgings. In 1968 fledging success was determined for all basic color phases at the north point (Table BN-7). These limited data indicate that there may be differential mortality of the various morphs. More work is neces- sary to determine if these differences exist every year and what the exact causes of death are and their possible correlation with down color. e7T9 Table BN-6. Percentages of various Brown Noddy color phases on Green Island, Kure Atoll, .1968. Number Percent Plumage Description Recorded of Sample I. White Body Down Subtotal 76 28 sil White body down Head and abdomen white 4 15 White body down with gray edgings — Me Ane: II. Medium Gray Body Down: Subtotal OL 302 Gray body down Forehead white, abdomen white 5} legal Forehead white, abdomen light 2 0.8 1/2 of head white, abdomen white 18 6.8 1/2 of head white, abdomen light 3 Lei Top of head white, abdomen white 22 S08 Top of head white, abdomen light 6 2E38 Head white, abdomen white AD 1565 Head white, abdomen light 2 ONS: Gray body down with white edgings 4 1.5 IIL. Deep Gray Body Down Subtotal 80 30n2 Lore white, abdomen white 6 28 Lore white, abdomen light 9 B32 Forehead white, abdomen white 7 26 Forehead white, abdomen light 12 Mh 1/2 of head white, abdomen white 14 543 1/2 of head white, abdomen light iy 6.4 Top of head white, abdomen white 4 165) Top of head white, abdomen light 6 2,3) Head white, abdomen white 2 0.8 Head white, abdomen light 3 esa IV. Black Body Down Subtotal 8 Br Dark head, abdomen light iL 0.4 Lore white, abdomen light 3 Iga Forehead white, abdomen white 2 0.8 Forehead white, abdomen light 2 0.8 Total 265 Cari lO@n© 280 Table BN-7. Nesting success of the four basic color phases of Brown Noddies on Green Island, Kure Atoll, 1968. Color Phase: White Gray Deep Gray Black Number hatched 17 e4 ala 5 Number fledged 10 14 9 4 % fledged 58.8 58.3 81.8 80.0 Banding and Movements Robbins banded 72 adult Brown Noddies and the POBSP banded 493. Table BN-8 summarizes the recapture of these birds at Kure in subse- quent years. Since little effort was expended catching this species, these recapture rates are low. One adult Brown Noddy banded on 5 May 1964 was captured on Eastern Island, Midway Atoll, on 19 July 1965. One thousand two hundred and thirty-three nestling Brown Noddies were banded at Kure Atoll (3 in 1959, 53 in 1962, 1 in 1963, 173 in 196K. 22k in WOES) 9G eam OGG.) Oil in OG andi5 Ol jn91968)). Six, of the 1962 cohort were recaptured (2 in 1964, 1 in 1965, 3 in 1966 includ- ing one previously captured in 1964, and one in 1969); 6 of the 1964 cohort in 1969; and 1 of the 1965 cohort (in 1967). In addition, 3 of the 1964 cohort were found on other islands: one on Eastern Island, Midway Atoll, in 1966; one on Lisianski in 1967; and one at Babelthuap, Palau Island, Caroline Islands, some 2,900 nautical miles away, on 21 May 1965. Two nestling Brown Noddies banded on Eastern Island, Midway Atoll, in 1962 by Robbins were recaptured at Kure Atoll, one in 1966 and the other in 1969. BLACK NODDY Anous tenuirostris Status Abundant visitor with peak populations present from May through September; maximum estimate 2,000. Recorded in all months but generally absent from late December through mid-March. Populations Since the number of Black Noddies varied considerably each day and throughout the year, it was difficult to determine if the difference between earlier estimates (Table BIN-1) and recent POBSP estimates (Table BIN-2) actually represented an increase in the size of the roosting popu- lation. POBSP data indicate that most early visits occurred at times of year when this species was probably absent from the atoll. 281 ‘pounqjdeo spatq jo esequeorted ayy ST eanStj puoodas ayy pue SATTe uaeq aAeY OF UMOUY SP4TQ JO asvequeoted ayy squeseadedt aamsTy 4S4TA x 0° 00T = = == =F == = a2 oo oe = ET 6961 (920) (©°O) (O7O) 9°0 9°0 9°0 0°O0T a5 a2 Sc a5 aS ae OS 1&1. 996T (eeZ) (©) (O20) (SPS) Oe °c Q’d 9°S 0° O0T -- -- -- -- == -- 9€ GO6T (oot) (Oo) (ro) (ror) wy O°T 1 9°T 4° eT 2° et 0°OOT -- -- -- -- -- Ctr2 HO6T (es) (@°@) (ro) (rer) (era)! Wo) Crd Crd E°? 6° ST 2° Qt Gui 0° O0T -- -- -- -- tt €96T (6:3) (@°O) (@°o) Gem) Urs) (@@) (ore) 6x 6°? 6°2 Corin TE? dat TOA 0°0?g 0°OOT -- -- -- GE ZI6T hice) (eo) (oo) (ee) Ure) Gor) @o) Wo) Wo (0°0) I? Vie, Nee 2° 9OT 6° QT 9°Td Que 9° Te 9°T2 Oi 0° O0T LE 6S6T -u papueg qeaR 696T R96T L96T +961 €96T Z96T T96T pornydecay 1e8z *x¥69-6661 ‘(seBequeored se pesseidxa ) aZay} pernydeoer pue [TOVV emmy ye pepueq SeTppoON UMoud FINpe Jo Seiwa sanydeosy 9° Q-Nd eTqeL 282 Table BIN-1. Previous records of Black Noddies on Green Island, Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 O (Munter, 1915). 1923 April 17-22 fairly (Wetmore, ms.). common 1957 June 5 ; yh (Kenyon and Rice, 1958: 190). 1958 May 9 6 On beach (Rice, pers. corr.). 1959 October 3-8 7 (Robbins, 1966: 53). 1960 March 28 iL (Robbins, 1966: 53). 1961 January 19-21 ©) (Robbins, 1966: 53). September 12-14 20-30 Roosting in Scaevola (Udvardy and Warner, 1964:.2). 1962 February 2-4 O (Robbins, 1966: 53). August 6-8 50 (Robbins, 1966: 53). 1963 February 3-7 O (Robbins, 1966: 53). POBSP estimates were based on counts made in conjunction with shorebird, or booby and frigate censuses. The latter were more accurate because they were made at dusk as the noddies returned to roost. Allow- ing for slight discrepancies in census techniques, it appeared that there was little change in population size from 1964 to 1969. Both adult and young Black Noddies roosted on the island. In 1967, 36.7 percent of the birds handled were immmaturés or subadults. At least three adults were actively molting and had bare brood patches, indicating that they had recently bred. The Black Noddies at Kure were probably a post-breeding population from Midway Atoll, where thousands breed, mainly in winter. Annual Cycle Black Noddies were generally absent from late December through mid- March. Small numbers were present in late March and April, except in March 1964 when large Black Noddy flocks were seen on two nights flying over the lagoon. After April, the population increased until a peak was reached in: early June (1967, 1968), late June (1966), and August (1964, 283 Table BIN-2. POBSP semi-monthly estimates of Black Noddies on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 1-15 - 3 O - 2 - iL 16-31 - 1 iL - - - O February 1-15 ~ O O ©) 0 - O 16-28 ©) O O - - os ih March 1-15 - O O ~ - - O 16-31 - 1,000 10 - * ©) 20 April 1-15 oe * 25 - - - 200 16-30 - 250 10 15 - - 300 May 1-15 100 250 15 1, 000 450 ~ 1, 000 16-31 - 250 NS 1, 000 COO 1,250 850 June 1-15 - 250 55 1, 000 2,COO. 1, 25 850 16-30 * US 110 2, 000 2,000 1,250 * July 1-15 - 500 360 479 A.OOO 1,250 - 16-31 - 750 500 700 = 1,250 - August 1-15 - 1, 000 1, 000 1, 000 - 500 - MWO=3)IL - 1, 000 - 800 S 800 - September 1-15 - 600 - 1, 200 - 900 i= 16-30 * 400 - 1, 200 - 500 - October 1-15 * 100 - - - x - 16-31 * 50 - - - * ~ November 1-15 200 25 - - - * - 16-30 15 0 O. = = * s December 1-15 "(5 O 1 - - * = 16-31 300 ©) - 2 = 6 a * Birds present, number unknown. 284 1965). Large influxes of Black Noddies were noted 10 to 14 May 1966, ca. 3 June 1967, and 12 to 15 May 1969. By October the population began to decline until a low point was reached in late November. No indication that this species bred at Kure was noted during POBSP studies. Ecology During the day Black Noddies roosted commonly on the beaches with Brown Noddies. They also fed on small fish along the water's edge, especially in the lagoon. A few were present in Scaevola and on man- made structures. At night they roosted in Scaevola, mainly along the southwest beach and in the southwest portion of the island (Fig. BIN-1). On some nights a few also roosted in the casuarinas, on the radar reflector and on the pier. In 1966 when the large number of noddies arrived, they roosted along the beach-Scaevola ecotone but later moved further inland. Some individ- uals roosted along the northwest and southeast beaches after the population peaked. Banding and Movements The POBSP banded 401 Black Noddies at Kure Atoll (Table BIN-3). Three adults have been recaptured: one banded on 11 August 1964 was captured on 5 June 1966 at Kure, another banded on 18 July 1966 was re- captured on Whale Island, French Frigate Shoals, on 6 June 1967, and a third banded on 10 May 1966 was found at Midway Atoll on 11 December 1967. Robbins banded 5 adults in August 1965, but none was recaptured. Table BIN-3. Black Noddies banded at Kure Atoll by the POBSP, Year Banded Adults Subadults Totals 1963 17 © Ly, 1964 62 @) 62 1965 16 10 26 1966 180 13 193 1967 36 20 56 1969 47 O 47 EO ney ee eee ee eet nee Oe Totals 358 AB KOL Recaptured at Kure were eight Black Noddies banded by POBSP personnel on other islands or atolls: Southeast Island, Pearl and Hermes Reef (3), Lisianski (1_), Whale Island, French Frigate Shoals (3), and Sand Island, 285 “TTOV eany ‘pUBTST USeT JO SsoqTs SuTpsetq Ute], S1TyM pUe Bere SUTYSOOL APPON YoeT_ utew Usa] HUM @ APPON 01¢ : “T-NT@ emMsT wT 286 Johnston Atoll (1). One of these birds, a subadult banded on Whale on 5 June 1967 was captured at Kure on 7 July 1967. The others were all. recaptured at least one year after banding. In addition, 16 adults banded in January and February 1964 by Eugene Kridler on Sand Island, Midway Atoll, were captured in the fol- lowing months: August 1964 (1), May 1966 (8), June 1966 (3), May 1967 (1), and May 1969 (2). WHITE TERN Gygis alba Status Uncommon visitor and spring-summer breeder: ca. 5 to 6 pairs annually. Present all year with peak populations from April through August. Breeds from April until at least September. Populations White Terns were first recorded from Kure Atoll by Lt. John T. Diggs of the U.S.S. Hermes on 15 September 1918. Other earlier observers (Table WI-1) saw them infrequently and in small numbers. During the POBSP survey of the atoll the population size remained fairly constant (Table WI-2). The maximum estimate made in July 1968 resulted from an influx early in the month of at least 25 adults that re- mained until early August. Since no more than 12 White Terns bred in any one year, most of the population represented non-breeding birds, probably from Midway Atoll. Table WI-1. Previous records of White Terns on Green Island, Kure Atoll: Population Date of Survey Estimate Breeding Status, Remarks, References 1915 March 28 0 (Munter, 1915). 1918 September 15 2 Offshore (R.G. 45, Nat. Archives, Report of Commanding Officer U.S.S. Hermes to Commandant 14th Naval District). 1923 April 17-22 O (Wetmore, ms.). 1957 June 5 i Offshore (Kenyon and Rice, 1958: 190). 1958 May 9 6 Over lagoon (Rice, pers. corr.). 1959 October 3-8 2 (Robbins, 1966: 53). 287 Table WT-1. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1960 March 28 0 (Robbins, 1966: 53). 1961 January 19-21 © (Robbins, 1966: 53). September 12-14 rare (Udvardy, 1961: 46). 1962 February 2-4 © (Robbins, 1966: 53). August 6-8 10 5 nests (Robbins, 1966: 53). 1963 February 3-7 O (Robbins, 1966: 53). Annual Cycle White Terns were present throughout the year with peak populations from April through August. POBSP data suggest thac some, ii not aii, of the breeding population left the atoll during winter, as very few terns were seen at this time. The breeding cycle was constructed from data from the 19 nests that were found. Only the 1967 and 1968 data were sufficient to compare yearly differences in the timing of the cycle. Egg laying began in mid-April and continued until at least 25 June. In 1967 three of four eggs (75 percent) were laid the first week of May, while one year later 67 percent were laid between 26 April and 11 May. When all data are combined, there was a peak period of egg laying the last week in April and first week of May (Fig. WT-1). The first egg hatched ca. 23 May. Hatching continued until ca. 30 July, with hatching peaks in 1967 and 1968 five weeks later than laying peaks. White Terns fledged 42 to 48 days (1965 and 1968 data, n=3) after hatching. Thus fledging occurred from early July through early September, although one nestling may have been present in October since an adult carrying a fish was seen on 21 October 1964. Nesting Success Table WI-3 summarizes White Tern productivity on the island. Although only a few young were produced each year, they represented a relatively large percentage of the total eggs laid. The most accurate success data were for 1968 when all six eggs that were found hatched, and four of the young fledged. 288 Table WI-2. POBSP semi-monthly estimates of White Terns on Green Island, Kure Atoll, 1963-69. 1963 1964 1965 1966 1967 1968 1969 January 1-15 - O O - - O 16-31 ~ ik 5 x ie iL February 1-15 - * ilk 3 4 - 2 16-28 - * iL2 ~ - - 2} March Tesi) = 15) D) = = = 5 15-31 - 15 8 - 10 4 20 April 1-15 - 20 13 - ~ - 20 16-30 - 20 15 10 - - 2 May 1-15 6 20 15 15 25 - 20 16-31 - 25 iS m5 25 30 30 June 1-15 - 25 15 iby 25 30 50 16-30 - 25 AG} ib) 25 30 * July 1-15 = 20 I. 15 25 DY) ~ 16-31 - 15) 20 iS - DD = August 1-15 - 30 29 aS = * - 16-31 - 30 - 6 - * - September 1-15 - LO - 12 = ¥% Z 16-30 * 10 - 8 = * a October 1-15 * 10 - - - * = 16-31 * 10 - - - 4 - November 1-15 10 5 - - - 4 - 16-30 O 5 3 - 7 2 2 December 1-15 1 5 1 - - 3 - 16-31 0 5 - O - 2 - *Birds present, number unknown. Number of Eggs J F M A M J J Figure WI-1. Number of White Tern eggs laid each semi-monthly period on Green Island, Kure Atoll, 1967-68 (combined data). LaplewWwi=5. Productivity On Waite. Terns ony Green island. Kure Atoll 1964-69. Maximum Max imum Maximum Number Egg Nestling Number Percent Year of Eggs Count Count Fledged Fledged 1964 mn 1 4 y -- 1965 3 2 2 2 66.7 1966 2 2 1 1 FO. 1967 5 2 yx TF 80.0 1968 6 6 6 4 66.7 1969 il 1* 0 2 -- * Number remaining in late June. Ecology Figure BIN-1 shows the general location of all White Tern nests found on the island. In 1968, 67 percent of the nests were located in the same general area as in 1967, in one case on the same bush, indicating a certain amount of nest-site specificity. Seventeen of these nests were on Seaevola while one was on Tourne- fortia. In all cases the single egg was placed on a branch where there was enough level area, or there were supporting projections, to prevent the egg from rolling off. In 1968 three nests were in crotches of limbs, two in depressions, and one between two branches. No materials were added to the nests. Most nests were surrounded by continuous Scaevola with a considerable amount of open area near the nests, but three were in open areas of 290 scattered Scaevola and Verbesina. The average height off the ground was 23.5 inches (n=4) in 1967 and 37.5 inches (n=6) in 1968, with a range of 20 inches to 6.5 feet. In 1967 four nests were at least 10 feet from the beach-Scaevola ecotone. Non-nesting White Terns roosted in Casuarina, Scaevola, Tourne- fortia and man-made structures such as the radar reflector. The main roosting area was in the casuarinas behind -the barracks. Usually they roosted alone or in twos, but occasionally small flocks were found in the easuarinas. Banding and Movements The POBSP banded 60 adult White Terns (2 in 1963, 17 in 1964, 8 in 1965, 1 in 1966, 2 in 1967, and 30 in 1969) and Robbins banded one in October 1959. Only two of these birds were recaptured--one 1964 bird in 1969 and a 1965 bird in 1966. Eleven nestlings were also banded (2 in 1964, 2 in 1965, 4 in 1967, and 3 in 1968). None was recaptured. No White Terns banded on other islands were found at Kure. HORNED PUFFIN Fratercula corniculata During the winter of 1962-63 several Horned Puffins were found on the Northwestern Hawaiian Islands. Fisher (1965) reported that in January 1963 one live bird was found at Kure Atoll. Robbins informs me (personal communication) that this bird was found on 27 January by a Navy pilot and that he retrieved it as a specimen on 3 February along with 2 more found that day. In addition, another one was found dead on 5 February and another the next day, thus accounting for the five carcasses reported in Robbins (1966). The POBSP found one washed up on the beach on 20 February 1963. Horned Puffins have also been recorded in the Northwestern Hawaiian Islands from Midway Atoll, Pearl and Hermes Reef, and Laysan (Clapp and Woodward, 1968). SHORT-EARED OWL Asio flammeus (flammeus? ) Short-eared Owls have been seen several times at Kure Atoll (Table SEO-1). They were recorded as early as 6 October and as late as 29 April. Usually a single owl was seen flying over the island, mainly over the central plain, but on three occasions two were recorded. The few pellets that were found indicated that they fed on Polynesian rats (Rattus exulans). Based on a single specimen, the owls at Kure are from the Eurasian and North American population (Asio f. flammeus), rather than the closer Hawaiian population (Asio flammeus sandwichensis) (Clapp and Woodward, 1968). 29g The temporal distribution of the records (mainly fall and winter) supports this evidence or origin as that is the time to expect northern migrants in the Hawaiian area. Table SHO-1. Observations of Short-eared Owls on Green Island, Kure Atoll. Date Number Observer Remarks and References 1962 February 2-4 2 C.S. Robbins (Robbins, 1966: 53). 1963 October 6 iL POBSP Flying over central plain. November 13 ill POBSP Near the central plain; flushed from ground. December 11, 1 H.-. Fisher (fisher, 1965: 357). 1964 February 12 a POBSP Collected. March 24 il POBSP On ground near central plain. November 15 1 POBSP Flying over north roost. December W=15 1 POBSP Seen on 3 different occasions. December 24 2 POBSP Under Scaevola. 1965 January 29, 305 Bit 1 POBSP Feeding on rats on 29 January. February 10 IL POBSP -- March 17 il POBSP -= 1966 December 31 a POBSP -- MOST Hebruary 9, LO 1 C.S. Robbins Flying over central plain. 1968 November 14-26 1 POBSP Seen several times. Decemberss/ a POBSP Flying over central plain. December 19 1 POBSP Flying over runway. 1969 February 2 iL POBSP Flying over lagoon. February 9 aL POBSP Flying over Scaevola near south antenna field. 292 Table SEO-1. (continued) Date Number Observer Remarks and References 1969 February 10 2 POBSP Flying over south antenna field. March 18 all POBSP Sitting in south antenna field. April 29 al POBSP Flying over runway. SKYLARK Alauda arvensis pekinensis On 29 September 1963 two sparrow-like birds with white outer tail feathers were seen. Ludwig shot one of these near the east end of the runway, but lost it in a dense Scaevola-Ipomoea tangle. Another individual of this type was seen flying overhead on 5 and 6 October. On 7 October a Skylark, which Ludwig subsequently shot, was seen with a group of American Golden Plovers along the road to the transmitter building. It proved to be a female. This is the only record for the Northwestern Hawaiian Islands (Clapp and Woodward, 1968). BARN SWALLOW Hirundo rustica On the evening of 25 September 1964 two Barn Swallows, believed to be an adult and immature, were seen sitting on an antenna wire and fly- ing over the central plain. Both were subsequently shot, but the specimens fell into dense vegetation and could not be found. Barn Swallows have also been recorded in the Hawaiian Islands from Midway Atoll (Clapp and Woodward, 1968). WATER PIPIT Anthus spinoletta japonicus Ludwig collected a very fat female Water Pipit on 25 October 1963 as it ran along the edge of Scaevola on the north beach. This is the only record for the main or Northwestern Hawaiian Islands (Clapp and Woodward, 1968). RED-THROATED PIPIT Anthus cervinus Ludwig shot a winter-plumaged Red-throated Pipit as it flew over the Scaevola along the east beach on 26 September 1963. This is the only record for the main or Northwestern Hawaiian Islands (Clapp and Woodward, 1968). 293 HOUSE SPARROW Passer domesticus Woodward observed a male House Sparrow around the barracks on three different occasions, 18 to 20 June 1966. On at least one night it roosted in the easuarinas along the road to the pier. Possibly this sparrow came on a cargo plane which had arrived three days earlier. This widespread species was introduced to Hawaii before 1870, and is now common on all the main islands of the Hawaiian Group, but this is the only record for the Northwestern Hawaiian Islands (Clapp and Woodward, 1968). SNOW BUNTING Plectrophenax nivalis townsendi Sibley shot a female Snow Bunting on 10 March 1963 along the runway. This is the only specimen for the Hawaiian Islands (Clapp and Woodward, 1968). Sand Island Avifauna Chandler §. Robbins (personal communication) saw a small group of Black-footed Albatross at the west end of Sand Island on 25 March 1960. On 12 September 1961 twelve Bristle-thighed Curlews and a number of Brown Noddies were seen on Sand Island (Udvardy and Warner, 1964). POBSP observers, in their infrequent visits to the island, recorded an additional seven species (Table SI-1). Only the Brown Noddy bred there. Since no nocturnal observations were made, it was not determined to what extent birds utilized Sand Island as a roosting site. Specimens Table S-1 lists the avian specimens known to have been collected at Kure Atoll by the POBSP, Chandler S. Robbins, and Alexander Wetmore. They are all in the collection of the United States National Museum. O€ OT 0 SS €9 fe) et ual 9T Se ci -¢ 002 ©0ds = 00¢ -- dS O O 0 0 0 QT O Ci 0 g rs OL 0 0 © O © 0 Z O O ¥*L O O ual + 62 Se asnsny Ane oune qsn3sny g96T L96T 996T 294 oO ioe) OO € 8c ZJoequeaon Atne AeW YOTeW G96T °99-796T S‘TIOFV eany “‘pueTsI pueg uo spatq jo sayzeurqse dSd0d et OS S O q Cts © @) ©) O T Ge 0 0 0 9€T OST S9 OOT SOT 0 € 0) O O S g O 6 OT O @) © O T 9 O T O€ as T ©) ©) € g @) 0 @) @) @) Oce temeaw es OL TE €T “peop punoy x 00¢ o Jaquaaon Zaqo1sQ ysnsny Len 96T AppoN Yoetd Sunox Ss33q AppoN umMoig SUTTZapues auo04s -uIny], Appny T9TIFEL SUT TapueM qoog umoig kqoog pooes-anTg IO7EM -Zeoysg £009 *T-IS eTqeh 295 Table S-l1. Avian specimens knawn to have been collected at Kure Atoll. Number of: Species Skins Skeletons Alcoholics Totals Black-footed Albatross Tl ie Laysan Albatross 9 16 Black-footed x Laysan Albatross ib ale Northern Fulmar 1 4 Bonin Petrel 35 1 53) Kermadec Petrel al ile Murphy's Petrel 1 a Wedge-tailed Shearwater 34 ho Sooty Shearwater 6 9 Christmas Shearwater 6 7 Leach's Storm Petrel IL 2 sooty Storm Petrel 16 ite Red-tailed Tropicbird 19 23 Blue-faced Booby 12 1 oh Brown Booby ail ili Red-footed Booby IL 20 Great Frigatebird 28 29 Lesser Frigatebird Black-crowned Night Heron Emperor Goose European Widgeon Pintail tine der Duck Peregrine Falcon Dotverel American Golden Plover Ruddy Turnstone Pintail Snipe Common Snipe Bristle-thighed Curlew Wood Sandpiper Wandering Tattler Lesser Yellowlegs Sharp-tailed Sandpiper Pectoral Sandpiper ' Dunlin Long-billed Dowitcher Western Sandpiper Bar-tailed Godwit (og ae Sanderling Red Phalarope Ring-billed Gull Herring Gull Slaty-backed Gull bh FPOrRPNDFPRFPHERPRPEOANDEUN FEF ONMPEPRPORPFREHEH HR GDOCOCDCCOCODOCOOCHONDCOHPNDODCOOCOCOCOPWHEHNNOCOHPWHOONWODO FPOrRPNFEP RPE RPP OANEPANDEHO FREPRPORPEHPEEH DOOOOOOOD OOO ODO OOOGDO OOOO OOOO OOOOKXONM OF OOrOO NOCH © 296 Table S-l. (continued) Number of: Species Skins Skeletons Alcoholics Totals Glaucous-winged Gull Black-legged Kittiwake Arebic Tern sooty Tern Gray-backed Tern Brown Noddy Black Noddy White Tern Horned Puffin Short-eared Owl Skylark Water Pipit Red-throated Pipit Snow Bunting HN PRERPPPpNMo FWrH DV. OOS GOON Cr OC: CO C1 CCC CS OM OAIO SO © BN PRPEPEPP FN OW FYE DO Wi ON Totals 356 28 Lo KURE REPTILES Four species of reptiles were recorded from the atoll. ‘Green! sea Turtles were occasionally seen swimming in the lagoon and a small popu- lation of Stump-toed and House Geckos was resident on Green Island. There was one hypothetical record of Hawksbill Turtles. Species Accounts HAWKSBILL TURTLE Eratmochelys imbricata Morrell (1841) reported that two Hawksbill Turtles were seen 13 to 14 July 1825. The validity of this record is uncertain. GREEN SEA TURTLE Chelonia mydas Morrell (1841) found Green Turtles abundant at Kure in 1825. Although few of the early visitors to the atoll reported them (Table GT-1), they probably killed them for food and were thus responsible for these reptiles' scarcity today. During POBSP studies, turtles, presumably Green, were occasionally seen, generally swimming in the lagoon (Table GT-2). They were found only four times on land and there was no indication of breeding at the atoll. eg7 Table GT-1. Previous records of Green Sea Turtles at Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1825 July 13-14 great (Morrell, 1841). abundance 1859 late June or plenty (Brooks, 1860). early July 1870 October 29 to ? 4 large turtles captured on sandspit 1871 January 4 (Read, 1912). 1881 December 30 to 2 Collected 6 turtles and found 2 rotten IGS2n January, 1 eggs (Hornell, 1934). 1886 July 15- 2 Turtles available as food (Hawaiian August 18 Almanac and Directory, 1887). 1936 April 5-13 il 1 turtle seen (R.G. 37 Report on the Sur- vey of Pearl and Hermes Reef and Kure Ocean Island). aplenGi-2. POBOP observations or Green -~Sea Turtles at Kure Atoll, 1963 -69. Date of Observation Number Remarks 1963 October 29 IL -- November 6 1 In lagoon. 1964 May 29 iL On Sand Island; caught, tagged, photo- graphed and released. July 7 a Offshore near the west end of the runway. August 13-28 rg Numerous tracks and diggings found along southeast beach. October 19 JL 2' long; offshore near the west end of the runway. : LOG He Mayael: 2 Small; off east beach. July 20-31 3 In lagoon; 2-3' diameter. 298 Table GT-2. (continued) Date of Observation Number Remarks 1967 June 10 al 20"; in lagoon along west beach. June 29 all 20"; swimming in lagoon near the pier. July 2 ales ca. 3' long; in lagoon; caught, photo- graphed, tagged and released. duly 43 1 Off west point. 1968 July 31 al Off southeast beach. August 17 aE Ofi pier. August 26 2 Off southwest beach. November 11 all On Sand Island; 250-300 pounds. December 1s i On northwest beach. 1969 January 25 2 Off west point. January 27 3 Off southeast beach; 20-24" carapace. February 16 1 Off northeast beach; 18" long. February 21 ail Off southeast beach. March 2 ij: Off northeast beach. March-April 2 Occasionally sighted, largest individual not over 24", May ? Small individuals continued to be sighted around reef. STUMP-TOED GECKO Gehyra mutilata HOUSE GECKO Hemidactylus frenatus Two species of geckos, the Stump-toed and the House, were recorded on Green Island. Based on the specimens collected, the former species was the more common. Undoubtedly they were accidentally introduced from Midway Atoll. Geckos were seen in small numbers around the barracks, the trans- mitter building, and the casuarinas along the southwest beach. They were 299 noticed more commonly in late spring and summer than during the rest of the year. Nothing was known of their life history at the atoll. KURE MAMMALS Nine species of mammals were recorded from Kure Atoll. Two, the Bottle-nosed Dolphin and the Spinner Dolphin, were found in the lagoon or just outside the reef, while the Sperm Whale and Goose-beaked Whale washed up on the beach. The Squirrel Monkey, the Pig, and the Domestic Dog were brought to the LORAN station as pets. The Polynesian rat was probably accidentally introduced by the Polynesians, and the Hawaiian Monk Seal evolved in the Hawaiian area. Species Accounts SQUIRREL MONKEY Saimiri sp. A male Squirrel Mohkey was released by the Coast Guard in late 1961. It remained on Green Island in a semi-wild state until January 1967 when it disappeared. Before it disappeared it was frequently seen on the is- land, and in 1965 it began sleeping on the top of the radar reflector where it could be fed by hand. It was observed taking an albatross egg, feeding on Scaevola berries and leaves, and chasing roosting Red-footed Boobies and Great Frigatebirds. Apparently it only rarely disturbed the birds and, in fact, appeared to be afraid of some, such as breeding Sooty Terns. POLYNESIAN RAT* Rattus exulans Status Abundant resident, populations ranging from 20 to 70 animals per acre in 1964 to 1965. Most common in summer and fall. Populations Polynesian rats were first recorded at Kure Atoll by George H. Read (1912) of the U.S.S. Saginaw, who reported, "Rats more in evidence of late. At first small and timid they are now growing larger and bolder." All biologists visiting the atoll prior to POBSP investigations also record- ed these mammals. Although unproven, it is probable that the Polynesians accidentally introduced the rats long before Europeans discovered the atoll. * Wirtz (ms.) studied this species intensively at Kure from 1963 to 1965. Pertinent data are summarized here from his paper. Readers are referred to His work for details. 300 Population estimates were made from March 1964 to May 1965 and in April 1966, by live trapping in a 6.94-acre study area of mixed habitats adjacent to the north antenna field. Figure PR-1l summarizes the 1964-65 data. In late April 1966 there were an estimated 340 rats in che area. Mean monthly density from March 1964 to May 1965 was 45 animals per acre, with a range of 20 to 77. Projecting for the whole island gave an average total population of 6,480, fluctuating from 2,880 to 11,090 animals. Although little or no quantitative data were collected from 1966 to 1969, observations indicated that rats were extremely abundant in 1966, almost absent in 1967, and common again, although less so than in 1966, in 1968 and 1969. Annual Cycle Polynesian rats were most abundant during the fall. They were un- common from January to May and then began increasing in June, reaching a peak in September or October. In fall 1963 there were an estimated 7,200 rats, or 50 per acre, on the island. From December 1963 to February 1964 the population declined to an estimated 30 animals per acre, and then increased to a peak of 75 rats per acre after the breeding season. Following this peak, numbers declined to about 20 rats per acre by 1965. Breeding occurred from January through September, with most litters being produced from March through August. The average litter size was 4.07. Ina sample of 100 live-trapped adult females, 7 produced 2 litters in a year, 1 produced 3, 47 produced 1, and 45 produced none. The observed pregnancy rate in the study area was about 90 litters per season, or about 360 new rats per breeding season. Ecology Polynesian rats occurred in all the vegetated areas on the island. They appeared especially abundant in the central plain and adjacent areas. The garbage area near the barracks was also attractive to them, and during the fall when they were abundant, they would often be found inside the buildings. The diet of Polynesian rats consisted of 62 percent plant material, 30 percent insects, and 8 percent vertebrate flesh. They were known to have eaten the seeds of Boerhavia, the tender shoots and heads of Era- grostis,; the berries of Scaevola, and the seeds of Lepidium, Tribulus, and Sicyos. Insect families identified from fragments in stomachs were Scarabaeidae, Elateiidae (larvae), Noctuidae (larvae), Blattidae, and Formicidae. The relationship of rats and birds was discussed earlier in this, paper. °GO6T ACN-796T UOTeN “TTOUY amy ‘pueTSI user) moO eare Apnas azroe 16°9 @ UT SsZey ueTsouxTOg JO azts uotTyeTndog “*T-Yq amstyq SO G961 ; v96L Ww \/ Ww J f q.N O S Vv f [ Ww Vv WwW ool Ost 002 OSV 00S OSS JOQUNN $O SJONPIAIPUl 302 GOOSE-BEAKED WHALE Ziphius cavirostris On 25 July 1966 a dead Goose-beaked Whale was found washed up on the northwest beach. This animal had been dead for about a day and large chunks of flesh were missing, probably as a result of sharks. Photographs and measurements were taken and the skull saved. Ziphius cavirostris ranges in the Pacific Ocean from the Bering Sea to Baja California, eastern Siberia and Japan, and off the Hawaiian and Midway Islands (Hershkovitz, 1966). SPERM WHALE Physeter catadon On 23 April 1964 Coast Guard personnel found two Sperm Whale man- dibles on the reef. These bones were collected and returned to the lab where they remained during the rest of the POBSP study period. Another Sperm Whale was recorded on 8 July 1967 when the remains of a 30- to 40-foot individual washed up on the south beach near the channel to the lagoon. This animal was photographed, but no bones were collected: This species is found in all seas from the Arctic to Antarctic Oceans (Hershkovitz, 1966). HAWAITAN SPINNER DOLPHIN Stenella roseiventris BOTTLE-NOSED DOLPHIN Tursiops truncatus Rice (1960a) saw about le Bottle-nosed Dolphins within 2 kilometers of the reef on both 5 June 1957 and 9 May 1958. Dolphins were recorded infrequently in the lagoon or beyond the reef during POBSP studies. Due to the unfamiliarity of most observers with ce- taceans, most of these mammals were unidentified. However, two species, the Hawaiian Spinner Dolphin and the Bottle-nosed Dolphin, were definitely identified by Robert L. Brownell. The latter species was seen on 16 November 1964 when 50 to 60 followed a boat in the lagoon, and on 21 September 1968 when several were seen off the south point. The former species was recorded twice: on 14 August 1968 when three schools totaling 80 to 100 individuals were seen east of Sand Island and 28 August 1968 when they were seen in the lagoon. Table PO-zlists the remaining sight- ings of unidentified cetaceans. Tursiops truncatus is widely distributed in temperate and tropical seas (Rice and Scheffer, 1968), and Stenella roseiventris is found in the tropical eastern and central Pacific Ocean (Robert Brownell, personal com- munication). 303 Table PQ-1. POBSP observations of unidentified cetaceans at Kure Atoll, 1964-69. Date of Observation Number Seen Remarks 1964 January 1 25+ East of Green Island. September 18 school Within 100 yards of eastern reef. September 25 school Close to eastern reef. October 2 u Off eastern reef. October 4 several Off northeast reef; leaping out of water. November 27 20 Swimming slowly up eastern edge of thie eceer.. 1965 January 21 12 Swimming northeast along outer edge Chi iSSit , May 24 50t In lagoon near Sand Island. 1969 January 18 2 Near Sand Island. April 16 B02 Off south side of Green Island. May 7 50t Off reef beyond Sand Island. HAWATTAN MONK SEAL Monachus schauinslandi Status Common permanent resident; 150 to 200 animals. Population apparently decreasing. Pups usually born from February to June. Populations Hawaiian Monk Seals were first recorded from Kure Atoll by Morrell in 1825 (Table MS-1). Most other earlier visitors also noted these mam- mals, but unfortunately did not quantify their observations. Undoubtedly seals were common in the 1800's, but the population probably fluctuated drastically with the repeated killings by the crews of wrecked ships. For example, the crew of the Parker reportedly killed 60 seals during their stay on the island in 1842-13 (Whalemen's Shipping List, November 7, 1843). Lt. Commander Montgomery Sicard, captain of the U.S.S. Saginaw, reported (in Annual Report of the Secretary of the Navy on the Operation of the Department for the Year 1871) that "I commenced by sending out parties to 304 kill seal...but after about a month I found that, owing to the rapid diminuation of the seal, I was obliged to cut the allowance down, and only killed one seal...per day for the whole crew." Thus at least 60 Monk Seals were killed in 1870. Kenyon and Rice placed the Kure population at 128 in winter 1956-57 and 142 in winter 1957-58. Based on the number of seals tagged and the number of previously tagged seals seen, the island population was 139 in 1964 and 141 in 1965. Wirtz (1968) estimated the total atoll population as 200 for both years. From the fall of 1963 to the summer of 1965, 112 adults, 22 subadults, and 71 juveniles (57 of them born at Kure) were tagged; thus the previous estimates were probably accurate. Besides these total population figures, population estimates were made at weekly or semi-monthly intervals by counting seals on Green Island as an observer walked around the island. These counts varied yearly. In 1964 and 1969 only those seals seen on Green Island were counted; in 1965 and 1968 all seals seen on Green Island, and Sand Island as seen from Green Island, were enumerated and the counts separated. However, in 1966 and 1967, although the same method was used, the counts were not separated. Finally, several ground counts were made on Sand Island. These data are summarized by month and year in Tables MS-2 to 5. These data suggested that Monk Seals were decreasing as most counts made from 1967-69 were lower than previous counts. Especially significant were the 1969 counts when very few seals hauled out on Green Island. This decrease probably resulted from the increased disturbance by man and by the station dogs who often chased the seals back into the water after they had hauled out. Continuous harassment of this type will lead to the dis- appearance of this mammal from Green Island. Kenyon (in prep.) discusses in detail disturbance as a population control factor. Annual Cycle Hawaiian Monk Seals were present throughout the year, with peak popu- lations on Green Island from December through May. When Sand Island washed away, all seals hauled out on Green Island. POBSP data suggest that after the main pupping season, adult seals moved either to Sand Island or away from the atoll. Actual copulation was not observed so it was not known when the breed- ing season began; it probably started in late summer or fall. The first pups were born on 20 February 1964, 14 February 1965, and 18 January 1969, and the last ones in late July 1964 and 9 June 1965. Figure MS-1 shows the number of pups born each semi-monthly period in 1964 and 1965. Pups spend about 5 weeks ashore before leaving (Wirtz, 1968). 305 Table MS-1l. Previous records of Hawaiian Monk Seals at Kure Atoll. Population Date of Survey Estimate Breeding Status, Remarks, References 1825 July 13-14 2 "the shores...were lined with sea elephants" (Morrell, 1841: 218). 1837 July 9 consider- (Hawaiian Spectator, July 1838). to able 1838 2 number 1842 September 24 Killed ca. 60 seals (Whalemen's Ship- to 2 ping List, November 7, 1843). 1843 May 2 3870 October 29 "main source of food will be the seal" to 7 (Read, 1912: 32). 1871 January 4 1910 January 23 ? Number of sea lions (Log of Thetis, RG. 26, U.S. Nat. Archives)’. 1915 March 28 large Hauled out on the beach (Munter, 1915: number 8X5) 6 1923 April 17-22 4O-50 At least one pup born (Wetmore, ms.). 1934 June 23 50-60 (R.G. 26, U.S. Nat. Archives, Cruise report for Itasca for month of June 1934). 1936 April 5-13 2 Photographs of U.S.S. Oglala (R.G. 80, U.S. Nat. Archives). 1949 Summer 20-30 (Bailey, 1952: 19). LOHL ANvleavishe 2 30-40 Aerial survey (Bailey, 1952: 19). October 12 Cas O (Bailey, 1952: 19). 1956 December 9 to 128 25 pups born in spring of 1957 (Kenyon 1957 June 5 and Rice, 1959: 221). 1957 December 18 to 142 25 pups born in spring of 1958 (Rice, 1958 June 28 1960b; 379). 1959 September 28 53 (Robbins, 1966: 54). October 3 59 (Robbins, pers. comm.). 306 Table MS-1. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1960 March 25 40-60 (Robbins, pers. comm.). 1961 September 12-14 65-70 (Udvardy and Warner, 1964; 3). Table MS-2. Summary of POBSP Hawaiian Monk Seal Counts on Green Island, Kure Atoll, 1964-69. Number Average of Number Date Counts seen Range January 1964 2 28.0 22-34 1965 4 IE) 8-74 196 4 ale 1- Total 10 aay ea 1-74 February 1964 4 By 23 24-42 1965 4 22r3 9-35 1969 4 1550 1-2 Total i 19.2 1-42 March 1964 4 ob .5 21-30 1965 4 46.7 31-71 1967 al: 10.0 - 1969 4 Bp 2-8 Total 13 eH. 2-71 April 1964 1 26.0 - 1965 4 53.0 32-65 196 5 6ise 4-8 Total 10 26. 4-65 May 1965 5 4o.4 25-61 1969 4 We 10-20 Total 9 26.6 10-61 June 1964 2 15 12-25 1965 3 30.3 23 -38 1968 4 120 5-17 196 3 VES} 11-15 Total 12 18.3 -38 Table MS-3. Date January February March April 307 Table MS-2. (continued) Number Average of Number ‘Date Counts seen Range July 1964 5 220 17-26 1965 3 30.0 23-38 1968 3 550 0-12 Total am 55) 0-38 August 1964 4 16.0 9-23 1965 a sO - 1966 ae ILL GO - 1968 2 2.0 o-4 Total 8 NOKS 0-23 September 1964 mM Tag) 9-19 1966 2 PAlLg®© alr Sile 1969 4 0.8 O-2 Total 10 9.1 O=-31 October 1964 6 Tok 2-13 November 1964 5 Pit 13-36 December 1964 5 41.8 16-78 iL Summary of POBSP Hawaiian Monk Seal counts on Green and Sand Islands, Kure Atoll*, 1965-68. Number Average of Number Counts seen Range 1965 3 6251 48-74 1965 a 530 = 1965 4 CeO 51-85 1965 4 80.3 ID 2 1966 2 3350 22-4h 1967 il 24.0 - Total ie FO) of 22-92 308 Table MS-3. (continued) Number Average of Number Date Counts seen Range May 1965 3 60.3 - 52-69 1966 4 3653 25-46 1967 4 30.0 22-36 Total 11 10.6 22-69 June 1965 3 BO 32-47 1966 2 28.0 24-32 1967 2 16.0 15-17 1968 4 Iae3 G3 Total It ep 7-47 July 1965 2 29.5 26-33 1966 2 19.0 16-22 1968 2 1265) 8-17 Total 6 20s 8-33 August 1965 i EO - Table MS-4. Summary of POBSP Hawaiian Monk Seal Counts on Sand Island, Kure Atoll*, 1965-68. Number Average of Number Date Counts Seen Range January 1965 3 2050 0-50 February 1965 iL 18.0 - March 1965 4 Ok .3 14-0 April 1965 4 29.8 27-33 May 1965 3 10.0 8-13 June 1965 3 9.6 9-11 1968 y 2-6 309 Table MS-4. (continued ) Number Average of Number Date Counts Seen Range July 1965 2 6.0 =H 1968 2 50 = Total 5 05) 5-7 August 1965 aL nO - * Made from Green Island. Table MS-5. POBSP Hawaiian Monk Seal ground counts on Sand Island, Kure Atoll, 1964-69. Date of Count Number Recorded 1964 May 29 30 August 13 47 1965 March 27 45 May 24 Ike) July 30 26 1966 August 3 Sul August 25 25 1967 June 29 13 1968 July 4 oh August 14 37 1969 January 18 eS) February 16 30 {Nopoolll sy2 34 iNjopasi IL ALS) 18 June 12 5) Breeding Success Table MS-6 summarizes Hawaiian Monk Seal productivity at Kure Atoll for 1964 to 1969. Only the 1964 and 1965 data are complete. Wirtz (1968) reported that the annual reproductive rate was 15 live pups per 100 adults, that about 19 percent of the adult females bred in successive years, and that only 56 percent of the adult females had pups in either of the 1964 or 1965 seasons. 310 0 1964 BB 1965 ZA Total y) y Y y A VY hae e y Ohno a Aa WG Band 2 A Boma mY : AW YG liv 2 6 ° y) Y) y) y) y DU Cla \e E Aw BIGGIE E A MHAIP 2 Awe BIAIe AY BM BiZiv nw © wiv iBoe Aim WieiPmibb ig Ai Wit iWiB il” ,\I, MIB IB IB IZ A;iIM IMI WIP lig . Y) y) Y) V) Y) . —7AIW IW IW ID IB IB IA January February March April May June Figure MS-1. Number of Hawaiian Monk Seal pups born each semi- monthly period at Kure Atoll, 1964-65. Table MS-6. Productivity of Hawaiian Monk Seals at Kure Atoll, 1964-69. Year Green Island Sand Island Total 1964 28 3 Syl 1965 29 2 Bal 1966* 8 . 8 1967* 13 1 14 1968* 7 2) 12 1969 5** 5x 10 *Incomplete coverage. **Minimum counts. Ecology Hawaiian Monk Seals were found either hauled out on the beaches or swimming in the surrounding waters. At night, and sometimes during the day, they hauled out into the Scaevola where they slept. They were more common along the lagoon beach than the ocean beach. several seals that appeared to have been bitten by sharks were found. They either had circular holes in the back or greatly lacerated skin. SILL Movements A subadult male tagged on 4 October 1963 was found at Midway Atoll on 4 November 1964 by Harvey Fisher. An adult male, tagged on 16 October 1963 and last noted at Kure on 27 March 1964, was found on Lisianski 12.March 1965. DOMESTIC DOG Canis familiaris In 1843 the crew of the Parker found a dog on Green Island that apparently had been there since the Gledstanes wrecked in 1837. (Whale- men's Shipping List, November 7, 1843). During POBSP studies from 1 to 3 dogs have been kept as pets by the crew of the LORAN station. A female mixed cocker spaniel was present from 1961 to the winter of 1966-67 when she died. In early 1964 she was bred with a mongrel from Midway and produced five pups. One of these pups was kept until August 1964 when it was sent to Midway. In 1967 a new pup was brought to the atoll and during the winter of 1967-68 another arrived. Both animals remain on the atoll at this writing. Another dog was present from January to May 1969. Generally, the dogs did not harm the wildlife on the island, although they did catch rats. However, the pup born in 1964 killed at least 7 adult Christmas Shearwaters and 7 nestling Brown Noddies; in 1968 at least 5 adult Christmas Shearwaters and a few Brown Noddy chicks were killed by the dogs. The effect of the dogs on seals is discussed in the Hawaiian Monk Seal account. PIG , Sus scrofa A pig was brought to the atoll in January 1966 to be cooked at a luau, but, instead of cooking the pig, the Coast Guard kept it as a pet until August 1966 when it was sent to Midway Atoll. This animal confined most of its wanderings to the area surrounding the barracks and caused no damage to the native flora or fauna. ACKNOWLEDGMENTS A work of this scope must of necessity be the product of many people. Without the effort of the numerous POBSP personnel who labored at Kure Atoll this study would have been impossible. I thank them all, especially William 0. Wirtz II whose initial organization of the field operations made it easier for those who followed. The Hawaii Division of Fish and Game, Michio Takata, Director, give the POBSP permission to work at Kure, a state wildlife refuge. 32 Ronald L. Walker and David H. Woodside of that organization aided in the field work. Living quarters and food were provided by the U.S. Coast Guard, 14th District. The officers and men of the Kure LORAN station aided the field party in various ways. C.W. Hymes, Federal Records Center, Suitland, Maryland, and Harry Schwartz, Modern Military History Division, U.S. National Ar- chives, Washington, D.C., aided me in the historical research. Dale W. Rice and Alexander Wetmore permitted me to use their unpublished mielidy notes. I am grateful to the following POBSP personnel who contributed to this manuscript: Jane P. Church, Roger B. Clapp, Thomas F. Dana, Mae H. Esterline, and Philip C. Shelton for commenting on early drafts; Anne Keenan Poulson for drafting the figures; and Philip Ss: Humphrey for encouraging and supervising the entire POBSP field and writing effort. I am especially indebted to Chandler S. Robbins, Migratory Bird Populations Station, Laurel, Maryland, for providing me with his un- published field notes and Kure banding data, and for reading the entire manuscript. His comments and suggestions greatly improved the final work. The camera copy was typed by Barbara B. Anderson with funding through a contract with the Bureau of Sport Fisheries and Wildlife, Department of the Interior (contract number 14-16 -008-596, February 3, TO7B)ie | Sits) LITERATURE CITED Aldrich, J.W., C.S. Robbins, and D.W. Rice. (ms.). Unpublished Report - Investigations of bird hazards to aircraft on Midway Island - 1956. Dated 9 February 1957. American Ornithologists’ Union. 1957. Check-list of North American birds. 5th ed. Lord Baltimore Press, Baltimore. xii + 691 pp. Amerson, A.B., Jr. 1968. Tick distribution in the central Pacific as influenced by seabird movement. J. Med. Ent. 5(3): 332-339. Archives of Hawaii. Interior Department (Kingdom of Hawaii). Bound volumes of letters. Book 28, p. 298. Report of James H. Boyd relative to the annexation of Moku Papapa to Hawaiian Island. Dated 29 September 1886. Ashmole, N.P. 1963. The biology of the Wideawake or Sooty Tern Sterna fuscata on Ascension Island. Ibis 103b: 297-364. Bailey, A.M. 1952. The Hawaiian monk seal. Denver Mus. Nat. Hist., MuUShuePcit mAs. Se) PD ---1956. Birds of Midway and Laysan Islands. Denver Mus. Nat. Hist., MulsrueeCit eee) SON ppl. Beardsley, J.W. 1966. Insects and other terrestrial arthropods from the Leeward Hawaiian Islands. Proc. Haw. Ent. Soc. 19: 157-185. Blanchard, B.D. 1941. The White-crowned sparrows (Zonotrichia leuco- phrys) of the Pacific seaboard; environment and annual cycle. Univ. Cale, tld; YOOILs “Os TYAS ja. Brooks, N.C. 1860. Islands and reefs west-north-west of the Sandwich Islands, Pacific. Naut. Mag. 29: 499-504. Bryan, E.H., Jr., et al. 1926. Insects of Hawaii, Johnston Atoll and Wake Island. B.P. Bishop Mus. Bull. 31: 94 pp. Bulter, G.D., Jr.,and R.L. Usinger. 1963. Insects and other arthropods from Kure Island. Proc. Haw. Ent. Sec. 18: 237-244. Christophersen, E. and E.L. Caum. 1931. Vascular plants of the Leeward lsilandsweHawaiise Bb. Bishop) Mus), Bull. Gil Ul pp. Clapp, R.B., V.M. Kleen, and D.L. Olsen. 1969. First records of Emperor Geese from the Northwestern Hawaiian Islands. Elepaio 30: 51-52. Clapp, R.B. and R.L. Pyle. 1968. Noteworthy records of waterbirds from Oahu. Elepaio 29: 37-39. 314 Clapp, R.B. and P.W. Woodward. 1968. New records of birds from the Hawaiian Leeward Islands. Proc. U.S. Nat. Mus. 124 (No. 3640): 39 pp. Clay, H.F. 1961. Narrative report of botanical fieldwork on Kure Tsiands | AtolbkarResw pulls woe oeppe Dana, T.F. 1971. On the reef corals of the world’s most northern atoll (Kure: Hawaiian Archipelago). Pac. Sci. 25: 80-87. Edmondson, C.H., W.K. Fisher, H.L. Clark, A.L. Treadwell, and J.A. Cushman. 1925. Marine zoology of the tropical central Pacific. B.P. Bishop Mus. Bull. 27: ii and 148 pp. Ely, C.A. and R.B. Clapp. (ms.). A Natural History of Laysan Island. Farrell, N. (ed.). 1928. John Cameron's odyssey. The MacMillan Co., New York. 461 pp. Findlay, A.G. 1870. A directory for the navigation of the North Pacific Ocean. Richard Holmes Laurie, London. 1,007 pp. Fisher, H.I. 1965. Bird records from Midway Atoll, Pacific Ocean. Conder 16/7395 —3bia Fleet, R.R. 1972. Nesting success of the Red-tailed Tropicbird on Kure Atoll. Auk 89: 651-659. Fowler, H.W. and S.C. Ball. 1925. Fishes of Hawaii, Johnston Island, and Wake Island. B.P. Bishop Mus. Bull. 26: 31 pp. Great Britain Hydrographic Department. 1946. Pacific Islands Pilot 3: 306-307. Gross, M.F., J.D. Milliman, J.I. Tracey, and H.S. Ladd. 1969... Marine geology of Kure and Midway Atolls: a preliminary report. Pac. Sci. PEO AU (Ceo Hawaiian Spectator. Ocean Island. July, 1838. p. 336. Hershkovitz, P. 1966. Catalog of living whales. U.S. Nat. Mus. Bull. Oh6. Honolulu Almanac and Directory. 1887. Wreck of the Dunnottar Castle-- annexation of Ocean Island. p. 99-101. Hornell, J. 1934. Log of the schooner Ada on a fishing cruise in the North Pacific, 1882. Mariner's Mirror 20: 436-437. Sul) Keller, G.V. 1969. Electrical resistivity measurements, Midway and Kure Atolls. Geol. Sur. Prof. Paper 680-1. Kenyon, K.W. In prep. Man versus the monk seal. Kenyon, K.W. and D.W. Rice. 1958. Birds of Kure Atoll, Hawaii. Condor 60: 188-190. ---1959. Life history of the Hawaiian monk seal. Pac. Sci. 13: 215-252. Kepler, C.B. 1967. Polynesian Rat predation on nesting Laysan Alba- trosses and other Pacific seabirds. Auk 84: 426-430. ---1969. Breeding biology of the Blue-faced Booby Sula dactylatra personata on Green Island, Kure Atoll. Publications of the Nuttall Ornithological Club, No. 8: 97 pp. Lack, D. 1966. Popiilation studies of birds. Clarendon Press, Oxford. 341 pp. Ladd, H.S., J.1. Tracey, Jr., and M.G. Gross. 1967. Drilling on Midway Atoll, Hawaii. Science 156: 1,088-1,094. Lamoureux, Charles H. 1961. Botanical observations on Leeward Hawaiian Esends.. Atoll Res. Budalis 792. 7 pp. Lord, W. 1967. Incredible victory. Harper and Row, New York. 331 pp. Maa, T.C. 1968. Records of Hippoboscidae (Diptera) from the central Pacific. J. Med. Ent. 5: 325-328. Medeiros, J.S. 1958. Present status of migratory waterfowl in Hawaii. Wildlife Manage. 22: 109-117. Morrell, B., Jr. 1841. A narrative of four voyages to the south seas, north and south Pacific ocean, Chinese Sea, Ethiopia, and southern Atlantic Ocean, Indian and Antarctiie Ocean from the year 1822 to 1831. Harper Bros., New York. 92 pp. Munter, W.H. 1915. Report of destruction of bird life on Laysan Island. An. Rep. Coast Guard for 1915: 130-140. Nelson, B. 1968. Galapagos: islands of birds. William Morrow and Co., New York. 338 pp. Pilsbury, H.A. 1917. Marine molluscs of Hawaii, IV-VII. Proc. Acad. Nat. Sci. Phila. 69: 309-333. Read, G.H. 1912. The last cruise of the Saginaw. Houghton Mifflin, Boston and New York. 128 pp. 316 Rice, D.W. 1960a. Distribution of the bottle-nosed dolphin in the Leeward Hawaiian Islands. J. Mammal. 41: 407-408. ---1960b. Population dynamics of the Hawaiian monk seal. J. Mammal. 41: 376-385. ---(pers. corr.). Letter to P. Woodward dated 2 December 1969. Rice, D.W. and K.W. Kenyon. 1962. Breeding distribution, history, and populations of north Pacific albatrosses. Auk 79: 365-386. Rice, D.W. and V.B. Scheffer. 1968. A list of the marine mammals of the world. U.S. Fish and Wild. Serv., Spec. Sci. Rep. - Fish No. 579. Robbins, C.S. 1966. Birds and aircraft on Midway Island, 1959-63 investigations. ~UcS. Fish and Wild: Serv., (spec. Sci.) Rep. .—nWaslddeiske Wo. 85< vi + 63 pp. ---(ms.). Field notes for 26-28 September 1959, 25 March 1960, 17 Janu- ary 1963, 6-10 February 1966, and 9-12 February 1967. Rowan, W. 1929. Experiments in bird migration -l1. Manipulation of the reproductive cycle; seasonal histological changes in the gonads. Boston Soc. Nat. Hist., Proc. 39: 151-208. Sandwich Island Gazette. Another disaster, November 11, 1837. Informa- tion, December 22, 1838. Schreiber, R.W. and N.P. Ashmole. 1970. Séabird breeding seasons on Christmas Island, Pacific Ocean. Ibis 112: 363-394. Seckel, G.R. 1962. Atlas of the oceanographic climate of the Hawaiian Islands Region. U.S. Fish and Wild. Serv., Fishery Bull. 193: 371-427. Sharp, A. 1960. The discovery of the Pacific islands. Clarendon Press, Oxforde pels appre Sibley, F.C. and R.B. Clapp. 1967. Distribution and dispersal of cen- tral Pacific Lesser Frigatebirds (Fregata ariel). Ibis 109: 328-337. Sibley, F.C. and R.W. McFarlane. 1968. Gulls in the central PaciRier Rac. Sci. 922: 3i=30. Standen, R.S. (ms.). Unpublished thesis. An explanatory description of the sand islands of Kure, Midway, and Pearl and Hermes Atolls: Hawaiian Islands. Sab Steele, C.W. 1967. Fungus populations in marine waters and coastal sands) of the Hawaiian, line, and Phoenix islands Pac. Sei. 2s SESS SL Temperance Advocate and Seaman's Friend. Whaleship Parker wrecked on Ocean Island. June 27, 1843. The Friend. Loss of the U.S.S. Saginaw. January 1, 1871. sudays Riel « 1966. Marine benthic algae from the leeward Hawaiian garowyoe /MG@ALIL Ieaie Jewel, AIS aS) Foysin Udvardy, M.D.F. 196la. Additions to the checklist of Hawaiian birds. Elepaio 21: 83-90. ---1961b. The Harold J. Coolidge Expedition to Laysan Island, 1961. Elepaio 22; 43-7. ‘Udvardy, M.D.F. and R.E. Warner. 1964. Observations of the birds of French Frigate Shoals and Kure Atoll. Atoll Res. Bull. 103: 3 pp- + 4 figs. United States National Archives. Washington, D.C. Record Group 2. og Om athe! U.S.s). LEoguois 905) LOX GH Waa Wooo Melosia aS hog of the U.S<.Si: Ramsay 1935 Log of the U.S.S. Childs 1940 Log of the U.S.S. Saury Saury 1941 Log of the U.S.S. Spearfish 1941 ---Record Group 26. Log of the Thetis 1910. Cruise report for the Itasca. For the month of June 1934. ---Record Group 37. Aircraft Advanced Base Report of the U.S.S. Childs Report of the survey of Pearl and Hermes Reef and Ocean Island. ---Record Group 45. Log of the U.S.S. Lackawanna. May 20, 1867 to November 24, 1867 1918 report of Commanding Officer U.S.S. Hermes to Commandant 14th Naval District. ---Record Group 80. Log of the U.S.S. Marblehead 1928. Photographs from the U.S.S. Oglala April 1936. United States Navy. 1871. Annual Report of the Secretary of the Navy for year 1871. 318 United States Navy Operational Archives. War Diary of U.S.S. Preble. Walker, F.D. 1909. Log of the Kaalokai. The Hawaiian Gazette Co., Ltd., Honolulu. 64 pp. Wetmore, A. 1925. Bird life among lava rock and coral sand. Nat. Geog. Mag. 48: 77-108. ---(ms.). Field notes taken on the 1923 Tanager Expedition (original in the possession of A. Wetmore). Whalemen's Shipping List. 1843. Whaleship Parker wrecked on Ocean Island November 7, 1843. Wirtz, W.O., II. 1968. Reproduction, growth and development, and juvenile mortality in the Hawaiian Monk Seal. J. Mammal. 49: 229-238. ---(ms.). Unpublished thesis. Ecology of the Polynesian Rat, Rattus exulans, on Kure Atoll, Hawaii. 132 pp. Wolfson, A. 1952. Day length, migration, and breeding cycles in birds. Sci. Monthly 74: 191-200. * U, S. GOVERNMENT PRINTING OFFICE : 1972 O - 478-373 a aa Tine b en “a -_= ,, eet | fe i o of & WS ‘ =o UO oni =» oe | a S = Ea = — ‘hy Y A e $, = a Fy a E | 2] 3 =e Sf NG JS §-§ GG sat ING J a) a y FO BN a a DW a 2 SS 4 RIES. SMITHSONIAN — i: NOLNLILSNI_ NVINOSHLINS, S31YV¥E17_ LIB RARIES |, SMITHSONIAN IN te ” = = x xX i: = < * ee = =< = ‘S =| = AS A ze \, 4 Zz = So ‘e) WG K <= OF ae 3 3 5 BQ F 2 Come 2 g 2 S = WO § EB! \. = i= x 2 = YS 2 = . 5 |S Shei . 3 aa JISNI- NVINOSHLIWS’S3 1yvy g 11 LIBRARIES SMITHSONIAN _ INSTITUTION | NOILOLILSNI_NVINOSHLIAS) igs 5 we | «( zZ ws a ws a ¢ 2 Ng Z z yp 3 : : EP 4 WN = <3 e < = = > = Vy, \. = ¢b%.= =i Ly, ® = S x 9 NN 5 GBS B Ye ca 2 8 zw NS HE ? 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