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THE NATURAL HISTORY OF LAYSAN ISLAND, NORTHWESTERN HAWAIIAN ISLANDS By Charles A. Ely and Roger B. Clapp Issued by THE SMITHSONIAN INSTITUTION Washington, D.C., U.S.A. ATOLL RESEARCH BULLETIN NO. 171 THE NATURAL HISTORY OF LAYSAN ISLAND, NORTHWESTERN HAWAIIAN ISLANDS By Charles A. Ely and 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 Washington, D.C., U.S.A. December 31, 1973 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 TABLE OF CONTENTS Page LIST OF FIGURES... ccccccccccccccccccvcccvccccvccsceseccccces lil ILLS OI IWASILOS 5 og Gocco cep ooo bUuDOO Db OnoO Ob ODNUoUdasc0n0000E so owal LIST OF APPENDIX TABLES ..... pabDa0De Sboccodo soo ds00000000000 ix EMER CIIUCHEI ON eRe Sena a ONO ors @) 01s eelsleie «a cies 0)0 syels ee seccececel: TENS CRIUPIEIUN 5 on Gigs opild bo uo bot ODS nb a Seo JE aor ec cccee 00006006 3 HITS HORNA safe ol ol cueke) silale’ aieus =) ai si ehelchoNehelcler oven suonehencieyarsyola exeleyereys) seliekes ove sucpeler Rss GPE AUE MON tercel Gr erovtays grove e eves tie’ oie. 6 one ee snares SiemEROPoTER PROT Manan iate 64. LAYSAN ISLAND FAUNA........... PSI arc acan cen RnR PORTE REAPER NRT OLS Pn EOCUC GNOME sence octet acca eee Seite ce sine cal fBsigccl Ss togeyetvetcte crore eiatevecn Cit ec ale a ore ET ee eee oid ald acre ne OO MEAG ILS 5 5 Soo ood DOC ODOC ONO OOO dO CODD ODDO OOOe DOOD D ODO OG00D0Oee REP GCsperetelelatelelstolevelteleiel ers DOOD CCOOODCOORGADOUoSOI06H00000 50088) SHE CESMACCOUMitc Sieperetsroncletel enelelelelsherehelelerereierelake) oreo sieterelclclelelelclelers OS Babys GISie ves orci s\ahele, ave. eve @eeeee etaleielalelstalelshotelaielelalaieialaieleloialcreretoreiae 5) Diomedea nigripes..... eecielerereleraiclc/ciaiarersvolcnslcncicisteehsio) Diemedea -immutabilise ey ewe cc cle ecm eles vce ee clOl Lelong 15 BeS)WeyplaN FDOINEIIEE AAR RAR Gpate- ooBOocodn coun os) Bulwer la Sb Ulwe Tleivsrer ee te } See atts Rare beges mot oe St nol) taogab i Ode j ae 4 fexy. r f ee * ey a J + ; oy 3 ’ 4 y Sy : 4 La ey } 4 ‘ ’ 0 s i { : wu Nal Pi " =| af i h fit 7 t Be Rea ' vit : i aa i Pee Shoes . , se ‘ were =) ee ee ee mm Sut : * Ys oy =F aD BS { ; i. ay i . 4 Dor fi j Nae i Ya Ey 1 eae at x f THE NATURAL HISTORY OF LAYSAN ISLAND, NORTHWESTERN HAWAIIAN ISLANDS 2 by Charles A. ley and Roger B. @tappe/ INTRODUCTION Laysan Island, the largest of the Northwestern Hawaiian Islands, is located in the central Pacific at latitude 25°42'41"N, longitude 171°44'06"W. It is approximately 115 nautical miles east of Lisianski, 202 miles northwest of Gardner Pinnacles and 709 miles northwest of Honolulu, Oahu, Hawaii (Figure 1). Laysan is almost due south of the Pribilof Islands and roughly equivalent in latitude to Monterrey, Mexico and Miami, Florida. It is roughly rectangular in shape and about 1.4 Square miles in area with a lagoon occupying about one-fifth of the island interior. lLaysan is a coral island ringed on its periphery by sand dunes; the beach crest and inland slopes are well vegetated. Laysan has the most remarkable biota of any island in the North- western Hawaiian Islands. It remained relatively undisturbed until the late 19th century and even the first decade of human occupancy had little apparent effect on island life. Introduction of rabbits (about 1903), however, proved very nearly disastrous--more so than the famous raids by Japanese feather poachers a decade later. Attempts to exterminate the rabbits in 1912-1913 failed and only the timely arrival of the Tanager Expedition in 1923 saved the island from complete devegetation. Even the extermination of the rabbits and replanting of vegetation were too late to save three of the five endemic birds. Today the vegetation and most animal populations have regained levels similar to those re- corded early in the 20th century. Laysan was declared a part of the Hawaiian Islands Bird Reservation by presidential executive order in 1909 and is now known as the Hawaiian 1/ Paper Number Th, Pacific Ocean Biological Survey Program, Smithsonian Institution, Washington, D.C. 2/ Present address: Department of Zoology, Fort Hays Kansas State College, Hays, Kansas 67601. 3/ Present address: Department of Zoology, University of Maryland, College Park, Maryland 20742, LIWAVH » IMVIOOHY invw ig IVNV¥1 ea *SpueTST UeTTeMey ou °T aanstq noly NOLSNHOS * vinv» « YovHIIN VOHIN * 4ayDIN * SIVOHS 31V9Idd HDN3Ud * SJIDVNNid Y3NGYVS * NVSAV1, INSNVISIT® 4338 S3W43H GNV 1aV3d . AVMAGIW © 3 Islands National Wildlife Refuge. During the 1950's and 1960's Laysan was visited primarily by biologists from the State of Hawaii Division of Fish and Game under contract to the Bureau of Sport Fisheries and Wildlife of the U.S. Department of Interior. In 1964 a federal refuge manager was assigned to Hawaii and direct responsibility for inspection, patrol and management of the refuge was assumed by the Bureau. In February 1967 Laysan was declared a "natural area" to be maintained as free from outside or non-natural disturbance as possible and island visits are restricted to official or scientific business. The Pacific Ocean Biological Survey Program (hereafter referred to as POBSP) of the Smithsonian Institution, Washington, D.C., made 13 trips to Laysan during the period 1963 to 1968. A number of these visits were made in conjunction with regular inspection trips by the Bureau of Sport Fisheries and Wildlife. These visits, together with additional surveys made by personnel of the Hawaii Division of Fish and Game and the Bureau of Sport Fisheries and Wildlife, form the basis of this paper, which summarizes biological information concerning Laysan Island. DESCRIPTION General Nature Laysan is roughly rectangular in shape with the long axis slightly east of North. The Tanager Expedition survey (by Major Chapman Grant) determined maximum dimensions of one and four-fifths mile long (9,375 feet) by just over one mile wide (5,580 feet). Warner (ms.: 5) used this map to calculate a total area (including the lagoon) of almost 913 acres (about 1.43 square miles). Bryan (1954: 4) had previously given the area as 1.56 square miles. Although island dimensions have been variously reported by different writers (e.g., Brooks, 1859: 500 [3 miles long, 2 1/2 miles wide]; Fisher, 1903a: 772 [3 miles long, 1 1/2 miles wide]), an appreciable change in size during historical times is unlikely. Aerial photographs taken in 1939, 1943 and 1966 (Figs. 2-4) agree in most particulars with the outline map prepared by the Tanager Expedition in 1923. Laysan is a coral island capped by large sand accumulations and with a large salt water lagoon in its central depression. The island is probably the flattened top of a once massive volcanic peak formed perhaps during the Miocene, since eroded far below the present sea level, and subsequently built up by the action of coral, other marine invertebrates and calcareous algae. The depth of the coral deposits capping Laysan has not been determined but is probably considerable. (Borings on Eniwetok and Bikini reached the volcanic bedrock at 4,222 and 4,610, and 2,556 feet, respectively [Weins, 1962: 92].) Relatively shallow water extends some distance in all directions from the present island surface and then drops off rapidly to an average depth *ydersozoud AAeN °S*°N TePTOTSTO “6C6T ysnsny G SpueTs~T ueskeT go ydearSoj0ud [etazay *g amnesty *ydersSoj0ud AaeN °S°N LetotsrO °€r6T Ae Qe ‘pueTsI uesfeyT fo ydersoqoyd [etsey °€ eansTad *ydergozoud AAeN °S*N TetotssO °996T Azenuer ‘puets] uesfeT Jo yderZoj0yd Tetszey "+ eInST A of 1,800 fathoms between Laysan and neighboring islands. The 100- fathom line encloses an area of approximately 210 square miles (Warner, 1963). As Warner (loc. cit.) pointed out, "the topography of Laysan suggests that the island was at one time a small atoll with a central lagoon which is now nearly filled with sand and coral fragments." He described its present configuration as "a great.ringed sand dune... heaped upon the coralline rock base whose subsurface is approximately five feet above mean high water mark." The island rises abruptly to a height of 15 to 18 feet (Bryan, 1942: 185), then rises more gradually to a crest of 30 to 40 feet. The maximum crest elevations (just over 40 feet) occur in the sand dune area at the north end of the island with crests of 30 feet elsewhere; lowest on the south. The beach crest varies in width from a narrow strip on the northwest and east to a width of about one-eighth mile on the southwest and west. Inside the beach crest the island slopes gradually (more steeply on the east) to the lagoon--the major physical feature of the island interior. The lagoon lacks an opening to the sea, is just above sea level and varies considerably in shape and area with water level. The island is presently well-vegetated except for coastal dunes, most of which are stabilized, and two "blow-out" areas inland from the beach crest on the west side. The original flora was the most varied in the Northwestern Hawaiian Islands chain. Destruction of the vege- tation by man and introduced rabbits resulted, by 1923, in near desert conditions, the extinction of several species, and the movement of sand which altered some physical features of the island interior and partially filled the lagoon. At present the vegetative cover has returned to some- thing approaching the original condition. Vegetation extends generally from the beach crest to a narrow, vegetation-free zone near the lagoon which apparently marks the zone of maximum salt impregnation. In some areas, particularly the north, sand dunes extend over the crest and part way down the inward slope. The fresh water pond once located near the southwest end of the lagoon has not been reported since 1915 and presently even brackish water exists only after periods of heavy rainfall. A few remnants of the guano period remain (several piles of phosphate rock, the rusting tram line), as do scattered debris from later visits. A HIRAN team which surveyed the island in 1961 left a tall concrete bench mark above the campsite near the northwest landing. The island has been uninhabited since 1906 except for brief periods by feather poachers in 1909 to 1910 and 1914 to 1915 and by the Schlemmers from 12 July to 2 December 1915. Physical Features Offshore structures and reef plate The fringing reef surrounding the island varies from 100 to 500 yards in width and is most extensive at the northwest end of the island. 8 Inside the reef is a narrow, shallow channel nearly encircling the island except for the south and southeast sides. A small boat can navigate most of this channel at high tide. A natural opening on the northwest reef provides a safe boat entrance during most weather and other less satisfactory breaks in the reef are present near the southwest and northeast corners of the island. Outside the reef the depth increases gradually, with generally under 20 fathoms at least five miles from the reef. The reef consists of coral, calcareous algae and the remains of shelled marine invertebrates. On the east side the rocks and stones are cemented together but with a dislocation or fracture extending across the bottom of the lagoon and crossing the reef near the south- east corner (Elschner, 1915: 30). Elschner (1915) described the reef plate or plate rock in some detail. It formed the bottom of the lagoon depression and extended seven to ten inches deeper than the water level of the lagoon and probably a foot lower than the average ocean level. It was strongly phosphatized and formed by the caking and cementing together of dis- integrated coral substances through the influence of guano solutions. The result was a fine conglomerate with the pores closed and clogged to such a degree that it was nearly impenetrable to water. In areas this plate was on the surface but in others partly covered by a more or less phosphatized sand or soil. Both the plate and the remainder of the lagoon basin were penetrated with horizontal stripes of brown phosphate between layers of white undecomposed carbonate. soil, fresh water, and guano The island surface consists primarily of more or less phosphatized coral sand. The phosphatized sand consists of round oolitic grains, which are covered with a layer of brown phosphate and contain in such cases undecomposed centers of carbonate. Between these grains there are larger pieces of limestone and splinters of bone. The thickness of the phosphate coating varies considerably. When phosphatization is nearly complete (high percentage of phosphate), the phosphate is fre- quently a very fine brown powder (Elschner, ies} Schauinsland (1899: 89) reported the finding of a "peat" deposit as follows: "In the north part of the island just below its highest elevation on a deeply situated spot where Scirpus laevigatus was abundantly growing, I excavated because there was presumably a ‘coal deposit’ there. Underneath a layer of humus I first found sand, then a hard substance like marl, and finally peat (somewhat mixed up with the sand) consisting of the remains of long plants (Scirpus?) situated rather uniformly together in significant amounts." In 1961, Woodside (ms. c) reported that members of his party dug small, shallow holes to water level near the lagoon to examine the underlying layers. No peat was found but what appeared to be a dark-colored soil was noted. 9 Laysan has had no permanent fresh water supply since at least the late 19th century. Paty (1857: 42) reported finding, on 1 May 1857, an abundance of "tolerable good fresh water" by digging two feet "not a 100 yards from the salt"(on shore of the lagoon). Two years later, also in May, Brooks (1860: 500-501) dug a well and found "very good water." Either the tastes of the individuals involved were different or potable fresh water was no longer present in quantity by 1896. Schau- insland (1899: 72) stated that the water on the island itself was "briny" and that they relied solely on rain water collected from the roofs. Wilder (1905: 392) made similar observations in September 1905. At that time a well sunk inland provided water (presumably somewhat brackish) for washing while drinking water was collected from roofs and stored in cement cisterns. In 1911, Dill and Bryan (1912: 9) observed that the pump was rusted out and the well partly filled with sand. All 20th century visitors have relied on imported water supplies or on frequent, usually brief, rain showers. It seems likely that the fresh water lens described by Warner (1963: 9) may have been more prevalent during the period of early island exploration. In recent years this fresh water lens, developing on occa- sion as a result of high rainfall, has provided the only fresh or brackish water on the island. McKernan (in Warner, 1963) indicated that the fresh water lens might occur also above the hypersaline water of the lagoon when rainfall and ground seepage are sufficiently high. The exposure of the fresh water lens in low depressions was un- doubtedly the source of small fresh water ponds or puddles noted by several observers up to the present time. The largest of these was a small pond of slightly brackish to fresh water (Fisher, 1903a: 773) near the southwest corner of the lagoon. This pond was permanent in nature through at least 1915. By the arrival of the 1923 Tanager Expedition, however, it had filled with wind-blown sand and disappeared. Warner (1963: 9) noted a number of brackish water depressions and small ponds during wet seasons. These were highly favored by Laysan Teal. Similar observations were made by the POBSP, usually at the southeast edge of the lagoon. Schauinsland (1899: 19) described the guano observed by him in 1896 as clean and odorless. It was present “partly as a more or less dusty or sandy, rather thick form under the surface (brown or white guano) and partly as solid rock several meters deep which must be broken with pick and shovel (the so-called 'rock guano')." He also reported finding whole granules of beautiful pure crystal. Elschner (1913) reported that Laysan guano occurred as a fine phosphatized coral sand or as a soft, loose phosphate sandstone of low grade. Elschner (1913, 1915) described the processes of guano formation on Pacific islands and described in some detail the deposits on Laysan about the turn of the century. These deposits were formed under conditions of moderate rainfall and resulted in a guano intermediate in nature between 10 excrement and mineral. Due to the great amount of leaching and chemical changes, most of the nitrogen was lost and phosphorization occurred, re- sulting in a phosphate guano, as contrasted with the nitrogeneous guano of arid regions. His map (Fig. 5) showed the greatest guano deposits at the south end of the lagoon with smaller deposits northwest, north, and northeast of the lagoon. A "tram" line ran to the south digging with a spur to the north end of the lagoon. Part of the former was still present though partly covered with sand in 1967 (Fig. 6). 1/2 1 sea mile we a Ncrev-ice Figure 5. Map of Laysan Island in 1915 showing location of guano deposits. Redrawn from Elschner (1915). Various accounts indicate that the greatest amount of digging was at the south end of the lagoon and the large flat mined area is a con- Spicuous landmark today. Also remaining are several piles of broken phosphatic rock some six feet in height. Sahcuinsland (1899: 33) stated that the guano level extended to below sea level and on the east side of the island extended into the ocean. Bones and "petrified" eggs occurred in this layer and some of the latter are now in the Bernice P. Bishop Museum collections. Elschner further stated (1913) that the high grade qualities con- tained up to 75%, in small areas even 80%, phosphoric lime (based on dry substance) and up to 1.25% nitrogen. All grades containing more than iit -yoateus °9°a faq ydeaZoqyoud qsdod ‘°996T eunr HI ‘pueTSI jo uotTq10d ule7SemMugnos uo ynoMOTG Apues UT eUTT Wey tomtos FO Ssjueuuled SUOTe FSeMUI1OU BSUTYOOT Be em cane —™ Vaddaae Aes IEA. °Q oinsTa 12 60% phosphoric lime had been completely exhausted by 1917 so that only lower-quality types of 40-50% CazP20g remained on the island. Schau- insland (1899: 89) stated that in 1896 the raw guano shipped from Laysan contained an average of 25-30% phosphoric acid and presented analyses of two samples. "Brown guano” (lying on the surface) contained 11.5% Po0s and 48.64% CaO; "light-colored rock guano" contained 36.99% Po05 and 33.3% CaO. Two samples reported by Elschner (1913) are of considerably higher quality. An analysis of these contained 72-80% tribasic phos- phoric lime and 0.82% nitrogen. Lower grade materials (soft stone) contained 48.20% Ca3Ps0g, 38.0% CagP20g and 1.02% nitrogen. The beaches When approached from the west, Laysan appears as a flat island with white sandy beaches capped by low green vegetation and with con- spicuous white sand dunes rising from the north end of the island. The west beach is narrow, 25 to 50 yards in width, and rises rapidly to the beach crest. The crest is bordered seaward by a narrow band of low Scaevola 10 to 60 yards in width. Inland the vegetation grades into the extensive bunch-grass zone. Most of the beach is open sand. Some 600 yards south of the northwest landing is a projecting coral block and a group of large boulders which extend from the shoreline to the beach crest. Two similar, but smaller, areas occur farther south. Just north of the southwest corner of the island the reef reaches the beach and the deeper water there provides a landing area. At the southwest end of the island a 20-foot high coral shelf (Fig. 7) extends southeastward along the shore for approximately 250 yards. The remainder of the southwest beach consists of waveswept reef and massive boulders--most near the waterline but some scattered up to the beach crest. The reef is very close to shore along the southeast part of the island and is characterized by large boulders chiefly along the reef. The southwest and southeast beaches proper are from 50 to 100 yards wide and rise gradually to a low crest. Scaevola occurs in scattered clumps and several sandy strips extend up and over the crest for distances of 150 to 200 yards (notably near the southwest corner). The east beach is much more extensive and is from 250 to 400 yards wide, sloping gradually to the beach crest. Although some Scaevola occurs on the beach crest, most of the crest is bare, with sand extending some distance over much of the inner slope. In season, beach morning glory and Boerhavia cover parts of the slope. The most extensive beaches occur on the north and northeast. These slope gradually to the beach crest some 150 to 200 yards inland. The beaches are usually bare sand except for scattered large rocks, mostly on the shoreline. The sand is usually compact but in some areas, especially near the shore, is more loose. Most of the large sand dunes present inland from the north beach are stabi- lized, especially those inside the beach crest. The beach crest is generally unvegetated and bare sand extends well down the interior slope of the island (Fig. 8). In the north, the bare sandy area (as measured from aerial photographs) extends various distances from 250 yards to about one-third of a mile before meeting stable vegetation. ° Taq Teuyang o7m°q Aq ydeasojyoud qsdod *)961 eunr ‘ueskeqT Jo pue ySemuynos 94e greys Tetop °) eansT a 14 ‘ueuyoeH °q°o Aq udersoq0ud qSd0d “°JUSTI-pTw 4e ST BAOTS JnUOD0D YAION *J96T UOTeN SyooseT go yseeujyzou ysnl 4utod wor yoeeaq yseeyj,a20U pue Y4tOU preMO4 SUTYOOT Pe °9 aINnsT A ale The island interior Visitors to Laysan usually camp at the site of the former guano headquarters on the northwestern part of the island where a large ironwood tree now marks the natural passage through the reef. Most of the few conspicuous landmarks can be seen from the beach crest near this point. The general impression is of an expanse of tall, coarse buneh grass in the foreground sloping gently to a long flat plain con- taining a shallow lake (Fig. 9). The inner slopes are generally covered with clumps of grass and capped with scrubby Scaevola except where bare sand extends over the beach crest. Scaevola forms a continuous zone along the west beach crest. The lagoon varies in size and shape with season and water level and is usually bordered by bare saline mud flats, particularly on the west and northwest sides. A ring of prostrate vegetation and sedges occurs between the mud flat and the bunch grass zone. On the east side of the lagoon the mud flat is much more narrow and the island rises rapidly to the beach crest. A small group of coconut trees is present a few hundred yards seaward from the northwest end of the lagoon (Fig.10). A smaller grove at the southeast end of the lagoon is barely visible from the campsite. Also barely visible are several patches of Pluchea bushes at the north- east and south ends of the lagoon. ‘Two large areas of bare sand ("blow- outs") are present on the inner slopes west of the lagoon. During dry periods the lagoon is separated by a narrow strip of vegetation from a small, shallow "pond" to the northeast. A sand bar (its size varying with water level) crosses the lagoon at about its northern third. During periods of high water, generally in winter, the lagoon floods and water may extend over the low vegetation surrounding the lagoon and even ex- tend to the northwest coconut grove. During major winter storms, heavy waves may also break over the low southern beach crest and drain inward to the lagoon. Winter storms also shift sand and destroy vegetation inland and near the lagoon. Available aerial photographs from 1939 through 1966 show two very distinct bands of light (less dense) vegetation west of the lagoon and extending the entire length of the island, roughly parallel to the island contours. These bands average about 75 yards in width and alternate with three bands of dark (denser) vegetation. These bands are within the bunch grass zone and are not easily seen from the ground. Lamoureux (1963: 12) determined that the less dense areas were due to more widely spaced Eragrostis clumps in nearly pure stands. The relationship of these bands to soil conditions, moisture, or other factors has not yet been determined. Perhaps these zones were associated with sand dunes while the island was denuded or perhaps they date far back into the island's history. These bands of vegetation were not mentioned by early visitors but could have been easily overlooked from the ground. Two "blow-out" areas of unconsolidated sand are the most prominent landmarks on the western side of the island. These areas, in the south- western and south-central portions of the island, are clearly visible on 16 *996T eunr Tz ‘uoyTeUS *o°a &q ydexZoj0ud qgaod ‘esouko STTAYSTzquTa (pUMoazZe10g |yeTpemMT uT) pue eSnIITp BTAeyAeOg JO Soyozed pate,7eoSs YIM STLTGVTACA STJSO1selq ST UOTJeJedaA JUeUTMOg *pueTST jo Jejeupzed 4semygaou uo 9074 euTZenSse) Teeu TeT [Td ayetouod WorZ 4YSeayynos SuTyYooT © 17 aAOIG SODOD Y41ON *uogTeus °O°d Aq ydersoj0yd qsqod ‘“punodcsezoj-ptm ut *996T aune Tg Suoose~T fo uotyaz0d uxayyaZ0U paemod YSeeygnoSs BSuTYyooT ne SLL TO es OT eamnsTa 18 aerial photographs taken in 1939, 1943 and 1966. They vary considerably in size and shape but are always centered in the zones of less dense vegetation. In the May 1939 photographs, one such blow-out is nearly continuous with the southwest beach and extends inland (northward) for about 1,000 yards or nearly one-third the length of the island. Photo- graphs taken four years later show that most of this area has revegetated, particularly at the southern end, and the area of bare sand covers an area of only about 200 by 100 yards. The extent of unstabilized sand is probably affected by season and by local rainfall and consequent condi- tion of the vegetation. Activities of nesting birds also affect the vegetation rather drastically at times. The January 1966 photographs show a large blow-out crossing the center strip of dense Eragrostis and extending inland almost to the lagoon flat. Its largest dimensions are approximately 560 by 300 yards. The blow-out did not appear this extensive when viewed from the ground in September 1967. The only other noticeable landmark on the western side of the island is the remains of the old tram line now badly rusted and largely covered with sand. The most prominent feature of the south end of the island is the very flat, almost circular site of the main guano diggings. This area is about 300 yards in diameter and portions still lack permanent vegetation. Also remaining from the guano period are one large "rock" pile about six feet in height, several smaller rock piles and numerous scattered loose rocks. The vegetation here is predominately Sesuvium and heliotrope rather than bunch grass. Also present near the south end of the lagoon are the clumps of Pluchea bushes and the small grove of coconut trees mentioned earlier. The 20-foot high inland cliffs mentioned by Schauinsland (1899: 33) are no longer present. Either they have been destroyed or perhaps covered by sand dunes. The lagoon The central lagoon area (lagoon proper and bare shore ) is about one mile long with a maximum width of about one-third of a mile. The actual water area fluctuates considerably with season and rainfall. Both Palmer (in Rothschild, 1893-1900: x) and Fisher {1903a: 773) estimated the lagoon at 100 acres. Warner (ms.), using the Tanager Expedition map, calculated a lagoon area of 204 acres. During at least some winters (e.g., 1961-62) the lagoon floods its shores and extends to the northwest coconut grove. Contours are most variable on the shallow western side and on the north end where fluctuations in rainfall cause temporary ponds and sand bars. In 1859 Brooks (1860: 501) reported a maximum lagoon depth of five fathoms in the center and a coral bottom. Schauinsland (1899: 20) described the lagoon as two to three, sometimes five, fathoms deep with clear water. By 1923 (Wetmore, ms.), it was "generally 3-5 feet deep 19 with a maximum depth of 15 feet." A similar depth was noted by Warner (1963: 4) near the eastern shore. He also noted that the sand bottom and changes in contours of the eastern shore were evidence of appreciable encroachment by shifting sand dunes, probably during the denuded period of the early 1920's. The reef plate mentioned earlier greatly reduces the percolation of lagoon waters as shown by the lack of any apparent tidal fluctuation and by the high salinity of the lagoon water. Salinities of 12 to 15% were found by Schauinsland (1899: 20) in 1896 and by Elschner (1915: 33) in 1914. Warner (1963: 6) reported similar concentrations ranging from 12% (June 1958) to 14% (July 1959). Samples taken by Walker from the surface over flooded vegetation and mud flats in December 1963 after a period of heavy rainfall varied from 5.9 to 8% while two samples taken from "flooded springs" were 4.3 to 5.3%. These lower values are probably typical during periods of high rainfall and flooded conditions. Some amount of flooding probably occurs during most winters. In some years (e.g., 1961-62), heavy rainfall after the Laysan Albatrosses have laid results in heavy egg mortality. Large amounts of vegetation bordering the lagoon may also be killed. Warner (1963: 4) also noted that during winter storms, waves sometimes wash over the east slopes of the island as evidenced by debris deposited on the lake flats. HISTORY Laysan in the early 1800's It is generally agreed that Laysan was originally discovered and named by an American ship, but no details are presently available con- cerning the actual discovery. The visit of the Russian ship, the Moller, in 1828 is usually listed as the earliest visit but we know of at least two that undoubtedly occurred prior to that. In the summer of 1828 J.N. Reynolds, who had been delegated this task by the Navy, visited New England whaling ports and gathered from various whaling masters ships' logs, etc., information on the location of islands in the south seas (Stackpole, 1953: 461). His report, sub- mitted September 1828, but not published until 1835 (Reynolds, 1835), con- tained several entries which bear on the history of early visits to Laysan. Since whalers were normally in the Pacific for over a year, and since his information was compiled in mid-1828, it is likely that all entries per- taining to Laysan were the result of visits made no later than 1827. On page 7 of his report Reynolds places "Laysan's island" at 25°50'N and 171°51'W. The position currently listed for Laysan is 25°46'N, 171°44'wW (Office of Geography, 1956: 46). On page 23 Reynolds related that "Captain Briggs discovered an island west and north of Sandwich Islands, in 25°47' north, longitude 20 172° west. The island is low, with not more than 60 feet in any part from the water, 3 miles long and 2 across it.” This island was almost certainly Laysan. Examination of the tables in Starbuck (1878) reveals that the only ship listed as whaling in the Pacific prior to 1828, and with a captain named (John) Briggs, was the Wilmington and Liverpool Packet of New Bedford which made two voyages, on either of which Laysan may have been discovered. On the first voyage the ship left New Bedford on 12 April 1821 and returned on 27 December 1823. On her second voyage the ship left on 1 December 1824 and returned on 8 March 1827. On a third voyage under the same captain the ship left on 25 August 1827 and did not return until 24 June 1830. Since Briggs must have been absent when Reynolds compiled his report, Reynold's information must have been obtained from some other individual. Briggs' discovery of Laysan must have occurred no later than 1826 and possibly as early as 1821 or 1822. Stackpole (1953: 304) relates that "The Lyra...[a new Bedford whaler] discovered a reef and an island which was probably the island of Lysan [sic], northwest of Oahu, but was wrecked not many miles dis- tant from it a few years later (in August 1830)."1* Capt. Stanikowitch of the Russian vessel Moller visited Laysan on 24 March 1828 and, not knowing of its previous discovery, named it Moller after his vessel. C. Isenbeck, the ship's surgeon, made a few observa- tions of the island and its biota which were passed to and later published by F.H. von Kittlitz (1834). Unfortunately, either the original notes were unclear or their translation was incorrect concerning several bird species. An English translation of parts of the Kittlitz paper was published by Rothschild (1893-1900). The description of Laysan in 1828 (Rothschild translation) follows: On March 12 (24) Herr Isenbeck landed on Moller (Laysan) which was originally a coral-island with a long reef round it. It seems that it was raised higher and became a real island from the accumulations of the birds' excrements. It is covered with a strong bushy kind of grass and partly with low shrubs, between which a few pigmy palms had grown up. Although there was no fresh water on the island, there were not only sea-birds but also several land-birds, as the following list will show. Most of the larger birds were already breeding, or had paired at least. 1850-1874 In 1857 Captain John Paty explored much of the leeward chain in the Hawaiian schooner Manuokawai. On 1 May he landed on Laysan and annexed it to the Hawaiian Kingdom. An account of this visit was published by Paty (1857: 40) with the following description: *Footnotes to the history section, numbered consecutively, begin on page 59. al Laysan Island--W. by N. 3/4 N. from Honolulu 808 miles. This is a low sand island, 25> to 30 feet highs 3 miles long and 1-1/2 broad. The surface is covered with beach grass, and half a dozen small palm trees were seen. It has a lagoon in the centre (salt) 1 mile long and half a mile wide, of salt water, and not a hundred yards from the lagoon, abundance of tolerable good fresh water can be had by digging two feet, and near the lagoon was found a deposit of guano. The island is "literally covered" with birds; there is, at a low estimate, 800,000. Seal and turtle were numerous on the beach, and might be easily taken. They were evidently unaccustomed to the sight of man, as they would Scarcely move at our approach, and the birds were so tame and plentiful, that it was difficult to walk about the island without stepping upon them. The gulls lay enormous large eggs, of which I have a specimen. A bank of rocks and sand extends off to the South and West 6 or 8 miles or more. Good anchorage can be found on the West side of the island in from 4 to 20 fathoms, by selecting a sandy spot to anchor upon, half to 2 miles from the beach. The best landing is about one-third of the distance from the Northern to the Southern point of the island, where there is a very smooth sand beach. Lt. J.M. Brooke, commanding officer of the U.S. Schooner Fenimore Cooper, visited Laysan on 14 January 1859 and took soundings, positions, and physiographic data which were later incorporated into Hydrographic Office charts. Three hours were spent on Laysan and "six small turtles and a variety of sea birds were taken."2 Captain N.C. Brooks of the U.S.S. Gambia visited Laysan in May 1859 and described the island as follows: Laysan Island.--Laysan Island is in lat. 25°46'N., long. 171°49' W., is 3 miles long and 2-1/2 broad, and covered with a luxuriant growth of shrubs. It is sur- rounded by a reef about half a mile from the land. Outside of this reef there is a bank five miles wide, on which I found from fourteen to nineteen fathoms water. There is a boat passage inside the reef nearly the whole way round the island. Good landing can be found anywhere, excepting on the South and §.E. sides; good anchorage anywhere on the West side,--the best, however, is about half a mile from the S.W. point, in from eight to twelve fathoms water. It can be approached from any point of the compass, no dangers existing within half a mile of the reef. On the east end of the island I found the remains of a wreck, but saw no signs of a camp. 22 There is a lagoon on the island about one mile long and half a mile wide, with five fathoms water in the eentre, and coral bottom. On the shores of this lagoon I found salt of good quality. There are five palm-trees on the island, and I col- lected twenty-five varieties of plants, some of them splendid flowering shrubs, very fragrant, resembling plants I have seen in gardens in Honolulu. I saw on the beach trunks of immense trees. The island contains about fifty acres of good soil. It is covered with a variety of land and sea birds; some of the land varieties are small and of beautiful plumage. Bird's eggs were abundant. Near the N.W. point of the island I found a stick about two feet long, and at the foot of it a bottle contain- ing a paper, but could not decipher the writing. From the East point, where the wreck lies, to a decayed palm-tree on the shore of the lagoon, in a direct line, I planted potatoes, Onions, and pumpkins. The soil on which [I planted them embraces every variety, and appears to be adapted to vegetation. There is a very small deposit of guano on this island, but not of sufficient quantity to warrant any attempts to get it. Dug a well and found very good water. The reefs here abound in fish and turtle (Brooks, 1860: 500-501; see also Brooks, 1859). Material from this log was quoted in nearly the same form by Roth- schild (1893-1900: i-11). Some Visits to Laysan in the 1880's and 1890's Laysan was visited twice in 1882 by the fishing schooner Ada sailing from Nagasaki, Japan. During the first visit, 26 to 30 January, 104 turtles were taken, 61 of them from beaches during the course of one day. Two hundred and seven beche-de-mer were also collected, but shark-fishing was a failure. During this visit the crew found a board on which "was an appeal to voyagers not to take the turtle away” (Mansbridge in Hornell, 1934: 432). The board was repainted to express the same sentiments. The ship returned to Laysan on 3 May and captured another 26 turtles (Hornell, 1934: 432-433). The schooner General Siegel (which wrecked shortly thereafter on Midway) visited Laysan while on a sharking expedition about late September 1886. A fair haul of sharks was made, some turtles and a few seals were killed; the crew spent a week ashore (Farrell, 1928; 253-254). John Cameron visited Laysan about fall of 1893 while on a shark- fishing voyage on the sloop Ebon. He noted the presence of two pigs and 23 indicated that albatross had begun to return to the island, an event which suggests the visit was made no earlier than October (Farrell, 1928: 399). Cameron revisited Laysan during the summer of 1894. His stay of about a month was broken by a visit to nearby Lisianski (Farrell, 1928: 414). ‘Laysan Becomes a “Guano Island” In 1890 George D. Freeth, an Englishman who had visited Laysan as early as 1864, and George N. Wilcox, who had previously managed a guano operation on Jarvis Island, inveigled a Honolulu firm, Hackfeld and Company, into financing the North Pacific Phosphate and Fertilizer Company, which was to work guano deposits on Laysan (Anon., 1939: 9-10). Previously Freeth had captained the vessel Akamai to Laysan where he had taken possession of the island, hoisted the Hawaiian flag, and left two men on the island in February 1890 to hold possession. On 13 March he returned to Honolulu with "reports [of] good guano deposits on Lycan [sic] Island." (The Friend, April 1890: 29; see also Lyons, 1890: 90). On 29 March Freeth and Charles N. Spencer got the Hawaiian Kingdom to grant them3 the right to mine phosphate deposits on Laysan and Lisianski Islands for a period of 20 years with a royalty of 50 cents per ton to be paid to the Hawaiian Government. These rights were later transferred to the guano company which was incorporated on 23 May 1890.4 On 10 July Freeth departed for Laysan for the purpose of systemati- cally examining the guano deposits. With him on the schooner Kaalokai was Captain Rosehill, who was to become a foreman of the laborers on Laysan, and A.B. Lyons. The party arrived at Laysan on 16 July after a six-day run from Honolulu. Lyons later published some notes taken on this visit which con- sisted primarily of observations of birds, but which also included an interesting description of the island. July 16. Here we are at Laysan Island we have brought the schooner inside the reef under the lea of the island, and are laying in quiet water within two hundred yards of the shore. The only indication of land this morning at day break was the flocks of sea birds which we could see in every direction, although we were in reality only twenty miles from the island. The land lies so low that it can be seen from the deck of a vessel only a few miles.... There was nothing indeed, particularly inviting in the land itself. A beach of white shell sand, a steep bank, also of sand, with little vegetation--beyond a strip of nearly level land scantily covered with coarse bunch grass and low shrubberry,--that was all we could ah see as we approached the shore. Not quite all, for there rests over the land perpetually a cloud of sea fowl, and these you can see at a glance hold undisputed possession of the island.... The island is quite small, barely two miles long by a mile and a quarter wide, of the familiar ring form, with a small closed lagoon. In its highest part the land may be as much as 35 or possibly 40 feet above high tide mark. Although the island is surrounded with reefs, there is very little rock to be seen above the water level, except on the south-east coast where there is a rampart of sand- stone rising ten feet or more perpendicularly from the water. The rock ig all a shell sandstone containing a very little coral, and even on the reefs little living coral is to be seen. The soil of the island consists of a peculiar kind of white sand, made up partly of fragments of sea shells, but largely of bits of egg shells and the bones of sea birds.... A rough calculation puts the bird population of the island at about 800,000; it may reach 1,000,000. They have not yet learned to fear man excessively, and are in fact no more shy than barn door fowl, so that it is very easy to study their habits. The flora of the island I find interesting, although somewhat disappointing. I gathered only twenty-one species of flowering plants, nearly all of them Hawaiian or cosmopolitan plants. The seeds of most if not all of them have floated to the island in sea-water. Among them should be mentioned the loulu palm, the maia pilo (caper) and Koali (convolvulus) and a stunted species of sandal- wood (Lyons, 1899: 90-91). At the first meeting of the North Pacific Phosphate and Fertilizer Company in October 1890, Wilcox was elected president and Spencer vice- president. One of the Hackfeld family was elected treasurer, no doubt to protect their investment; Freeth was appointed resident superintendent for Laysan (Anon., 1939: 10). Freeth thereupon hired a chemist, a foreman, and eight laborers and chartered the Inter Island Steam Navigation Company's S.S. Pele to transport them and their supplies to Laysan. When they reached the island, about November 1890, all but the railway equipment was landed before the party was driven from the island by one of the Northwestern Hawaiian Islands’ notorious winter storms. All returned to Honolulu, arriving there on 9 December (Anon., 1939: 10). From 1891 through 1903 guano operations were conducted directly under the aegis of the North Pacific Phosphate and Fertilizer Company, and much guano was removed by a series of ships, some of Hawaiian registration and others of American or German registration (Bryan, 1942; 186). 25 On 25 June 1892 the fertilizer company renegotiated its agreement with the Hawaiian Government. In the Memorandum of Agreement drawn up between C.N. Spencer (then Minister of the Interior of the Hawaiian Kingdom) and the company, the government agreed not to raise royalties on the guano, and gave permission for the phosphate to be made into fertilizer in processing plants other than those on the Hawaiian Islands. On 31 March 1893, Sanford B. Dole, President of the Provisional Government of Hawaii, signed an act, effective on publication, which confirmed the earlier Spencer-Freeth franchise and which authorized the lease of Laysan and Lisianski to the guano company. On 17 April that year the lease was executed by the Minister of the Interior, J.A. King.© By terms of this lease these islands were rented to the guano company at a dollar a year until 29 March 1910. (The lease was eventually sur- rendered to the Hawaiian Government on 31 December 1908.7) On 15 February 1894 the fertilizer company leased, for a period of 2> years at a rental of a dollar a year, other islands of the Northwestern Hawaiian group (Morrell [sic], Ocean [sic], Pearl and Hermes Reef, Midway, and French Frigate Shoals). The company was also granted the exclusive right to mine the phosphate deposits thereon, provided a royalty of 50 eents a ton was paid to the Hawaiian Government. A clause within this contract stated that operations had to begin within five years of the signing of the contract or the right to the deposits would be withdrawn. Since no guano was mined on any of these islands, the right must have been automatically withdrawn on or about 15 February 1899. On 3 April 1894 the North Pacific Phosphate and Fertilizer Company changed its name to the Pacific Guano and Fertilizer Company, and in 1896 Max Schlemmer became superintendent of operations on aysan.8 In April 1891 the first shipment of guano, 80 tons that sold at $15 a ton, was removed from Laysan by the chartered schooner Mary E. Foster, Captain Berry commanding; 90 tons were taken from the island by the same ship in May.9 More guano cargos were shipped later in the year via the Foster and the clipper Elizabeth Nicholson. By 30 November, 1,017 tons (valued at $1,975.20) had been removed (Anon., 1939: 12-13). Thomas (ms.) gives the most detailed picture of guano operations when they were at or near their peak of productivity. Laborers mined the guano, consisting mostly of a hard, conglomerated, phosphate of lime, with picks, crowbars, shovels, and sledges. This material was placed on cars on the narrow guage railway and pulled by mules to storage sheds where the guano was kept until a ship arrived. A small amount of brown guano (bird droppings and soil) was also collected and sifted, but it was a small proportion of the total amount of guano shipped from the island. When ships came in the guano was carried from the storage sheds in barrows holding about a ton out onto the wharf that extended from the west side of the island. At the end of the wharf the guano was dumped into a chute which deposited the material in lighters. These lighters in 26 turn transferred their cargo to the clipper ships that were anchored between two buoys offshore. As much as 100 to 125 tons per day could be loaded under favorable conditions (cf. Bailey, 1956: 2) Guano was shipped only from April through September but a care- taker remained on the island throughout the winter. Schauinsland (1899: 32) related that the caretaker left on the island one winter was found dead when the island was revisited seven months later.10 He was found sitting at the table where he had been working on his journal and was subsequently buried on the west beach. On the few occasions when ships arrived at Laysan during periods of high surf, 11 landings were made at the "winter landing" on the north coast of the island. Figures 11-17 are from the guano period. Scientific Work on Laysan in the 1890's The Rothschild Expedition The first intensive scientific collecting expedition in the North- western leeward Hawaiian Islands was sponsored by Walter Rothschild (Tring Museum) in 1891. Henry C. Palmer and his assistant, George C. Munro, arrived on Laysan 16 June aboard the schooner Kaalokai (Capt. F.D. Walker) and stayed with Captain Freeth, then manager of the guano operation. They departed 27 June. Ornithological results of this visit, a portion of Palmer's diary, and a summary of earlier observations were published by Rothschild (1893-1900); more popular accounts of the visit were written by Munro (1930, 1941-43, 1946) and Walker (1909). Island conditions were described, a biological survey made and numerous birds collected, including four species new to science (see Appendix Table 2). Visit by J.J. Williams Between mid-November 1892 and late January 1893, J.J. Williams, a Honolulu photographer, spent about a month on Laysan taking photographs and collecting birds. He collected "several barrels of stuffed birds” and a about 300 photographs (Pacific Commercial Advertiser, 3 February 1893). Collection of birds in 1895 In September 1895 a small collection of birds was made on Laysan by W.H. Hall who collected birds for the Bernice P. Bishop Museum (Bryan, 1901: 259). None of his specimens has specific dates on the labels and we have been unable to discover anything else about his visit. Schauinsland's visit in 1896 Dr. and Mrs. H.H. Schauinsland spent the summer of 1896 with the guano company on Laysan. They arrived on the barque H. Hackfeld at the beginning of the guano season, anticipating a short visit, but when the magnitude of their planned survey became obvious, they decided to remain ‘IT Setaeg ‘eg dnozy pazooey saatyory TeuotzenN ‘yderso 0d SoTATES SFTLTPLIM pue USTA °S°N “SSO7JeqTY Touwes419 UCTSSTuMOD SaTTOYsSTT °“S°N FO estnazo ZutInp ute[sequreuy “Wd Aq uderSoj0"ug °*Zo6T LEW ynoqe uesfeT uo eare ButTpeoyT ouenH “TT eins 28 “IT setzeg ‘gg dnozy ptoosy SsaaTyory TeuoTzeN Syders0j0ud SOoTAIES eFTIPTIM pUe UST °“S°N “SssorpeqTy teweeys UOTSSTUMOD SeTTeysTA °S°N FO estnszo Sutinp utetszequeyp “Wa Aq yderso070uUd *Z061 ut uesfeyT uo suotjezedo oumns jo aseg 29 d “TT Sataeg *‘eg dnozH prooey saATyoLy Wie Stare ie ue USTH °S°N “SSOrZeqTy Teureseqs9 uoTsstTmmop ee a8 Re ee eae . *q £q uderSoj0uq “east ys Bup~peosad ut umMoy Ee aes Beg es pee lea Be Me teaeots “3061 AEN Jnoge ‘uooBse, jo pus uteyy -Uja0U UO TeM04 Worms TIC TT ‘i cA ‘masnW doustd *d eoTureg Aseqanoo ‘zeydeas -ojf0yd umouyun Aq yderZo0104g °s,06Q9T ATzee sutTamp Atqeqoid ‘uesfe] uo ZuTSStp ouwenyH “tT etnsTA 30 31 -o10yd uMouyUun Aq yderso70uUg “n—n~Touoy ‘umesny doyst@ °d sotuateg Aseqanos ‘zayderg °S,0691 ATte9 Sutanp Atqeqord ‘uesfeT uo SuTsstp oueny “CT aati ‘n—Tntouoy ‘umesny, douystg °qd sotuaeg Aseqymoo ‘zeyderZ030yd umouyun Aq yderZ0j,0ud °S,0691T ATtee Sutanp Atqeqoid ‘uesfey uo varie SuTpeot pue peys oueny “*OT emMsta Be 33 *n—njTouoH ‘umesnw doystg °q aotuszeg &saqyamoo ‘zaydersoq30ud umouyun Aq ydeaZoj0ud payepun ‘*szeqazenbpeay uotzeredo oueny: */T ammsta 3h until the departure of the last guano vessel on 24 September. Schau- insland’s report (1899) is wonderfully detailed and includes data on geology, meteorology, and biology. Extensive collections were made of many biological taxa and were subsequently sent to Germany. Corres- pondents on Laysan apparently sent bird specimens to the Bremen Museum for several years afterward. Many of these collections were subsequently reported by various German collaborators (see Appendix Table 3). Homicide on Laysanl® Life on Laysan during the guano mining era was not always peaceful as is attested by a case of homicide that occurred in 1900. On the evening of Saturday, 11 August, forty Japanese laborers became involved in a fracas with the overseers of the guano mining operation. We give below an extract of the original story of the shooting reported under the banner headline "Laysan Island's Story of Blood." War has been declared, waged, and ended on Laysan Island...four against forty--those were the odds, four white men fighting desperately against forty infuriated Japanese. And the white men conquered. [On the evening of Saturday, 11 August 1900]...the forty Japanese rose in a body, determined to annihilate all the white people and run things to suit themselves. Captain Spencer [the manager of the guano-mining operation]...called his son and Captain Spilner, late of the Honolulu mounted patrol and the engineer, Luhrs...and together they went out to talk to the mob which had gathered between the white men's house and the Japanese quarters. [Captain Spencer asked what the trouble was whereupon]...the leaders of the mob answered defiantly, cursing...and shouting.... They threatened the white men's lives..., were carrying flags and waving them excitedly and were armed with knives, clubs, stones, and cutlasses made of hoop iron sharpened. They made a movement towards the platform.... Old Captain Spencer...raised a six shooter in either hand. qi "The first man who steps upon this platform shall die! shouted Captain Spencer. "Shoot away!" cried the mob and at a signal from the leader, charged all together for the platform. Though they moved quickly, Captain Spencer's trigger fingers moved quicker. Eight times his revolvers spoke and they spoke to the point. Pistols in the hands of the other white men also had something to say. Two... Japanese dropped dead...[who] were the leaders...[and] three others fell helpless, sorely wounded. (Pacific Commercial Advertiser, 8 September 1900). How much of this lurid account was true? Subsequent testimony given at a pre-trial hearing varied considerably according to various witnesses, and the excited yellow journalism with which the case was reported tends to obscure the facts. Events following the shooting seem fairly clear, however. ap) Events following the killings After the fusillade, the Japanese dispersed to their quarters and the white men returned to their house, where, according to later testimony by Spencer, the whites awaited a further attack. The following day the surviving 3913 Japanese were rounded up at gunpoint and incarcerated aboard the barque Ceylon which had recently arrived at Laysan. Subse- quently the two dead Japanese were buried and on 16 August the Ceylon sailed for Honolulu with all concerned except Spencer's son and the engineer. After an uneventful passage the Ceylon arrived in Honolulu on 7 September. After being questioned by the police, Spencer and Spilner were arrested. The pre-trial examination followed shortly thereafter. Spencer was formally charged with murder and Spilner was held as a witness for the prosecution. On the llth testimony for the prosecution was begun with statements by two of the Japanese, Higuchi Shiro, and Oguma, who cooked for the Japanese. Their testimony was followed by that of Kinoshita who had been standing near Goto when Goto was shot. His testimony was largely corroborated by witnesses the next day. Thursday, 14 September, Tanaka, the luna (foreman) of the Japanese, and Captain Spilner testified for the prosecution. Spilner's testimony was particularly damaging to Spencer's case. Although avowing himself a good friend, Spilner implied that Spencer fired on the Japanese without warning. Letters, however, entered as evidence for the defense, written at an earlier date by Spilner to Dr. Averdam, manager of the Pacific Guano and Fertilizer Company in Honolulu, rather clearly established the extent of Spilner's friendship. Therein Spilner implied that Spencer was a drunkard and an incompetent, wasteful manager, and that he, Spilner, would be a fit replacement. Spilner concluded his testimony on the 17th and the last of the prosecution witnesses, a carpenter named Wahlers, told his story.1 The defense then began to introduce its witnesses. The first to testify were five sailors from the Ceylon who served primarily as character witnesses for Captain Spencer. Three of them, though, had recently spent six weeks on Laysan when they had been stranded there by the wreck of the barque McNear, 16 and presumably knew something of condi- tions on the island preceding the uprising. Captain Spencer then took the stand and gave what appears to be a fairly credible version of the incidents. On the 18th the defense briefly interrogated its last witness, Dr. Averdam, who repeated some hearsay evidence concerning a conversation with the carpenter and who strongly rebutted Spilner's assertions about Spencer. After the summing up by the attorneys on 21 September, all charges against Spencer were dropped. 36 What really happened on 11 August 19007 The facts of the case are hard to ascertain in spite of, or perhaps because of, all the evidence offered at the trial. The following account, derived primarily from testimony by Shiro, Wahlers, and Spencer, is prob- ably as close to the truth as one can get this many years after the event. On the 9th of August, Spencer, his wife, and 12 new laborers arrived at Laysan on the Ceylon. During Spencer's absence Spilner had been in charge, with Tanaka acting as luna. With Spencer's arrival Spilner was demoted to luna and Tanaka to common laborer, a tremendous loss of face for the latter. On the morning of the day the incident occurred, part of the labor force went to dig guano and the others went to the wharf. Those on the wharf did not work, however, and first Spencer's son and then Spencer himself, talked to them, telling them that they must go home if they would not work. It should be noted here that apparently only Tanaka spoke both English and Japanese. Thus, one can only speculate as to the actual degree of communication between the two parties on the day of the shooting. Hither then or shortly thereafter according to Shiro, Tanaka and a delegation went to Spencer to ask him about alternating work ashore with work afloat. Testimony by Wahlers, Spencer, and Spilner indicates that Tanaka warned them that the Japanese were going to "pull down the house and make us all sore." It seems clear that Spencer had thrown a number of Japanese out of Tanaka's house earlier in the day and that he had taken a bottle of gin from Goto. When Spencer refused the original request and another to increase wages, it seems evident that the Japanese on the wharf brought in those from the guano fields. We suspect that much of the following may have been fomented by fear of collective and individual "loss of face" by the various Japanese con- cerned. Using the excuse that they had not received their day’s ration and were "weak with hunger,"17 the Japanese moved at dusk en masse for a confrontation with Spencer. Few if any bore arms. Spencer, aware that trouble was afoot, took the two revolvers that he had received from the Captain of the Ceylon and with the three others went to meet them. Shortly after they mounted the platform, they were joined by the unarmed carpenter. When the Japanese arrived, demands were made for more food and for higher pay. Since Tanaka was doing the translating, it is not clear whether the laborers actually expressed the demands or whether the demands were Tanaka's idea. In any case, four or five of the Japanese who stood on the steps of the platform began to press forward. Sit Spencer told them to stop or he would shoot. The Japanese pressed On and Spencer then fired three shots, one into the air and two into the crowd, and told the others to shoot .18 Spilner fired two shots and each of the others fired at least one. All testimony considered, it is not actually known who killed the Japanese. We suspect that Spilner, who held some animosity toward the Japanese, may well be the real villain. In his testimony he stated that he had fired one shot in the air and one into the house. Spencer, however, later testified that Spilner had told him after the shooting that, "My two shots counted all right--they got their men.” Schlemmer's Attempts to Lease Laysan In 1904, with guano deposits nearly depleted and with little profit to be gained from continued operations, the Pacific Guano and Fertilizer Company relinquished its rights to guano operations, and on 2/7 April sold "everything on Laysan...excepting houses...to Max Schlemmer...for $1750" (Anon., 1939: 19). On 6 May Max Schlemmer received an agent's commission from the Pacific Guano and Fertilizer Company.19 Schlemmer evidently believed that this document and the bill of sale conferred on him full rights to Laysan (see below), but since he wanted to make a coconut plantation of the island, he wished further to confirm his legal title. Thus, on 25 March he had applied to the Hawaiian Land Commissioner for lease to Laysan, Lisianski, and French Frigate Shoals for a period of 99 years.©20 No action was taken on Schlemmer's proposal at the time, presumably because Schlemmer had left for Laysan about the end of April, possibly aboard the schooner Robert Lewers.2l On his return to Honolulu Schlemmer pressed the issue. In his pro- posal to the governor Schlemmer had agreed, in addition to the limitation on bird killing, to: (1) pay a royalty of 50 cents per ton for all guano shipped, (2) maintain his residence on Laysan and keep a caretaker there should it be necessary for him to leave the island, (3) maintain a schooner of not less than fifty tons which could be used to bring shipwrecked persons to Honolulu from the Northwestern Hawaiian Islands, and (4) plant not less than one thousand coconut trees a year for a period of ten years. In the same letter he proposed that he kill the following numbers and kinds of birds on Laysan and that he turn over the skins to the Territorial Govern- ment which would sell them and retain 10 percent of the proceeds as a royalty: 38 Number Number Number Number Number Number Number Number Number Number Number Number Number Number Number Number Number 17. Variety Could be killed as follows: Black Widacks [Wideawakes = Sooty Terns].......5,000 a Blue Widacks [= Gray-backed Terns]........-.....2,000 a Large Black Birds [= Brown Noddies]..........++++200 a Small Black Birds [= Black Noddies]..............200 a Tropical Birds [= Red-tailed Tropicbirds]........200 a love) Birds) [i= Whaitcel ler sillemisicieieleieleielelieieieiaieieieet One Four large kinds of Mutton birds [= Bonin Petrels, Christmas Shearwaters, and pre- sumably the two color phases of the Wedge- ta Lleds Shearwajten |] ssi :) (g° 1) asdod qsnsny (9°€) (9° €)asdod Atn¢e (S°EL) (4° T) Masa (o°S)asdod (1°1)asdod aun aSdod asdod asdod (T°9T) (€°C) Masa (O°C)MASH (O°T)MASd (L°t)MISd (2°h)dSaod (6°) Misa yore (6°T) (6°T)dsdod Arenzgadg 996T G96T +961 €96T UUON x%° MIS pue dsqod au Aq pueTs~T ueskey go sfaaamns quecey °T eTqel 967 8967 L96T 58 2S) HISTORY SECTION FOOTNOTES 1. The source Stackpole cites (which we have not seen) is the "Journal" of Captain Stephen Reynolds, date cited 24 September 1824, that is in the Peabody Museum, Salem, Massachusetts. The date in the citation certainly implies that the island the Lyra discovered was sighted earlier than 1828. 2. Log of the U.S. Schooner Fenimore Cooper, Rec. Group 37, U.S. Nat. Archives, Washington. 3. See Hawaiian Star Bulletin, 1 April 1893. This lease was not to the North Pacific Phosphate and Fertilizer Company as Bryan (1942: 186) states. That firm did not yet exist. 4. Hawaiian Star Bulletin, 25 June 1890. 5. Act published in the Hawaiian Star, 1 April 1893. 6. Copy of agreement between J.A. King and the North Pacific Phosphate and Fertilizer Company, 17 April 1893, Rec. Group 126, U.S. Nat. Archives, Washington. 7. Copy of Surrender of Lease to the Commissioner of Public Lands of the Territory of Hawaii, Rec. Group 126, 31 October 1908, U.S. Nat. Archives, Washington. 8. Schlemmer had been foreman of the Japanese laborers as early as 1894 (Farrell, 1928: 414). 9. Bryan (1942: 186) stated that active guano digging began in 1892 but this statement is evidently in error. 10. Farrell, 1928: 400, 402, 459, states that the man's name was "Kosten, or something resembling that," and that he died in late 1893 or early 1894. 11. Periods of inclement weather often made it hazardous for the ships offshore awaiting their cargos. The American barque Albert was nearly beached by a stiff northeast wind in 1902. In 1905 the schooner C. Kennedy was driven ashore and totally demolished. During that summer, the C.L. Woodbury, which was taking a load of guano to Honolulu, met with heavy seas and had to return to Laysan for repairs (Anon., 1905). 12. This account is derived from articles in The Honolulu Pacific Commercial Advertiser of 8, 12-15, 18-19, 21-22 September 1900. 13. Subsequent testimony reveals that Goto, one of the two Japanese killed, was not shot dead but died of his wounds aboard the Ceylon two days later. The Ceylon was the primary carrier of supplies to Laysan and guano to Honolulu from 1900 until 3 July 1902 when it broke up as a result of bad weather and sank with a cargo of guano. The crew took to boats and made it back to Laysan 4 days later (Thrum, 1902: 154). 60 14. Spencer's first account of the shooting, given to Honolulu reporters, is somewhat at odds with his later testimony. We believe that Spencer somewhat exaggerated his first account as he thought of himself as a hero who had stemmed a bloody uprising by the "yellow peril.” 15. Although not mentioned in the first account of the homicides, Wahlers had been on the platform with the four others when the shooting occurred. 16. The McNear was lost on Dowsetts' Reef 14 May 1900 while en route to Laysan. The 33 persons aboard set out in the ship's boats on the 15th for Laysan, about 60 miles away, and arrived there 36 hours later (Lydgate, 1914; 138). 17. The various testimonies make it clear that none of the Japanese was suffering from want of food. 18. Testimony conflicts as to who fired the first shot. The Japanese agreed that Spencer's son fired the first shot but Spencer himself claims to have done so. 19. Copy of Agent's Commission from Pacific Guano and Fertilizer Company, 6 May 1904, Rec. Group 126, U.S. Nat. Archives, Washington. 20. Schlemmer to G.R. Carter, Governor of Hawaii, 14 April 1904, State of Hawaii, State Archives, Honolulu. 21. Bryan (1942: 186) stated that "On May 1, 1904, the schooner Robert Lewers made a last trip to Laysan for the final cargo of guano for Hackfeld and Company...." We suspect that "Hackfeld and Company" was in fact the Pacific Guano and Fertilizer Company which had been originally financed by Hackfeld and Company and which had several of the Hackfeld family on its board of trustees (Anon., 1939: 10). 22. Schlemmer to Carter, 17 December 1904, State of Hawaii, State Archives, Honolulu. 23. Carter to William Dutcher, President of the National Audubon Society, 29 March 1905 (Dutcher, 1905: 306). 24, American vice-consul in Japan to the Assistant Secretary of State of the United States, 3 April 1909, Rec. Group 126, U.S. Nat. Archives, Washington. 25. An article, originally published in the Japan Times (cf. The Pacific Commercial Advertiser, 20 April 1909) was apparently a cover story for these operations. It stated that the Suminoye Maru, which had sailed from Japan for "shark fishing in the Hawaiian waters” in late November 1908, had been disabled by a storm and had drifted to Laysan where it arrived on 4 January 1909. About a month later the eighteen shipwrecked "sailors" were rescued by the Niigata Maru, which had drifted to the island under the same conditions. It is unlikely that either of these ships was at Laysan for any reason other than feather poaching, since both vessels are on the 61 list of seven reported by the American vice-consul. 26. It seems likely that this ship was the one that delivered the poachers to Laysan in late April 1909. 27. The C. Kennedy, captained by Schlemmer, went aground on Laysan on 3 March 1905. The vessel was totally destroyed but no one was killed. The erew was rescued on 23 March by the U.S. Gunboat Petrel which was en route from Midway to Honolulu (Lydgate, 1914: 138; Thrum, 1905: 189). 28. Copy of agreement between Schlemmer and Genkichi Yamanouchi, dated 22 December 1908, Rec. Group 126, U.S. Nat. Archives, Washington. Subsequent events make it clear that the Japanese had no intention of mining guano but intended instead to harvest plumage for the millinery trade. Only part of the Japanese operations on the Northwestern Hawaiian Islands is known in detail; it seems likely that feather harvesting occurred on the various islands on several occasions of which we know nothing. 29. Schlemmer had received a Police Constable's Commission from the Hawaiian Government for Oahu and the outer islands on 13 May 1907. The authority was probably granted primarily for the purpose of preventing poaching (implied in a letter from Carter to H.A. Isenberg, 15 December 1904, State of Hawaii, State Archives, Honolulu). 30. Partial copy of lease between Schlemmer and the Commissioner of Public Lands of the Territory of Hawaii, 8 February 1909, Rec. Group 126, U.S. Nat. Archives, Washington. 31. W.F. Frear, Governor of Hawaii, to the Secretary of the Interior, 30 April 1909, Rec. Group 126, U.S. Nat. Archives, Washington. 32. About $54,250 worth of material was shipped to Japan, $51,250 worth was seized by the Thetis, and about $25,800 worth was destroyed on the island. 33. We have found no evidence that the Japanese knew their operatians were illegal. It seems likely that the original feather gathering party did not know of the Presidential Order since it was promulgated only about a month before they landed on Laysan. 34. W.A. Bryan to T.S. Palmer, 30 May 1910, Rec. Group 22, U.S. Nat. Archives, Washington. 35. D.B. Kuhns to T.S. Palmer, 14 November 1910, Rec. Group 22, U.S. Nat. Archives, Washington. 36. Log of the U.S. Revenue Cutter Thetis, Rec. Group 26, U.S. Nat. Archives, Washington. 37. Log of the U.S.S. Iroguois, Rec. Group 4, U.S. Nat. Archives, Washington. 38. Nutting to T.S. Palmer, Bureau of Biological Survey, 26 September 1910, Rec. Group 22, U.S. Nat. Archives, Washington. 62 39. Henshaw to Nutting, 25 April 1912, Rec. Group 22, U.S. Nat. Archives, Washington. 4O. Nutting to Henshaw, 29 April 1912; Henshaw to Nutting, 2 May 1911, Rec. Group 22, U.S. Nat. Archives, Washington. 41. Dill to Henshaw, 20 December 1911, Rec. Group 22, U.S. Nat. Archives, Washington. 42, Henshaw to Dill, 31 May 1912, Rec. Group 22, U.S. Nat. Archives, Washington. 43, T.S. Palmer to G.R. Salisbury, 18 November 1912, Rec. Group 22, U.S. Nat. Archives, Washington. 4h, Log of the U.S. Revenue Cutter Thetis, Rec. Group 26, U.S. Nat. Archives, Washington. 45, Salisbury to T.S. Palmer, 20 March 1913, Rec. Group 22, U.S. Nat. Archives, Washington. 46. Log of the Thetis, Rec. Group 26, U.S. Nat. Archives, Washington. 47, Most information concerning this visit is from Schlemmer and Schlemmer (ms.), a log of the Helene, and a diary kept by Schlemmer and his son Eric. 48, Henshaw to W.A. Bryan, 1 July 1915, Rec. Group 22, U.S. Nat. Archives, Washington. 49, J.H. Brown to the Captain Commandant of the Coast Guard, 13 December 1915, Rec. Group 26, U.S. Nat. Archives, Washington. 50. See also Log of the Thetis, Rec. Group 26, U.S. Nat. Archives, Washington. 51. Log of the U.S.S. Pelican, Rec. Group 24, U.S. Nat. Archives, Washington. 52. Gregory (1925: 10) stated that "During July [1924 Wilder] took advantage of an invitation extended by officials of the United States Navy to visit Laysan and Midway Islands--a trip which yielded collections and notes on land and marine fauna." Despite considerable searching in the files of the Bureau of Biological Survey in the National Archives and in the files of the Bernice P. Bishop Museum, Honolulu, we have been unable to find any of his observations. 53. Warner (1963: 7) reported that "living rabbits were reportedly seen on Laysan by a visitor to the island in 1926." We have no other record of this visit. 54. Log of the U.S.S. Marblehead, Rec. Group 80, U.S. Nat. Archives, Washington. 55. An unpublished four-page report by Wilder (ms. b) gives a little 63 additional data and a few notes on birdlife which have been incorporated into this report. 56. Log of the U.S.C. and G.S.S. Pioneer, Rec. Group 27, U.S. Nat. Archives, Washington. 57. Log of the U.S.C.G.S. Reliance, Rec. Group 26, U.S. Nat. Archives, Washington. 58. Log of the U.S.C.G.S. Reliance, Rec. Group 26, U.S. Nat. Archives, Washington. 59. Most information on such visits may be found in the scientist's logs and narrative reports of each cruise. These data are filed at the Bureau of Commercial Fisheries in Honolulu. 60. Svihla (1959: 227) gives the result of a seal count made during this visit. He implies incorrectly that Laysan was visited during January. 61. The BSFW assumed direct responsibility for inspection, patrol, and management of the refuge in 1964 when a refuge manager was assigned to Hawaii. 64 VEGETATION The first fully documented botanical collection from Laysan was made by Schauinsland in 1896 .* Subsequently, the following collections were made: J.O. Snyder, May 1902; W.A. Bryan, April and May 1903 and April 1911; D.T. Fullaway, December 1912; Fullaway and E.L. Caum, April 1923; G.P. Wilder, August 1930; G.D. Butler, April and July, 1959; C.W. Daniel, July 1959; C.H. Lamoureux, September 1961; J.W. Beardsley, June 1962; R.T. Tsuda, December 1963; C.R. Long and A. Young, September 1964; P.C. Shelton, June 1966; and possibly others for which data are not available. Several highly useful papers have been published on the flora and vegetation of Laysan. Schauinsland (1899) dealt at length with conditions on the island in 1896 and reported the first collection of specimens on which additional notes were published by Bitter (1900). Christophersen and Caum (1931) reported conditions in 1923 when the island had been largely de- nuded of vegetation by rabbits. Their annotated list included valuable material on most specimens collected up to that time. The most recent thorough summary of the vascular flora was by Lamoureux (1963) whose account, as well as that of Christophersen and Caum, should be consulted for historical accounts of the vegetation not included here or in the history section of this account. Lamoureux also gave a brief annotated list of the vascular plants recorded from Laysan through 1961; discussed vegetative associations; and presented a vegetation map which, in general, if not in detail, well represents the present distribution of the primary associations. Lamoureux's account has been supplemented recently by Tsuda (1965) who gave brief observations on the vascular flora as observed in December 1963. *Two previous collections reportedly were made. "25 varieties of plants” were collected in 1859 by Brooks (1859: 501) and "twenty-one species of flowering plants" were collected by Lyons (1890: 91) in 1890. The loca- tions of these collections, doubtfully still in existence, are unknown. acelin ee 65 His paper primarily summarized current knowledge of the algal flora of Laysan and should be consulted by those interested in that aspect of the island's botany. Other recent work includes that by St. John (1970) who designated three new species of Sicyos from Laysan. One of these, Sicyos semitonsus and Pritchardia sp. (regarded by some as probably a distinct species), are the only vascular plants endemic to Laysan. Two varieties, the still extant Cyperus pennatiformis var bryanii, and the extirpated Santalum cuneatum var laysanicum were regarded by Lamoureux (1963: 8) as endemic. Fosberg (1962: 34), however, does not consider the latter distinct from plants found on Oahu. Although relatively little new information has been added recently, we nevertheless think it worthwhile to update knowledge of the island flora and to present a brief annotated list of native and introduced vascular plants. At present 42 species of vascular plants are known from Laysan, four more than reported by Lamoureux (1963: iba The difference consists solely of plants introduced by man. Two (onion and potato) probably would not have successfully colonized the island in any case. The grass, Cenchrus echina- tus, is a tenacious weed and although believed to have been extirpated in September 1969, may well be found by future observers. The fourth, Conyza bonariensis, a weedy composite, seems well established despite efforts at eradication and will probably become much more numerous on Laysan in ensuing years. Twenty-four species were present on Laysan in 1961 (Lamoureux, 1963: 2)** whereas 27 were present in 1969. The additional species resulted from the unintentional introduction of Conyza bonariensis, by the intentional intro- duction of Chenopodium oahuense (formerly an abundant species on Laysan) and by the reappearance of Solanum nigrum. Present data are insufficient to justify any major emendation of Lamoureux's vegetation map (Lamoureux, 1963: Fig. 1) but examination of recent (1966) aerial photographs (Fig. 4) and personal ground observations suggest two changes. The large, predominately bare, sandy area on the southwest portion of the island is not indicated on his map. Secondly, Scaevola is somewhat better developed at present, particularly on the east side of the island, than his map suggests. *Only 36 species are included in Lamoureux's annotated list. Both Cyperus laevigatus L. and Sicyos microcarpus Mann were evidently omitted through oversight. Specimens of the latter were recently attributed to distinct species by St. John (1970). **Lamoureux's annotated list reports only 22 species for reasons listed in the preceding footnote. 66 Annotated List The following list of species closely follows the able summary of Lamoureux (1964), particularly for species no longer present on Laysan. For completeness we have included all species recorded as present on Laysan whether or not the species ever became established. Considerable attention has been given to details of plant introduction, whether de- liberate or accidental, since the future composition of the terrestrial flora may well be dependent on these events.* Species formerly established on Laysan but no longer growing there are indicated by asterisks. Known introductions not known to have become established are enclosed in brackets. Names of species currently present on Laysan are unadorned. GRAMINEAE *Cenchrus agrimonioides var. laysanensis F. Br. Specimens were collected in 1896, 1902, 1903, and 1911 but the species had disappeared from Laysan by 1923 (Christophersen and Caum, 1931: 10). In December 1963, seeds collected on Kure Atoll in September 1961 were planted north of the north coconut grove and near the northwest edge of the lagoon (Tsuda, 1965: 26-27). These seeds evidently did not germinate as none was found by visitors during the next few years. *(?)Cenchrus echinatus L. This species was probably introduced by military personnel in the 1960's. A single plant found near the campsite on the northwest side of the island was destroyed in March 1969 by BSFW personnel. Two more plants flowering in the same general area in September 1969 were also destroyed. Cynodon dactylon (L.) Pers. Bermuda grass, first collected on Laysan in 1903 (Lamoureux, 1963: 2), was introduced from Honolulu by employees of the guano company and was scat- tered over the island by 1905 (Wilder, 1905: 392). It was not found in 1911 or 1923 but in 1930 Wilder collected a specimen from a patch growing at the edge of the lagoon. Specimens were subsequently collected in 1959, 1961, 1963 and 1964. At present, Cynodon occurs around the lagoon with dense stands along the northern perimeter (Fig.25) and with poorest development on the west side. [Melinus minutiflora Beauv. ] Molasses grass was planted in 1930 (Wilder, ms. b) but was not found by subsequent observers. *Species other than those included in the list below were certainly intro- duced. Existing records of the 1924 and 1930 plantings by Wilder are known to be incomplete. Figure 26. Looking southeast toward the lagoon, dense stand of Cynodon dactylon much infiltrated by Ipomoea pes-caprae in foreground; Eragrostis variabilis and Cocos nucifera beyond. POBSP photograph by P.C. Shelton, 21 June 1966. Sesuvium portulacastrum (left), Cyperus laevigatus (center) and Heliotropium curassavicum (center and right) along west shore of lagoon. POBSP photograph by P.C. Shelton, 21 June 1966. 68 Eragrostis variabilis (Gaud.) Steud. This bunchgrass, one of the dominant plants on the island prior and subsequent to denudation of the island vegetation by rabbits, reached its nadir in 1923 when no living plants were found (Lamoureux, 1963: 2). Seeds and rhizomes were planted at that time but even as late as 1936 the plant was evidently not very abundant (Coultas, ms.). Not until recent years has Eragrostis regained a good measure of its former abundance on the island. At present it forms dense stands on the inner slopes of the island, par- ticularly on the western side (Fig.24). Eragrostis was collected in 1896, 1902, 1903, 1911, 1930, 1959, 1961 1963 and 196}. j é i ' *Lepturus repens (Forst.) R. Br. This small bunchgrass was formerly common growing near the beaches, particularly on the north side of the island (Schauinsland, 1899: 99). It was collected in 1896 and 1903 but not found subsequently. In 1923 plants from Pearl and Hermes Reef were introduced by the Tanager Expedition (Christophersen and Caum, 1931: 14) but albatross pulled up much of the grass (Wetmore, ms.). *Sporobolus virginicus (L.) Kunth. This grass was found growing near the ocean shore in 1896 (Schauinsland, 1899: 99). Specimens were reported collected in 1896 and 1903 (Christophersen and Caum, 1931: 22). (The 1903 specimen listed by Christophersen and Caum was apparently misdetermined and is actually Cynodon dactylon). CYPERACEAE Cyperus laevigatus L. This rush-like plant now grows in dense stands (Figs. 26,3) around the perimeter of the lagoon as it did in 1896. Specimens were collected in 1896, 1903 and 1911 but no Cyperus was found in 1923. This species was next Sener by Wilder in 1930 and more recently in 1959, 1961, 1962, 1963, and 1964. Cyperus pennatiformis var. bryanii Klukenthal This sedge, collected in 1896, 1902, 1903, and 1911 was absent in 1923 but was thereafter collected in 1959, 1962, 1963 and 1964. Schauinsland (1899: 98) found this species confined to the vicinity of the lagoon but widespread in this area. In 1903 Bryan considered it not at all common and saw only a few bunches near the southwest corner of the lagoon. It presently has much the same distribution as in 1903, being largely confined to an area near the southern end of the lagoon (Tsuda, 1965: 24, Fig. 7; Fig. 27). Tsuda's specimen data indicate that he found this species growing only in a 69 Figure 27. Figure 28. Stand of Cyperus pennatiformis near south end of lagoon. POBSP photograph by P.C. Shelton, 14 June 1966. Casuarina tree inland from campsite on northwest shore of island. POBSP photograph by P.C. Shelton, 21 June 1966. 7O small area about 50 meters southwest of the coconut grove present near the southeastern perimeter of the lagoon. Fimbristylis cymosa R. Br. Fimbristylis, first collected in 1930, may occur on Laysan as the result of plantings made in 1923 (Lamoureux, 1963: 3) or thereafter. This sedge is presently widely distributed over the drier portions of the island, being abundant at times on the slopes of the western interior (Fig.9 ). Recent collections were made in 1959, 1961, 1962, 1963, and 1964. PALMAE Cocos nucifera L. Coconuts were planted on Laysan on several occasions but relatively few survived for very long. In 1902 four palms that had been imported from Honolulu were seen by Thomas (ms.). By 1905 the number had been reduced to two, thought by Wilder (1905: 392) to be about 12 years old. At this time another tree was planted and 20 sprouted nuts were left with the island manager for planting. Attempts to establish coconuts continued. In 1912 the Bureau of Biological Survey planted 100 sprouted nuts, surrounding them with woven wire guards (Salisbury, ms.). Evidently none long survived for only two trees were found by the Tanager Expedition (Christophersen and Caum, 1931: 13} 9 Figs.19,21). The latter are most likely the same trees mentioned by Wilder in 1905, and subsequently seen by Bailey (1956: 24) in 1912 and by Coultas (ms.) in December 1936.* Most recently young trees were planted in 1959 (Lamoureux, 1963: 3). By March 1961 two small groves were well established. One, comprised of "some 13" trees,was near the northwest corner of the lagoon; the other was comprised of seven trees near the southeast edge of the lagoon (Woodside and Kramer, ms.). All survived through December 1963 when Tsuda (1965: 23) found 12 to the northwest and seven to the southeast. By September of the following year only seven of the northern and three of the southern trees were thriving. In 1969 the remaining groves consisted of nine northern and two southern trees (BSFW, cf. Figs.25, 35, 36). [Phoenix dactylifera L. ] Date palms were planted in 1930 by Wilder (ms. b) but did not survive. [Pritchardia pacifica Wendl] This species was planted in 1923 but was not found subsequently. Greg- ory (1931: 16) states that Pritchardia was planted during Wilder's 1930 visit *Gregory (1931: 16) indicated that Wilder planted coconuts during his 1930 visit but Wilder's report of his visit (ms. b) fails to mention this species. tL but this species does not occur in the list appearing in Wilder's (ms. b) report of his trip. *Pritchardia sp. A small fan palm was seen on Laysan by Isenbeck in 1828, Paty in 1857, Brooke and Brooks in 1859, Lyons in 1890 and by Munro in 1891 (Lyons, 1899: 90; Lamoureux, 1963: 3). At least five trees up to 15 feet tall were present in 1859 but Schauinsland (1899: 99-100) found only numerous dead stumps and roots, some in the northern part of the island and others not far from the lagoon on the southeast part. Christophersen and Caum (1931: plate VI) include a photograph of two still living trees. This photograph, listed as having been taken between 1891 and 1896 by an unknown photographer, may have been one of the series of photographs of Laysan that was taken by J.J. Williams in 1892 and/or 1893. This palm has been thought to be the same as that occurring on Nihoa (Pritchardia remota) or alternatively, to be a distinct species (Lamoureux, 1963: 3). LILIACEAE *Allium sp. Onions were planted in 1859 (Brooks, 1859: 501) but did not survive. A few plants, evidently growing from garbage left by the HIRAN operation, were found growing on Laysan in June 1962. These were all uprooted (Kramer and Beardsley, ms.). CASUARINACEAE Casuarina equisetifolia L. Tronwoods were introduced to Laysan on at least three occasions. In September 1905 Wilder (1905: 392) planted a box of ironwood trees, one of which may have survived to be recorded in 1923 by the Tanager Expedition (Christophersen and Caum, 1931: 13; Fig.2l) and by recent visitors. In 1923 two pounds of ironwood seeds were sown (Wetmore, ms.) and in 1930 Wilder planted an unstated number of trees (Wilder, ms. b; Gregory, 1931: 16). Some of the latter evidently survived for some years since Coultas (ms.) found five trees in December 1936. One was near the old buildings and the other four were near the north side of the lagoon. At present, the single remaining tree (Fig. 28) often suffers some damage from wind and salt spray of winter storms but is usually luxuriant and supports the largest Black Noddy colony on the island. Specimen material from this tree was collected in 1961, 1963, and 1964. ie SANTALACEAE *Santalum ellipticum var. littorale (Hillebr.) Skottsberg The Laysan sandalwood, listed by others as Santalum freycinetianum Gaud. and as §. cuneatum var. laysanicum Rock, was first collected in 1896 when it was found along the shoreline. At that time it was most abundant on the northwest side and was the largest plant on the island growing to as much as 2.5 meters tall (Schauinsland, 1899: 98). It was subsequently collected in 1902, 1903, and 1912 but by 1923 the only remaining plants were found in a small patch along the southwestern side of the island (Christophersen and Caum, 1931: 10, pl. VII). Some of the leafless stumps were trying to sprout but evidently did not long survive as none was found by subsequent observers. POLYGONACEAE [Cocecoloba uvifera L. | Gregory (1931: 16) stated that this species was planted by Wilder in 1930. CHENOPODIACEAE [Atriplex muelleri Benth. ] Seeds of the Arizona salt bush were sown by Wilder (ms. b) in 1930 but no plants were found subsequently. Chenopodium oahuense (Meyen) Aellen In 1896 this shrub was second in abundance only to Eragrostis (Schau- insland, 1899: 98). Specimens were collected then and in 1902 and 1903. In 1903 Chenopodium was still a common plant but between then and 1911 disappeared from the island. (Coultas, ms., noted seeing a few small bushes in December 1936 but there are no specimens to verify this report.) In recent years a number of efforts were made to reestablish the aweoeo on Laysan. In June 1962 HDFG personnel planted seeds from Nihoa on the northwestern interior slope, a little more than halfway from the usual campsite to the lagoon shore. In December 1963 Tsuda found no evidence that any of these seeds had germinated. During Tsuda's visit other Cheno- podium seeds, obtained on Nihoa in December 1961, were planted at two locations in the northwestern portion of the island (Tsuda, 1965: 26-27, see Cenchrus agrimonioides above). Subsequent observations by the POBSP and others suggest that this planting also failed. Other seeds, from French Frigate Shoals, were subsequently introduced by BSFW personnel. A number were planted near the northwest shore of the lagoon in March 1966 and seeds were broadcast around the campsite on the northwestern rim of the island in September 1966. A single plant was found growing in the latter area in September 1967. In March 1969 a single plant observed in this area was producing seed and two thriving plants were found there the following September (BSFW). 73 AMARANTHACEAE *Achyranthes splendens var. reflexa Hillebr. In 1896 Schauinsland (1899: 97) found a small patch approximately 100 paces in diameter on the northwest side of the island near the beach. Specimens were subsequently collected by Bryan in April 1903 (Christophersen and Caum, 1931: 26) but the plant was not found by subsequent collectors. In December 1963, seeds, collected on Kure Atoll in September 1961, were sown north from the northernmost coconut tree and on the northwestern side of the island near the lagoon edge (Tsuda, 1965: 26-27), but the introduction was unsuccessful. *Amaranthus viridis L. Schauinsland found individuals of this species along stagnant pools in the southern guano fields and scattered among the Chenopodium at the north end of the island near the lagoon (Bitter, 1900: 135). Specimens were collected subsequently in 1902 and 1903, the latter being found along the tramway and in the guano beds (Christophersen and Caum, 1931: 26). NYCTAGINACEAE Boerhavia diffusa L. Except for 1923, when only dead plants were seen, this plant has ap- parently always been a major constituent of the flora. It is now, as in 1896, widely distributed over the island (Fig.9 ). Specimens were collected in 1896, 1902, 1903, 1930, 1959, 1961, 1963 and 1964. ATZOACEAE Sesuvium portulacastrum L. This is a common species forming luxuriant mats in low areas bordering the lagoon (Figs.26,29; cf. Bailey, 1956: 46; Christophersen and Caum, 1931: pl. VIY. First collected by Schauinsland in 1896, it was the only native species at all abundant in 1923 (Lamoureux, 1963: 40; Fig.20). After 1896 specimens were collected in 1902, 1903, 1911, 1923, 1959, 1961, 1963 and 1964. Sesuvium covered large areas just above the high water mark of the lagoon in 1930 but none was collected (Wilder, ms. b). PORTULACACEAE Portulaca lutea Sol. This species is present in most collections from Laysan through 1964 .* *Lamoureux (1963: 4) indicates that this species was not collected by Wilder in 1930 but Wilder's (ms. b) report states that it was collected. Perhaps the specimen or specimens were later lost. Th In 1896 it was mostly found in scattered localities in drier parts of the island (Schauinsland, 1899: g4). By 1923 only a small patch within a Sesuvium patch east of the lagoon remained (Christophersen and Caum, 1931: 11; Fig.18). At present it is once more widely distributed over the island. Portulaca oleracea L. This species, first collected in 1959, is apparently a recent intro- duction (Lamoureux, 1963: 9).* Specimens were subsequently collected in 1961 and 1963 (Lamoureux, op. cit.; Tsuda, 1965: 25). The latter was noted as occurring on the southwestern side of the island near the beach. CRUCIFERAE *Lepidium bidentatum var. o-waihiense (C. & S.) Fesb. This species was collected only by Schauinsland (1899: 94), who found a single stunted shrub 30 cm. high on the east side of the island near the beach. Seeds from Kure Atoll were sown in two localities (see Cenchrus agrimonioides above) in December 1963 (Tsuda, 1965: 26-27) but the intro- duction failed. CAPPARTDACEAE Capparis sandwichiana DC The puapilo was evidently first seen by Brooks in 1859 and was present on the island through at least 1903 (Lamoureux, 1963: 5). It was not found again until collected in 1930 by Wilder and was collected subsequently in 1959, 1961, and 1964. In 1896 it occurred abundantly primarily on the west side of the island (Schauinsland, 1899: 94). In 1903 it occurred in patches all over the higher parts of the island; Wilder found it on higher elevations of the sand dunes. All recent collections were from the west side of the island usually within or close to the bordering fringe of Scaevola south of the Casuarina tree. LEGUMINOSAE [Caesalpina crista L.] Single seeds were found washed up on the beach in 1923 (Christophersen and Caum, 1931: 13) and in 1963 (Tsuda, 1965: 26). [Canavalia ensiformis (L.) DC] This species was planted by Wilder in 1930 but did not survive. * Lamoureux (1963: 4) points out that specimens formerly attributed to this species by Schauinsland, Bitter, and Christophersen and Caum are in fact examples of Portulaca lutea. 1) Figure 29. Cyperus, Sesuvium portulacastrum, and Tribulus cistoides along southeastern side of lagoon. POBSP photograph by P.J. Gould, October 1966. 76 [Dioclea altissima (Velloso) Rock] [Dioclea violacea Mart. ] [Entada scandens (Roxburg) Benth. ] Seeds of all three species have been found on Laysan. In 1923 two seeds of D. altissima were found on the north beach and a single seed of E. scandens was found on the south side of the island, halfway between the lagoon and the shore (Christophersen and Caum, 1931: 13). In 1963 a seed of D. violacea was found on the beach by Tsuda (1965: 26). [Haematoxylon campechianum L. ] [Leucaena leucocephala (Lam. de Wit)] Both species were planted by the Tanager Expedition in 1923 (Lamour- eux, 1963: 6; Wetmore, ms.) but neither survived. [Mucuna gigantea (Willd.) DC] [Mucuna urens (L.) DC] [Mucuna sp. ] Seeds of the above plants have washed up on Laysan with some frequency. M. gigantea was found in 1923, 1930, and 1963 and urens was found during the latter visit (Christophersen and Caum, 1931: 26; Wilder, ms. b; Tsuda, 1965: 26). A sprouting seed of Mucuna sp. was found in 1962 (Kramer and Beardsley, ms.) and Shelton collected a seed from the shore of the lagoon in 1966. ZYGOPHYLLACEAE Tribulus cistoides L. First collected in 1896, this species has appeared in most subsequent collections. In 1896 this plant was found throughout the island, particularly in drier areas but by 1923 only tiny seedlings and numerous seeds could be found (Christophersen and Caum, 1931: 11). In 1930 it grew in many places on the island and at present can once again be found commonly throughout the island (Tsuda, 1965: 26, Fig. 8 ; Fig.29). Collections were made in 1896, 1902, 1903, 1911, 1923, 1930, 1959, 1961, 1963 and 1964. EUPHORBIACEAE [Aleurites moluccana (L.) Willd. ] Candlenut seeds were found on Laysan on at least three occasions. In 1902 Snyder found several seeds in the interior of the island (Fisher, 1903: 788) and more were found along the beaches in 1923 (Christophersen and Caum, 1931: 13). In 1966 Shelton collected a single seed from the shores of the lagoon. MALVACEAE *Hibiscus tiliaceus L. Hau was introduced early in the 1900's, possibly in 1905. At that Hol time Wilder (1905: 392) planted branches. In 1923 three trees were grow- ing near the buildings left by the guano workers (Christophersen and Caum, 1931: 13). During that visit ten plants, a packet of seeds, and 40 seed branches were planted (Wetmore, ms.) but none was found on the island during later visits. [Thespesia populnea Sol. ] Several pounds of milo seeds were sown by the Tanager Expedition in 1923 and several trees were planted in 1930 by Wilder, but the species never became established. CONVOLVULACEAE Ipomoea indica (Burm.) Merr. In 1896 Schauinsland (1899: 96) found this species dispersed over the island except in the vicinity of the lagoon. It was evidently considerably less common than the following species. Snyder and Bryan collected specimens in 1902 and 1903 but the species was apparently not seen thereafter until 1959 ees collected by Daniel. Subsequently it was collected in 1961, 1963 and 1 : At present this morning glory is apparently largely confined to low areas near the lagoon. Despite an extensive search of other areas in 1963, Tsuda (1965: 24) could find but one small patch--on the southwest side of the island, halfway between the lagoon and the beach. Ipomoea pes-caprae (L.) Sw. Beach morning glory, first noted by Munro in 1891 (Lamoureux, 1963: 5), was in 1896 found everywhere in higher places along the beach (Schauinsland, 1899: 96). By 1903 the species was evidently much less common and was ap- parently absent by 1911 (Christophersen and Caum, 1931: 11). In 1923 only two seeds were found. At that time a half-pound of seeds was sown (Wetmore, ms.). By 1930 beach morning glory was well reestablished, particularly in sandy, more elevated areas, and extended on the southeast almost to the edge of the ocean (Wilder, ms. b). In 1959 the first collections since 1903 were made and specimens were since collected in 1961, 1963 and 1964. It is now once again widespread, occurring on almost all areas of the island (Fig. 30 ; Lamoureux, 1963: Fig. 4). [Convolvulus sp. ] Wilder (ms. b) planted Convolvulus in 1930 but the introduction failed. HYDROPHYLLACEAE Nama sandwichensis var. laysanicum Brand Nama was first reported by Schauinsland (1899: 96) who found it Figure 30. Black-footed Albatross chick in dense growth of Ipomoea pes-caprae. POBSP photograph by A.B. Amerson, Jr., 10 March 1964. Figure 31. Nama sandwicensis near north end of island. North Cocos grove in mid-background. POBSP photograph by P.C. Shelton, 21 June 1966. 79 distributed around the island on the higher parts of the beach. Like many other species it decreased in abundance under the onslaught of the rabbits and was not found in 1923. It now approaches its former abundance and is found most abundantly on the wide sandy beaches of the northern end of the island (Fig.31 ). Specimens were collected in 1896, 1903, 1911, and from 1959 through 1964. BORAGINACEAE [Cordia subcordata Lam. ] Trees planted by Wilder (ms. b) in 1930 did not survive. Heliotropium curassavicum L. In 1896 seaside heliotrope was confined to the water-free part of the lagoon. It was abundant there through at least 1903 (Christophersen and Caum, 1931: 11) but was not found again until collected by Wilder in 1930 (Lamoureux, 1963: 5). It appears in all recent collections through 1964 and now occurs both on the outer beach and in the inner band of vegetation sur- rounding the central lagoon (Figs. 26,32). Tournefortia argentea (L.f.) This species was first found on Laysan in September 1961 when a small tree was found growing at the top of the northwest beach between the Nama and Scaevola associations (Lamoureux, 1963: 12). In June 1962 the plant ap- peared in excellent condition but in December 1963 appeared all but dead, likely from the effect of salt spray. On closer examination a few new shoots were seen (Tsuda, 1965: 25, Fig. 4). On subsequent visits (March and Septem- ber 1964) the tree was thought to be dead but it still survived in December 1967 an Specimen material from this plant was collected in 1961, 1963 and 1964. Two additional small trees were found alive in recent years. Kridler (BSFW) noted in December 1967 that two trees south of the northwest landing were thriving and in bloom. At least one of these trees, and probably both, had been present more than a year earlier (Fig. 33). LABIATAE *Phyllostegia variabilis Bitter Specimens found by Schauinsland in 1896 were later described as this species by Bitter (1900: 437). In 1896 it was scattered near the beach of the west and east sides (Schauinsland, 1899: 97). It was still present in small patches in 1903, mostly on the windward side (Christophersen and Caun, 1931: 11). It disappeared from Laysan before 1911. 80 | Figure 32. Cyperus laevigatus and Heliotropium curassavicum near north- { western corner of lagoon. POBSP photograph by P.C. Shelton, 21 June 1966. Figure 33. Small Tournefortia at top of west beach south of northwest campsite. POBSP photograph by P.C. Shelton, 15 June 1966. | 81 SOLANACEAE [Nicotiana glauca Grah. ] Tree tobacco was introduced by Wilder in 1930 but did not long survive. Nicotiana tabacum L. Introduced early in the 1900's, tobacco was first collected by W.A. Bryan in 1911. In 1923 a rather large patch was found near the southern end of the lagoon and spreading through the southern and southwestern por- tions of the island (Christophersen and Caum, 1931: 13; Fig.22). Specimens were collected subsequently in at least 1930, 1959, 1963 and 1964. At present scattered plants can be found in many locations in the interior of the island (Fig. 34) and apparently most abundantly on the west side. *Solanum nelsoni Dunal In 1896 Schauinsland (1899: 96) found this species on small sand dunes in a few places near the north beach. The only further record of this plant, once described as Solanum laysanense (Bitter, 1900: 433-435), was a few patches observed in the same area in 1903 (Christophersen and Caum, 1931: 12). Seeds collected on Nihoa were planted in the northwestern interior of the island in June 1962, and others, collected on Kure in September 1961, were planted in this area and north of the northern coconut trees in December 1963 (Kramer and Beardsley, ms.; Tsuda, 1965: 26-27). Subsequent observers found no evidence that any of these survived. Solanum nigrum L. The black nightshade was first collected on Laysan (as Solanum nodi- florum Jacq.) by Wilder in 1930 who found it growing along the eastern end of the lagoon above high water. It has been collected by many recent ob- servers but is apparently limited in its distribution on the island. Beardsley collected a specimen in 1962 as did Tsuda in 1963. The latter (Tsuda, 1965: 26) found but two plants growing near Scaevola along the trail leading from the beach to the campsite on the northwest side of the island. In 1964 Young collected a specimen from the same area and Long collected two other specimens, one growing in association with Eragrostis and Ipomoea on the southwest side of the lagoon and another growing in an open area on the west slope of the interior in association with the above and Fimbristylis. In 1966 Shelton collected a plant from among Scaevola along the western rim of the island. *Solanum tuberosum L. Potatoes were planted in 1859 (Brooks, 1859: 501) but did not survive. Several found growing in 1962 were uprooted. These were probably intro- duced during 1961-62 by military personnel (Kramer and Beardsley, ms.). 82 Figure 34. Nicotiana tabacum (left foreground) and Conyza bonariensis (right foreground) at top of west seaward slope. POBSP photograph by P.C. Shelton, 14 June 1966. Figure 35. Scaevola taccada, Cyperus laevigatus, and Ipomoea pes- caprae along small slough near southeast corner of lagoon. South Coeos grove in mid-background. POBSP photograph by R.B. Clapp, 18 March 1968. 83 CUCURBITACEAE [Cucurbita pepo L.] Pumpkins were planted in 1859 (Brooks, 1859: 501) but did not survive. Sicyos atollensis St. John St. John (1970) recently examined specimens of Sicyos from the north- western Hawaiian Islands and described six new species which include all Sicyos occurring on these islands. S. atollensis, first collected by Schauinsland in 1896, was subsequently collected in 1911, 1930 and 1962. The 1896 and 1903 specimens are evidently those formerly attributed to Sicyos hispidus Hillebrand by Christophersen and Caum (1931: Bile Sicyos laysanensis St. John This species was first collected by W.A. Bryan in April 1903 and sub- sequently listed as Sicyos sp. by Christophersen and Caum. Subsequent collections were made in 1911, 1959, 1962, 1963 and 1964* (St. John, 1970: 442), It has been collected at many localities near the lagoon and may be the most abundant species of Sicyos on Laysan judging from the greater frequency with which it has been found in recent collections. [Sicyos nihoaensis St. John] Seeds of this species, at the time believed to be Sicyos microcarpus Mann, were brought from Nihoa to Laysan in June 1962 and were planted inland from the campsite on the northwestern perimeter of the island (Kramer and Beardsley, ms.). Subsequent observers found no evidence that these seeds ever germinated. Sicyos semitonsus St. John This species was described from specimens collected from a flat area at the northwest end of the lagoon by Long on 19 September 1964. Details of the distribution on Laysan of this and other species of Sicyos must await a detailed examination of a considerably larger series of specimens than has been available previously. Sicyos sp. Several specimens from at least September 1964 and June 1966 have not been critically re-evaluated since the publication of St. John's paper in 1970. ¥st. John (1970) lists a specimen (C.R. Long #2,370) as both the holotype of Sicyos semitonsus (p. 448) and as an example of Sicyos laysanensis (p. 42). 84 GOODENTACEAE Scaevola taccada (Gaertn.) Roxb. Scaevola” was in 1896 limited to the zone along the beach, abundant along the west beach, more stunted along the east side (Schauinsland, 1896: 95-96). By 1923 the growth had been reduced to three "exceedingly poor” patches, one southeast of the lagoon halfway between shore and lagoon, one on the north side of the island, and another in the northeastern corner of the island (Christophersen and Caum, 1931: 12). During this visit and in 1930 Wilder planted seeds and plants, respectively (Wetmore, ms.; Wilder, ms. b3; Gregory, 1931: 16). At present this species has largely recovered its former abundance and in fact may be more widespread than in 1896. It is now found in abundance in a well developed band around the outer perimeter of the island just inland from the beach (cf. darker vegetation in Figs.2-4 ). Scattered plants are found inland on the west side of the island to within about 50 yards of the lagoon. It occurs in greater abundance in the eastern interior, occasionally forming well developed stands near the lagoon edge (Fig.35 ). The recovery of this species no doubt played an important role in the recovery of the seabird populations since it is the principal nest site for populations of Great Frigatebirds, Red-footed Boobies, and Black Noddies and provides as well the necessary cover for nesting Red-tailed Tropicbirds. COMPOSITAE Conyza bonariensis (L.) Cronq The hairy horseweed was probably introduced by the HIRAN operation of the early 1960's. It was first found growing just south of the campsite on the northwest side of the island. All plants seen at that time were up- rooted (Tsuda, 1965: 23). Specimens were subsequently collected from the same area in 1964 and 1966. Despite continued effort in recent years, BSFW personnel have been unable to eradicate the plant and it continues to survive and spread (Fig. 34). In September 1969 numerous dry plants retaining much seed were seen (BSFW). *Lipochaeta integrifolia (Nutt.) Gray In 1896 it occurred in a band just oceanward from the association of Sesuvium, Heliotropium and Cyperus bordering the central lagoon (Schauinsland, *We have records indicating that specimens were collected in 1896, 1902, 1911, 1923, 1930, 1959, 1963 and 1964. Lamoureux (1963: 6) states that the species "is represented in all collections from Laysan" which implies that it was also collected in 1903 and 1961. Christophersen and Caum (1931: 38), however, fail to list any specimens from 1903. 85 1899). Specimens were collected by Schauinsland and by Bryan (in 1903) who found it growing in the fine guano around the edge of the lagoon. It was not reported by later observers and presumably is one of the species that vanished under the onslaught of rabbits. Seeds from Kure were sown in two localities (see Cenchrus agrimonioides, above) in December 1963 (Tsuda, 1965: 27), but no plants were reported by later visitors. Pluchea indica (L.) Less. Pluchea is a recent introduction, first collected by Butler in April 1959 and subsequently in 1959, 1961, 1963 and 1964. It occurs in many areas around the lagoon, being largely absent from the western border and reaching its maximum development at the northwest (Fig.36)and northeast corners of the lagoon. GUTTIFERAE [Calophyllum inophyllum (L.) Sol] LECYTHIDACEAE [Barringtonia asiatica (L.) Kurz] COMBRETACEAE [Terminalia catappa L.] [Terminalia myriocarpa Huerck & Muell.-Arg. ] [Conocarpus erecta L. ] The five species listed above were all unsuccessful introductions to Laysan. All five were planted in 1930 and Calophyllum and Barringtonia were planted in 1923 as well (Wetmore, ms.; Wilder, ms. b). 86 dsdod *punoisyoeq ut sAorg $0909 320K *996T euNr Te ‘uo4TeYS °o°a kq yders0z0ud *“uooseT FO yaz0U" BoTPUT eeyonTd jo pueig “QE sanst ya 87 LAYSAN ISLAND FAUNA Introduction Laysan has long been the most familiar biologically of the North- western Hawaiian Islands and was one of the first to be exploited. Its relatively easy access, large area, fresh water supply, and natural re- sources permitted profitable occupancy for over a decade. lLaysan also had the most remarkable biota of any island in the chain with a very high degree of endemicity including four (of 27) plants, five birds, and a number of insects. The combination of tolerable living conditions at the guano headquarters and the remarkable biota early attracted several biologists and their reports soon made Laysan synonymous with teeming colonies of fearless seabirds--a veritable paradise for biologists. Unfortunately island occupancy also resulted in a continuing conflict between the native biota and man and his introductions. In less than 35 years this conflict resulted in the destruction of two endemic plants (and nine other native species), three endemic birds, and a number of the endemic insects. Other populations, notably those of seals and turtles, were gravely depleted and only the timely arrival of the Tanager Expedition prevented complete destruction of the vegetation and the en- tire endemic biota. Reports published prior to and including Schauinsland's suggest that the island flora was little affected by man through 1896. One species, the endemic palm, Pritchardia sp., had become extinct and only one introduced species, Amaranthus viridis, was yet present. Schau- insland'’s description of the fauna also agrees with those of Palmer and Munro and again suggests minimal disturbance by man. He listed 5 endemic land birds, 17 breeding seabirds, and 18 transients and acci- dentals. Three of the endemics later became extinct. Subsequently many additional transients and accidentals were recorded. However, the number of species of breeding seabirds has not changed although great population changes have occurred in the last seventy-five years. Seabird colonies flourished throughout the 19th century. Although the island was occupied during the last part of this period, most birds were not adversely affected by the guano operation. In fact this occu- pancy may have protected the island from the ravages of feather harvest- ers. Some populations were, however, being affected before termination of the main guano operation in 1904 and feather hunters did great damage in 1909, 1910, and 1915. The designation of Laysan as a bird reservation in 1909 and subsequent patrolling and inspection by U.S. Revenue Cutters probably prevented even greater destruction. Ironically the most lasting damage to the Laysan biota resulted not from feather raids but rather from the introduction of rabbits by a well-meaning island manager. Rab- bits were introduced about 1903 and were already causing significant damage to the vegetation by 1911. Efforts to eliminate all rabbits in 191é2 and 1913 failed and not until 1923 was a second attempt made. By this time Laysan was a near desert, three endemic birds were near 88 extinction and two had vanished forever (a third became extinct within a month). Elimination of rabbits and artificial planting started the island on the way to recovery and by 1936 conditions were superficially near normal. Breeding bird populations fluctuated with changes in island condi- tions and environment. For example, the Laysan Albatross numbered in the many hundreds of thousands, perhaps a million or more, in 1891. By 1911 the population was reduced to under 200,000 breeding birds with continued drastic decreases to under 50,000 in 1915 and under 30,000 in 1923. Even though the number of non=-breeding birds is unknown, the magnitude of the population decline is nevertheless readily apparent. By 1950 and the return of favorable habitat the population had in- ereased to over 100,000 birds and by the 1960's had reached at least a half million birds. Another dramatic example is the Laysan Teal which survived a population low of one in 1930 but which was recently believed to number at least 600 birds. Birds Fifty-nine species have been recorded from Laysan but over half of these are accidental or irregular visitors. The original 22 breed- ing species (Tables FI-1 and FI-2) represented six orders: Procellarii- formes (7), Pelecaniformes (5), Anseriformes (1), Gruiformes (1), Charadriiformes (5), Passeriformes (3). The remaining transient and accidental species (Tables FI-3 and FI-4) are predominantly shorebirds, gulls, and ducks. The 17 breeding seabirds occur also on most of the other North- western Hawaiian Islands but Laysan has the largest total seabird population of the group and many species have their largest populations there. Among the exceptions are the Bonin Petrel, most abundant on nearby Lisianski, and the Great Frigatebird, most abundant on Nihoa Island. All breeding species exhibit a definite annual cycle which, however, for some species may vary a month or more from year to year. Five species, Black-footed Albatross, Laysan Albatross, Bonin Petrel, Sooty Storm Petrel, and Black Noddy, have their peak breeding season in winter and spring while the other species are spring and summer breeders. Some, notably the White Tern and Blue-faced Booby, have far more extended seasons than others with a few birds breeding at all seasons of the year but with the main population breeding at a peak period as shown in Table Hie The partial temporal replacement of species utilizing similar nest- ing areas has been discussed by numerous writers including Schauinsland (1899) and Richardson (1957). Thus the Bonin Petrel and the two alba- trosses are well into their breeding cycles before the arrival of the Wedge-tailed Shearwater. Competition for the same space is reduced as a result of nest loss and reduced space requirements of the former as the nesting cycles progress. The marked specifie requirements for nesting sites have also been discussed at length, notably by Fisher (1903a). 89 Table FI-l1. Present status of breeding seabirds on Laysan. Species Status and Numbers Primary Breeding Period Black-footed Albatross Laysan Albatross Bonin Petrel Bulwer's Petrel Wedge-tailed Shearwater Christmas Shearwater Sooty Storm Petrel Red-tailed Tropicbird Blue-faced Booby Brown Booby Red-footed Booby Great Frigatebird Sooty Tern Gray-backed Tern Brown Noddy Black Noddy White Tern Abundant (40, 000 - 80, 000) Abundant (500,000 - 600,000) Abundant (200, 000) Common ~ (10,000 - 20,000) Abundant (200,000 = 400, 000) Common (6,000 - 10,000) Common (2, 000 - 3,000) Common (4,000) Common (1,000 - 2,000) Uncommon (100 - 200) Common (2,000 = 3,000) Common (5, 000 - 8,000) Abundant (2, 000, 000) Common (12,000 - 40,000) Common (20,000 - 40,000) Common (5,000) Common (1,500) Mid-November through mid- July Mid-November through early July Mid-January to late June Late May through August June through November Mid-April through late September Late December to June Late April through early October Late March through September Late March through October February through Septem- ber March through October Early April to early September March through early August March through September November through July May through August 90 Table FI-2. Status of endemic Laysan birds. Species Previous Status* Present Status Laysan Teal Uncommon to common Uncommon (500-700) (ee) Laysan Rail Abundant (2,000) Extinct, ca. 1923** Laysan Millerbird Common to abundant Extinct, prior 1923 (under 300) Laysan Honey-eater Uncommon to common Extinct, April 1923 (300) Laysan Finch Abundant (2,700) Abundant (10,000) *First statement gives relative abundance during the late 19th century; numerical estimate in parentheses is from 1911 just before major habitat destruction occurred. a **Birds transplanted to Midway survived until 1944. Table FI-3. Status of regular shorebird species on Laysan.* Months Recorded Species Status i ob na Emon b Golden Plover Abundant X—————-X X X X X——X ? X Ruddy Turnstone Abundant X———__ > xX Ke Bristle-thighed Curlew Common 2 ha aaa Wandering Tattler Common X——_—_———X X X XK ————X ? X Sanderling Uncommon X X. x xX x Dr tie 6 *Solid lines indicate period of greatest abundance. ?No specific records, probably because of no recent visits. 91 Table FI-4+. Months of occurrence of accidental and vagrant species on Laysan. Total © 2 Species Records & = Sooty Shearwater Pelagic Cormorant Emperor Goose Mallard Pintail Common Teal [Garganey Teal ]* American Widgeon Shoveler Bufflehead Harlequin Duck Semipalmated Plover Black-bellied Plover [Whimbrel ] Greater Yellowlegs Lesser Yellowlegs Sharp-tailed Sandpiper Pectoral Sandpiper Baird's Sandpiper Dunlin Unidentified Sandpipers Dowitcher sp. Marbled Godwit Bar-tailed Godwit Red Phalarope Northern Phalarope [Glaucous-winged Gull] Glaucous Gull Herring Gull Unidentified large gulls Bonaparte's Gull Black-legged Kittiwake Blue-gray Noddy Horned Puffin = PMP PWWHE ND FFE PAWEPPWWHEE PEPE DINE ND PwWwHHeE Ps PS > bd bd bs ta ta Pd PS PS a ("winter") X *The occurrence of species listed in brackets has not been adequately confirmed. Thus the Black Noddy, Great Frigatebird, and Red-footed Booby utilize the branches of woody vegetation while the Red-tailed Tropicbird and Christmas Shearwater nest under the deep cover and the Brown Noddy chooses the more open edges. Likewise the Sooty Tern and Laysan Alba- tross nest on the surface of the more open bunch grass association while ge the Bonin Petrel and Wedge-tailed Shearwater burrow in the same area. Black-footed Albatrosses prefer to nest on sandy beaches whereas Laysans prefer open areas inland from the beach, especially the flat area near the lagoon. Numerous other examples are cited by Fisher. The relatively large number of species of endemic land birds formerly present (five) was probably made possible by the large island area, the varied and stable vegetation, and perhaps also the presence of a fresh or mildly brackish pond. The three extinctions were clearly the result of devegetation and with the return of stable vegetation the surviving teal and finch have probably returned to their original numbers (Table FI-2). By far the majority of the transient and accidental species recorded from Laysan are strong-winged shorebirds and ducks that breed in the arctic or subarctic and migrate southward for the winter months. Five of these shorebirds (Table FI-3) are of regular occurrence in consider- able numbers and winter regularly throughout the central Pacific. Others, such as the Pintail and the Shoveler, occur regularly in the Northwestern Hawaiian Islands and probably visit Laysan during most years. Most of the transients, however, are stragglers into the central Pacific and occur sporadically in small numbers during the fall months. Since rela- tively few accidentals are seen during the spring months, most must either move farther south during the winter or succumb to the environment. Observation of living gulls on one trip up the leeward chain in 1963 and the finding of dead gulls on a subsequent trip suggest the latter. The presence of a large central lagoon and extensive mudflats undoubtedly makes Laysan an attractive spot for ducks and shorebirds. Surprisingly few oceanic birds and vagrant land birds have been reported, probably because of inadequate coverage of the island for an extended period of time. Continued occupancy of the island by an alert ornithologist would undoubtedly result in a marked increase in the number of accidentals recorded. Mammals The native mammalian fauna is limited to one species, the Hawaiian Monk Seal, Monachus schauinslandi, which uses the sandy beaches bordering Scaevola for hauling grounds. The monk seal was considered "numerous" by Paty in 1852 but was hunted almost to extinction during the last half of the 19th century. With protection the population has increased greatly and it is again a regular part of the Laysan fauna. Numerous domestic animals were imported to Laysan during the period of human occupancy. Among these were hogs, mules, cows, horses, a donkey (?), guinea pigs, and rabbits. All except the rabbits were either removed when the island was deserted or else failed to survive. The rabbits multiplied tremendously and had nearly destroyed the island vege- tation by 1923 when the last ones were believed killed. 93 Reptiles Two species occur on Laysan--the Green Sea Turtle, Chelonia mydas, and the Snake-eyed Skink, Ablepharus boutoni--and another, the Fox Gecko (Hemidactylus garnoti), formerly occurred there. Both the lizards were probably introduced to the island by man but the turtle is native to Laysan. Turtles still use the island as hauling grounds but probably only a very few breed there. They were abundant on Laysan during the 19th century and were heavily utilized by visiting seamen and fishermen. Numbers were greatly reduced and there has been little increase in the population since. Species Accounts Birds In the following species accounts (birds), the common and scientific names and the sequence of species are from standard references, primarily the AOU Checklist (1957)--and for the Procellariiformes and Laridae, King (1967). A standard format is employed for each species as indicated below: Status: Intended to provide a very brief summary of the occurrence and activity of each species while on Laysan. Included are; A. Relative Abundance: For breeding seabirds the following scale is used: 1) abundant--peak populations in excess of 50,000 indi- viduals; 2) common--peak populations of about 1,000-50,000 individuals; 3) uncommon--populations of less than 500 individuals. These limits were chosen because estimated breeding populations of all species fall easily within one of these categories. A different scale is used for transient shorebirds and endemic land birds because of the much smaller numbers in- volved: 1) abundant--peak populations in excess of 1,000 individuals; 2) common--peak populations of 100-1,000 individuals; 3) uncommon--regular in occurrence but peak populations less than 100 individuals. B. Status: Two categories are used: 1) breeder--a species breeding on the island but most individuals absent during some part of the non-nesting season; varying numbers of non-breeding birds (local, from other islands, or both) may be present during the non-nesting season; 2) migrant--species visiting the island only during the non-reproductive season; may visit the island only briefly in transit elsewhere or remain for a substantial period, usually during the winter months. C. Maximum recent estimate: These are maximum (conservative) estimates during the last decade. All extreme estimates have been re- evaluated and some have been omitted from the text. All such estimates are enclosed in brackets in the appropriate table of observations or wherever mentioned in the text. Whenever available, estimates made by the POBSP are used but some estimates were provided by the BSFW. gh D. Period present: The inclusive period of usual presence on Laysan is indicated, together with duration of stragglers if appro- priate. Period of absence is also provided. E. Nesting period: The period of major breeding activity is given together with tates appropriate) the extent of minor breeding periods. F. Nesting area: A summary statement includes both the usual nesting habitat and the general areas utilized on the island. G. Nest: A summary statement gives the usual nest site and, if pertinent, the type of nest built. Populations: All available data (published and unpublished) are summarized in a chronological table which is placed at the end of the species account. Where available, actual counts or population estimates are presented; when unavailable, more qualitative data are cited. Where sufficient data exist, the variation in population estimates is discussed, particularly where historical changes can be documented or strongly indicated by available data. Factors affecting these changes (e.g. feather raids and habitat destruction) are discussed. Annual Cycle: A generalized annual cycle based chiefly on obser- vational data is presented for each species. Where actual observations are lacking (as in November with no recent trips), interpolated data based on incubation period and fledgling period are included. Variations in the nesting cycle are discussed as are such topics as breeding peaks, and, when available, actual dates of egg-laying, hatching and fledging. Ecology: A. Breeding: Both historical and current data are presented and whenever possible the two are compared. Details concerning pre- ferred nesting areas and nest sites are included. Where possible, known changes in breeding ecology are presented and analyzed together with environmental changes. B. Non-breeding: Utilization of the island by any non-breeding birds is discussed whenever data are available. Specimens: Nearly all known Laysan museum skins were examined but a detailed listing or analysis is impractical because of the large num- bers involved. A tabular summary indicates the current distribution of study skins in a number of major museums. The locations of a few additional specimens (including skeletons, alcoholics, and nests and eggs) are noted briefly in the text. The tabulation of study skins (divided into three categories--adult males; adult females; other, including unsexed, chicks, etc.) is intended to indicate the most profitable sources of specimens for future work. A more detailed list- ing of Laysan specimens, providing additional data, is on file at the U.S. National Museum and can be obtained upon request. 95 Banding and Movements: A brief summary of all POBSP banding is presented in tabular form (often broken down into age and sex classes) by date of banding. When available, banding data from HDFG and BSFW personnel are included, especially if POBSP bands were used. Inter- island movements are presented in summary form and are also listed in Appendix Tables. These include both birds banded on Laysan and recaptured elsewhere and birds banded elsewhere and recaptured on Laysan. Ina few cases birds are known to have made several movements, often involving several different islands. No detailed analysis of interisland movement is attempted. BLACK-FOOTED ALBATROSS Diomedea nigripes Status Abundant breeder; maximum recent estimate of breeding population about 67,000. Present from early November through early August; absent remainder of year. Most nesting is from mid-November through mid-July. Nests in open areas, especially the outer beaches. Populations No population estimates are available for the entire period preceding the feather raid of 1909 to 1910 but numbers (Table BFA-3) apparently never approached those of the Laysan Albatross. Dill and Bryan estimated 85,000 breeding birds in 1911 and commented that this species suffered less than did the Laysan Albatross. Two years later Bailey estimated a population of 15,444 breeding birds (based on nest count) and following the 1915 raid Munter estimated 20,000 birds. Both Bailey's and Wetmore's estimates of the breeding populations (the latter, 18,800) are considerably less than those during the last two decades. The largest recent estimates (Table BFA-3) are those of Rice and Kenyon whose work suggests a breeding population on the order of 65,000 birds. Of the more accurate recent estimates, only one (near 40,000 birds in December 1963) was made at a comparable time of year. All the other recent estimates were made in March or later in the year and probably reflect less than maximum populations due to nest mortality prior to censusing. Allowing for annual variation in the size of the breeding population (which evidently may be considerable), the more accu- rate recent estimates of the number of young (March, 1964, 1969; May, 1958; June, 1966, 1967) rather consistently suggest maximal breeding populations on the order of 25,000 to 30,000, occasionally perhaps as large as 40,000. Thus we feel that the breeding population is somewhat smaller than that indieated by Kenyon and Rice. We rather suspect that the 1957 estimates were somewhat excessive or the result of a particularly large nesting population that season. 96 Annual Cycle Adults arrive in early November and egg laying begins shortly after the birds return to the island. Bailey (1956: 37) found eggs common in late December 1912. He found the first nestling on 21 January 1913, indicating that the first egg was laid about 16 November. This agrees well with Schlemmer's 1915 observations (Table BFA-3). Willett's observations indicate eggs may be laid through mid-January. Young begin fledging in June and all are gone by September. From 5 to 12 August 1965 only 25 emaciated young remained, indicating that the majority of the birds had left the island by the end of July. Peak numbers of adults occur on the island during the winter, and they decrease rapidly after the chicks hatch. By late spring only a few adults are on the island at any one time (more at night), and by early summer they visit the island only infrequently to feed the young. No birds have been found on the island in September or early October. Ecology Breeding: Nesting is typically on open sand beaches and to a lesser extent in large openings among Scaevola and Eragrostis. In 1891 Palmer (Rothschild, 1893-1900: 55) found the main concentration of birds on the south beach. Fisher (1903a: 790) found them on the sandy beaches on the north, east, and south sides, with only a few on the west side and in the interior. Other early observers (Dill and Bryan, 1912: 17; Bailey, 1956: 373; Wetmore, ms.) found the same pattern of distribution. However, Munter (1915: 139) in April 1915 found the young that had survived a raid by the feather poachers principally on the southwestern, southern, and south- eastern beaches. The distribution of birds in recent years has been similar to that of the early part of the 20th century, with the major concentration along the northern beach. Non-breeding: Non-breeding birds roost on the beaches among the nesters. Before they fledge, young birds gather in small groups along the outer edges of the beach, where they fan their wings and practice flying, especially during strong winds. Specimens Fifty-seven Black-footed Albatross skins from Laysan are currently distributed in museums as indicated in Table BFA-l. ‘Twenty-five addi- tional mounted specimens are distributed as follows: AMNH (2 adult males and 1 adult female in Laysan exhibit); BPBM (2 adult males, 2 adult fe- males, 1 immature); CMNH (2 adult males and 2 adult females in Laysan exhibit); SUI (12 in Laysan exhibit); MCZ (1 immature). At least 6 skeletons (BPBM, 1; UCLA, 1; USNM, 4+) and 26 eggs (BPBM, 14; MCZ, 12) are also preserved. zi Table BFA-1. Locations of Black-footed Albatross skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 4 3 2 a BPBM ) O 7 7 CMNH 1 ale O 2 DMNH 0 2 D if MCZ i (0) (0) 1 UMMZ 0) 2 2 4 USNM (non-POBSP) 13 y 6 23 (POBSP) @) ) @) ©) Other* 3 1 O 4 Totals 22 13 22 Si *Dickey Coll. (2 ct"); Law Coll. (10°); U. Minnesota (1 9). Banding and Movements Various agencies have banded e535 Black-footed Albatrosses on Laysan (Table BFA-2). Bands from two Black-footed Albatrosses banded as locals on Whale-Skate, French Frigate Shoals, were subsequently recovered on Laysan. Three birds banded on Laysan were later recovered at sea (Appendix Tables 4-3a and 4-3b). Table BFA-2. Black-footed Albatross banded on Laysan. Period of Survey Bander Adults Young Age Unknown Total 1957 June-July HDFG 0 200 200 1958 June BSFW - - goo* 900 1965 July POBSP O 685 0 685 1967 June POBSP O 600 0 600 Totals 0 1,485 900* 2385 *These birds were probably all or mostly young. 98 Table BFA-3. Observations of Black-footed Albatross on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1828 24 Mar. common Young (Isenbeck, in Rothschild, 1893- 1900: iii). 1891 16-27 June fairly Young (Rothschild, 1893-1900: 55). numerous 1896 24 June- ? Breeding (Schauinsland, 1899: 101). 2h Sept. 1902 16-23 May ? Less abundant than Laysan Albatross; young (Fisher, 1903a: 790). 20 Nov. 2 14 eggs collected by Schlemmer (MCZ). 1903 18-24 Apr. ra At least 9 skins and 1 skeleton col- lected by W.A. Bryan (AMNH, BPBM). 1905 20-21 Mar. & 2, including 1 juvenile, collected (MCZ). 1906 18-20 Nov. 2 12 fresh eggs collected (MCZ). 1911 2h Apr.- 85 , O00* Estimate based on nest density* and 5 June colony area computations (Dill and Bryan, 1912: 17). 1912 12 Dec.- 28, 000 Count in Feb.: 7,506 occupied and 1913 11 Mar. (15, 444) 216 abandoned nests. First eggs hatched 21 January. First estimate includes 5,000 young and 7,500 un- employed birds (Bailey, 1952a: 39, 55). All eggs laid by 15 January (Willett, ms.). TOTS, ae Sayre 20, 000 Young (Munter, 1915: 139). 14 July no more than (Schlemmer and Schlemmer, ms.). 100 seen 26 Oct. i First bird arrives (Schlemmer and Schlemmer, ms.). 28 Oct. 4 (Schlemmer and Schlemmer, ms.). 2 Nov. Tak Count (Schlemmer and Schlemmer, ms.). 99 Table BFA-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1915 6 Nov. 500 Estimate (Schlemmer and Schlemmer, msiee) ie 8 Nov. ? Arriving in large numbers (Schlemmer and Schlemmer, ms.). 13 Nov. ? lst egg found (Schlemmer and Schlem- mer, ms.). 24-25 Nov. 2 350 eggs pickled (Schlemmer and Schlemmer, ms.). 1916 Q Feb. ? Nesting; more numerous near shores of island (Munter, ms.). 1918 8-10 Sept. O @ueessamse): 1923 8-13 Apr. 2 Young (Wetmore, ms.). 29 Apr.- 18,800" 4,700 young, based on count (Wetmore, 14 May msi.)). 1930 2-18 Aug. O (Wilder, ms.b). 1936 7-18 Mar. countless Young; 2 banded with brass rings thousands (Trempe, ms.). 12 Dec. 2 "Some on North and East beaches” (Coultas, ms.). 1950 23 June numerous Fully-feathered young (POFI). 1951 12 May 2 Young (POFI). late June- 36, 480% Estimate (from transect censuses) of early July ca. 18,240; most, if not all, young (Brock, 1951b: 18). 1955 10 Feb. ? Ca. half as numerous as Laysan Alba- tross. More with young than eggs (POF). 1957 7 Jan. 64, OOO Estimate of 32,128 nests from aerial photos (Rice and Kenyon, 1962: 375). 25 June- 2 Estimate of ca. 9,000 young, based on 3 July transect census (Woodside, ms. b). 100 Table BFA-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1957 8-12 July few (Labrecque, 1957: 17). 28 Dec. 67, 000* Estimate of 33,523 nests from aerial photos (Rice and Kenyon, 1962: 375). 1958 2] May- ? "Few adults present at any one time;" 4 June estimate of 8,700 young, 1/2 to 2/3 grown (Warner, ms.). 1959 28 Apr.- ? Nesting; young, most on NW part of 1 May island (Kramer, ms.). 1961 7-8 Mar. ” Young (Woodside and Kramer, ms.). | 4-10 Sept. 0 None present (Woodside, ms.c). 1962 14-19 June 2 Downy and flying young (Kramer and Beardsley, ms.). 1963 11-13 Feb. abundant Young (POBSP). 3-10 Dec. 38,666 Evidently a count; eggs (Walker, ms.b). 1964 10-11 Mar. 37, 000= Ca. 10,700 young on outer beach; less 42,000 than 1,000 elsewhere (BSFW, POBSP). 16-20 Sept. 0 (BSFW, POBSP). 1965 6-11 Mar. 30, 000*- Ca. 10,000 to 15,000 young, 1/4 to 45), OOO 1/3 grown (POBSP). 17-21 July 2 Ca. 4,500 large young (POBSP). 5-12 Aug. ? Ca. 25 emaciated young. No adults seen (POBSP). 1966 26-31 Mar. ? Estimate of 10,000 young, 3/4 to nearly full grown (BSFW). 10-16, 1,500° Estimate of 10,000 young, based on par- 20-21 June tial count of 7,652 young 3/4 to nearly full grown (POBSP). 17-18 Sept. O (BSFW) . 20-23 Oct. ©) (POBSP) . 1O1 Table BFA-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References . 1967 18-19 Mar. ? Estimate of 10,000-20,000 young (BSFW, POBSP). 7-12 June ? Estimate of 10,000 young based on partial count of 8,732 young; some losing down (POBSP). 5-11 Sept. ) (POBSP) . 13 Dec. thousands Nesting (BSFW). 1968 17-19 Mar. 10, OOO= An estimated 5,000-10,000 young 20, 000% present; many unemployed birds (BSFW, POBSP) . 1969 26-29 Mar. ? Estimate of 14,694 young based on 159 transect censuses (BSFW). 9 Sept. 0) (BSFW). *Estimate is of the number of breeding birds. +Estimate is of breeding population prior to estimated mortality observed by Wetmore. Rice and Kenyon (1962: 375) give the post-mortality figure. #The figure given by Rice and Kenyon for this visit (1962: 375) is evidently the result of a misreading of Brock's paper. LAYSAN ALBATROSS Diomedea immutabilis Status Abundant breeder; maximum recent estimate 500,000 to 600,000. Present from late October through August or early September; absent during remainder of year. Most nesting is from mid-November through early July. Nests over most of the island with the major concentration around the lagoon. Populations It is evident from the photographs (Rothschild, 1893-1900: plate 38) and general comments of the earliest visitors to Laysan that the Laysan 102 Albatross was abundant on Laysan in the 1890's and early 1900's. For example, in 1891 Palmer (Rothschild, 1893-1900: 57) reported that the species "literally covers the island...the young in some places being as thick as they could stand." During this period (no specific year given) Max Schlemmer, the island manager, estimated that two million birds were present. No subsequent estimates are this high. Although some albatrosses were killed and parts of the colony dis-= rupted by guano operations on Laysan, it is probable that the presence of guano workers prevented the massive kills that occurred on other islands during this period. The spectacular decline in population oc- curred later and was prohably the result of feather raids in 1909 to 1910 and 1915. Dill's estimate of 180,000 birds in late April 1911, one year after the first raid, was based on a nest count during mid-season but neverthe- less represented a major and sudden deciine from the great numbers present at the turn of the century. Numbers were still very low during the 1912-1913 season and in February Willett and Bailey estimated only about 24,600 breeding birds with considerable nest loss. Following the 1915 feather raid, Munter reported the Laysan Albatross to be the "chief sufferer" among the 150,000 to 200,000 dead birds found on the island. He estimated that only 40- 50,000 live birds (with very few young) remained on the island. When visited in 1923, Laysan was a near desert. Wetmore (ms.) calculated an adult breeding population of 13,600 based on a chick count and a 50 percent nest loss. Unless a large number of unsuccessful breeders had departed the island prior to his arrival, this represents the lowest level ever reported for Laysan. By 1936 the island and ap- parently the albatross population were recovering nicely and today the population is considerably larger than in 1911. It is difficult to determine accurately the population size today for we lack estimates of the number of nests at the peak of the nesting season, the amount of yearly nest loss, and the number of non-nesting birds pres- ent. The data suggest, however, maximal breeding populations on the order of 300,000 to possibly as many as 500,000 birds (Table LA-3). Since some available estimates include all birds, including non- nesters using the island, and others include only the number of birds present at one time, it is of little value to compare these figures (e.g., 5 to 12 August 1965 and 7 to 12 June 1967). ‘The only comparable figures are nest counts and these show considerable yearly variation. In March 1965, 15,000 nestlings were banded and an estimated 20,000 to 25,000 were present, while in June 1966 an estimated 150,000 were present (91,403 actually counted). This most likely represents a difference in nest loss rather than a change in the population and demonstrates the difficulty in determining long range population trends with the available data. 103 Annual Cycle According to Fisher (1903a: 789), who probably received his information from Schlemmer, Laysan Albatross begin to arrive about 25 to 26 October and remain until the following August. Only one recent record, from September 1961, falls outside these limits. Evidently most of the population is present by December. Egg laying begins by at least 20 November (Bailey, 1956: 44) and probably continues no later than late December. The earliest hatching date is 23 January 1913 (Bailey, 1956: 44). Schlemmer (in Fisher, 1903a: 789) reported that the eggs hatched in February, suggesting a peak hatching period in this month, and a peak egg laying period in December. Young birds begin to leave the island by late June and most have left by the end of July. However, a few weak young, which probably soon died, were found 4 to 10 September 1961. Largest populations are present during the winter. After the chicks hatch, one adult spends most of its time at sea, and as the chicks mature both adults spend an increasing amount of time away from the island. By May and June relatively few adults are present during the day, though larger numbers may come in at night to feed the chicks. Visits become less and less frequent and few adults are present at any one time in July and August. By September only a few late-maturing or deserted chicks remain. Ecology Breeding: lLaysan Albatross nest over the island in various habitats, but only a few use dense vegetation or open beaches. Fisher (1903a: 786) found them distributed over the whole island, with the single exception of the beaches. "The flat plain surrounding the lagoon is their favorite habitat, and we found them here in the greatest numbers. This great colony extended all the way around the lagoon, but certain portions were more congested than others. The largest single colony of young is on the south side of the lagoon, where the ground has been leveled off in past years by the phosphate-rock diggers." Dill and Bryan (1912: 15) found them only "along the shores of the lagoon and on a small area at the south end of the lagoon." Munter (1915: 139) found the few young that remained after the raid of feather poachers "in the central parts of the island around the lagoon." Recent observers found the main concentration of nestlings around the lagoon but birds were also found on the east, north, and south beaches, several open areas on the southwest section, and along the beach perimeter. "The nest is made by the female by merely scraping together the earth or mud wherever she is resting, and building it into an elevated 104 ring within which her single egg is deposited." (Schlemmer, in Nutting, 1903: 325-326). This embankment proves of great service when the lagoon floods. In December 1912, Bailey (1956: 44) observed the loss of several thousand nests by flooding, and thought that many moré would have been lost, except for the dikes erected by the incubating birds. Flooding of nests around the lagoon apparently occurs during most winters. Young birds, as they mature, congregate in open areas and along the beaches during the day, seeking shade during the warmest periods and often retiring to denser vegetation at night. As they mature, an increasing amount of time is spent exercising on the beaches and by early August the older and stronger birds have left the island. Perhaps up to 3 percent of those which survive to mid-summer starve to death on the island, ap- parently as the result of desertion or insufficient feeding by the adults. Non-breeding: Data concerning the activities of unemployed birds on Laysan are lacking. Specimens One hundred six Laysan Albatross skins from Laysan are currently distributed in museums as indicated in Table LA-l. Thirty additional mounted specimens are distributed as follows: AMNH (3 females in Laysan exhibit); BPBM (2 adult males, 2 adult females, 1 chick); CMNH (2 adult males, 2 adult females in Laysan exhibit); MCZ (1 adult male, 1 adult female, 1 nestling); SUI (15 birds in Laysan exhibit). Also preserved are at least 8 skeletons (BPBM, 1; USNM, 6; Carnegie Mus., 1), a head (DMNH) and 22 eggs (BPBM, 10; MCZ, 12). Table LA-1. Locations of Laysan Albatross skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 6 6 abil 23 BPBM 0 0 10 10 CMNH 1 z 2 4 DMNH 3 4 @) 7 SUL 3 g {i 1g) UMMZ 2 2 3 T USNM (non-POBSP) 12 1 5 32 (POBSP)* O 0 0 0 Other* 3 () ik h Totals 30 37 39 10 *one hybrid. *Brit. Mus. (Nat. Hist.) (1); Law Coll. (10); Leningrad (10); Acad. Nat. Sci. Phila. (1c). 105 Hybrid (Laysan X Black-footed) Albatrosses have been reported from Laysan on several occasions and three specimens are preserved. Data for the three skins (all males) are as follows: BPBM, taken by Bompke, May 1905; CMNH 156076, taken by Willett 22 February 1913; USNM 494113, taken by the POBSP on 7 March 1965. Another believed to be a hybrid was seen in March 1968 by Karl W. Kenyon (pers. comm.). Banding and Movements Various agencies have banded 22,237 Laysan Albatross on Laysan Island (Table LA-2). Four Laysan Albatross banded elsewhere (Kure, 1; Lisianski, 1; Pearl and Hermes Reef, 2) have been recorded from Laysan; and 16 birds banded as nestlings or locals on Laysan have been recovered elsewhere (2 at Pearl and Hermes Reef; 1 at Midway; 2 at Kure; 1 at Japan and 10 at sea) (Appendix Tables 4-4a and 4-4b). A 17th bird originally banded on Kure was recaptured on Laysan and then later recaptured on Kure. At least 142 of the 15,000 chicks banded in March 1965 died before fledg- ing, and their bands were recovered from the carcasses. Table LA-2. lLaysan Albatross banded on Laysan, Period of Survey Bander Adults Young Age Unknown Total 1957 June-July HDFG O 200 0 200 1958 June BSFW ©) 0 2, 000* 2,000 1964 Mar. BSFW 0 200 0 200 1965 Mar. POBSP 0 15, 000 0 15,000 July POBSP O 1,807 O 1,807 Aug. POBSP O 151 0 151 1966 June POBSP O 1,979 @) 1,979 1967 June POBSP ) 900 ) 900 Totals O 20, 237 2,000 22, 237 *These birds were probably all or mostly young. Table LA-3. Observations of Laysan Albatross on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1828 2 Mar. ? (Isenbeck, in Rothschild, 1893-1900: ris EETS) 9 1890 16 July ? Young (Lyons, 1890: 90). 1890's 2, 000, 000 (Schlemmer, in Nutting, 1903: 322). 106 Table LA-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1891 16-27 June ? Nearly fledged young (Munro, 1942a: 72). 1896 24 June- 2 (Schauinsland, 1899: 101). 2k Sept. 1902 16-23 May "thousands" Young 2/3 grown (Fisher, 1903a; 786; 1940a: 9). Nov. 2 8 eggs collected by Schlemmer (BPBM). 1903 Apr. % 11 skins and 1 skeleton collected by W.A. Bryan (BPBM). 1904 11 May 2 1 collected by Schlemmer (MCZ). 15-16 Nov. ? 12 fresh eggs collected by Schlemmer (MCZ). 1905 18 Mar.- b 3 collected, including a nestling, by Apr. Schlemmer (MCZ). 1906 May ? 1 collected by Schlemmer (AMNH). 1911 24 Apr.- 180, 000* Ca. 90,000 young based on count of nests 5 June (Dill and Bryan, 1912: 16; Dill, 1916b: 173). 1912 22 Dec.- 34, 000 Count in Feb. of 9,201 occupied and 3,120 1913 11 Mar. (25,000)* abandoned nests. First estimate includes 4,600 non-nesting birds. First egg hatched 23 January (Bailey, 1952a: 55; 1956: 44). OMS 3 Mjoie, 40, 000- Few young (Munter, 1915: 139). 50, 000 22 Aug. @ 1 seen (Schlemmer and Schlemmer, ms.). 28 Oct. ? First two arrived (Schlemmer and Schlemmer, ms.). 29 Oct. 2 8 more seen (Schlemmer and Schlemmer, ms. )e 2 Nov. 10 Counted (Schlemmer and Schlemmer, ms.). 107 Table IA-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1915 8 Nov. 300 Estimate (Schlemmer and Schlemmer, ms.). 12 Nov. 600 More arrived than total for past two weeks (Schlemmer and Schlemmer, ms.). 1916 9 Feb. 2 Nesting (Munter, ms.). 1918 8-10 Sept. 2 Few seen (Diggs, ms.). 1923 8-13 Apr. 2 Young (Wetmore, ms.). 29 Apr.- 14 May 13,600+ 3,400 young (Wetmore, ms.). 1930 2-18 Aug. ? A few unfledged young still present (Wilder, ms.b). 1936 7-8 Mar. countless Ten banded with brass rings (Trempe, thousands ms.) 12 Dec. "abundant". (Coultas, ms.). 1950 23 June abundant Well-feathered young (POFT). 1951 12 May ? Young (POFI). late June- 207 , 800# Count of ca. 103,900 (most, if not all). early July young (Brock, 1951b: 18). 1955 10 Feb. ? Ca. twice as numerous as Black-footed Albatross. Ca. 80% of nests contained young, ca. 10% eggs (POFI). 1957 7 Jan. 263 , O0O* An estimate of 131,328 nests made from and/or aerial photos (Rice and Kenyon, 1962: 28 Dec.** 3) 25 June- z Estimate of 45,000 young based on trans- 3 July ect census; estimate of 215,000 young based on banding of chicks and density/ area relationship (Woodside, ms.b). 8-12 July "most numer- Half-grown to almost mature young ous...bird on (Labrecque, 1957: 17). the island." 108 Table LA-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1958 27 May- ? Ca. 67, 000 1/2 grown young based on 12 June transect censuses. A considerable num- ber of unemployed birds still in area (Warner, ms.). 1959 28 Apr.- y Estimate of ca. 69,000 young (based on 1 May 2 of same transect lines [Kramer, ms.] as used in 1958). 20-27 July 2 "Downy, but full grown chicks" noted (Udvardy, 1963: 191). 1961 7-8 Mar. 2 Young (Woodside and Kramer, ms.). 4-10 Sept. 2 10-20 young, most weak and emaciated, one adult seen 4 Sept. (Woodside, ms.c). 1962 14-19 June ? Young. Many apparently almost ready to take flight (Kramer and Beardsley, ms.). 1963 1113 Feb. thousands Mostly with small downy young; few eggs (POBSP) . 3-10 Dec. 497, 948 Evidently a count; most birds on eggs (Walker, ms.b). 1964 10-11 Mar. 500, 000- An estimated 150,000 young (BSFW, POBSP). 600, 000 16-20 Sept. 0 (BSFW, POBSP). 1965 6-11 Mar. 75, 000- An estimate of 20,000 to 25,000 1/4 to 100, 000 1/2 grown young based on 15,000 young banded (POBSP). 17-21 July ? An estimated 10,000 downy to almost fledged young (POBSP). 5-12 Aug. 5 Less than 1,000 young (POBSP). 1966 26-31 Mar. 300, 000 Estimate includes 150,000 non-nesting adults, ca. 75,000 young present (BSFW). 10-16, 2 An estimated 150,000 young from 3/4 20-21 June grown downy chicks to almost fully fledged birds (based on a partial count of 91,403 young and 4,246 adults). An estimated 5,000 to 6,000 adults on the island at any one time (POBSP). 109 Table LA-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1966 17-18 Sept. O (BSFW). 20-23 Oct. ) (POBSP). 1967 18-19 Mar. thousands Ca. 30,000-60,000 young (BSFW, POBSP). 7-12 June 200, 000 Ca. 25,000 young (based on count of ca. 16,500). Estimate includes all adults using the island (POBSP). 5-11 Sept. O (POBSP). 13 Dec. ? Nesting (BSFW). 1968 17-19 Mar. 120, 000- Ca. 60,000-100,000 young present (BSFW, 200, 000% POBSP). 1969 26-29 Mar. ? Estimate of ca. 77,300 young based on 159 transect censuses (BSFW). 9 Sept. O (BSFW). *Estimate is of the breeding population. **It is not clear which date or dates this estimate is for. +Estimate of breeding population prior to estimated 50% loss of young. Rice and Kenyon (1962: 375) calculated 6,800 breeding pairs based on Wetmore's post-mortality estimate. #The figure given by Rice and Kenyon for this visit (1962: 375) is evi- dently the result of a misreading of Brock's paper. BONIN PETREL Pterodroma hypoleuca Status Abundant breeder; maximum recent estimate: "several hundred thousand." Present from mid-August to late June or July; absent remainder of year. Most nesting is from mid-January to late June. Nests in burrows chiefly under the Eragrostis association between the lagoon and the beach crest. 110 Populations This is one of the most difficult species to census accurately because of the large numbers involved and its burrowing and nocturnal habits. Dill and Bryan and Willett estimated populations of 160,000 and 100,000 in 1911 and 1912 to 1913 respectively (Table BP-2). They did not detail the basis for these estimates but Willett's discussion indicates that his was little more than a guess. Willett (1919: 61) described considerable nest loss when burrows were filled with blowing sand in February 1913 but no quantitative data were given. It is likely, however, that the species sustained heavy losses throughout the period when too little vegetation was present to hold the sand during high winds. No nesting occurred during Wetmore's visit (April 1923) and only a few were seen. The larger recent estimates, especially those for September 1966 and 1967, indicate that the population has regained its earlier size (Table BP-2). Annual Cycle Bonin Petrels appear to have an unusually synchronous and regular breeding cycle. Schauinsland (1899: 50) described in detail the arrival of the first birds on 17 August 1896. His statement that these birds arrived “over a period of several years from 15 to 18 August every year without fail," probably was based on information obtained from island residents. The single recent August visit found a few birds on the 5th, with a marked increase by the 12th. Throughout the fall the birds apparently return to the island each night, dig burrows, and court. Although Kridler noted copulation in September (BSFW, 1966), eggs are not laid until January. Max Schlemmer told Fisher (1903a: 793) that "eggs are laid about the first of January, but the birds arrive in vast numbers months before." Willett (1919: 60) said "laying commenced the first week in January and was at its height about 20 January." A marked change in behavior accompanies the beginning of incubation. Willett (1919: 60) noted that "the air at night fairly swarmed with the birds" from the time of his arrival, 22 December, until 7 January, after which the birds were still abundant but the numbers in the air decreased considerably. These high numbers immediately before egg laying may in- dicate that the birds do not leave the igland for an intensive feeding period just before laying. Hatching probably occurs as early as late February, with a peak from early March through perhaps early April. On most March visits observers reported eggs and/or small young; no eggs have been found later than March but the presence of unfledged young in July 1957 implies that they may be present through late April. Some young may fledge in early May but 111 most fledge from mid-May to mid-June. All are gone from the island by late June or early July. During June they spend considerable time out- side their burrows at night. Personnel visiting Laysan in June 1966 and 1967 found noticeable mortality of young Bonin Petrels, but no quantita- tive data were obtained. Few adults are present in June, and probably are usually absent in July. As long as adults are on the island, some may be seen courting and digging, but whether these are young non-breeding birds, birds that lost eggs or young earlier, or normal breeding adults is unknown. Ecology Breeding: Fisher's (1903a: 793) description of the nesting area of Bonin Petrels on Laysan is equally appropriate today: "The long burrows in which the birds nest honeycomb the sandy soil over all the region cov- ered by coarse bushy grass (Eragrostis), or from the edge of the plain surrounding the central lagoon to the divide overlooking the sea." The only additional information from recent visits is that on the inner or lagoon side of the nesting area, birds nest in cover composed of both Ipomoea and Eragrostis. Greatest numbers occur on the west side of the lagoon where the Eragrostis belt is widest. The breeding habitat of this species widely overlaps that of the Wedge-tailed Shearwater, the other major burrower on the island. Although the breeding cycles of the two species are staggered so that the shear- water eggs are not laid until petrel chicks are well grown, there are periods in spring and fall when adults of both species are digging and courting at the same time. Observations of relationships between these two species (e.g., competition for sites, use of each other's burrows ) are few. Willett (1919: 61) thought that the shearwaters used petrel burrows later in the season, but he did not document his information. Fisher (1903a: 793) wrote "the burrows are quite long, 6 feet at least, and usually turn either to the right or to the left after the first few feet." Willett (1919: 61) found them to be 6 to 10 inches in diameter at the mouth and 7 or 8 feet in length, with the nest cavity at the end fairly well lined with grass and leaves. He also found one bird nesting under an overturned basket. In March 1964 Walker found one on an egg under a pile of boxes. Nests examined in March 1968 were invariably well lined with grass and sedge. Non-breeding: As far as is known, non-breeding birds in the popu- lation use the same habitat as the breeders. Specimens One hundred fifteen Bonin Petrel skins from Laysan are currently distributed in museums as indicated in Table BP-1. Six additional mounted specimens are distributed as follows: AMNH (1 adult in Laysan exhibit); BPBM (1 male); CMNH (1 adult); SUI (3 in Laysan exhibit). Also preserved are at least 2 skeletons (BPBM, USNM) and 4 skulls (USNM). 112 Table BP-1. Locations of Bonin Petrel skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 10 9 1 20 BPBM 9 3 6 18 CMNH y 1 ) 5 MCZ 5 2 iL 8 SUI O O AL 1 UMMZ in 4 0 8 USNM (non-POBSP) 16 ani, 3 36 (POBSP) 4 10 2 16 Other* 2 1 O 3 Totals Bn 47 1 1G) *Brit. Mus. {Nat. Hist.) (1 2); Law Coll. (10°); Moseley (1c). Banding and Movements The POBSP banded 3,895 Bonin Petrels on Laysan: 399 adults in February 1963; 500 adults in March 1964 and 352 adults in September 1964; 2,544 adults in March 1965 and 100 young in June 1966. One adult, 723-60810, banded at Southeast Island, Pearl and Hermes, on 1 March 1963, was recaptured on Laysan on 19 September 1964. None banded on Laysan has been recovered elsewhere. Table BP-2. Observations of Bonin Petrels on Laysan. Population 3 Date of Survey Estimate Breeding Status, Remarks, References 1891 16-27 June z 4 molting birds found in burrows. Freeth, the island manager, said the breeding season was over and that they occurred in large numbers during their breeding time (Rothschild, 1893-1900: 49). Nearly all, both old and young, had left the island (Munro, 1941c: 17). 1896 24 June- ? First returned 17 August; 1,000's ok Sept. present by 19 August (Schauinsland, 1899: 50). vals} Table BP-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1902 16-23 May great Most abundant procellariid on the numbers island. Young partially molted into juvenal plumage (Fisher, 1903a: 793). 1903 Apr. ? 10 skins and 1 skeleton collected by Bryan (AMNH, BPBM). 1907 20-21 May ? 8 collected by Schlemmer (MCZ). 1911 24 Apr.- 160, 000 Young nearly fledged (Dill and Bryan, 5 June 1912: 18). 1912 22 Dec.- 100, 000 Laying began during first week of Janu- 1913 11 Mar. ary and reached its peak about 20 January (Willett, 1919: 60); egg noted 9 January (Willett, ms.); 1 laid as late as 25 January (Bailey, 1956: 58). 1918 8-10 Sept. 250, 000 Estimate considered conservative; 2nd most abundant species (Diggs, ms.). 1923 +=8-13 Apr. 2 Seen occasionally in evening (Wetmore, ms.). 1936 12 Dec. ? 3 seen; others in holes (Coultas, ms.). 1957 2> June- 2 Post-nesting; 1 young (Woodside, ms.b). 3 July 8-12 July ? (Labrecque, 1957: 18). 1958 27 May- ? Young present but not in large numbers. 4 June All examined had some down, but flight feathers were in final stages of dev- elopment (Warner, ms.). 1961 7-8 Mar. very Digging burrows and mating; eggs noted abundant (Woodside and Kramer, ms.). 4-10 Sept. ? Great numbers at night; none during day; much digging (Woodside, msc ). Appeared every night in increasing numbers; not nesting; inspecting bur- rows, pairing, or, in a few cases, excavating burrows or cleaning old ones (Udvardy, 1963: 193). 114 Table BP-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1962 14-19 June 2 Only 1 pair seen; no eggs or young found (Kramer and Beardsley, ms.). 1963 11-13 Feb. tens of Several eggs were moderately incu- thousands bated (POBSP). 3-10 Dec. B 1 burrow dug up contained 2 mature birds not breeding (Walker, ms.b). 1964 10-11 Mar. 5, 000 Ca. 2,000 nests; 1 bird on egg (BSFW, POBSP). 16-20 Sept. 2,500 Many paired and excavating burrows (BSFW, POBSP). 1965 6-11 Mar. 30, 000- Burrow digging; eggs, and small 50, 000 young (POBSP). 17-21 July 0 (POBSP) . 5-12 Aug. 2, 000 Only a few on 5 August, but many more by 12 August; burrow digging noted on 11 August (POBSP). 1966 26-31 Mar. thousands Many birds digging burrows (BSFW). 10-16, 500% Estimated 3,000 young; most molted 20-21 June into juvenal plumage; ca. 30-40% still with tufts of down (POBSP). 17-18 Sept. several Many fresh burrows; copulating hundred (BSFW). thousand 20-23 Oct. 20, 000 Courtship and burrow digging; no eggs or young found (POBSP). 1967 18-19 Mar. 10, 000 Courtship and burrow digging; 1 heav- ily incubated egg found (BSFW, POBSP). 7-12 June 10, 000 Young present; most near fledging (POBSP) . 5-11 Sept. 75,000 Most birds digging burrows or sitting on surface; no eggs found in burrows examined; present only at night (POBSP). i Table BP-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1967 13 Dec. 2 Only 1 moribund adult seen during di- urnal visit (BSFW). 1968 17-19 Mar. 10, 000 Conservative estimate; 4 burrows con- tained small downy young (BSFW, POBSP). 1969 26-29 Mar. thousands Large downy young (BSFW). *Estimate is of the number of adults present. BULWER'S PETREL Bulweria bulwerii Status Common breeder; maximum recent estimate: 20,000. Present from late March or April through September or perhaps early October; absent remainder of year. Most nesting is from late May through August. Nests most commonly in shallow burrows or in crevices beneath rocks on the south and southwest beaches and on the ground beneath Scaevola on the island rin. Populations The two earliest numerical estimates (1,000 in April-June 1911; 750 in May 1923, see Table BuP-3) are considerably smaller than the largest recent estimates (3,000 in June-July 1957; 10,000 and 20,000 in June 1966 and 1967), but the variability of recent estimates and the limited number of early estimates make it clear that we cannot definitely establish any change in population levels. The extreme variation in recent estimates demonstrates the difficulty of obtaining good numerical estimates of this petrel. Some of this vari- ability is probably due to differences in the birds' behavior during different periods of the breeding cycle. The three largest estimates were made early in the breeding season when courting and burrowing birds are most conspicuous; all smaller estimates were made later in the season when burrows are easily overlooked and when adults become much more secretive and difficult to observe. Annual Cycle Bulwer's Petrels begin to arrive at Laysan in late March or April. A few eggs may be laid in the last week of April (1903, 1923) but the 116 data suggest that most egg laying occurs during early June. A few eggs may be laid as late as the third week of June (1959, 1961, 1967). Young may hatch as early as early June but most hatch in mid-July. Fledging may begin as early as mid-August but the peak fledging period is probably from mid- to late September. Late fall observations are insufficient to determine exactly when the last birds depart but a few birds may fledge as late as early October. Ecology Breeding: bulwer's Petrels have nested in at least four distinct hatitats on Laysan: (1) under rocks, especially on the south and south- west sides of the island, and in favorable areas throughout the island such as the rock piles south of the lagoon (1891, 1911, 1961, and most recent visits); (2) under Scaevola in small depressions on top of fallen leaves (September 1964, August 1965, June 1966 and 1967); (3) in burrows under Eragrostis, particularly when it is covered with Ipomoea, as in a belt on the west shore of the lagoon (August 1965, June 1966 and 1967); (4) under old and partly buried sheets of roofing iron (July 1957, May and June 1958). Birds are most conspicuous among the rocks where they were reported both early in the century and in nearly all recent reports. In this situation they may scoop out depressions less than a foot deep or crawl into cavities more than four feet deep under large boulders. POBSP notes and reports indicate that birds nesting under Scaevola may be as numerous as those nesting elsewhere. Eragrostis was used to a much lesser extent but birds were apparently common there during peak breeding periods. Nesting under vegetation was not noted previously but whether birds were overlooked or whether this is a recent development is not known. If the latter is true, this could account for apparent larger populations at the present time. Use of artificial sites (roofing, boards, etc.) is now less frequent, probably because most of these have been covered with sand or have rotted or rusted away. Non-breeding: In June 1967 two groups of at least 1,000 birds each were sitting around on the southwest and south portions of the island at night. Such clubs have not been previously observed for this species. Like other petrels, they are rarely observed in the open during the daytime. Specimens Sixty-eight Bulwer's Petrel skins from Laysan are currently distri- buted in museums as indicated in Table BuP-l. Three additional mounted specimens are in the Laysan exhibit at SUI. Also preserved are at least 3 skeletons (BPBM, 1; USNM, 2); 2 alcoholics (USNM) and 1 egg (BPBM). Banding and Movements The POBSP and BSFW banded 478 Bulwer's Petrels on Laysan (Table BuP-2). No interisland movements were recorded. 117 Table BuP-1. Locations of Bulwer's Petrel skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 8 y ) 12 BPBM 2 1 @) 3 CMNH 0 iL 0 i DMNH 3 3 0 6 MCZ ye 5 1 10 SUL 3 2 © 5 USNM (non-POBSP) iy 9 0 26 (POBSP) 2 al 0 3 Other* 1 T 0 2 Totals Nea ne Hemmer aOR, og *Basel Museum (1), Bonn (1). Table BuP-2. Bulwer's Petrels banded on Laysan. Period of Survey Bander Adults Young Age Unknown Total 1958 June BSFW 6) 0 138 138 1964 Sept. POBSP 2 15 0 17 1967 June POBSP 200 0 @) 200 Sept. POBSP 14 68 41 123 Totals el 3 7g Tf Table BuP-3. Observations of Bulwer's Petrels on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1891 16-27 June ? Breeding season appeared to be over; not many were seen ashore during day, while many came at night (Rothschild, 1893-1900: 51). Munro (1941b: 2), however, stated that he found them in- cubating eggs. 1896 24 June- ? Not mentioned by Schauinsland (1899) 2h Sept. who nonetheless collected 10 specimens (AMNH) . 1902 16-23 May fe) (Fisher, 1903a: 794). aLAKS) Table BuP-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1903 28 Apr. ? 3 specimens and 1 egg collected by W.A. Bryan (BPBM and AMNH). 1904 18-20 May ? 2 skins collected by M. Schlemmer (MCZ). 1907 18-20 May ? 10 skins collected by M. Schlemmer (MCZ). 1911 2) Apr.- 1, 000 First fresh egg, 2 June; other nests 5 June with fresh eggs found subsequently (Dill and Bryan, 1912: 18). 1912 22 Dec.- ©) (Bailey, 1956). 1913 11 Mar. 1923 8-13 Apr. 0 (Wetmore, ms.). 17 Apr. ? 2 pairs found under buildings (Ball, ms.); 5 found in crevices in guano (Dickey, ms.). 26 Apr. % Egg laid (Ball, ms.). 29 Apr.- 750 Common and in pairs by 29 April, becom- 14 May ing more numerous thereafter (Wetmore, ms.). 1957 25 June- 3, 000 Adults (Woodside, ms.b). 3 July 8-12 July a Quite a few in burrows (Labrecque, 1957: I) 1958 27 May- locally Egg-laying had just begun. All eggs 4 June common checked were fresh or nearly fresh (Warner, ms.). 1959 20-27 July ? Adults incubating (Udvardy, 1963: 193). 1961 7-8 Mar. 0 (Woodside and Kramer, ms.). 4-10 Sept. few Full-grown young observed (Woodside, ms.c). "Almost or quite full-sized chicks," with complete plumage but some down on neck (Udvardy, 1963: 193). 1962 14-19 June ? 1 pair; not nesting* (Kramer and Beardsley, ms.). 119 Table BuP-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1963 11-13 Feb. 0 (POBSP) . 1964 10-11 Mar. 0 (BSFW, POBSP). 16-20 Sept. ? 9 downy young and ca. 10 adults; estimated 50 half-grown to nearly fledged young (BSFW, POBSP). 1965 6-11 Mar. 0 (POBSP) . 17-21 July 1,800 Ca. 200 young present; 2 newly hatched young seen (POBSP). 5=12 Aug. 2, 000 Estimated 500 small young present (POBSP). 1966 10-16, 10, 000 Burrow excavation to incubated eggs; 20-21 June estimated 3,000 burrows with eggs (POBSP). 20-23 Oct. 0 (POBSP) . 1967 18-19 Mar. 20 Heard calling (BSFW, POBSP). 7-12 June 20, 000 Fresh to moderately incubated eggs; estimated 5,000 burrows with eggs (POBSP) . 5-11 Sept. 1,600 Completely down-covered young to down- less near fledging young; ca. 65% of burrows with young more than 90% clear of down (POBSP). 1968 17-19 Mar. ? None positively identified; none in usual areas (BSFW, POBSP). 1969 9 Sept. ? 2 seen (BSFW). *Hggs probably present but overlooked by observers. 120 SOOTY SHEARWATER Puffinus griseus Status Accidental; one record: April 1906. Observations Bailey (1956: 57) reported the only known record from Laysan, a male collected 17 April 1906 by Bompke and now in the Bishop Museum (BPBM 4536). These shearwaters commonly migrate through the Hawaiian area and specimens, mostly carcasses that have washed up on the beach, are known from Midway and Kure Atolls in the northwestern Hawaiian chain (Clapp and Woodward, 1968: 8). WEDGE-TATLED SHEARWATER Puffinus pacificus Status Abundant breeder; maximum recent estimate [1,000,000]. Present from March through early December; absent remainder of year. Most nesting is from June through November. Nests in burrows, chiefly under the Eragros- tis association between the lagoon and the beach crest. Populations The maximum recent estimate (1,000,000 June 1967) is probably ex- cessive since no other recent estimates, made independently by a series of different observers, have exceeded 200,000 birds (Table WT-3). This nocturnal, burrowing species is difficult to census, however, and more accurate appraisals of the Laysan population must await lengthier, more detailed studies. Maximal populations probably are more on the order of several hundred thousands than one million. The earliest available population estimates, those by Dill and Bryan (100,000), and Wetmore (77,500) are probably not significantly different from most large estimates made recently. Annual Cycle Most observations indicate that birds begin returning in March after an absence during the late fall and winter. During March, April, and May they engage in a long period of courting activities and burrow prepara- tion which culminates with egg laying in early June. Eggs are laid through perhaps early July and eggs hatch from the last week of July through mid- or possibly late August. Young fledge from early November through mid-December. However, two observations do not fit this breeding regime. Dill and Bryan reported that young had "nearly fledged" by 4 June 1911 and Kramer found "only two downy young" at the end of April 1959 (while otherwise 121 indicating that the breeding season was beginning). We think both observations are probably erroneous in view of the consistency of many other observations by many other observers. Kramer, who failed to men- tion the presence of Bonin Petrels in his report, may have confused the young of that species with those of the Wedge-tailed Shearwater. Ecology Breeding: In May 1902 Fisher (1903a: 791-792) found most Wedge- tailed Shearwaters in "a zone perhaps 50 yards wide around the lagoon, some distance seaward from the bare flood plain" and noted that they were rare elsewhere on the island. They burrowed among the tall bunch- grass and in the open “among matted juncus [Cyperus] and succulent portulaca." Fisher noted that burrows were usually at least 3 feet long--often longer and very rarely shorter. In April to June 1911 Dill and Bryan (1912: 17) found them "on nearly every part of Laysan, with the exception of the beaches and the hard shores of the central lagoon." In May 1923 (Wetmore, ms.) they were on the open sand in many areas of the island but more abundant near the lagoon. POBSP personnel found them burrowing in a wide variety of habitats. Some birds burrowed under the hardpan in open areas on the southwest portion of the island. These strong-roofed burrows can withstand a considerable amount of weight and may be used several years in succession. Others dig burrows in loose sand under rocks, particularly at the north, southwest, and south ends of the island (Fig. 37). Most burrow beneath the varied vegetation associations such as the belt of Scaevola around the island and the belt of Ipomoea near the edge of the lagoon. Most POBSP survey parties, however, reported greatest densities of nesting birds under Eragrostis, particularly on the inner slope of the west side of the island. Burrow depths in such areas were three feet or more as opposed to those found under rocks, some of which were no more than a foot deep. Occasionally eggs were found being incubated on the surface of the ground. Non-breeding: Clubs (aggregations of roosting birds) were noted by Dill and Bryan (1912: 17) and by POBSP observers. Dill and Bryan noted during April to June 1911 that "at times a dozen or more of these birds congregate." POBSP observers found such aggregations on most visits to Laysan. In September 1964 groups of up to 50 birds were found on the open beaches of all but the west side of the island and clubs were seen in open areas of the interior in July 1965. In August 1965, October 1966, and September 1957 clubs containing from 30 to several hundred birds were seen in open areas bordering the lagoon, notably the northwest cor- ner (Fig.38), in open areas of the interior such as the blowouts at the southwest corner of the island, and on the beaches, particularly the north beach. Notes from the August visit indicate that larger clubs were found inland and that the aggregations along the lagoon were decidedly smaller. No such difference was noted in clubs seen in the same area in September 1967. 122 Figure 37. Downy young Wedge-tailed Shearwater at nest site under rock on southwest beach, September 1967. Photo by D.I. Hoff. Figure 38. Wedge-tailed Shearwater club near northwest corner of lagoon, September 1967. Photo by R.B. Clapp. 123 Data from June visits suggest that clubs are smaller and less numerous earlier in the breeding cycle. No clubs were seen in June 1966 and those present in June 1967 were considered smaller than those observed later in the nesting season. These clubs probably contain non-breeders or birds that failed in early nesting attempts but criti- eal observations to determine the status of these birds are lacking. Specimens Sixty-nine Wedge-tailed Shearwater skins from Laysan are currently distributed in museums as indicated in Table WIS-l. Three additional mounted specimens are distributed as follows: SUI (1 in Laysan exhibit); BPBM (2). Also preserved are at least 8 skeletons (BPBM, 2; USNM, 6); 1 alcoholic (BPBM); and 3 eggs (BPBM). Table WIS-1. Locations of Wedge-tailed Shearwater skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 6 4 2 12 BPBM 6 3) 5 14 DMNH a Al, O 2 MCZ ik O O 1 SUL 2 2 O 4 UMMZ O O 1 aL USNM (non-POBSP) 14 14 4 32 Other* O O 3 3 Totals 30 2 iy 9 *Hachisuki (1); Yale U. (1); Indiana U. (1). Banding and Movements The POBSP banded 7,392 adult Wedge-tailed Shearwaters on Laysan (Table WIS-2). One, 615-18119, banded on Lisianski as an adult on 22 August 1964 was recaptured on Laysan on 8 August 1965. On 16 September 1964 a Wedge-tailed Shearwater with an orange streamer (indicating that it had been banded on Johnston Atoll) was seen but not captured. None from Laysan has been recovered elsewhere. 12h Table WIS-2. Wedge-tailed Shearwaters banded on Laysan by the POBSP. Period of Survey Number Banded 1964 Sept. 700 1965 Mar. 25 July 1, 000 Aug. 4,000 1966 June 396 Oct. 100 1967 June 800 Sept. 371 Totals 75392 Table WIS-3. Observations of Wedge-tailed Shearwaters on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1828 24 Mar. ? Probably present; based on Isenbeck's vague description that was later identified by Rothschild (1893-1900: v). 1890 16 July ? Probably present (Lyons, 1890: 91). 1891 16-27 June very scanty (Rothschild, 1893-1900: 47). Eggs numbers being laid 16-18 June (Munro, 1941la: 2). 1896 2 June- 2 Nesting (Schauinsland, 1899: 101). 24 Sept. 1902 16-23 May abundant Second in numbers to Bonin Petrel; egg-laying began early June (Schlemmer vide Fisher); preparing burrows in May but no eggs found (Fisher, 1903a: 791-792) . 1903 Apr. ? 10 specimens and -2 eggs collected by Bryan (AMNH, BPBM). 1904 10 May 2 1 collected by Schlemmer (MCZ). 1906 17 Apr. 2 At least 1 collected by Bompke (BPBM). 1911 24 Apr.- 100, 000 Young nearly fledged (Dill and Bryan, 5 June LOU ee) 1912 22 Dec.- ? 1 half-grown young in burrow late Dec- 1913 11 Mar. ember; first adults returning 10 March (Bailey, 1956: 55). Only bird seen during winter [Bailey's half-grown young?] was a dying bird on 24 December (Willett, ms.). 125 Table WIS-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References mgs) |. SNpY possibly (Munter, 1915: 139). 250, 000 1918 8-10 Sept. 150, 000 (Diggs, ms.). 1923 8 Apr. thousands Sitting in pairs (Dickey, ms.). 29 Apr.- 77,500 Copulating and preparing burrows 14 May (Wetmore, ms.). 1930 2-18 Aug. 2nd most Mating and burrowing (Wilder, ms. b). abundant species 1936 7-8 Mar. ? Some present in burrows (Trempe, ms.). aes 4 1 adult seen; hundreds of dead young (Coultas, ms.). 1950 23 June numerous In burrows (POFI). 1951 12 May 2 Nesting (POFI). late June- 6, 290 Diurnal census of shearwaters, largely early July Wedge-tailed (Brock, 1951b: 18). 1957 25 June- 40, 000 1 or more nests with eggs found (Wood- 3 July side, ms. b). 8-12 July ? (Labrecque, 1957: 18). 1958 27 May- ? Most copulating and preparing burrows. 4 June “After inspection of hundreds of bur- rows" a single egg found 1 June (Warner, ms.). 1959 28 Apr.- ? Most beginning to nest; only 2 downy 1 May young seen (Kramer, ms.). 1961 7-8 Mar. not many Not breeding, no eggs found (Woodside and Kramer, ms.). 4-10 Sept. ? Many adults present during day; large numbers arriving at night. Young about 1/4 grown (Woodside, ms. c). 126 Table WIS-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1962 14-19 June usual Egg laying just begun (Kramer and abundance Beardsley, ms.) 1963 11-13 Feb. 0 (POBSP). 3-10 Dec. 2? Young nearly feathered; ca. 150 dead young seen on beach (Walker, ms. b). 1964 10-11 Mar. 50-100 25-30 seen; courtship behavior ob- served (BSFW, POBSP). 16-20 Sept. 25, 000 Many non-breeding birds present; nearly all burrows contained small young but a few half-grown chicks seen (BSFW, POBSP). 1965 6-11 Mar. 100-200 Beginning to return (POBSP). 17-21 July 10, 000 Second in abundance only to Sooty Terns. All burrows examined contained eggs in varying stages of incubation (POBSP) . 5-12 Aug. 150, 000 Ca. 25,000 newly hatched chicks and 25,000 near-hatching eggs; many eggs hatched during survey (POBSP). 1966 26-31 Mar. thousands No eggs found (BSFW). 10-16, 20-21 June 200, 000 Egg-laying just begun; much burrowing activity (POBSP). 17-18 Sept. many Thousands of large downy young in bur- thousands rows (BSFW). 20=23 Oct. 200, 000 Ca. 40-70,000 young present. All seen were beginning to assume juvenal plumage (POBSP). 1967 18-19 Mar. very few Not breeding (BSFW, POBSP). 7-Le June [1, 000, 000 j Ca. 10,000 burrows present. All eggs observed were fresh or slightly incu- bated. Estimate probably excessive (POBSP). I2sT/ Table WIS-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1967 5-11 Sept. 85, 000 Small to large downy young; most burrows contained medium-sized downy young (POBSP). 13 Dec. ? No adults; 1 bedraggled immature found in surf (BSFW). 1968 17-19 Mar. 100 Scattered birds; little moaning; no breeding (BSFW, POBSP). 1969 26-29 Mar. thousands Adults digging burrows; no evidence of egg laying (BSFW). 9 Sept. many 3 downy young (BSFW). thousands CHRISTMAS SHEARWATER Puffinus nativitatus Status Common breeder; maximum recent estimate: 10,000. Present from mid-February through late October; absent remainder of year. Most nesting is from mid-April through late September. Nests throughout the island, chiefly on the surface under dense vegetation or in a shal- low trench under debris, vegetation or boulders. Populations Population estimates for the early part of this century are about ten times larger than recent estimates (Table CS-3). This may indicate a decline in population but may also reflect the absence of recent visits during April and May--the early part of the breeding season when birds are presumably most conspicuous. Recent summer estimates suggest a population of 3,000 to possibly 10,000 birds at that season. Annual Cycle Bailey noted that the first birds arrived on 13 February 1913, but none was seen on the only other February visit (11 to 13 February 1963). Birds were reported on all March visits. Eggs may be laid as early as 5 to 8 March (1901), but no eggs were found on the six recent March visits (1964 to 1969).* Most egg-laying occurs from late April *On some visits, as in March 1967, the presence of eggs may have been overlooked. 128 through mid-May. Laying is essentially finished by the end of May but possibly a few eggs are laid in early June. Hatching may begin in the last week of May but the peak is from mid-June through early July. Young may fledge as early as the last week in July, but most fledging occurs from early September through early October, with a very few late young probably fledging in early November. Birds probably are absent from the island from sometime in November until mid-February. Christmas Shearwaters were not mentioned on any of the three December visits. Ecology Breeding: At night, early in the breeding season, concentrations of birds have been noted around the lagoon and open Eragrostis areas. Most nesting occurs under the Scaevola rimming the island (especially the north and west sides) and among various rock outcrops and boulders (particularly along the outer beaches). Smaller numbers of nests are scattered throughout the island. Nests are occasionally reported under Eragrostis and regularly in the Ipomoea areas but never on the open beaches. Essentially the same habitat was utilized by birds early in the century (Fisher, 1903a: 792) but during the denudation of the island the rocky outcrops must have been of major importance. Eggs are usually laid on the ground under dense Scaevola and also in shallow unlined trenches under other vegetation, boards, or rocks. Schauinsland (1899: 54) mentioned burrows underground but was probably referring to burrows of P. pacificus. Fisher (1903a: plate 8) shows a bird incubating an egg at the edge of vegetation without overhead shelter. Nests are easily overlooked early in the cycle but nearly fledged chicks are commonly seen outside the nest burrows and along the edges of vege- tation at night. Non-breeding: At night roosting birds are usually found in open areas at the edge of vegetation. Most birds leave the island by dawn and by mid-morning the remainder retire to the shade of rocks or vegetation and disappear into shelter as the day advances. Specimens Eighty-four Christmas Shearwater skins from Laysan are currently distributed in museums as indicated in Table CS-l. Four additional mount- ed specimens are distributed as follows: BPBM (1 immature male); CMNH (1 adult in Laysan exhibit); SUL (2 birds in Laysan exhibit). Also preserved are at least 7 skeletons (BPBM, 3; USNM, 4); 1 alcoholic (USNM); and 11 eggs (BPBM, 1; USNM, 10). Banding and Movements The POBSP and BSFW banded 317 Christmas Shearwaters on Laysan (Table CS-2). No interisland movements were recorded. 129 Table CS-1. Locations of Christmas Shearwater skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 7 3 5 15 BPBM (0) 2 8 10 CMNH IL i al 3 DMNH 3) 1 (0) 4 MCZ 2 3 0) 5 SUL O 2 0) 2 UMMZ O 4 6) ye USNM (non-POBSP) 13 ah a 38 (POBSP) ©) 0) iL 1 Other* i 1 O 2 Totals 27 1 il 8 Shaw Coll. (1'c); Utah St. Univ... (1 9). Table CS-2. Christmas Shearwaters banded on Laysan. Period of Survey Bander Adults Young Age Unknown Total 1964 Mar. BSFW @) ) 8 8 Sept. POBSP 2 88 @) 90 1967 June POBSP 100 ) ) 100 Sept. POBSP 58 23 38 119 Totals 160 111 317 Table CS-3. Observations of Christmas Shearwaters on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1891 16-27 June ? Nesting (Rothschild, 1893-1900: 45). Near-hatching eggs and recently hatched chicks (Munro, 1941c: 16-17). 1896 24 June- ? Nesting (Schauinsland, 1899: 101). 24 Sept. 1902 16-23 May ? Eggs (Fisher, 1903a: 792-3). Of 4 eggs collected 1 was fresh and 3 were almost ready to hatch (USNM). 130 Table CS-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1903 Apr. 2 5 specimens and 1 egg collected by W.A. Bryan (AMNH, BPBM). 1904 24 May 2 1 collected (Schlemmer?) (MCZ). 1907 20 May 2 4 collected (Schlemmer?) (MCZ). 1911 24 Apr.- 75,000 Fresh eggs, first 2 weeksof May (Dill 5 June and Bryan, 1912: 17). 1912 22 Dec.- z First pair arrived 13 Feb.; abundant 1913 11 Mar. by 17 Feb.; "nest building and pre- paring to breed" by 9 Mar. (Bailey, 1956: 5 Dp 57). CTS ae Ss epre 50, 000 (Munter, 1915: 139). 1916 9 Feb. 2 1 seen (Munter, ms.). 1918 8-10 Sept. 75,000 (Diggs, ms.). 1G23) Se OuNpr.. 2 Only a few dozen seen by day but num- bers becoming larger at night (Dickey, msi) 12 Apr. ? Becoming more common daily. Bird col- lected with fully formed egg in oviduct (Dickey, ms.). 8-13 Apr. fairly (Wetmore, ms.). common 12-14 Apr. ? 7 fresh eggs collected (Dickey, ms.; USN) . 29 Apr.- 2,000 1 egg found 7 May (Wetmore, ms.). 14 May 1957 25 June- 10, 000 (Woodside, ms. b). 3 July 1958 27 May- ? Eggs under 2 weeks old (Warner, ms.). 4 June 1959 28 Apr.- < 100 None found nesting, no pairs noted 1 May (Kramer, ms.). 1961 7-8 Mar. ? Paired, few with eggs (Woodside and Kramer, ms.). 131 Table CS-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1961 4-10 Sept. ? Well-developed young; many had de- parted (Woodside, ms. c). 1963 11-13 Feb. fe) (POBSP) . 1964 10-11 Mar. 30-50 Courtship behavior (BSFW, POBSP). 16-20 Sept. 250-450 Ca. 200 chicks; most nearly fledged (BSFW, POBSP). 1965 6-11 Mar. 1, 000- Pairs; no eggs (POBSP). 2,000 17-21 July 2,500 Ca. 500 large chicks (POBSP). 5-12 Aug. 3, 000 Ca. 500 large chicks (POBSP). 1966 26-31 Mar. several No nesting activity (BSFW). hundred 10-16, 1, 000 Ca. 200 incubated eggs (POBSP). 20-21 June 17-18 Sept. 15 Adults; 3 full-grown young with scant down on nape (BSFW). 20-23 Oct. 50 Nearly fledged chicks (POBSP). 1967 18-19 Mar. 100 Pre-nesting (BSFW, POBSP). 7-12 June 6, 000 Lightly incubated eggs to medium- sized young; of 36 nests tabulated, 2h (67%) contained eggs and 12 (33%) contained small downy chicks; esti- mated 2,000 nests (POBSP). 5-11 Sept. 2, 000 Medium-sized downy young to nearly fledged young; most nests contained large downy young (POBSP). 1968 17-19 Mar. 400-500 Many apparently paired; no nesting (BSFW, POBSP). 1969 26-29 Mar. 7,500 Seemed more abundant than on previous March visits (BSFW). 9 Sept. ? -More common than on earlier visits (BSFW) . 132 SOOTY STORM PETREL Oceanodroma tristrami Status Common breeder; maximum recent estimate: 2,000 to 3,000. Present from late October through June; absent remainder of year. Probably nests from late December until June. Nests in burrows in small, local colonies under the zone of dense vegetation (usually Eragrostis and Ipomoea) or under guano hardpan near the lagoon. Populations No highly accurate population data are available because of this species' secretive nature and the paucity of scientific visits during their peak nesting period. It occurs in small, local, scattered colonies, and on Laysan is often unsuccessful as a breeding species (Table SSP-2). The several hundred nests present in 1913 suggest a population of a thousand or more, which is roughly the same as recent estimates. (Willett's [ms.] estimate of 20,000 breeding birds was almost certainly excessive.) Annual Cycle Birds evidently return to the island in October after an absence of almost five months. Egg laying begins in mid- or late December and con- tinues through at least January. The only definite record of egg-laying is for late December and the first two weeks of January 1913. Hatching probably begins in early February and continues through at least early March. In mid-March 1968 one medium-sized downy young was found while other burrows contained only the adult birds and their empty nests. Fledging probably occurs from April through June. Ecology Breeding: Fisher (1903a: 795) reported (vide Max Schlemmer) that these petrels nested in burrows under scattered coral boulders on the southwest side of the island. In view of more recent observations, we suspect that this comment may, in fact, have referred to Bulwer's Petrels. During the winter of 1912 to 1913, Bailey and Willett (Bailey, 1956: 61) found two colonies, one on low ground at the north end of the lagoon, and the other in the southwest corner. Burrows averaged 5 inches in diameter and 2 1/2 feet in length; the nests were composed of rootlets, weed stems, and leaves (Willett, 1919: 61). Many observers have reported dead birds in the vicinity of the lagoon. Much of this mortality seems to be the late season loss of young typical of many seabirds. Bailey (1956: 61) recorded a considerable nest loss from flooding and shifting sand during the winter of 1912 to 1913. Since flooding of the lagoon is common, it may have an important limiting effect on storm petrel populations. USS) In recent years most birds have been found near the lagoon in the Eragrostis-Ipomoea zone. In 1965 birds were restricted to an area within 100 yards of the lagoon. In March 1968 a small colony (100 yards in diameter) was present at the south end of the lagoon under dense Ipomoea and hardpan about 200 yards from the lagoon border. A calling bird was also heard at the northwest end of the lagoon. Specimens Forty-five Sooty Storm Petrel skins from Laysan are currently distributed in museums as indicated in Table SSP-1. An additional mounted specimen is in the Laysan Exhibit at SUI. Also preserved are a skeleton and a head at USNM. . Table SSP-1. Locations of Sooty Storm Petrel skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 3 2 2 Wa BPBM 3 2 Jk 6 CMNH ale 2 (©) 3 UMMZ 4 6 0 10 USNM (non-POBSP) 6 5 i 12 (POBSP) 2 2 @) 4 Other* 2 il 0 3 Totals al 20 mn 5 *Law Coll. (200); Mus. d'H. Naturelle (1 9). Banding and Movements One hundred one Sooty Storm Petrels have been banded on Laysan: 91 adults and 1 chick by the POBSP in March 1965, and 9 adults by the BSFW in March 1966. No interisland movements were recorded. Table SSP-2. Observations of Sooty Storm Petrels on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1896 24 June- 0) Remains of skeletons found; skin sent oh Sept. to Bremen later in year after birds returned to nest (Schauinsland, 1899: 101; Rothschild, 1893-1900: 308). 134 Table SSP-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1902 16-23 May hardly 1 hurt or sick bird with a trace of common down, and up to a dozen dead ones (Fisher, 1903a: 795). 1903 23-29 Apr. % At least 9 (6 adults) collected by W.A. Bryan (AMNH, BPBM). 1911 24 Apr.- not A few dead and dying fledged young; 5 June common 2 adults (Dill and Bryan, 1912: 18). 1912 22 Dec.- 20, OOO* Substantial breeding colony (Willett, NeW ILL Wwe, 1919: 61). Eggs laid in late December and early January (Willett, ms.). Several hundred nests destroyed by rising lagoon waters; others by sand storms (Willett, 1919: 61; Bailey, 1956: 61). 1918 8-10 Sept. ©) (Diggs, ms.). 1923 2/7 Apr. ie Nearly fledged young found dead (Ball, ms.). 1936 12 Dec. apparently One found in one of many burrows not many examined (Coultas, ms.). 1957 2 June- 2 Post-nesting (Woodside, ms. b). 3 July 1961 7-8 Mar. Z 1 dead week-old chick (Woodside and Kramer, ms.). 4-10 Sept. 0 (Woodside, ms. a). 1963 11-13 Feb. 12 No nests (POBSP). 3-10 Dec. % Not nesting (Walker, ms. b). 1964 10-11 Mar. e 1 dead bird (BSFW, POBSP). 16-20 Sept. 0 (BSFW, POBSP). 1965 6-11 Mar. 2, 000- Young near fledging; adults digging 3, 000 burrows (POBSP). 17-21 July 0 (POBSP). 135 Table SSP-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1965 5-12 Aug. O (POBSP) . 1966 26-31 Mar. 2 1 nearly full-grown chick; 9 adults banded (BSFW). 20-21 June ? 1 carcass (POBSP). 17-18 Sept. ) (BSFW). 20-23 Oct. 1,500 No nests; numbers apparently in- creasing from night to night (POBSP). 1967 18-19 Mar. several No nests (BSFW, POBSP). 7-12 June 10 3 or 4 dead birds; 1 fledged chick (POBSP). 5-11 Sept. ) (POBSP). 1968 17-19 Mar. 200 1 local colony; others possibly present; only a single medium- sized downy young (and adults) found in the several burrows examined (BSFW, POBSP). 1969 26-29 Mar. ? 2 adults found in a nest-burrow (BSFW). 9 Sept. O None found in colony at southern end of lagoon (BSFW). *Estimate of number of breeding birds by Willett (ms.). RED-TAILED TROPICBIRD Phaethon rubricauda Status Common breeder; maximum recent estimate: 4,000. Some present in all months but only small numbers present from late fall through winter. Most nesting is from late April through early October. Nests on the ground under Scaevola or other vegetation, chiefly around the perimeter of the island. 136 Populations Population figures for Red-tailed Tropicbirds (Table RTTB-3) are probably less reliable than those for any other large, primarily diur- nal seabird nesting on the island because nests are usually hidden beneath dense shrubbery and are easily overlooked by survey parties. Recent estimates suggest that populations are larger now than during the period about 1915 through 1930 but the paucity of estimates from 1890 through 1913 makes it impossible for us to determine whether popu- lations of that period differed from those of today. It is very likely that the tropicbird population was greatly reduced when the island was denuded of vegetation since almost no nesting sites were then available. In May 1923, Wetmore noted, "They are hard put here to find any shelter for nests and I doubt if many will succeed in breeding successfully here." His count of 80 birds is the lowest ever recorded in the period from May to July. Annual Cycle There probably is no time when Red-tailed Tropicbirds are not present on Laysan, but very few are present in winter. Numbers apparently begin to increase in late March or April with peak numbers present from mid-May to August. The population begins to decline in September, and by late October there are few birds on the island. Population size closely corresponds with the amount of nesting activity. Minimal numbers are present in winter and early spring when few birds are nesting. Laying may begin as early as early January (1913) but in most years nesting probably begins in March (1958, 1962, 1965-67, 1969); however, in 1923 and 1911, it apparently did not begin until May. The peak laying period is probably most often from the last week of April through early June but eggs may be laid through early October (1912). It seems likely that future visits during the winter months will reveal that at least a few eggs or young may be present in any month. Eggs may hatch from mid- February through mid-November and young may fledge from mid-May through the end of January. Peak periods of hatching and fledging rather consis- tently fall between early June and mid-July and late August and early October, respectively. Despite the extended breeding period, the population as a whole apparently follows a distinct annual cycle with most breeding occurring from late April through early October. Ecology Breeding: Fisher (1903a: 795) found Red-tailed Tropicbirds under bushes, including the now extinct Chenopodium oahuense, and stated that ".,.not infrequently several will congregate beneath colonies of Fregata..." In 1923 when vegetation was almost non-existent, Wetmore (ms.) found the few birds present nesting about rock piles and inside buildings. 137 Presently most nesting occurs under Scaevola and is therefore con- centrated around the perimeter of the island. In June 1966 the largest concentrations were found on the western side. In September 1967 birds were considerably more abundant on the west, northwest, north, and east sides of the island than they were on the south and southeast sides. Birds often nest in loose groups of 5 to 20 or more, with 30 or TK) yards between the groups. Eggs are laid in shallow scrapes without nesting material but some plant debris may be present incidentally. Specimens Eighty-six Red-tailed Tropicbird skins from Laysan are currently distributed in museums as indicated in Table RITB-1. Twenty-three of these are dataless "Japanese trade skins," at least in part from the 1909 to 1910 feather raids. Six additional mounted specimens are dis- tributed as follows: BPBM (2); CMNH (1 in Laysan exhibit); SUI (3 birds in Laysan exhibit). Also preserved are at least 3 skeletons (BPBM, 2; USIM, a 1 alcoholic (BPBM); 1 skull (USNM); and 12 eggs (BPBM, 6; USNM, - Table RETB-1. Locations of Red-tailed Tropicbird skins from Laysan. Adult Adult Museum Males Females Other Totals AMMN 6 4 4* ak BPBM 1 3 6 10 CMNH iL iL 3x* 5 DMNH a 5 Oxx 8 MCZ 4 3 @) ii SUL 1 2 ) 3 UMMZ 1 ) 1O*%* 13 USNM (non-POBSP) 8 6 2 16 Totals 23 2 39 8 *Includes 7 "Japanese trade skins." **"Japanese trade skins." ***Includes 11 "Japanese trade skins." Banding and Movements The POBSP and BSFW banded 638 Red-tailed Tropicbirds (Table RTTB-2). One adult, 705-13327, banded on 10 September 1967, was recovered at sea ca. 15°00'N, 117°30'W on 27 February 1968. In addition, an adult banded on Laysan 31 May 1958 by Dale Rice was recaptured there by the POBSP on 19 September 196}. 138 Table RITB-2. Red-tailed Tropicbirds banded on Laysan. Period of Survey Bander Adults Young Age Unknown Total 1958 June BSFW @ O 125 125 1964 Sept. POBSP 35 65 0 100 1965 Mar. POBSP 2 @) ) 2 Aug. POBSP 62 37 0 99 1966 Mar. BSFW 3 ) 0 3 June POBSP y 0 0) 4 1967 June POBSP 98 O ) 98 Sept. POBSP 90 iLL 0 207 Totals 29 219 125 638 Table RITB-3. Observations of Red-tailed Tropicbirds on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1890 16 July a Incubating eggs, few of which had hatched (Lyons, 1890: 91). 1891 16-27 June ? Breeding; eggs (Rothschild, 1893- 1900: 34). 1896 24 June- 2 Breeding (Schauinsland, 1899: 101). ok Sept. 1902 16-23 May fairly Most nests had heavily incubated common eggs (3 collected had begun incu- bation [USNM]). 1 downy nestling seen (Fisher, 1903a: 796). 1903 Apr. ? 3 specimens and 5 eggs collected by Bryan (BPBM). 1905 2 May 2 1 collected by Schlemmer (MCZ). 1907 4-21 May a 6 collected by Schlemmer (MCZ). 1911 24 Apr.- 300 Very few seen first 3 weeks; common 5 June later and nesting (Dill and Bryan, Gs TO) 1912 22 Dec.- ? "Nesting in small numbers from late 1913 +11 Mar. December into March." Adults rarely seen in December and January but plentiful by mid-February. A juve- nile noted on 24 December; large young noted on 29 December and 9 January; eggs on 4 and 19 January (Bailey, 1956: Table RTTB-3. (continued) Population aS) Date of Survey Estimate Breeding Status, Remarks, References 1912 22 Dec.- 62). Rather plentiful by 11 March 1913 11 Mar. (cont'd.) (Willett, ms.). 1918 8-10 Sept. 30 (Diggs, ms.). 1923 8 Apr: ? 3 or 4 pairs with eggs on the south- west ridge (Dickey, ms.). 8-13 Apr. fairly Seeking nest sites (Wetmore, ms.). common 29 Apr.- 14 May 80 Eggs (Wetmore, ms.). 1930 2-18 Aug. few in Evidently nesting (Wilder, ms. b). number 1936 7-8 Mar. ? Ca. 12 seen in air and several others found with eggs. 5 banded with brass rings (Trempe, ms.). 12 Dec. ) (Coultas, ms.). 1950 23 June 2 Nesting (POFI). 1951 late June- 200 Diurnal census (Brock, 1951b: 18). early July 1957 25 June- 1,000 (Woodside, ms. b). 3 July 8-12 July Eggs to nearly full-grown young (Labrecque, 1957: 18). 1958 27 May- ? Fresh eggs to young about a month 4 June old; most nests with heavily incu- bated eggs or newly hatched young (Warner, ms.). 1959 28 Apr.- ze Nests with eggs, including 1 that 1 May was pipped (Kramer, ms.). 1961 7-8 Mar. v4 A few seen flying; none on ground 4-10 Sept. not at peak abundance (Woodside and Kramer, ms.). Only large young (Woodside, ms. c). 140 Table RTTB-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1962 14-19 June abundant Young in all stages (Kramer and Beardsley, ms.). 1963 11-13 Feb. several (POBSP). 3-10 Dec. ? A fully feathered young found 5 December was gone by 9 December (Walker, ms. b). 1964 10-11 Mar. Ti No nests (BSFW, POBSP). 16-20 Sept. 4.00 Eggs to fledging young; most nests with nearly fledged immatures; esti- mated 160 nests present on island. Sample count of 80 nests: 6 (8%) with eggs; 8 (10%) with small downy young; 34 (43%) with medium-sized or large downy young; 32 (40%) with dependent immatures (POBSP). 1965 6-11 Mar. 7-10 3 seen on ground but no nests found (POBSP). 17-21 July 2, 000 Partially incubated eggs to fledg- ing young; most eggs heavily incu- bated; estimated 1,000 nests with young (POBSP). 5-12 Aug. 2,500 Eggs to fledging young. Sample count of 80 nests: 25 (31%) with eggs; 14 (18%) with small downy young; 33 (41%) with medium-sized or large downy young; 8 (10%) with dependent immatures. Estimated 200 nests with eggs; 800 young (including flying immatures) (POBSP). 1966 26-31 Mar. very Nests with eggs but no young found common (BSFW). 10-16, 1, 000 Estimates 400 nests; 88 of 90 (98%) 20-21 June nests in sample count contained eggs, 1 contained small young, and 1 contained large young (POBSP). 17-18 Sept. ? Eggs to nearly full-grown young (BSFW). 41 Table RTTB-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1966 20-23 Oct. 50 Several nearly fledged young; no nests with eggs or small young (POBSP). 1967 18-19 Mar. 50 Several nests with eggs. Ca. 30 seen in flight on 18 March (BSFW, POBSP). 7-12 June 4,000 Estimate includes all birds using the island; eggs to large downy young; most nests contained eggs. Sample count of 87 nests: 69 (79%) with eggs; 16 (18%) with small downy young; 2 (3%) with medium-sized or large downy young. Estimated 1,000 nests (POBSP). 5-11 Sept. 1,500 Eggs to fledging young; 117 young banded. Less than 5% of nests with eggs; ca. 5% with small downy young; ca. 10% with medium-sized or large downy young; ca. 85% with dependent immatures (POBSP). 13 Dec. 4 No check made of breeding status (BSFW). 1968 17-19 Mar. 100 Apparently just returning; none found nesting (BSFW, POBSP). 1969 26-29 Mar. 500 Most incubating eggs (BSFW). 9 Sept. common A few nests examined contained eggs to nearly fledged young (BSFW). BLUE-FACED BOOBY : Sula dactylatra Status Common breeder; maximum recent estimate: 2,000. Present and may breed throughout the year but most nesting occurs from the end of March through September. Nests chiefly around the island perimeter and around the central lagoon, normally in areas of open sand or scant vegetation. The nest is a scrape in bare sand, occasionally with a little added vegetation. 42 Populations No 19th century population estimates are available and more recent data (Table BFB-3) are too scanty to determine whether any significant population changes have occurred on Laysan during the 20th century. Recent estimates show nearly as much variation within a single year (e.g., 1965) as for various years. This annual variation is probably correlated with the stage of the nesting cycle and the number of non- nesting birds present. Munter found over a hundred Blue-faced Boobies killed by feather poachers; undoubtedly many more were destroyed in the 1909 raid as white birds were favored by the feather hunters. Thus, there probably was some population decline during the 1908-15 period. Since this species nests on bare sand, its nesting habitat probably was not greatly affected by the loss of vegetation. However, Wetmore (ms.) noted that it usually nested where some faint trace of vegetation remained. Annual Cycle Blue-faced Boobies oecur on Laysan throughout the year, but peak numbers are evidently present in the summer, coincident with the largest number of nests. Blue-faced Boobies have bred during all months, although perhaps not in every month in most years. Although the initiation of the breeding cycle may vary by a month or more from year to year, the population as a whole exhibits a distinct annual breeding regime. Eggs may be laid as early as February (1911, 1923, 1963, 1966, 1969) and even December (1912) but the number laid during these months is only a small proportion of the total laid during the entire season. In most years peak egg laying probably occurs during the last week of March and the first two weeks of April, with at least a few eggs being laid through early July. Peak hatching and fledging periods usually occur during the last three weeks of May and last three weeks of September, respectively. Late-fledging young may be present through mid-December but in most years breeding is essentially completed by mid- or late October. Ecology Breeding: Many observers noted that Blue-faced Boobies nested pri- marily around the outer perimeter of the island and several noted that they tended to nest more on the east than on the west. Palmer (Roth- schild, 1893-1900: 26) noted that "they invariably frequented the shore and never went inland." Fisher (1903a: 796), more specific in his ob- servations than others, reported that these boobies were "most plentiful on the northeast, east, and southern exposures, where the littoral slope 143 is broadest, but on the west side, where a little bluff replaces the seaward slope, the birds (were) absent." Neither Palmer nor Fisher reported any nesting on the inner slopes of the island, but Dill and Bryan (1912: 19) found some nesting on the interior slope of the east side of the island. In 1923 Wetmore found these birds nesting on bare sand but believed that most preferred to nest where some trace of vegetation remained. More recently, POBSP observers found these boobies nesting in many areas of the island. Most of the population nested around the island perimeter or along the central lagoon and only a relatively small pro- portion were found on the inner slopes. Nests on the perimeter were usually on the littoral slope and could be found on both sides of the Scaevola belt. Those on the seaward side of the Scaevola were usually in little openings; those on the inland side usually in the sand-Eragrostis association just behing the Scaevola. The former nesting situation was found most frequently on the north, east, and south sides of the island, while the latter appeared to be more common on the west side. Some also nested at the edges or within the sandy southwest blowouts. Surveys from August 1965 and June 1966 and 1967 reported that the most dense nesting concentrations occurred along the central lagoon or in the fringe of vegetation on the north and east beaches. In the former area the birds nested on the sand or mud margin, or in more open areas among the Cyperus. On one survey (March 1965) it appeared that the south end of the lagoon held a more dense concentration of nesting birds than did other areas around the lagoon. The eggs were laid on the bare sand, usually with no semblance of a nest; occasionally there may be a little vegetation scratched about the eggs or young (Fisher, 1903a: 796). Non-breeding: Non-nesting birds roost in open areas mainly on the beach and around the lagoon. These roosting flocks or clubs were seen on most POBSP visits. In September 1964 two clubs of 15 and 8 individuals, respectively, were present on the northwest beach. In March 1965 a flock of about 30 birds was found on the south beach. On the night of 10 September 1967 three clubs were found: one with 22 birds in an open sand area on the edge of the southwest beach; another with 50 birds on the south beach; and a third of about 175 birds on the southeast beach near the margin of vegetation. Data are insufficient to determine seasonal variation in the size and number of these clubs. Little is known about the age composition of the clubs but in September 1967 more than 90 percent of the birds in them were adults. yy) Specimens Forty-nine Blue-faced Booby skins from Laysan are currently dis- tributed in museums as indicated in Table BFB-l. Five additional mounted specimens are distributed as follows: CMNH (1 male in Laysan exhibit); SUI (4 birds in Laysan exhibit). Also preserved are at least 3 skele- tons (BPBM, 2; USNM, 1); 1 alcoholic chick (USNM) and 24 eggs (BPBM, 11; MCZ, 6 clutches; USNM, 4). Table BFB-1l. Locations of Blue-faced Booby skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 6 6 4 16 BPBM 2 2 6 10 CMNH db 1 O 2 DMNH 1 iL O 2 SUL O 3 }] 5 UMMZ O AL O aL USNM (non-POBSP) 4 5 3 12 Other* il O O dL Totals 15 19 15) IS) AUe Otmnlonidar (Gl cir Banding and Movements The BSFW and POBSP banded 618 Blue-faced Boobies on Laysan (Table BFB-2). Twenty-two birds originally banded or handled on Laysan were later reported elsewhere (Johnston Atoll, 1; French Frigate Shoals, 1; Lisianski, 20); 26 birds banded elsewhere (Johnston Atoll, 5; Gardner Pinnacles, 1; French Frigate Shoals, 4; Lisianski, 11; Pearl and Hermes Reef, 3; Midway, 1; Kure, 1) were recaptured on Laysan (Appendix Tables 4-5a and 4-5b). 145 LE co SOTTO OO OVSTO ClO) (© e STeTON bé osy pepueg sseTO xeg/asy yoem TO Sazoqumy OOoNHMOnH 6T SOFC ClO CleTe@ Qstore@ =! SSOWOHAOOSGOS COLE OO OLOlOlon@ Ot AHN &é be bé SUTNPV bd BUTTISON oemzeuMT yTnpeqng Tejoyqng 4 4TnpyY STe10L 92 G2 4M "TEN B96T t t u 4das Ze 9€ x eune 196T LT GT Al "400 OT t Mu eum 996T @) @) . “ony 0 @) i ATor 0 € ~ “TeW SQ6T 62, 69 ni) eades TT a u “TEN 496 0 @) dsd0od “qed €96T 0 0 MISd eunf QC6T 66 pele) Tepueg SAING q1npy 4Tmpy go potsag ‘ueskey uo pepueq Satqoog peoej-enTg ‘°*2-dad eTaeL 146 Table BFB-3. Observations of Blue-faced Boobies on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1828 24 Mar. i Probably with fresh eggs; based on description by Isenbeck (in Kittlitz, 1834; Rothschild, 1893-1900). 1890 16 July ? (Lyons, 1890: 90). 1891 16-27 June few (Rothschild, 1893-1900: 26). 1896 24 June- ® Breeding (Schauinsland, 1899: 101). 2h Sept. 1902 16-23 May @ Eggs and young present in about equal numbers; most eggs heavily incubated; 2 collected (USNM); young from newly hatched to about a week old (Fisher, 1903a: 796). 1903 Apr. % 2 specimens and 8 eggs collected by Bryan (BPBM). 1904 1-3 May ® 3 single eggs (incubation begun); 3 sets of 2 eggs (2 beginning incuba- tion; 1 advanced) (MCZ). 1911 24 Apr.- 85* In first week of May colony of 45 birds 5 June had large downy young and a few well- incubated eggs; on 5 June another colony of ca. 20 pairs had fresh eggs (Dill and Bryan, 1912: 19). 1912 22 Dec.- 200 Nest with egg found 24 December; 1913 11 Mar. several more in early January; 30 nests counted 18 February; first naked young found 26 January (Bailey, 1956: 71; Willett, ms.). 1915 3 Apr. 350 Nesting; 100 or more had been killed by feather poachers (Munter, 1915: 193). 1916 9 Feb. 4O Count (Munter, ms.). 1918 8-10 Sept. a5 No eggs or young found (Diggs, ms.). 1923 9 Apr. 2 Hatching eggs noted (Dickey, ms.). 8-13 Apr. ? Eggs to young about a week old (Wetmore, Tasty) Fe ee eee oe ee ! . : | Table BFB-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1923 21 Apr. 2 45 nests in one colony on SE side, largest colony on island (Ball, ms.). 2k Apr. ? Ca. half with eggs, many with hatch- ing eggs, and a few with two-week-old young (Dickey, ms.). 29 Apr.- 160 Fresh eggs to well-grown young; some 14 May just laying (Wetmore, ms.) 7 May ? 2 heavily incubated eggs collected (USN) . 1930 2-18 Aug. 6 No nesting reported (Wilder, ms. b). 1936 7-8 Mar. 2 Only eggs found; 6 banded with brass rings (Trempe, ms.). 12 Dec. 50-100 Had just finished nesting (Coultas, ms.). 1950 23 June common Medium and "full-sized" young (POFI). 1951 12 May 2 "Young in down" noted (POFI). late June- ? Not distinguished from other species early July of booby in census; 2,940 boobies (3 species combined); diurnal count (Brock, 1951b: 18). 1955 10 Feb. 100 Most seen at south end of lagoon (POFI). 1957 25 June- 300 1 nest with 2 eggs noted (Woodside, 3 July ms. b). 1958 27 May- 2 Most nesting pairs with naked young to 4 June young about a month old; most incubated eggs checked were rotten (Warner, ms.). 1959 28 Apr.- ? Most birds with eggs; a very few with 1 May young (Kramer, ms.). 1961 7-8 Mar. usual Few eggs present; most birds paired number (Woodside and Kramer, ms.). 4-10 Sept. ? Only large young and immatures (Wood- 147 side, ms. c). 148 Table BFB-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1962 14-19 June B Mostly 1/2- to 3/4-grown young; several nests with eggs (Kramer and Beardsley, ms.) 1963 11-13 Feb. small 1 nest with egg (POBSP). numbers 3-10 Dec. ? 1 full-grown unfledged bird and 1 im- mature; adults seen (Walker, ms. b). 1964 10-11 Mar. 100-150 Estimated 10 nests; all with eggs (20)* (BSFW, POBSP). 16-20 Sept. 420 Ca. 50 fledged immatures and 10 sub- adults; also 2 large flightless young (BSFW, POBSP). 1965 6-11 Mar. 400-500 -Nocturnal estimate; 40-50 nests, all (80-100)* with eggs (POBSP). 17-21 July 800% Estimated 400 young, half- to full- grown (POBSP). 5-12 Aug. 1, 000 Estimated 200 young--mostly from 1/2- (400)* to full-grown; a few recently hatched; no eggs (POBSP). 1966 26-31 Mar. 250 A few nests with eggs (BSFW). 10-16, 500 Fresh eggs to nearly fledged young; 20-21 June (300)* ca. 50 nests with eggs; 100 nests with young, mostly small chicks; ca. 100 flying immatures. Presumably from previous breeding season (POBSP). 17-18 Sept. a Some nearly fledged young (BSFW). 20-23 Oct. 250 Almost all young fledged; no eggs (POBSP). 1967 18-19 Mar. 150 1 nest with egg (BSFW, POBSP). 7-12 June [2,000] Estimated 300 nests; most with eggs, (600)* recently hatched or small downy young. Of 200 nests counted, 33% contained eges, 17% recently hatched young, 33% small downy young, and 17% medium-sized downy young (POBSP). 149 Table BFB-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1967 5-11 Sept. 500 2 large downy young and ca. 100 de- (200)* pendent immatures; no eggs or small young (POBSP). 13) We. at least No nests found but immatures in- several capable of flight seen (BSFW). hundred 1968 17-19 Mar. 150 Most were prenesting; most were paired; (30)* 6 nests with fresh or slightly incu- bated eggs seen (BSFW, POBSP). 1969 26-29 Mar. 64xx 32 nests counted, a few more probably present. of 15 nests examined were those of prelaying birds. Of the re- maining 11 nests, 9 (82%) contained eggs, 1 (9%) contained an egg and a chick, and 1 (9%) contained a small chick (BSFW). 9 Sept. ts Some large downy young and several flying immatures seen (BSFW). *Estimate is of number of breeding birds (birds with eggs or unfledged young). **An estimate of breeding birds. Possibly no more than 48 birds were with eggs or young. BROWN BOOBY Sula leucogaster Status Uncommon breeder; maximum recent estimate: [250]. Present in all months but most numerous from May or June through October. Most breeding occurs from late March through October. Nests in small colonies, usually in openings in Scaevola, on the west central part of the island. The nest, usually a substantial mass of vegetation, is on the ground. Populations Only in recent years have more than a very few been noted. Prior to 1923 it was recorded as a rare visitor (only 3 definite records) and in 1923 only a single nesting pair was present (Table BB-2). Small 150 breeding populations have been reported regularly since 1957. Recent POBSP observations indicate a minimum breeding population of 40 birds, although in some years (e.g., 1967) probably fewer birds breed. Popu- lations may reach a maximum of about 100 birds when numbers are swelled by young and visitors from other islands. Annual Cycle Few birds occur on the island during the winter, and the early spring population remains low even though nesting may begin in March. Available data on breeding status during various visits suggest that Laysan Brown Boobies may nest in at least ten months of the year (March-December) and further observations may well reveal that breeding may occur in any month. It appears, however, that in most years, very few birds, if any, breed from late December through mid-March and that most birds in the population complete their breeding cycles between mid-spring and mid-fall. Interpolation from the presence of eggs and size of young on various visits suggests that peak periods of laying, hatching, and fledging usually occur respectively in May, late June and early July, and late September and early October. Ecology Breeding: Recent nesting has occurred in a relatively small area at the inner edge of the Scaevola zone on the west central part of the island. Here 6 to 25 pairs have been found in one or two loose groups. Substantial nests of grass, sticks, and other vegetable matter are built on the ground. Non-breeding: Non-breeding birds may occur almost anywhere on the island, but are most frequently seen flying over the reef or roosting on rocks along the shores. No large concentrations have been found, but in September 1967 eleven birds were seen together on a flat rock off the west beach. Specimens We know of but three Laysan specimens: an adult male at AMNH and an adult female and an adult male at BPBM. Banding and Movements The POBSP and BSFW banded 28 Brown Boobies on Laysan through 1969 (Table BB-1). The only recorded interisland movement is of an adult male, 737-30106, banded on Southeast Island, Pearl and Hermes Reef, on 26 February 1963. It was recaptured where banded on 19 June 1963 with a newly hatched young and subsequently was recaptured on Laysan, 7 March 1965. On 18 March 1965 it was captured on Seal Island, Pearl and Hermes Reef. oe oat appa lM | 151 STeIOL il ©) @) T @) T 0) ‘i "TEN Q96T TI © 0 TT 0 + Jr ii eune 196T € (0) 0) & 9 0) € € Z ) 0 Z S & ©) 0) STe}OL 32 gb SqTnpy 33 3d 99 zapueg BANG JO potted aay 3.14 CUT, Te8707GNS 7TOPV TINY qFIMpy pepueg sseTo xeg/asy yoem jo szequni ‘ueskey uo papueq Setqoog uMOZg “T-dd eTqeL 52 Table BB-2. Observations of Brown Boobies on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1828 24 Mar. 2 Possibly present; based on vague description by Isenbeck (in Kittlitz, 1834) identified by Rothschild (1893- 1900: iv). 1891 16-27 June ? Not found by Palmer despite intensive search (Rothschild, 1893-1900: 31); Munro (1942b: 6) states they were “very scarce." 1896 24 June- % Regular visitor, but not nesting 2k Sept. (Schauinsland, 1899: 101). 1902 16-23 May O Not seen despite intensive search (Fisher, 1903a: 797). 1903, 928 Apr. 2 1 specimen collected by Bryan (BPBM). 1911 24 Apr.- - Not mentioned in report (Dill and 5 June Bryan, 1912). 1912 22 Dec.- 1 Only 1 seen in 3 months (Bailey, 1956: 1913 11 Mar. 69). Seen on 21 January (Willett, ms.). 1923 11 Apr. al 1 seen in the surf 11 April (Dickey, Hits, )) 23 Apr. 2 Pair found nesting by Ball (Dickey, ms.) 24 Apr. 2 Birds incubating 2 eggs (Dickey, ms.). 5-7 May 2 Nest still active (Wetmore, ms.). 1936 12 Dec. O (Coultas, ms.). 1950 23 June few No nests (POFT). & | 1951 12 May 2 No nests (POFI). | late June- ce 2,940 boobies (3 species combined); H early July diurnal count (Brock, 1951b: 18). : : 1957 25 June- 50 (Woodside, ms. b). ; 3 July 153 Table BB-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1958 27 May- a few 1 pair incubating addled eggs; at 4 June pairs least 2 pairs with young ca. 3 weeks old (Warner, ms.). 1961 7-8 Mar. 20 Nests with eggs; no young (Woodside and Kramer, ms.) 4-10 Sept. 2 Many immatures, adults, and a few eggs or small young (Woodside, ms. c). Some nearly mature young (Walker, ms. a). 1962 14-19 June 2 More seen this year than in past; most nesting birds with recently hatched young (Kramer and Beardsley, ms.). 1963 11-13 Feb. 2 Evidently not nesting (POBSP). 3-10 Dec. 2 Present, not nesting (Walker, ms. b). 1964 10-11 Mar. 2 None nesting (BSFW, POBSP). 16-20 Sept. 19 Diurnal count; 5 nests; 3 1/2 to 2/3 grown young and 2 almost fledged young (BSFW, POBSP). 1965 6-11 Mar. 5-10 No nests (POBSP). 17-21 July [250] Ca. 6 nests with heavily incubated eggs or newly hatched young; estimated 50 young present (estimates probably excessive) (POBSP). 5-12 Aug. 50 Estimated 20 nests: 5 with eggs, 15 with young from newly hatched to full- grown (POBSP). 1966 26-31 Mar. 25 Most at night (BSFW).* 10-16, 80 Fresh eggs to nearly full-grown young; 20-21 June 5 nests with eggs counted; estimated 20 young present (POBSP). 20-23 Oct. 100 Most breeding birds with large de- pendent young; 1 nest with eggs; num- ber of flying immatures (POBSP). 154 Table BB-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1967 18-19 Mar. 15 No nests (BSFW, POBSP). 7-12 June 50 5 nests, 4 with eggs and 1 with a medium-sized downy young (POBSP). 5-11 Sept. 20 4 nests with large downy young; 3 dependent immatures (POBSP). 13 Dec. 3 All adults (BSFW). 1969 17-19 Mar. 4 Only 4 birds (apparently paired) present in colony (BSFW, POBSP). 1969 26-29 Mar. O (BSFW). *One or more active nests present but not seen by survey party. RED-FOOTED BOOBY Sula sula Status Common breeder; maximum recent estimate: 2,000-3,000. Most breeding is from February through September with smaller numbers present during remainder of year. Builds bulky, well-constructed nest in Scae- vola rimming the island and in woody vegetation near the inner lagoon. Populations Although observers in the 19th century considered the Red-footed Booby a common part of the breeding avifauna, no actual population estimates were published. More recent observations document a ten-fold increase in population size from the early part of the 20th century to the present time (Table RFB-3). All observers from 1911 through 1930 noted that very few breeding birds were present, with the lowest breeding population during spring or summer a mere 60 birds in 1930. It seems likely that these low popula- tions resulted from a combination of factors--in part from decimation by feather hunters and in part by destruction of habitat by rabbits intro- duced to the island. In 1923 Wetmore noted only two colonies nesting in low bushes and remarked further that a number of nests were placed on the ground about the stems of old tobacco plants. Although ground nesting rarely occurs on some bushless islands in the Phoenix Islands to the Be south (POBSP data), no ground nests were found from 1963 through 1969 on Laysan or on other Northwestern Hawaiian Islands. The ground nesting reported by Wetmore suggests the extreme lack of nesting habitat avail- able at that time. Recent numerical estimates indicate that maximum populations are present during the summer and fall. During winter most birds leave the island and the population decreases by about 90 percent. Early spring (March) populations have been quite variable, probably due to variations in the inception of the beginning of the breeding season. Annual Cycle Some birds are present during all months but numbers are generally low during mid-winter. A limited amount of egg-laying occurs during February during some years. Generally, however, egg-laying begins in March with the date of inception varying somewhat annually. In 1964 and 1965 about 20 percent of the occupied nests had eggs by the second week of March; in 1967 a similar stage was reached two weeks later. Egg-laying usually begins in March and probably reaches its peak in April or May. Small young are generally present from sometime in late ' April or May through June, most nests containing half-grown or larger young thereafter. The first young usually fledge about late June but the peak fledging period is usually in August. Some dependent young are present in September but the breeding season is essentially finished by the end of that month. Ecology Breeding: Fisher (1903a: 797) noted that most of the birds nested on the inner slopes of the island, "usually well down towards the lagoon." He further noted that these boobies always built in bushes and never on the ground. Only during 1923 when vegetation was barely existent has ground nesting been observed. Presently, Red-footed Boobies nest primarily in Scaevola and Pluchea bushes (Fig.39). Most of the Scaevola occurs along the rim of the island and it is here, particularly in the northwest quadrate and on the south, that most of the boobies nest. Considerable numbers nest also in the Pluchea and Scaevola sur- rounding the lagoon, especially in the east and southwest portions. While most of the boobies nest in colonies by themselves, others often nest among the Great Frigatebirds that utilize much the same habitat. Non-breeding: Unoccupied and transient boobies utilize the nesting areas for roosting. The Casuarina tree and the palms are other important roosting sites. mre RSS, Calas, Rha ee Figure 3 Red-footed Booby at nest in low Scaevola, June 1966. Photo by P.C. Shelton. Specimens Forty-six Red-footed Booby skins from Laysan are currently distributed in museums as indicated in Table RFB-1. Seven additional mounted specimens are distributed as follows: BPBM (2); MCZ (1 male, 1 female, 1 nestling); SUI (2 birds in Laysan exhibit). Also pre- served are at least 1 skeleton (BPBM) and 11 eggs (BPBM, 9; USNM, 2 clutches). Banding and Movements The POBSP has banded 860 Red-footed Boobies on Laysan (Table RFB-2). Red-footed Boobies frequently travel between islands. Eighty-four birds originally banded, or handled, on Laysan were subsequently recaptured elsewhere, making 89 movements to other islands. Several birds were recaptured on two or more islands. Recaptures were: Johnston Atoll (39); French Frigate Shoals (25); Lisianski (14); Pearl and Hermes Reef (1); Kure (7); Wake (2); Namu, Marshall Islands (1) (Appendix Table 4-6a). Thirty-six birds (38 movements) from elsewhere visited Laysan from the following islands: Johnston, 14; French Frigate Shoals, 14; Lisianski, 7; Midway, 1; Kure, 1; Wake, 1 (Appendix Table 4-6b). WT Table RFB-l. Locations of Red-footed Booby skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 4 2 5 Tal BPBM O 0 5) 5 CMNH 2 IL ©) 3 DMNH 2 O O 2 MCZ iL O O ale SUI 3 4 ) 7 UMMZ ale 1 O 2 USNM if 6 2 15 Totals 20 L 12 Table RFB-2. Red-footed Boobies banded on Laysan by the POBSP. Numbers of Bach Age Class Banded Period of Survey Adults Subadults Immatures Nestlings Totals 1963 Feb. 3 0) ) ) 3 1964 Sept. 140 2 57 a 200 1965 Mar. ake 0 1 O 243 Aug. 0 O 184 0) 184 1966 June 18 O @ O 18 Oct. 115 3 O 2 120 1967 June 30 3 0 O 38 Sept. 55 3 O O 58 1968 Mar. a 0 ) 0 1 Totals 11 He 3 O Table RFB-3. Observations of Red-footed Boobies on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1828 2h Mar. 2 A vague description by Isenbeck (in Kittlitz, 1834: 123-124) is apparently this species. 1891 16-27 June very Nesting (Rothschild, 1893-1900: 28). plentiful 1896 24 June- ? Nesting (Schauinsland, 1899: 101). 4 Sept. —S ee 158 Table RFB-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1902 16-23 May ? Most nests with eggs; a few small downy chicks (Fisher, 1903a: 797). 23 May % 5 eggs of advanced incubation col- lected (USNM). 1903 Apr. ? 4 eggs and 3 specimens collected by W.A. Bryan (BPBM). 1904 2 May 2 1 collected by Schlemmer (MCZ). 1905 3 May 2 1 collected by Schlemmer (MCZ). 1906 30 Apr.- ? 2 specimens collected. (One col- 28 May lected on 28 May was a nestling [Mcz]). 1911 24 Apr.- 125 "Not very numerous," increased to 5 June peak about 5 June. Nesting (Dill and Bryan, 1912: 20). 1912 late Dec. very few (Bailey, 1956: 66). OTS evar oO (Bailey, 1956: 66). 11 Mar. 100 Nest building (Willett, ms.). 1918 8-10 Sept. 12-15 In bushes at north end of lagoon. No eggs or young found (Diggs, ms.). 1923 8-13 Apr. ? Small colonies with eggs and small young (Wetmore, ms.). 29 Apr.- 80 Count (Wetmore, ms.). 14 May 2 May ? Many eggs still unhatched (Wetmore, ms.) 13 May ? Nests under construction and with eggs seen (Wetmore, ms.). 1930 2-18 Aug. 60* Seen in two colonies, one of them in Casuarina. Young nearly full grown (Wilder, ms. b). 159 Table RFB-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1936 7-8 Mar. ? Nesting; only eggs seen (Trempe, ms.). 12 Dec. 12 One pair nesting in Casuarina (Coultas, ms.) 1950 23 June common Medium and "full-sized" young (POFI). 1951 late June- 2 2,940 boobies (3 species combined); early July diurnal count.(Brock, 1951b: 18). 1955 10 Feb. 20 At south end of lagoon (POFI). 1957 25 June- 1,000 (Woodside, ms. b). 3 July 1958 27 May- 2 "Most fertile eggs had hatched" 4 June (Warner, ms.). 1959 28 Apr.- ? Most adults on eggs; no young seen 1 May (Kramer, ms.). 1961 7-8 Mar. few Some with eggs, no young (Woodside and Kramer, ms.). 4-10 Sept. v4 Many immatures observed; no eggs or small young seen (Woodside, ms. c). 1962 14-19 June 2 Eggs and downy young (Kramer and Beardsley, ms.). 1963 11-13 Feb. 100 Several nests with eggs; 1 full-grown immature (POBSP). 3-10 Dec. ? Present, breeding (Walker, ms. b). 1964 10-11 Mar. 500-700 250-300 nests; 10-20% with eggs; some nest building (BSFW, POBSP). 16-20 Sept. 500 Also an estimated 200 flying imma- tures; only 1 dependent young (large) seen (BSFW, POBSP). 1965 6-11 Mar. 2, 000- Many sitting on empty nests; ca. 200 3,000 nests with eggs; no chicks seen (POBSP). 17-21 July 1, OO0O* Ca. 500 nests with half-grown to nearly fledged young (POBSP). 160 Table RFB-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1965 5-12 Aug. 1, 000* Also an estimated 200 flying immatures. Ca. 400 nests with young half-grown to nearly fledged. A few very small chicks seen (POBSP). 1966 26-31 Mar. 600* Perhaps 300 nests, a few with eggs, others empty (BSFW). 10-16, 20-21 June 1,500 Fresh eggs to nearly fledged young; ca. 100 nests with eggs; 300 with young; a fledged immature present (POBSP) . 17-18 Sept. 500+ (BSFW). 20-23 Oct. 2,500 Only a few near-fledging young present. About Th of population composed of subadults and flying immatures (POBSP). 1967 18-19 Mar. 150+ Several nests built but no eggs seen (BSFW, POBSP). 7-12 June 1 OO Estimated 200-250 nests. 172 counted: 89 with eggs (52%); 19 naked chicks (11%); 51 small downy chicks (30%); 12 medium-sized chicks (7%); 1 large downy chick (1%) (POBSP). 5-11 Sept. 2,000 Ca. 30 large downy young; ca. 80 non- flying immatures; ca. 200 flying immatures (POBSP). 13 Dec. at least None nesting (BSFW). several hundred 1968 17-19 Mar. 100-150 Little nesting; 8 active nests seen (1 with eggs; 7 empty) (BSFW, POBSP). 1969 26-29 Mar. Lox Of 60 nests whose contents were examined, 20 (33%) were empty but active and 40 (67%) contained eggs (BSFW). 9 Sept. 2 Several downy young noted (BSFW). *Estimate is of the number of nesting birds. 161 PELAGIC CORMORANT Phalacrocorax pelagicus Status Accidental; one record: October 1896. Observations The only certain record from Laysan, or from any of the islands of the Hawaiian area, is a female specimen collected 22 October 1896, ap- parently by one of Schauinsland'’s correspondents (Rothschild, 1893-1900: 308). Schauinsland (1899: 101) listed the species as a "winter guest." GREAT FRIGATEBTRD Fregata minor Status Common breeder; maximum recent estimate: 8,000. Peak populations occur between March and September with smaller numbers during the remainder of the year and fewest during winter. May be found nesting in any month but most birds breed from March through October. Builds bulky nests in _ the Scaevola rimming the island (particularly on the west side) and near the interior lagoon. Populations Records from the last decade of the 19th century indicate that Great Frigatebirds, like many other species breeding on the island, were very numerous at the turn of the century but subsequently suffered a great population decline, caused in part by the depredations of feather hunters and in part by the destruction of the vegetation (Table GF-3). Observations of Munter (1915), who visited the island immediately following the feather poaching era, compared with those of Wetmore (ms.), who visited Laysan when vegetation had become greatly reduced, indicate that the loss of vegetation was by far the more important factor in the reduction of the Great Frigatebird population. In April 1915 Munter observed that some 2,000 frigatebirds had been killed by poachers but he also estimated a population of 30,000, the highest ever recorded for this species on Laysan. Dill and Bryan's (1912) estimate of 12,500 made in the spring four years earlier suggests that the population was indeed large and that Munter did not overestimate numbers present. On the other hand, in May 1923 Wetmore estimated that only about 1,500 frigatebirds were present. Recent estimates for March, when frigatebirds would presumably be less abundant than in May, suggest that the population has increased con- comitant with the reappearance of vegetation. However, even the largest recent estimate, which may be somewhat excessive, is less than two- thirds as large as that recorded by Dill and Bryan. See 162 Annual Cycle Although POBSP population estimates vary considerably from month to month throughout the year, and in the same month in different years, an annual pattern of population change is apparent. In only one instance (March 1965) are populations recorded in the spring as large as those recorded in the summer when maximal numbers have been reported. Populations of flying birds (adults, subadults and flying immatures) decrease during the autumn as post-breeding birds disperse from the island. Some of this apparent decrease in population during the fall may be the result of incomplete observation rather than a real phenomenon. The numbers actually utilizing the island may be greater than recorded since many adults would be feeding away from the island during any one census period. Nocturnal populations are usually larger during this period. During the winter, populations on the whole are considerably smaller than during the breeding season. More observations during the winter are required to further document the reduction in populations at that time. The breeding cycle, or at least its initiation, is somewhat variable from year to year, but on the whole a fairly consistent annual breeding regime occurs. Eggs may be laid as early as mid- or late January (1936, 1964, 1969) and as late as mid- or late May (1963, 1967) but the peak laying period is usually during March. Peak hatching and fledging periods usually occur during May and late September through late October, respec-= tively. A few young may still be fledging in November and December and young birds, most presumably from the previous breeding season, can be found on the island through at least part of the succeeding breeding season. Although populations breed on an annual basis, it is possible that individuals usually breed no more often than every other year. Ecology Breeding: All observers except Wetmore (ms.) report frigatebirds nesting exclusively in woody vegetation; in 1923 birds were nesting on the scanty vegetation remaining and on the ground and slightly elevated ridges. At that time many eggs and some 150 birds (chiefly immatures) were destroyed by sand storms. In June 1967 frigatebirds nested only in Scaevola with nest height averaging 2 feet above ground. In March 1968 heights of eleven nests measured ranged from 12 to 28 inches above ground (x = 19.9 in.). Nesting areas have varied with the distribution of woody vegetation. The chief nesting area from 1890 through 1918 was along the east side of the inner slope of the lagoon basin. Although details are lacking, birds probably nested in Chenopodium, then quite widespread on the island, as well as Scaevola. Birds were scattered in small colonies which in 1911 (Dill and Bryan, 1912: 20) "would cover about 6 acres if placed near together." All observers reported small colonies of from a few to 50 pairs. 163 Recently birds have nested throughout the island wherever Scaevola was present, particularly on the west side (Fig.40). Nests frequently occur in scattered aggregations whose sizes vary with the amount of Scaevola present. In March 1968 the numbers of nests in each of eight colonies were recorded. Seven colonies in low (under 4 feet) Scaevola contained 4 to 10 nests each; the eighth colony of 35 nests was in tall (over 5 feet) Scaevola. The more advanced stage of nesting in the tall Scaevola sug- gests that more elevated positions may be preferred. Non-breeding: Adults and immatures of various ages spend much time on or over the island and roost in the Scaevola in large numbers at night. A considerable amount of movement to and from the island continues after dark. Adults spend much time at the nest. Once the nest site is deter- mined, one bird, usually the displaying male, remains at the nest most of the time. After the chick hatches, the young bird is usually shel- tered by one parent for a considerable time. Figure 40. Young Great Frigatebirds in low Scaevola above the west beach, September 1967. Photo by R.B. Clapp. 164 Specimens Seventy-nine Great Frigatebird skins from Laysan are currently distributed in museums as indicated in Table GF-l. Eight additional mounted specimens are distributed as follows: BPBM (1 male); CMNH (1 male and 1 female in Laysan exhibit); MCZ (1 nest- ling); SUI (4 birds in Laysan group). Also preserved are at least 2 skeletons (BPBM); 2 alcoholics (BPBM, USNM); 3 skulls (USNM); and 19 eggs (BPBM, 7; USNM, 12). Table GF-l1. Locations of Great Frigatebird skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 2 5 7 14 BPBM 2 ae 5% 8 CMNH iL 2 i 4 DMNH aL 2 a 4 MCZ 3 @) 0 3 SUL 6 2 2 10 UMMZ 2 0) aL 3 USNM (non-POBSP) 10 y 14 28 Other** 2 2 1 5 Totals 29 18 32 79 *Includes 2 destroyed by dermestids. **Brit. Mus. (Nat. Hist.) (1c, 2 99, 1 imm.); U. Minnesota (1c). Banding and Movements The POBSP banded 514 Great Frigatebirds on Laysan (Table GF-2). Eleven birds banded on Laysan were later recaptured or recovered as follows: French Frigate Shoals, 2; Kure Atoll, 5; Philippine Islands, 4. One bird originally banded on Kure was recaptured on Laysan (Appen- dix Tables 4-7a, 4-7b). 165 Table GF-2. Great Frigatebirds banded on Laysan by the POBSP. Numbers of Each Age/Sex Class Banded Period of Adult Adult Subtotal Sub- Imma- Nest- Survey oo" 9° Adults Adults tures lings Totals 1963 Feb. 2 7 9 alt O O 10 1964 Sept. 2 8 10 ab 4 O 15 1965 Aug. ) ) 0 O 2 0 2 1966 June 23 13 36 2 0 0 38 Oct. 2 6 8 iL al 135 145 1967 June 139 63 203* 76 O 0 279 Sept. 15 2 17 8 O ) 25 Totals 183 99 283 89 7 135 514 *Includes one bird not sexed at time of banding. Table GF-3. Observations of Great Frigatebirds on Laysan. : Population Date of Survey Estimate Breeding Status, Remarks, References 1828 24 Mar. 2 On nests but no eggs (Isenbeck, in Kittlitz, 1834: 121). 1890 16 July ? Downy young, east of lagoon (Lyons, 1890: 91). 1891 16-27 June ? Numerous rookeries throughout island; eggs and young present (Rothschild, 1893-1900: 23; Munro, 1953: 56). 1895 Sept. ? 3 or 4 collected by Hall (BPBM). 1896 24 June- ? Nesting (Schauinsland, 1899: 101). 2h Sept. 1902 16-23 May ? Eggs and recently hatched young (Fisher, 1903a: 798). 23 May ? 3 heavily incubated eggs collected (USNM). 1903 22 Apr. ? 4 specimens and 4 eggs taken by Bryan (BPBM) . 166 Table GF-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1906 18 Apr.- ? 5 specimens taken including one nest- 30 May ling (on 30 May) by Schlemmer (MCZ, CMNH). TOT FAA — 12,500 Nesting (Dill and Bryan, 1912: 20). 5 June 1912 22 Dec.- 2,500- Common on 22 December but most birds 1913 11 Mar. 3, 000 were immatures; adults began arriving after lst of year. Nest building began in middle or late January; first egg found 4 February; in late February a "few over" 1,000 nests counted, ca. 200 containing eggs; an estimated 3,000 birds present by 11 March (Willett, ms., Bailey, 1956: 74). 1915 3 Apr. 30, 000 Eggs found (Munter, 1915). 12 Aug. ? 500 killed as they "have been killing young terns by the hundreds" (Schlemmer and Schlemmer, ms.). 1916 9 Feb. B No nests found; 2 large roosts present on the eastern side of the island (Munter, ms.). 1918 8-10 Sept. 16,000 Estimate includes young (Diggs, ms.). 1923) 5G) Apres iz Courting birds and nests with eggs (Dickey, ms.). 8-13 Apr. a Establishing colonies (Wetmore, ms.). 29 Apr.- 1,500 Fresh eggs, recently, hatched young seen; 14 May count of 1,377 birds (Wetmore, ms.). 10 Apr.- 9 fresh eggs collected (USNM). 4 May 18 May ? Day old nestling noted (Wetmore, ms.). 1936 7-8 Mar. many thou- Only eggs present but some eggs pipped. sands nesting - Some birds had evidently not yet laid; 4 banded with brass rings (Trempe, ms.). 2 SE - 150 Just completing nesting season (Coultas, mee) Table GF-3. (continued) 167 Population Date of Survey Estimate Breeding Status, Remarks, References 1950 23 June numerous Eggs and young (POFI). 1951 12 May 2 Eggs and young (POFT). late June- 9,011 Count, including young birds (Brock, early July 1951b: 18). 1955 10 Feb. 500 In ca. 5 colonies (POFI). 1957 2 June- 1,500 Adults (Woodside, ms. b). 3 July 8-12 July ? Most young about 2 months old (La- brecque, 1957: 18). 1958 27 May- 2 From heavily incubated eggs to young 4 June a month or more in age (Warner, ms.). 1959 28 Apr.- ? Some eggs; naked and downy young about 1 May equally evident; some courting seen (Kramer, ms.). 1961 7-8 Mar. very Courting males; eggs; no young seen numerous (Woodside and Kramer, ms.). 4-10 Sept. ? Mostly well-grown young and immatures; no eggs or small young (Woodside, ms. c). 1962 14-19 June z Mostly with downy chicks (Kramer and Beardsley, ms.). 1963 11-13 Feb. 100 Nesting? (Probably fledged young) (POBSP). 3-10 Dec. ? Most young able to fly; 2 with down on neck (Walker, ms. b). 1964 10-11 Mar. 3,500- 1,200-1,500 nests, most with eggs; a 4,000 few with recently hatched young. One 2/3 grown young seen (BSFW, POBSP). 16-20 Sept. 2,000 Mostly young 3/4 grown to nearly fledged and immatures; a few young 1/2 grown or less (POBSP). 1965 6-11 Mar. 4, 000- Ca. 500 nests with eggs (POBSP). 5,000 168 Table GF-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1965 17-21 July 3,500 Ca. 1,500 nests; from recently hatched to nearly fledged young; 200 flying im- matures* (POBSP). 5-12 Aug. 5,000 Ca. 1,500 young from small, downy chicks to flying immatures (POBSP). 1966 26-31 Mar. 3,500- No count; estimate based on March 1964; 4000 very few nests with eggs; no chicks (BSFW) . 10-16, 5,000 From heavily incubated eggs to flying 20-21 June immatures;* 1,000 nests--100 with eggs; most with small to medium-sized downy chicks (POBSP). 17-18 Sept. 2 Many immatures and large young in nests i! (BSFW). \ 20-23 Oct. 1,500 Most nests with near-fledged or fledged young; no eggs or downy young seen " (POBSP). ij 1967 18-19 Mar. 2, 000 On empty nests or eggs; many males seen i displaying (BSFW, POBSP). if 7-12 June 8,000 1,177 nests counted: of 1,177 nests ex- amined 460 (39%) with eggs; 202 (17%) recently hatched young; 440 (37%) small downy young; 75 (6%) medium to large downy young. Count believed to be about 90% complete (POBSP). -a 5-11 Sept. 3, 000 No eggs; 1 small downy young; ca. 500 large young, still downy or retaining some down (POBSP). is} Wee several Many immatures; no evidence of nesting thousand (BSFW). i 1968 17-19 Mar. 1,500 175 nests counted: 54 (31%) still empty; 121 (69%) with eggs; no downy young seen; flying young present; courtship noted (POBSP). *Probably young from a preceding breeding cycle. 169 Table GF-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1969 26-29 Mar. 2, 286* Count of 1,142 nests, a few more may have been present. Of 80 nests ex- amined, 32 (40%) were empty but active and 48 (60%) contained eggs. 9 Sept. hundreds Nesting status not checked (BSFW). *Estimate is of the number of breeding birds. EMPEROR GOOSE Philacte canagica Status Accidental; one record: March 1969. Observations On 27 March 1969 Olsen and Sincock flushed four Pintails and 11 Shovelers from the edge of the southeastern corner of the lagoon. Re- maining behind was an Emperor Goose that was carefully studied through a 7 x 35 binocular from a distance of about 75 yards. It was approached to within 30 yards before it flushed, alighting again at the north end of the lagoon. The following day Kridler saw the bird on the east side of the lagoon and confirmed the identification. It was seen again on 29 March by other members of the field party. Emperor Geese are among the least common of vagrant waterfowl in the Hawaiian area. They have been recorded only four times in the main Hawaiian Islands (Oahu, Kauai, and Hawaii), and only once each from Kure, Midway, and Laysan in the Northwestern Hawaiian Islands (Clapp, Kleen, and Olsen, 1969). MALLARD Anas platyrhynchos Status Rare visitor; three records: November 1896, January-February 1913, March 1968. 170 Observations Rothschild (1893-1900: 307) reported a specimen collected 7 November 1896, apparently by one of Schauinsland's correspondents. Schauinsland (1899; 101) later listed the Mallard as a "winter guest." Bailey (1956: 79) recorded the Mallard again when he and Willett visited Laysan. He reported that a male and female “were noted in flight on January 4 [1913], and we flushed them from the little fresh water pond on the southwest end on the 11th and February 8." Extracts from Willett's (ms.) report, however, state that the pair were seen from 22 December through 9 February. Both the male (USNM 24091) and the female (USNM 24092) were collected by Bailey on the latter date. On the morning of 18 March 1968 Clapp flushed a drake from the northernmost of the two small inlets at the southwestern corner of the lagoon. It was not seen subsequently. Mallards are occasional migrants to the main Hawaiian Islands (Bryan, 1958: 9) but are decidedly uncommon visitors to the Northwestern Hawaiian Islands. LAYSAN TEAL Anas laysanensis Status Endemic. Uncommon; maximum recent estimates about 500-700 birds. Occurs throughout the island but predominately in vegetation near the lagoon. Most nesting is from late April through August. Nest is a down-lined depression under vegetation. Populations The Laysan Teal was apparently never very common and its confiding ways and dependence on the lagoon increased its vulnerability. Ducks were killed for food and sport by members of the early guano operation (Warner, 1963: 11) but were apparently protected by some island managers. Nevertheless, Fisher estimated the population at under 100 in 1902 (Table LT-3). More were killed for food during the feather raids of 1909 and in 1911 Dill and Bryan estimated a total remaining population of only six adults (7 adults and 5 young according to Bryan on p. 28). Next year Bailey counted seven birds. At this time rabbits were seriously affecting the vegetation and teal were concentrated at the small fresh water pond near the southwest corner of the lagoon, always a favorite spot. In 1915 Munter counted 13 on the fresh water pond during a quick survey of the island on 3 April and Schlemmer recorded 18 birds on 26 July. Probably others had been killed by feather hunters earlier that year. Efforts to exterminate the rabbits in 1912-1913 failed but AL A/a perhaps slowed destruction of the vegetation for a time. However, almost no vegetation remained when Wetmore arrived in April 1923 and his maximum of 20 birds seen was probably the entire remaining popu- lation. By this time the fresh water pond had filled with drifting sand and even the lagoon was filling. The few brief surveys made during the next decade, as the vegetation became re-established, indicate an almost miraculous escape from extinction. Later counts suggest a steady increase in population even though the number of birds seen remained low. By 1936 the vegetation was re- covering and Coultas (ms.) counted 11 birds during a brief survey of the island. In June 1950 Brock (195la: 371-372) counted 26 adults and two broods during a walk around the lagoon and in the following summer he counted 39 birds. These counts are probably quite conservative because most teal remain under vegetative cover during the day. In 1957 22 adults were counted on a transect census by Woodside (ms. b) and the population was estimated at between 580 and 740 birds. Similar transect censuses conducted in 1958 and 1961 gave population estimates within the range of 600 to 700 birds which is probably about the present population. Population estimates of the POBSP from 1963 to 1968 are not comparable as these are visual estimates and not census figures. Potentially valuable data from banding ratios by BSFW and HDFG personnel are not presently available. Warner's statement (1963: 14) that a level of about 600 birds is at or near the saturation level is probably correct. Habits Most observers visiting Laysan commented on the teal and most of this literature (and his observations) were compiled by Warner (1963: 3-23) in the most useful single work on the species. The following account draws heavily from his compilation. As a result of recent interest in the teal and other endangered species by BSFW personnel, the POBSP made only cursory observations on the endemic species. Teal are largely terrestrial but can fly when pressed. They usually seem reluctant to fly and most flightsare for not more than a few hundred yards. When approached, birds usually walk into vegetative cover but less often paddle out onto the lagoon or take flight. Many observers have commented that teal rarely occur on the main lagoon and almost never on the ocean. However, in February 1963 Kramer (1963) reported over 60 birds, in flocks of 8 to 12, far out on the lagoon. Birds usually occur singly, in pairs or in small groups except when concentrated near the lagoon--here flocks of up to 55 birds have been observed (Walker, 1963: ms. b). During the guano period, teal were said to be very tame and even today are very curious and can often by approached closely with a little care. Human activity, particularly the killing of adults for food, un- doubtedly did grave harm to the population until early in the 20th ee century. Adults have no known enemies on the island and destruction of eggs (by Laysan Finches) and young (by frigatebirds) was probably negligible. The limiting factor throughout much of the teal's history, as now, must have been the environment. That the teal survived the destruction of vegetation in the 1920's is remarkable as the vegetation provided both cover and food. Wetmore (ms.) noted that birds rested during the day among rocks and at night walked inland to the lagoon margin and patches of vegetation. The rocks probably provided cover from sand storms and the lagoon and vegetation provided their food supply. Warner noted that Sesuvium, so heavily utilized when it was the major remaining vegetation on Laysan, is now "completely ignored" as a food source. He also believed that a much greater dependence on littoral feeding probably occurred during the critical devegetated period. Warner suggests that larger areas of fresh water habitat were available during the evolutionary development of the species. The fresh water pond present near the southwest end of the lagoon was heavily utilized by teal until its disappearance in the 1920's. More recently ducks have concentrated at the slightly brackish water available at several points on the island following heavy rains. Although captive birds utilize fresh water for both bathing and drinking, the Laysan population survives long periods without apparent harm when neither fresh nor slightly brackish water is available. Teal occur throughout the island but most are concentrated, during the daylight hours, at the beach morning glory near the lagoon. Most spend the day resting under cover but a few can usually be seen walking around the lagoon edge, in the sedges nearby or just loafing in either area. The teal is largely nocturnal and most feeding and other activity begins at dusk. Warner (op. cit., 11) determined that "movements fol- lowed a regular pattern and indicated favored routes for feeding activities.” He determined the home range to be about two acres--much smaller during the summer molting (and flightless) period. Woodside and Kramer (ms.) state "it appears that the home range (territory?) of each pair usually contains a strip of lagoon shore." Banding returns also suggest that the birds do little moving about the island. The Laysan Teal is primarily insectivorous, at least during the summer months. Warner (p. 15) describes in detail their nocturnal feed- ing on cutworms, both from vegetation and sifted from the sand beneath, and the pursuit of brine flies along the lagoon edge. Teal also sift sand around rotting carcasses, probably for fly and beetle larvae and pupae. Early observers noted crepuscular feeding on the myriads of small moths then so prevalent on the island. Teal also puddle around the edges of small potholes near the lagoon (in 1967 on the southeast end). Warner also observed feeding on littoral invertebrates, particu- larly at the south end of the island. The nesting cycle is apparently an extended one. Pairing has been observed by early March (Woodside and Kramer, ms.) and continues until at least mid-summer. Although numerous observers report "pairing activity" 173 and Warner noted copulation, there seems to be no published report of actual courtship behavior. Warner suggests that pair bonds may be formed in September after the late-summer molt. Some observers report consider- able aggression (territorial defense?) between birds while others (e.g., Woodside and Kramer, ms.) comment on its lack. In captivity birds are very aggressive both toward other species and their own. Breese (in Warner, p. 20) reported infra-specific aggression as one of the major factors in acclimating the birds to captivity. The nest is a shallow bowl of vegetation lined with bits of down and secreted under vegetation, usually Cyperus, Chenopodium or Scaevola. Kridler (1964) reported a nest site being prepared under a bush on 10 March. The eggs are greenish-white and large for the size of the bird (55 x 38 mm., Fisher). The usual clutch size is five or six. Few nests with eggs have been reported in the wild and the earliest hatching date is 19 May (Fisher, 1903a: 799-800). Interpolation of dates based on the sizes of chicks observed and an incubation period of 27 to 28 days would indicate that most egg laying occurs during May with a span from early May through probably July. Woodside (ms. c) reported "gravid females" in early September. In captivity (Lint, 1960: 7) the drake guards the incubating female (also observed on Laysan, 30 April [Kramer, ms.]) and the chicks remain in the nest with the hen for two days. In the wild, males sometimes attend the brood. Warner commented that it was rather common to find elements of two broods being cared for by a Single bird. Introductions A report of flightless ducks on Lisianski in 1828 is not convincing and is almost certainly erroneous. Wetmore (ms.) reported that 24 or 25 were brought to Honolulu by George D. Freeth about 25> years earlier and released about 1894 by a Mr. Whitney in the Kewalo marsh in the outskirts of Honolulu. The birds were strong and flew well but none was taken since. In 1958, thirty-six adults were transported to the Honolulu Zoo for acclimatization and were ultimately donated to selected aviculturalists in the United States and Europe (Warner, op. cit.). These aviaries were generally successful in rearing teal, and - preservation of the species seems assured. In March 1968 twelve birds were introduced (from Laysan) to Southeast Island, Pearl and Hermes Reef, by BSFW personnel. Three days later at least four of these birds had left the island or were dead on the island and at present none remains. Specimens Fifty-three Laysan specimens are currently distributed in museums as indicated in Table LT-1. Hight additional mounted specimens are distributed as follows: AMNH (2 in Laysan exhibit); BPBM (2); DMNH (2 = Laysan exhibit); SUI (2: 1 in Laysan exhibit; 1 in extinct bird case). 3 174 Banding and Movements A total of 525 Laysan Teal was banded since 1958, chiefly by BSFW and HDFG personnel. No natural interisland movements are known for this sedentary species. Table LT-1. Locations of Laysan Teal skins from Laysan. Adult Adult Museum Males Females Other : Totals AMNH T 10 2 19 BPBM 2 3 9 14 MCZ 4 2 @) 6 USNM (non-POSSP) 5 T 1 13 Other* O O He ne Totals 18 22 13 53** *Carnegie Mus. (1). **Additional specimens are said to be at Berlin, Bremen, Chicago, London, Los Angeles and Paris (Greenway, 1958: 168, 169). Table LT-2. Laysan Teal banded on Laysan. Period of Adults Sex/Age survey Bander Males* Females* Ge Young Unknown Totals 1958 June BSFW © ) ) @) Ol.xx Qh 1961 Sept. HDFG*** 119 ee 0 ae 0 204 1963 Dec. HDFG 5 12 ) O @) iy 1964 Mar. BSFW 58 Tah @) 0) 0) 99 Sept. BSFW y Te ) O 0 alas 1965 Mar. POBSP 0 0 5 0 O 5 1966 Mar. BSFW 27 i 0 0 0) 34 Sept. BSFW 1 2 fe) @) at mn 1967 Sept. BSFW 3 4 © 6 O 1g 1968 Sept. BSFW 8 8 O 8 O ok Totals 225 155 25 25 95 525 *Presumably adults except those in March and September 1966 for which age is not known. **36 of these birds were transported to the Honolulu Zoo. ***¥In conjunction with the Coolidge Expedition. 1) Table LT-3. Observations of Laysan Teal on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1828 2k Mar. small flocks Not breeding (Isenbeck in Rothschild, 1893-1900: v). 1891 16-27 June not very Breeding (Rothschild, 1893-1900: 20). plentiful 1895 Sept. ? At least 1 collected by Hall (BPBM). 1896 24 June- 2 Breeding (Schauinsland, 1899: 101). 24 Sept. 1902 16-23 May <100 Eggs hatching and small ducklings 19-21 May (Fisher, 1903a: 799-800). 1903 5-28 Apr. ? At least 15 collected by W.A. Bryan (AMNH, BPBM). 1907 16-20 May ? At least 7 collected by M. Schlemmer (MCZ, AMNH). 1911 24 Apr.- ? "Flocks of up to 6" (Dill in Dill and 5 June Bryan, 1912: 23). Only 7 adults seen in 1 day; 5 young seen (Bryan, loc. cit.: 28). 3 specimens collected (SUI, UMMZ). 1912 22 Dec.- T 3 pairs and an odd male (Bailey, 1956: 80). 1913 11 Mar. Not nesting by 11 March (Willett, ms.). 1on5 3 Apr. 13 On pond (Munter, 1915: 139). 26 July 18 Counted (Schlemmer and Schlemmer, ms.). 1916 9Q Feb. 35 On pond near south end of island (Munter, ms.). 1918 8-10 Sept. quite 1 flock of at least 20; others of from numerous 2-6 birds. No nests found (Diggs, ms.). 1923 8-13 Apr. dtr Small flocks and pairs, pre-laying (Wetmore, ms.). 29 Apr.- 20 18 seen in Sesuvium patch on NE side of 14 May lagoon on 23 April ( Ball, ms.). 6 col- lected (Wetmore, ms.). 1924 6 May 2(7) Only 2 seen (Wilder, ms. a). 176 Table LT-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1928 6 May (7?) 15) Seen in 1 flock by W.G. Anderson (Wilder, ms. b). 1930 2-17 Aug. iL A single female seen. A nest with eggs punctured by a curlew found (Wilder, ms. b). 1936 7-8 Mar. 4O=-50 Most seen in 2 flocks (Trempe, ms.). 12 Dec. 11* 2 collected (Coultas, ms.). 1950 23 June 26 Also 7 downy young counted around lagoon (Brock, 195la: 371-372). 1951 late June- 39 Count (Brock, 1951b: 17). early July 1955 10 Feb. < 200 "161 counted and more known to be on nests’ (POFI; Warner, 1963: 13). 1957 25> June- 580-740 Based on census; as many as 22 adults 3 July and 13 young seen feeding on flies at one time; 10 teal seen with broods of young (1 with 6, 2 with 4, 4 with 3, 1 with 2 and 2 with 1), copulation and a nest (or nests) with eggs observed (Woodside, ms. a, b). 400-600 Based on cursory transect census (Warner, 1963: 13). 8-12 July G ",,..perhaps a dozen...together...they were no doubt much more numerous...” (Labrecque, 1957: 18). 1958 27 May- 594 Estimate based on transect census. ! 4 June Actual count of 25 on census. No nests i found; behavior "suggested that egg i laying and possibly incubation were oc- curring" (Warner, 1963: 13; ms.). l 1959 28 Apr.- 600-700 150 counted along lagoon; 2 nests found i 1 May 30 April, 1 with 6 eggs and 1 with 4 1 eggs; an estimated 90-95% paired } (Kramer, ms.). i *Bailey (1956: 80) gives the number seen as 1}. 177 Table LT-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1961 7-8 Mar. 600-700 437 counted in a circuit of the lagoon over a period of a day and a half. No young nor nests found; pairing behavior observed (Woodside and Kramer, ms.). 4-10 Sept. 688-746 Low estimate based on transect census (Warner, 1963: 13). 21 teal actually counted (Woodside, ms. c). High estimate based on banding data; 11 young. from two weeks old to full grown seen at night; an additional 1 or 2 broods of 1 young seen during day (Wood- side, ms. c). 1962 14-19 June ? 1 duckling, perhaps 2 weeks old, seen; several females apparently heavy with eggs (Kramer and Beardsley, ms.). 1963 11-13 Feb. 500 300 counted (POBSP); 145 counted by Kramer (BSFW). 3-10 Dec. a Daily counts of 103-112 (Walker, ms. b). 1964 10-11 Mar. 400-500 "Prenesting" (POBSP). Count of 235 around lagoon on 10 March (BSFW). 16-20 Sept. 400 Count of 202, a few 1/3 grown ducklings seen (POBSP). 19-20 Sept. 400 Count of 257 (BSFW). 1965 6-11 Mar. 200-300 No nests found (POBSP). 17-21 July 200 Few seen (POBSP). 5-12 Aug. 150-200 Less than 50 large ducklings (POBSP). 1966 26-31 Mar. ? 169 counted around lagoon on 27 Mar. A brood of 3 ducklings less than week old seen (BSFW). 10-16, 300 215 counted on 13 June. 4 broods of 20-21 June 2 young seen (POBSP). 17-18 Sept. ey 126 counted; census considered unsatis- factory; no broods (BSFW). 178 | Table LT-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1966 20-23 Oct. 450 137 seen at about same time (POBSP). | 1967 18-19 Mar. @ No count made (BSFW). | 7-12 June 300 No nesting recorded (POBSP). 5-11 Sept. 300-400 Young in post-juvenal molt (POBSP). 13 Dec. 2 6 seen; no census made (BSFW). 1968 17-18 Mar. several No eggs or young found (BSFW, POBSP). hundred seen 1969 26-29 Mar. 476 Estimate derived from transect census. (14 teal actually counted during cen- sus). 123 counted on beatout census around lagoon (BSFW). 85 counted around lagoon on evening of 2 June. Most ducks apparently paired; a day old dead young found (BSFW). 2-3 June Lov) en) 74 counted on beatout census along lagoon. No broods of young seen (BSFW). 9 Sept. PINTAIL Anas acuta Status Uncommon but regular visitor; at least eleven records. Observations The most abundant anatid migrant to the Hawaiian area, Pintails have been reported from Laysan in larger numbers than any other species of vagrant or migrant duck (Table P-1). This species probably occurs on Laysan almost every year. Table P-l. Observations of Pintails on Laysan. Number When Observed Seen Remarks and References 1896 (77) ? Listed by Schauinsland (1899; 101) as "winter guest;" 3 specimens taken by him or his correspondents later de- posited in Bremen Museum (Rothschild, 1893-1900: 307). iy) Table P-l. (continued) Number When Observed seen Remarks and References 1903 19 Apr. @ Hitherto unreported specimen (9, BPBM 2981), collected by W.A. Bryan is in Bishop Museum. 1912 24 Dec. 15 Seen over the ocean by Willett and Bailey (Bailey, 1956: 83). 26 Dec. 8 (Bailey, 1956: 83). 27 Dec. 12 (Bailey, 1956: 83). 1958 1 June 1 Drake on mud-flats near lagoon (Warner, 1963: 6). 1963 7 Dec. 3 Males; on west side of lagoon (Walker, mcrae 5) 1966 26-31 Mar. a Female on east side of lagoon some time during survey period (BSFW). 17-18 Sept. 3 Males. 14 other migratory ducks also observed but not identified (BSFW); these may have been some of those sub- sequently seen by a POBSP survey team. 21 Oct. 19 On pond at north end of lagoon (POBSP). 1968 18 Mar. 3 Drake and hen seen under Scaevola with a few Laysan Teal near the southwestern corner of lagoon. 2 hens subsequently flushed from this area (POBSP). 1969 27 Mar. 5 Seen at southeast corner of lagoon (BSFW). 9 Sept. 63 Censused in central lagoon (BSFW). COMMON TEAL Anas crecca (GREEN-WINGED TEAL) Status Accidental; two records: October 1896; March 1969. 180 Observations The Common Teal was reported as a "winter guest" on Laysan by Schauinsland (1899: 101). Rothschild (1893-1900: 307) reported a female specimen collected by one of Schauinsland's correspondents on 27 October 1896. He listed this bird as Nettion carolinense [= A. crecca carolinensis] because A. c. crecca had not been taken on the Pacific coast of America. Bailey followed him by listing the record as A. c. carolinensis. Recently, on 27 March 1969, Kridler (pers. corrs) saw three other female teal on the lagoon. Since females of the two subspecies are very difficult to distin- guish, we believe that no subspecific designation should be attached to the specimen until it has been re-examined. Teal, either crecca or carolinensis, have also been reported from Midway Atoll in the Northwestern Hawaiian Islands (Clapp and Woodward, 1968: 12-13) and are occasional migrants to the main Hawaiian Islands (Udvardy, 196la: 84). [GARGANEY TEAL Anas querquedula] Status Hypothetical; one unconfirmed record. Observations Schauinsland (1899: 101) reported this species as a "winter guest" on Laysan but Rothschild (1893-1900: 307) later stated that he thought the bird seen by Schauinsland was "more likely" a Blue-winged Teal (Anas discors). Rothschild stated (1893-1900; 308) that, with the exception of two shorebirds, he had examined the specimens of the fifteen species (including the Garganey Teal) added to the Laysan list by Schauinsland. This clearly suggests that he had seen a specimen of this duck. Checklists of the Hawaiian area (Henshaw, 1902; Bryan and Greenway, 1944; Bailey, 1956; Bryan, 1958; and Udvardy, 196la make no mention of this record. Since Garganey Teal recently were reported from Midway Atoll (Clapp and Woodward, 1968: 12) and, since Blue-winged Teal have not been reported from the Northwestern Hawaiian Islands, we feel that Schauinsland's original identification could well have been correct. AMERICAN WIDGEON Mareca americana (BALDPATE ) Status Rare visitors; two records: October 1896; March 1964. 181 Observations The only specimen record from Laysan is an immature female col- lected 15 October 1896 and subsequently reported by Rothschild (1893- 1900: 307). On 10 March 1964 Walker (BSFW) saw two drake American Widgeons standing on a sandbar on the east side of the lagoon. This species has also been reported from Midway in the Northwestern Hawaiian Islands (Clapp and Woodward, 1968: 13) and is a regular visitor in small numbers to the main Hawaiian Islands (Udvardy, 1961: 85). SHOVELER Spatula clypeata Status Uncommon visitor; at least seven records. Observations Shovelers have been observed more frequently on Laysan than any species of vagrant duck but the Pintail. This is not surprising since these two species of ducks are the most abundant of those that regularly migrate to the main Hawaiian Islands; thousands of Shovelers and Pintails were reported from there by Medeiros (1958: 111). Table Sh-l summarizes known records of Shovelers from Laysan Island. Table Sh-1. Observations of Shovelers on Laysan. Number When Observed seen Remarks and References 1896 Oct.-Nov. ? Rothschild (1893-1900: 275, 307) lists records from October, November 1897 Jan. ? and January apparently based on specimens secured by Schauinsland or his correspondents; "winter guest” (Schauinsland, 1899: 101). 1913 30 Jan. 6 Flock seen by Willett (Bailey, 1956: 84). 1916 Nov. or Dec. 1 Shot by Eric Schlemmer who identified bird by its broad bill (Wetmore, ms.). 1961 7 or 8 Mar. 2 Pair on north shore of lagoon (Woodside and Kramer, ms.). 1963 7 Dec. fe) 5 males, 4 females on west side of lagoon (Walker, ms. b). 182 Table Sh-1. (continued) Number When Observed seen Remarks and References 1966 22 Oct. 3 On small pond north of lagoon (Clapp and Woodward, 1968: 14; POBSP). 1969 22 Mar. 14 At southeast corner of lagoon (BSFW). BUF FLEHEAD Bucephala albeola Status Rare visitor; two records; January 1897, December 1912-January 1913. Observations Schauinsland (1899: 101) recorded this species as a "winter guest" on Laysan. Rothschild (1893-1900: 307) reported a juvenile male col- lected 15 January 1897, apparently by one of Schauinsland's correspondents. Bailey and Willett saw a female Bufflehead over the beach on 2/ December 1912. On 3 January 1913 Willett collected a female (USNM 240989), probably the same bird, on a fresh water pond (Bailey, 1956: 84). HARLEQUIN DUCK Histrionicus histrionicus Status Accidental; one record: April 1906. Observations An immature female (BPBM 4538) taken by Bompke (Bailey, 1956: 84) on 17 April 1906 is the only record for any of the Hawaiian Islands. LAYSAN RAIL Porzanula palmeri Status Endemic. Formerly an abundant permanent resident. Extinct, probably since 1923. Nested in bunch grass and especially in the Cyperus fringe around the lagoon. Nesting apparently occurred from April through July. Populations The Laysan Rail, both abundant and conspicuous, was recorded by even the casual observer. General descriptions by Isenbeck in 1828 and 183 by a Captain Wood in 1872 were referred to this species by Rothschild (1893-1900: 10). Palmer (in Rothschild, 1893-1900: vii) purchased several live birds from a ship captain before his voyage up the Leeward Chain and the number of old, unlabelled specimens in the Bernice P. Bishop Museum suggests that others were captured as curiosities. Frohawk (1892: 247) used live birds shipped to Europe in his formal description of the species. Although all visitors reported the species, only two (prior to Wetmore) gave population estimates (Table LR-2). The rail was still “abundant” during the winter of 1912-1913 although rabbits were then making serious inroads on the vegetation. Two years earlier Dill and Bryan had estimated a total population of 2,000 birds. Munter estimated a population of 5,000 in 1915 and said they were "everywhere" on the island. The population probably dropped drastically as the vegetation disappeared. Wetmore found only one or two mummies from 8 to 13 April 1923 but reported that two singles were seen at different parts of the island during his temporary absence (14 to 28 April). He introduced eight birds from Midway on 29 April and later found several mummies believed to be from this stock. Subsequent to his introduction the party saw one or two birds at intervals at two points on the island. The species probably became extinct soon after this visit--at any rate the species was never again seen on Laysan and with the destruction of the Midway population in June 1944 the species vanished forever. A re-introduction to Laysan after the vegetation became re-established might well have saved the species. Laysan Rails were introduced to other islands on several occasions: to Midway 13 July 1891 (Munro, 1947: 24), and in 1904 [19057] (Wilder, ms. b), 14 March 1913 (Bailey, 1956: 89); to Lisianski in March 1913 (Bailey, op. cit.); from Midway to Pearl and Hermes Reef in 1929 (Baldwin, 1947: 14); from Midway to Laysan in 1923. However, only on Midway did rails become well established. The introduction to Lisianski and re- introduction to Laysan occurred after massive destruction of the vege- tation by rabbits; the Pearl and Hermes Reef population was probably destroyed by storms during the first winter; the Midway population survived until rats were introduced to the islands. Suggestions that rails were introduced on islets off Oahu are without documentation (Munro, op. cit.). Habits Paul H. Baldwin compiled nearly all available information on the Laysan Rail in a popular account (Baldwin, 1945) and later (Baldwin, 1947) in a scientific paper. The latter paper included both published material and data from personal conversations with several men who had observed the rail in life, either on Laysan or on Midway. The most informative original account is that of Fisher (1903a: 800-802). The following summary is from several sources but draws most heavily from Baldwin (1947). 184 The Laysan Rail was conspicuous and very active, running rapidly between bits of cover, creeping through vegetation or darting in and out of burrows. On Laysan, rails were most active during the morning and evening but on occasion were equally active throughout the day. Though flightless the wings were used for balancing when running, jump- ing, or fighting. Rails were inquisitive and comparatively fearless, readily entering buildings for such tidbits as meat scraps from the skinning table. Palmer (in Rothschild, 1893-1900: 10) comments on this inquisitiveness as follows: "I could always catch them by placing my net edgeways on an open space of ground, for they would immediately run up to see what it was.” Rails were virtually omnivorous, eating a wide variety of arthro- pods, eggs of seabirds (which they on occasion broke themselves), and bits of flesh from carcasses. Fresh water was used for bathing and drinking when available. Birds were quite vocal, calling intermittently when foraging and occasionally giving a "rattle" which seemed to be a territorial call. Frohawk (1892: 248) described a dusk chorus by several birds and lasting but a few seconds which sounded like a handful of marbles thrown onto a glass roof and then descending in a succession of bounds. Rails occurred over the entire vegetated area of the island. The preferred nesting area was the ring of "Juncus" (actually Cyperus) sur- rounding the lagoon but bunch grass was heavily utilized as well. Nest building began in April and nests were present into July. Nests were placed on the ground. Those in Cyperus were often a hollowed-out, re- arranged accumulation of dried stems; those in grass tussocks were bulkier masses of dried grasses and leaves, lined with shredded stems and young albatross down. The nature of the vegetative cover (especially in Cyperus) often "roofed" the nest. Eggs were recorded during May and June. The usual clutch was three, less frequently two. Eggs were pale olive buff with faint varied mark- ings of "pale clay color or raw sienna and faint lilac gray" (Fisher, 1903a: 801). On Laysan most chicks hatched in June. The downy chicks (all black with yellow bill and black legs and feet) were very noisy and probably stayed with the parents for a month. They soon learned to feed themselves and in five days could run as fast as their parents. Specimens One-hundred thirty-two Laysan Rail skins are currently distributed in museums as indicated in Table LR-l. Eight additional mounted speci- mens are distributed as follows: BPBM (1 male, 1 female); DMNH (2 males and 1 female in Laysan exhibit); SUI (3 birds in Laysan exhibit). Also preserved are at least 9 skeletons (1 a partial) at USNM; 3 nests (BPBM, 2; USNM, 1); and at least 12 eggs (BPBM, 6; USNM, 6)., 185 Table LR-l. Locations of Laysan Rail skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 18 11 16 45 BPBM 9 14 3 26 CMNH 2 2 O 4 MCZ al 2 O 3 UMMZ O O 16 16 USNM 21 14 0 35 Other** 2 aL O 3 Totals 53 Ly 35 132%%* *Includes 1 transplant from Midway. **Law Coll. (10, 19 [missing]); Acad. Nat. Sci. Phila. (1c). ***According to Greenway (1958: 235), additional specimens are at Berlin and London. Table LR-2. Observations of Laysan Rails on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1828 2k Mar. ? An inaccurate description by Isenbeck (in Kittlitz, 1834: 124) was applied to this species by Rothschild (1893- 1900: v). 1872 ? Finsch (in Hartlaub, 1893: 400) re- ported species on basis of description by Capt. Walter Edward Wood (in Roth- schild, 1893-1900: 10). 1891 16-27 June ? "Very plentiful," all over the island; 2 nests (Rothschild, 1893-1900: 10). 1896 24 June- 2 Present, apparently numerous (Schau- 2h Sept. insland, 1899: 42-43). 1 1902 16-23 May en) "Great numbers;" everywhere; many nests with fresh eggs; young apparent- ly begin to hatch in middle of June (Fisher, 1903a: 800-802). 23 May 4 clutches of eggs (1 partly incubated) and 2 nests collected (USNM). 186 Table LR-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1903 15-25 Apr. ? At least 19 birds and nest and egg taken by W.A. Bryan (AMNH, BPBM). 1904 10 May 2 2 collected by Schlemmer (MCZ). 1905 19 Sept. ? Mentioned in report (Wilder, 1905: 393). 1911 24 Apr.- 2, 000 "Fairly abundant;" nests (but no 5 June eggs); 2 chicks on 4 June (Dill and Bryans 12.20). 1912 22 Dec.- 2 "Abundant" all over island, about 1913 11 Mar. 100 transported to other islands (Bailey, 1956: 84-90). Not nesting by 11 March (Willett, ms.). At least 45 collected (CMNH, JEL, MCZ, UMMZ, USNM). IONS e503) Apr. 5,000 "Everywhere on the island" (Munter, 1915: 139). 1916 9 Feb. ? "Seen everywhere" (Munter, ms.). 1918 8-10 Sept. ? Often seen (Diggs, ms.). 1923 8-13 Apr. - 1 or 2 mummies found (Wetmore, ms.). 14 Apr. - 1 seen east of lagoon by Reno (Ball, ms.). 16 Apr. - Rail mummy found by Reno (Dickey, ms.). 18 Apr. 2 Two rails known to be alive, one seen in guano piles, the other in Sesuvium (Dickey, ms.; Wetmore, ms.). 23-28 Apr. 0? Storm with heavy winds, driving sand (Dickey, ms.). 29 Apr. 8 8 introduced frem Midway (Wetmore, ms.). 12 May 2 Two seen. 13 May 1 One seen. 187 SEMLPALMATED PLOVER Charadrius semipalmatus Status Accidental; one record: September 1967. Observations Ely saw a Semipalmated Plover late in the afternoon of 5 September 1967 as it fed with Ruddy Turnstones on the mudflats on the west shore of the lagoon. The following day Clapp saw this bird feeding by itself around the margins of the small pond north of the lagoon. He collected it later in the day as it fed along the muddy northwestern margin of the lagoon. The specimen (USNM 543043), an immature female, was not molting and had heavy fat deposits. It constitutes the first record from Laysan. These plovers are of fairly regular occurrence in the main Hawaiian Islands but have been recorded only rarely from the Northwestern Hawaiian Islands (Midway, Lisianski, Laysan). GOLDEN PLOVER Pluvialis dominica Status Abundant migrant; maximum recent estimate: 1,200. Probably present all year; recorded for all months for which data are available; most numerous in February, March, and September. Occurs both singly and in small to large flocks. Found over most of the island but most common along the lagoon shore. Populations and Annual Cycle Golden Plovers are most common during the spring and fall when large migrating flocks occur. The extent of the migration period is unknown but in 1966 very few were seen after 10 June when 233 were counted (Table GP-3). Our data suggest migration peaks in March and September. Populations are probably lowest during mid-summer (mid-June to early August) and moderate during the winter season. The length of time migrants stay on the island and the amount of daily population turnover are not known. Ecology Plovers utilize the area between the beach and lagoon to a much greater degree than do any other shorebirds, with the possible exception of the Bristle-thighed Curlew. Many birds occur singly in small open areas scattered throughout the vegetation and many set up and defend "winter" territories. Large flocks are sometimes present, such as the flock of 750 to 1,000 birds seen in a large open area near the south end 188 of the island in mid-September 1964. Golden Plovers occasionally flock with turnstones, tattlers, and curlews, but at least in September 1964 and 1967 most of them occurred in pure flocks. When feeding in the lagoon, plovers wade further from shore than do turnstones. Wetmore (ms.) saw them eating bird eggs and also noted that many were killed by sandstorms and alkali poisoning in April and May 1923. Specimens Fifty-one Golden Plover skins from Laysan are currently distributed in museums as indicated in Table GP-1. Two additional mounted specimens are in the Laysan exhibit at SUL. Banding and Movements Various workers banded 188 Golden Plovers on Laysan (Table GP-2). One banded on 9 March 1965 was recaptured there by Eugene Kridler on 17 September 1966. q Table GP-1. Locations of Golden Plover skins from Laysan. Museum Males Females Other Totals | AMNH 6 4 6) 10 BPBM 1 al 1 3 CMNH at al 0 2 DMNH i 0 ©) a SUI il fo) ih 2 | UMMZ, 10 8 0 18 USNM (non-POBSP) 6 al ) (POBSP) 4 mn 0 5 Other* 2 i @) 3 Totals 32 17 2 51 *Law Coll. (10, 19); Museo Parense (10). Table GP-2. Golden Plovers banded on Laysan. | j j i Period of Survey Bander Number Banded 1958 June BSFW a i 1961 Sept. HDFG 7a } 1963 Feb. POBSP 5 Dec. HDFG 17 189 Table GP-2. (continued) Period of Survey Bander Number Banded 1964 Mar. BSFW 23 Sept. POBSP h 1965 Mar. POBSP 4 1966 Mar. BSFW 29 Sept. BSFW 8 1967 Mar. BSFW \ Sept. POBSP 10 BSFW iL 1968 Mar. : POBSP TEL Total 188 Table GP-3. Observations of Golden Plovers on Laysan. Population Date of Survey Estimate Remarks and References 1890 16 July ? (Lyons, 1890: 91). 1891 16-27 June 2 Frequently seen (Rothschild, 1893- 1900: 11). 1896 2k June- 2 Regular visitor (Schauinslana, ou Sept. 1899: 101). 1902 16-23 May common Flock near lagoon; all in winter plumage (Fisher, 1903a: 803). 1903 29 Apr. ? 2 collected by Bryan (BPBM). 1911 24 Apr.- 2,000 (Dill and Bryan, 1912: 21). 5 June 1912 22 Dec.- flocks of 100 (Bailey, 1956: 91). 1913 11 Mar. or more 1915 19 Sept. 2 Eric Schlemmer killed 14 "plovers" with one shot (Schlemmer and Schlemmer, ms.). 1916 9 Feb. common (Munter, ms.). 1918 8-10 Sept. great Seen in large flocks along lagoon numbers (Diggs, ms.). 1923 8-13 Apr. ? Fairly common (Wetmore, ms.). 190 Table GP-3. (continued) Population Date of Survey Estimate Remarks and References 1923 29 Apr.- several Some in full breeding plumage and 14 May hundred others in winter dress (Wetmore, ms.). 1930 2-18 Aug. % Beginning to arrive (Wilder, ms. b). 1936 7-8 Mar. 2 Several flocks of 50-100 seen feeding. along lagoon (Trempe, ms.). 1950 23 June numerous (POFT). 1955 10 Feb. 50-100 Common along lagoon (POFI). 1957 25 June- 20 (Woodside, ms. b). 3 July 1958 27 May- uncommon Several in winter plumage were ap- 4 June parently residents in the area (Warner, ms.). 1959 28 Apr.- ? Several (Kramer, ms.). 1 May 1961 7-8 Mar. 300 Over entire island (Woodside and Kramer, ms.). 4-10 Sept. OO 71 banded (Woodside, ms. c). 1962 14-19 June ? Common throughout island (Kramer and Beardsley, ms.). 1963 11-13 Feb. 1, 000 Found in all habitats (POBSP). 3-10 Dec. 2 Count of 462; 10 (2%) on outer beaches; 452 (98%) in the interior (Walker, ms. b). 1964 10-11 Mar. 800- Count of 583: 183 (37%) on outer 1,000 beaches; 400 (63%) in the interior; many beginning to change into breed- ing plumage (BSFW, POBSP). 16-20 Sept. 1, 200 One flock of ca. 1,000 birds on bare island area (BSFW, POBSP). 1965 6-11 Mar. 1, 000 Entire island (POBSP). 17-21 July 50 Most around lagoon (POBSP). Reed 191 Table GP-3. (continued) Population Date of Survey Estimate Remarks and References 1965 5-12 Aug. 450 1 flock of 300 birds (POBSP). 1966 26-31 Mar. 1, 000 850 seen, primarily in lagoon; some molting into breeding plumage (BSFW). 10-16, 250 Count of 233: 158 (68%) on outer 20-21 June beaches, 75 (32%) in the interior; most had left by 11 June; many in breeding plumage (POBSP). 17-18 Sept. 900 Most using lagoon; 600 in one huge flock (BSFW). 20-23 Oct. 600 Count of 477: 27 (6%) on outer beaches; 450 (94%) along the lagoon; all in winter plumage (POBSP). 1967 18-19 Mar. 1, 200 635 counted, 560 at the northwest end of the lagoon and 75 around the beaches; concentrated around lagoon; about 1/4 in obvious prenuptial molt (BSFW, POBSP). 7-1le June 300 Over 200 birds, most in breeding plumage by lagoon (POBSP). 5-11 Sept. 1, 200 Primarily on mud flats, west side of lagoon (POBSP). 13 Dec. % 1 flock of 300; all in winter plumage (BSFW). 1968 17-19 Mar. 500 Count of 204 around lagoon in morn- ing (36 on north and east; 68 on south; 20 on southwest; 80 on north- west). Flock of 179 feeding in northwest corner in late afternoon (BSFW, POBSP). 1969 26-29 Mar. 990 Along lagoon (BSFW). 9 Sept. large Seen feeding along lagoon (BSFW). numbers 192 BLACK=-BELLIED PLOVER Squatarola squatarola Status Accidental; one record: December 1912-January 1913. } 7 q } % 4 i. 7 an Observations The only record of Black-bellied Plovers on Laysan was made by Bailey (1956: 90) who noted that two were seen along the lagoon on 28 December 1912,* and that one (USNM 240985), a male, was collected by Willett. The other, or another, was seen subsequently on 10 January 1913. ee a RUDDY TURNSTONE Arenaria interpres Status Abundant migrant; maximum recent estimate: 5,000. At least a few birds recorded during all months for which data are available; most numerous from February through April and in September. Occurs in most habitats but most common along the lagoon shore. Populations and Annual Cycle The Ruddy Turnstone is the most abundant shorebird on Laysan at all seasons, but for brief periods it may be equaled or exceeded in numbers by the Golden Plover. Peak populations occur from at least December through May and in September (Table RT-3); lowest numbers occur during the summer when most turnstones are on the northern nesting grounds. Daily population variations, possibly due to migration, were noted in April 1923, June 1966, and September 1967. The data are too scant to determine accurately migration periods, migration peaks, or the amount of population turnover on the island. Ecology Turnstones occur in a variety of habitats: tidal pools along the rocky beaches, sandy beaches, open areas in the upland part of the island, and along the shore of the lagoon. On four counts--September 1961, December 1963, March 1964, and September 1967--they were con- siderably more abundant along the shore of the lagoon than on the outer beaches. However, on various trips turnstone concentrations were found along different areas of the lagoon shore, probably because the wave action that apparently concentrates suspended materials, including food items, varies with wind conditions. In September 1964 and March 1965 birds were most common along the eastern shore; in October 1966 along *Extracts from Willett's report (ms.) state that the plovers were ac- tually first seen 27 December. 193 the northern shore; and in September 1967 and March 1968 from the northwest to southwest corners of the lagoon. The lagoon is probably the best feeding area on the island because of the concentration of fly larvae and brine shrimp. Dill and Bryan (1912: 21) found turnstones in the shallow water of the lagoon and about the fresh-water pond feeding on small black flies. In April and May 1923 Wetmore (ms.) found them feeding on Gray-backed Tern eggs, meat and fat scraps, flies, brine shrimp, and mollusks on coral that had been placed on the beach to dry. In September 1967 turn- stones were noted wading up to their breasts in the lagoon. At night turnstones roost along the lagoon, in the vegetation on the slopes of the island, and on the rock wall at the south end of the island. In September 1964 an increase in the number of birds along the outer beach in the evening was noted. Wetmore (ms.) reported many dead and dying turnstones with paralysis of the nictitating membrane and lack of control of feet, wings, and neck suggesting alkali poisoning. No similar mortality has been reported recently. Specimens Fifty-three Ruddy Turnstone skins are currently distributed in museums as indicated in Table RI-1. Four additional mounted specimens are distributed as follows: BPBM (1 female); MCZ (1 male, 1 female); SUI (1 bird in Laysan exhibit). Banding and Movements The POBSP and BSFW banded 79 Ruddy Turnstones on Laysan (Table RT-2). A bird with a broken wing, banded on Laysan in March 1965, was recaptured there on 10 June 1966; it was unable to fly but healthy. On 6 September 1967, three adults, 1103-00277, 1103-01393, and 1103-02152, banded on St. George Island, Alaska, on 3, 9, and 17 August 1967, respectively, were collected along the lagoon. Three additional turnstones color-marked on St. George were seen but not collected. 652-45015, of unknown age, banded on Laysan on 17 September 1964, was recaptured on St. George Island’in the Pribilofs, Alaska, on 18 August 1968. Table RI-1. Locations of Ruddy Turnstone skins from Laysan. Museum Males Females Other Totals AMNH 4 y () 8 BPBM ) eee (0) mn 194 ee Table RT-1. (continued) Museum Males Females Other Totals CMNH a 1 0) 2 MCZ 2 2 O 4 UMMZ 4 2 ©) 6 SUL O 2 ) 2 USNM (non-POBSP) 5 9 O 14 (POBSP) 4 6 iL aa Other* 1 1 O 2 Totals al 31 dL 53 caliecy) (Ofonlal, (Gl ey, al &))-, Table RT-2. Ruddy Turnstones banded on Laysan. Period of Survey Bander Number Banded 1964 Mar. BSFW 7 Sept. POBSP ho 1965 Mar. POBSP 7 1966 Mar. BSFW 5 Sept. BSFW 1 Oct. POBSP 1 1967 Sept. POBSP 1a, 1968 Mar. POBSP 7 Total 79 Table RT-3. Observations of Ruddy Turnstones on Laysan. Population Date of Survey Estimate Remarks and References 1891 16-27 June common (Rothschild, 1893-1900: 13). 1896 2h June- z "Regular visitor" (Schauinsland, 24 Sept. 1899: 101). 1902 16-23 May abundant Concentrated near lagoon. Flocks seen (Fisher, 1903a: 803). 1907 15 Oct. 2 6 collected by Schlemmer (MCZ). 1911 24 Apr.- 2,500 Large flocks of plovers and turnstones, 5 June especially along lagoon (Dill and Bryan, 1912: 22). Table RT-3. (continued) Population Date of Survey Estimate Remarks and References 1912 22 Dec.- common Seen daily (Bailey, 1956: 92). 1913 11 Mar. 1915 3 Apr. 5,000 Flock on lagoon shore (Munter, 1915: 140). 1916 9 Feb. common (Munter, ms.). 1918 8-10 Sept. great Seen in flocks along lagoon (Diggs, numbers ms.). 1923 8-13 Apr. common Considerable daily variation in num- bers due to migration. Many molting into breeding plumage (Wetmore, ms.). 29 Apr.- 3, 000- (Wetmore, ms.). 14 May 4,000 1930 2-18 Aug. great (Wilder, ms. b). numbers 1936 7-8 Mar. thousands Flocks of 75-100 feeding along lagoon (Trempe, ms.). 12 Dec. 3, 000- Along lagoon edge (Coultas, ms.). 5,000 1950 23 June numerous (POFT). 1951 12 May ? (POFT). 1955 10 Feb. ? Presence not noted but reference to "Sanderlings" probably referred to this species (POFI). 1957 25 June- 150 (Woodside, ms. b). 3 July 8-12 July ? Flocks of a dozen or so (Labrecque, NOs, 1S) . 1958 27 May- 2 Flocks of 10-20 seen commonly (Warner, 4 June Meee 1959 28 Apr.- small (Kramer, ms.). 1 May numbers 195 196 Table RT-3. (continued) Population Date of Survey Estimate Remarks and References 1961 7-8 Mar. 500-700 Feeding along lagoon (Woodside and Kramer, ms.). 4-10 Sept. 1,500 Count of 71 (5%) along the beaches ; i and 1,400 (95%) along the lagoon : (Walker, ms. a). ; 1962 14-19 June large (Kramer and Beardsley, ms.). numbers ; 1963 11-13 Feb. 5,000 3,000 counted around lagoon (POBSP). 3-10 Dec. 3,000 Count of 2,768: 427 (15%) on the outer beach and 2,341 (85%) along the lagoon (Walker, ms. b). 1964 10-11 Mar. 2,000 Count of 1,777: 189 (11%) on the outer beaches and 1,588 (89%) along lagoon (BSFW, POBSP). 16-20 Sept. 1,600 Most along east shore of lagoon (BSFW, POBSP). 1965 6-11 Mar. 3, 000- Most along lagoon (POBSP). 5,000 17-21 July 250 Most along lagoon (POBSP). 5-12 Aug. 4.00 1 flock of 400 birds (POBSP). 1966 26-31 Mar. 2,450 Some molting into breeding plumage (BSFW). 10-16, 500 Count of 493: 85 (17%) on outer beach, 20-21 June 408 (83%) in the interior; ca. 30% in full breeding plumage; most departed soon after 10 June (POBSP). 17-18 Sept. 2,000 An estimated 90% along lagoon (BSFW). 20-23 Oct. 550 A number in breeding plumage (POBSP). 1967 18-19 Mar. 800 638 counted (250 along beaches, re- mainder in northwest corner of lagoon (BSFW, POBSP). 7-12 June 300 No large flocks (POBSP). 197 Table RT-3. (continued) Population Date of Survey Estimate Remarks and References 1967 5-11 Sept. 5,000 Concentrated around lagoon (POBSP). 13) Weeq several Most on east shore of lagoon (BSFW). thousand 1968 17-19 Mar. 3,000 Count of 1,839 around lagoon on the morning of the 18th; 246 on north and east shores, 246 on south shore, 475 on southwest half of west shore, and 872 on northwest half. Considerably more. abundant on west shore in late afternoon (BSFW, POBSP). 1969 26-29 Mar. 1,580 Seen along lagoon (BSFW). 9 Sept. several . Feeding with lesser numbers of Golden thousand Plovers along lagoon shoreline. [WHIMBREL Numenius phaeopus | Status Hypothetical; one unconfirmed record: September 1961. Observations Woodside's report (ms. c) states that "One whimbrel was identified by Dr. Udvardy." We have been unable to obtain confirmation of this Sighting and so treat its occurrence on Laysan as hypothetical. The only other records for the Hawaiian area are two sight records from Midway Atoll (Udvardy, 1961a: 86). BRISTLE-THIGHED CURLEW Numenius tahitiensis Status Common migrant; maximum recent estimate: 150-200. Probably occurs in all months but most numerous during March, April, and September. Usually occurs singly or in small flocks. Occurs throughout the island in all habitats. Populations and Annual Cycle Probably small numbers of Bristle-thighed Curlews occur on Laysan throughout the year (Table BTC-3), with the largest populations (several Fe ne : hundreds) present during the March to April and the September migration i periods. The number of birds spending the winter or summer on Laysan { is unknown. ; : i) Ecology ? Curlews utilize all major habitats on Laysan and are common both : throughout the interior and along the rocky shores, mainly on the south beach. In September 1964, August 1965, and March and September 1967 the only concentrations were found along the rocky shores. This is probably the only shorebird on Laysan that is as common away from the lagoon as near it. Curlews usually occur singly or in small groups, but flocks of up to 48 individuals have been seen, usually near the south end of the island. The egg-eating habits of this species are well known. Bailey (1956: gi) found them opening albatross eggs that had been deserted and Wetmore (ms.) found them feeding on Gray-backed Tern, Red-footed Booby, and Great Frigatebird eggs. Curlews regularly patrolled the tern colonies for eggs. A curlew would empale an egg with its beak, then run away to devour it at its leisure. Wetmore also watched a curlew fly to an un- attended Red-footed Booby nest in a low bush and remove the egg. Another removed an egg from a frigatebird which momentarily raised up but did not leave the nest. In this instance the egg was carried off unbroken. He also reported that they ate fat that was discarded during bird skinning operations. Wilder (ms. b) recorded curlews eating Brown Noddy eggs and avidly devouring near-hatched embryos. Walker (ms. a) observed a curlew feeding on a recently killed Laysan Finch. Other foods include Scaevola berries and insects. Curlews also feed along the lagoon, in tide pools and among the inland vegetation. Dill and Bryan (1912: 21) reported that as many as 20 birds roosted on the roofs of the old buildings. Scattered individuals have been found roosting in the interior at night and on 10 September 1967 a flock of 20 birds was found roosting on the south beach. Specimens Seventy-one Bristle-thighed Curlew skins from Laysan are currently distributed in museums as indicated in Table BIC-1. Additional mounted specimens are distributed as follows: DMNH (1 male, 1 female in Laysan exhibit); MCZ (2 males); SUI (1 in Laysan exhibit); BPBM (1, possibly 3). Banding and Movements The POBSP and BSFW banded 50 Bristle-thighed Curlews on Laysan (see Table BIC-2). An adult, 645-12620, banded 31 May 1967 by the POBSP on Southeast Island, Pearl and Hermes Reef, was recaptured by the BSFW on Laysan on 27 September 1967. 199 Table BTC-1. Locations of Bristle-thighed Curlew skins from Laysan. Museum Males Females Other Totals AMNH 5 " 0 12 BPBM 3 3 a t CMNH 3 4 ©) "i DMNH 3 1 0) h MCZ it 5 @) 12 SUL 2 ) ) aL UMMZ 3 3 aL T USNM (non-POBSP) 8 8 0) 16 Other* 2 3 ) 5 Totals 35 3h 2 71 *Law Coll. (1c, 19); Leningrad (1 9); Coryndon Mus. (10, 1 9). Table BIC-2. Bristle-thighed Curlews banded on Laysan. Period of Survey Bander Number Banded 1958 June BSFW 2 1963 Feb. POBSP 2 1964 Mar. BSFW 3 Sept. POBSP 10 BSFW i 1965 Mar. POBSP 10 1966 Mar. BSFW 3 1967 Sept. POBSP 13 1968 Sept. BSFW 6 Total 50 Table BIC-3. Observations of Bristle-thighed Curlews on Laysan. Population Date of Survey Estimate Remarks and References 1890 16 July ? (Lyons, 1890: 91). 1891 16-27 June ? (Rothschild, 1893-1900: 18). 1895 Sept. ? At least 3 collected by Hall (AMNH, BPBM). 1896 24 June- ? Regular visitor (Schauinsland, 1899: 24 Sept. onl) |p 200 Table BIC-3. (continued) Population Date of Survey Estimate Remarks and References 1902 16-23 May 2 Small flocks (Fisher, 1903a: 803). 1903 1-20 Apr. ? 3 collected by Bryan (AMNH, BPBM). 1904 17 May, 2 1 specimen collected each date (MCZ). 6 Sept. 1906 26-27 Apr. ? 7 collected by Schlemmer (MCZ). 1907 18-22 May ? 6 collected by Schlemmer (MCZ). 1911 24 Apr.- 250 Along lagoon and beaches (Dill and 5 June Bryan, 1912: 21). 1912 22 Dec.- common Flocks of up to a dozen birds (Bailey, 1913 11 Mar. 1956: 94). TOT Sep Ts. 1,000 Scattered over island (Munter, 1915: 140). 1916 9 Feb. common (Munter, ms.). 1918 8-10 Sept. 1,000 On all parts of island (Diggs, ms.). 1923 8-13 Apr. common (Wetmore, ms.). 29 Apr.- 30 (Wetmore, ms.). 14 May 1930 2-18 Aug. ? (Wilder, ms. b). 1936 7-8 Mar. 2 Many groups of from 2-3 to 25-30 birds found mostly in grass and along shore of lagoon; 2 banded with blue celluloid rings (Trempe, 1936). 12 Dec. 150-200 (Coultas, ms.). 1950 23 June a few (POFT). 1951 12 May ? (POFI). late June- 50 Diurnal census (Brock, 1951b: 18)¢ early July 1955 10 Feb. ? Ca. 100 along the northeast shore (POFI). OTTO Gm Table BIC-3. (continued) Population Date of Survey Estimate Remarks and References 1957 25 June- 50 (Woodside, ms. b). 3 July 8-12 July 2 Flocks of up to 25 birds (Labrecque, 1957: 18). 1958 27 May- 15 Most at south end (Warner, ms.). 4 June 1959 28 Apr.- small (Kramer, ms.). 1 May numbers 1961 7-8 Mar. 75-100 (Woodside and Kramer, ms.). 4-10 Sept. 400 Count of 140: 132 (94%) on outer beaches; 8 (6%) in the interior (Walker, ms. a). 1962 14-19 June very (Kramer and Beardsley, ms.). common 1963 11-13 Feb. 20 (POBSP). 3-10 Dec. 90 Count of 75: 23 (31%) on outer beaches; 52 (69%) in the interior (Walker, ms. b). 1964 10-11 Mar. 150-200 Count of 73: 12 (16%) on outer beach; 61 (84%) along lagoon (BSFW, POBSP). 16-20 Sept. 100 Over entire island (POBSP). 1965 6-11 Mar. 100-200 (POBSP). 17-21 July 25 Scattered along beaches (POBSP). 5-12 Aug. 70 Flock of 48 (POBSP). 1966 26-31 Mar. 150 18 on outer beach (BSFW). 10-16, 60 Count of 51: 18 (35%) on outer beaches 20-21 June and 33 (65%) in the interior (POBSP). 17-18 Sept. ? 15 seen (BSFW). 20-23 Oct. 50 Count of 41: 31 (74%) on outer beaches; 201 10 (25%) near the lagoon (POBSP). 202 Table BTC-3. (continued) Population Date of Survey Estimate Remarks and References 1967 18-19 Mar. . 60-120 Flock of 48 (BSFW, POBSP). 7-12 June 20 (POBSP). 5-11 Sept. 100 Flock of 43 (POBSP). 13 Dec. 100 (BSFW). 1968 17-19 Mar. 100 Count of 41 around edges of lagoon on the morning of the 18th: 26 along north and east edges, 4 along south- western half of the west side, 7 along northwestern side, 4 along south end (BSFW, POBSP). 1969 26-29 Mar. 63 Counted along lagoon shoreline (BSFW). WANDERING TATTLER Heteroscelus incanum Status Common migrant; maximum recent estimate: 200-500. Probably present in all months; most numerous during February, March, August, and September. Usually occurs singly or in small flocks; most common around the lagoon and on the rocky beaches; rarely inland in vegetated areas. Populations and Annual Cycle All recent counts and estimates (Table WI-2) indicate that the tat- tler ranks behind the Golden Plover and Ruddy Turnstone in numbers at all seasons. Since tattlers show less flocking tendency than the other two species and utilize the rocky shoreline to a greater extent than do other shorebirds, counts were probably often low. The generally higher counts made recently are probably the result of more complete censuses rather than an increase in population. As with other shorebirds, peak populations are present in the spring and fall. Ecology Wandering Tattlers frequent the shores of the lagoon and rocky areas around the ocean beaches, particularly those at the southern end of the island. Unlike plovers and turnstones, tattlers seldom occur inland on the more-vegetated portions of the island but are usually quite common along the shores. On only one of five counts (October 1966) were they found to be more common on the outer beaches than around the lagoon. 203 Wandering Tattlers occasionally occur in turnstone and plover flocks. The POBSP recorded a large flock of 150 on the rocky southeast shore on 11 August 1965. Pure flocks of as many as 50 to 75 birds are occasionally seen (October 1966; March 1968). Fisher (1903a: 802) saw them wading in the lagoon feeding on flies and possibly brine shrimp. Wetmore (ms.) indicated that they caught fish on the reef. Specimens Twenty Wandering Tattler skins from Laysan are currently distributed in museums as indicated in Table WI-1. An additional mounted bird is in the Laysan exhibit at SUI. Also preserved is a skeleton at the USNM. Banding and Movements Ten Wandering Tattlers were banded on Laysan through 1969: 9 by the POBSP (5 in September 1964, 1 in September 1967, and 3 in March 1968); and 1 by the BSFW (September 1964). No interisland movements were recorded. Table WI-l. Locations of Wandering Tattler skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 3 5 O 8 BPBM O 2 O SUL aL O O ald UMMZ, O O 5 D USNM (non-POBSP) 3 O 0 3 (POBSP) iy ) o) i Totals 8 7 5 20 Table WI-2. Observations of Wandering Tattlers on Laysan. Population Date of Survey Estimate Remarks and References 1828 2h Mar. ? Probably present. Vague description by Isenbeck subsequently identified as this species by Rothschild (1893-1900: v) could apply to several species. 1891 16-27 June few (Rothschild, 1893-1900: 15). 1895 Sept. % Specimen which from label was probably collected by Hall (BPBM). 204 Table WI-2. (continued) Population Date of Survey Estimate Remarks and References 1896 24 June- ? "Regular visitor" (Schauinsland, 1899: 24 Sept. 101). 1902 16-23 May ? Least common of migrants. Few seen | each day. 1 in breeding plumage col- 3 lected 18 May (Fisher, 1903a: 802). 1911 24 Apr.- very Usually on the reefs or on large rocks 5 June few on the beach (Dill and Bryan, 1912: 21). 1912 22 Dec.- z Regularly seen; 7 collected (Bailey, 1913 11 Mar. 1956: 95). Very common around lagoon and on beaches (Willett, ms.). 1916 9 Feb. common (Munter, ms.). 1918 8-10 Sept. aL Only 1 observed (Diggs, ms.). 1923 8-13 Apr. 2 Single birds (Wetmore, ms.). 29 Apr.- 50 Flocks of 10-20 seen; along rocky shore 14 May and lagoon (Wetmore, ms.). 1930 2-18 Aug. a few (Wilder, ms. b). seen 1Q36 3 2 Deer 100 (Coultas, ms.). 1957 25 June- 20 (Woodside, ms. b). 3 July 1958 27 May- 5 Very uncommon and wary (Warner, ms.). 4 June 1961 7-8 Mar. 150 Most in lagoon; few on outer beaches (Woodside and Kramer, ms.). 4-10 Sept. 253 Count. 38 (15%) on outer beach and 215 (85%) along lagoon (Woodside, ms. c). 1962 14-19 June hundreds Least common of shore birds (Kramer and Beardsley, ms.). 1963 11-13 Feb. 100 All habitats (POBSP). 3-10 Dec. 59 Count. 22 (37%) on outer beach and 37 (63%) in the interior (Walker, ms. b). 205 Table WI-2. (continued) Date of Survey Estimate Remarks and References 1964 10-11 Mar. 125-150 Count of 120: 40 (33%) on outer beach and 80 (67%)}-along lagoon. One flock of 35 seen (BSFW, POBSP). 16-20 Sept. 200 West shore of lagoon; 132 counted (POBSP). 19-20 Sept. 200 (BSFW, POBSP). 1965 6-11 Mar. 200-500 Lagoon and rocky beaches (POBSP). 17-21 July 0 None seen (POBSP). 5-12 Aug. 350 Flock of 150; lagoon and rocky shores (POBSP). 1966 26-31 Mar. 4 (BSFW). 10-16, 20 Count of 12 scattered birds: 4 (334%) 20-21 June on outer beach, 8 (67%) in the interior (POBSP). 17-18 Sept. 7 (BSFW). 20-23 Oct. 150 Count of 147: 91 (62%) on outer beaches, 56 (35%) along lagoon. Flocks of 50 and 75. Nearly all birds in winter plumage (POBSP). 1967 18-19 Mar. 20 Most near lagoon (BSFW, POBSP). 7-12 June 20 Estimate (POBSP). 5-11 Sept. 250 Most common along the west shore of lagoon; small numbers in rocky areas of southwest shore (POBSP). 1968 17-19 Mar. 200 Count of 65 avound lagoon in morning (22 on north and east shores; 7 on south; 12 on southwest; 24 on northwest). Ca. 40- 50 feeding in mixed flock with plovers at northwest corner in afternoon (BSFW, POBSP). 1969 26-29 Mar. 135 Seen along shoreline (BSFW). 9 Sept. ? some seen feeding along lagoon shore (BSFW). 206 @ GREATER YELLOWLEGS Totanus melanoleucus Status Accidental; one record: October 1966. Observations : POBSP personnel collected one of two Greater Yellowlegs seen feeding with a group of Wandering Tattlers in the lagoon on 21 October 1966. The specimen (USNM 496780), a very fat immature female, constitutes the first record from Laysan and the first specimen from the Hawaiian area. The only other record from the Northwestern Hawaiian Islands is the sighting of a single individual on Midway in June 1941 (Donaghho, 1953- 1954: 49); at least seven sight records have been reported from the main Hawaiian Islands (Clapp and Woodward, 1968: 18). LESSER YELLOWLEGS Totanus flavipes Status _ Rare visitor; three records: October 1966, September 1967, March 1968. Observations On 21 October 1966 POBSP personnel saw a Lesser Yellowlegs feeding with two Greater Yellowlegs and a number of Wandering Tattlers. This bird was closely compared with the Greater Yellowlegs (Clapp and Woodward, 1968: 18). It called several times, giving the distinctive di-syllabic note of this species. On 5 September 1967 Clapp saw a Lesser Yellowlegs at dusk as it roosted among Wandering Tattlers at the northwestern corner of the lagoon. He collected two immature females the following day. One (USNM 543047) was shot as it stood in the shallow muddy pond north of the lagoon. The other (USNM 543048) was shot at the northwestern corner of the lagoon. Both yellowlegs were roosting alone when collected and both were less wary than the other species of shorebirds that usually frequent the island. USNM 543047 was molting lightly on the head and breast and had moderate fat deposits when collected. The following spring, on 18 March 1968, Kridler (pers. corr.) saw another Lesser Yellowlegs among plovers and turnstones on the west shore of the lagoon. The only other atolls in the northwestern chain of islands from which this species was reported previously are Midway and Kure (Clapp and Wood- ward, op. cit.; Clapp, 1968: 76-77). Sight records of Lesser Yellowlegs 207 have been reported frequently from the main Hawaiians and they are apparently regular but uncommon visitors to the Hawaiian area. SHARP-TATLED SANDPIPER Erolia acuminata Status Uncommon visitor; three records: 1896, October 1966, March 1969. Probably more common than the records indicate. Observations Two specimens were collected on Laysan, one by Schauinsland or his correspondents in 1896 (Rothschild, 1893-1900: 307), the other by POBSP personnel, 21 October 1966 (Clapp and Woodward, 1968: 24). The POBSP specimen (USNM 496697) is an immature male taken from a flock of sand- pipers also thought to be of this species (see Sandpiper spp. below). In addition, Kridler (pers. comm.) saw one or possibly two Sharp- tailed Sandpipers feeding along the edge of the lagoon on 27 March 1969. PECTORAL SANDPIPER Erolia melanotos Status Accidental; one valid record: September 1967. Observations The only definite record of this species from Laysan is two specimens collected 6 September 1967. Clapp collected an immature male (USNM 543046) as it roosted on the sandy margin south of the north pond; Ely collected an immature female (USNM 544003) at the southwestern end of the lagoon. The only other Laysan record is a doubtful sighting in February 1963 (see Sandpiper ssp. below). Recent records of this species from Midway and Kure Atolls (Clapp and Woodward, 1968: 23) suggest that this species may occur regularly on many of the low Northwestern Hawaiian Islands. BATRD'S SANDPIPER Erolia bairdii Status Accidental; one record: September 1967. Observations Clapp collected two Baird's Sandpipers 6 September 1967 as they foraged at the northwestern corner of the lagoon. These birds usually 208 fed apart from the flocks of plovers and turnstones but when flushed occasionally flew with flocks of turnstones. On several occasions they fed and flew with a Semipalmated Plover. Both sandpipers (USNM 543044, 543045) proved to be immature females. Neither was in molt and both had little fat. The specimens mentioned above and one collected on Oahu constitute the only certain records for this species for the Hawaiian area (Wood- ward and Clapp, 1969: 25). DUNLIN Erolia alpina Status Rare visitor; three records: 1896 or 1897, January 1913, March 1968. Observations Schauinsland (1899: 101) first reported the Dunlin from Laysan (calling it Tringa americana) as a "winter guest." It is not known whether he or his correspondents only saw these birds or if specimens were taken. Bailey (1956: 96) collected (USNM 240987) a female, the only Dunlin he saw on the island, on 20 January 1913 along the shore of the lagoon where he found Golden Plovers. On 18 March 1968 Clapp saw a Dunlin feeding with turnstones in a small pool near the southwest corner of the lagoon. He collected the bird the following day as it fed along the west shoreline of the lagoon. The specimen (USNM 543337) was a very fat male in heavy body molt. Dunlins have been reported from three other Northwestern Hawaiian Islands: Kure, Midway, and Pearl and Hermes Reef (Kenyon and Rice, 1957: 33; Clapp and Woodward, 1968: 24). They apparently occur there more regularly than early records indicate. SANDPIPER Spp. Observations On seyen occasions unidentified, or inadequately identified, small sandpipers were seen on Laysan (Table Ss-1). Some of these records were thought to be of Pectoral, Sharp-tailed, or Least Sandpipers; one (September 1967) record was probably of a fourth species. The December 1963 record of a "Least Sandpiper" is included here because it was not accompanied by either detailed notes or a specimen. Table Ss-l. Population 209 Observations of small Sandpiper Spp. on Laysan. Date of Survey Estimate Remarks and References 1961 1963 1965 1966 1967 7-8 Mar. 4-10 Sept e 11-13 Feb. 3-10 Dec. 17-21 July 20-23 Oct. 5-11 Sept. z ca. 70 Ca. 30 "Small sandpiper type birds may have been seen but [were] not identified." (Woodside and Kramer, ms.). Walker's (ms. a) notes 5 Sept. state that a small sandpiper-like bird was seen near the small pond just inside the lagoon shore. "It had a dark cap, light bill with a dark tip, light legs --pale yellow, light belly (white), about least sandpiper size, dark brown stripe down back, upper tail, light on either side." The observer thought that the bird might have been a Pectoral Sandpiper (POBSP). On 8 December, Walker's (ms. b) notes state that "Nixon Wilson saw a least sandpiper yesterday." No other details are given. Thought to have been Sharp-tailed Sand- pipers. The birds were seen ina compact flock along the shore of the lagoon (POBSP). Thought to have been Sharp-tailed Sand- pipers but observers disagreed on the identification. A Sharp-tailed Sand- piper was collected (POBSP). An unidentified "peep" was seen on the morning of 8 Sept.; two were seen the following day, and one on 10 Sept. All were apparently of the same species and no more than two individuals were probably involved. The birds were brownish above, white below with a faint wing stripe and with white showing to either side of the base of the tail. They appeared to be less than half the size of Ruddy Turnstones (POBSP). 210 DOWITCHER Sp. Limnodromus sp. Status Accidental; one record: September 1967. Observations Kridler (pers. comm.) observed a dowitcher along the shoreline of the lagoon on the morning of 24 September 1967. ‘The bird was observed from within 15 feet for over 15 minutes and when it flushed the white back was clearly seen. More than 15 sightings of dowitchers have been recorded from the main Hawaiian Islands, but only two from the Northwestern Hawaiian Islands (Clapp and Woodward, 1968: 20-21). None had been reported pre- viously from Laysan. MARBLED GODWIT Limosa fedoa Status Accidental; one record: October 1966. Observations Marbled Godwits were first recorded on Laysan when POBSP personnel observed two feeding in the northern half of the lagoon on 21 October 1966. One of these, an immature male with heavy fat deposits (USNM 496790), was collected the same day. This specimen is not only the first record from Laysan, but also the first record for the entire Hawaiian area (Clapp and Woodward, 1968: 17). BAR-TAILED GODWIT Limosa lapponica Status Rare visitor; four records: November 1896, September 1964, March 1965, October 1966. Probably more common than the number of records suggests. Observations The preponderance of fall records (Table BIG-1) of Bar-tailed Godwits, despite the greater amount of observation time in the spring, indicates that this species visits Laysan primarily in the fall. A similar preponderance of fall records from the Phoenix Islands (Clapp and Sibley, 1967: 124) suggests that it is primarily a fall migrant throughout most of the central Pacific. 211 Since all recent specimens of Bar-tailed Godwits from the north- western Hawaiian chain have been L. 1. baueri (Clapp and Woodward, 1968: 17; Fisher, 1960: 480), it seems likely that those seen on Laysan were of the same race. Table BITG-1. Observations of Bar-tailed Godwits on Laysan. Number Date Observed seen Remarks and References 1896 5 Nov. 1 Male specimen in the Tring Museum (Rothschild, 1893-1900: 307). 11 Nov. 1 Male specimen (AMNH 738526) formerly in the Bremen Museum. Ca. 1896 ? Several specimens (including the one above) were collected and deposited in the Bremen Museum (Rothschild, op. cit.); these godwits were listed as a winter guest by Schauinsland (1899: 101). 1964 19 Sept. il Seen on the west shore of the lagoon (POBSP). 1965 6-11 Mar. 2 Seen foraging together on the south and east beaches (POBSP). 1966 21 Oct. 1 Sight record (POBSP). SANDERLING Crocethia alba Status Uncommon migrant; maximum recent estimate: 30 to 35. Recorded from February to May, from August to October, and during December; most numerous during March and September. Occurs singly or in small flocks, mainly along the lagoon edge. Populations and Annual Cycle Sanderlings have been reported on most recent visits to Laysan and are apparently more numerous in March and September (Table S-1). A few birds probably occur throughout the winter but apparently there is no summering population. Ecology Sanderlings occur most commonly in sandy areas along the lagoon. In December 1963, however, Walker (ms. b) found them most abundant on the outer beaches. 212 They occur singly or in flocks of up to eight birds, occasionally in flocks of Golden Plovers or Ruddy Turnstones. Specimens We know of only one specimen from Laysan, a male (USNM 240988) taken by Willett on 29 January 1913. Table S-l. Observations of Sanderlings on Laysan. Population Date of Survey Estimate Remarks and References 1896 2h June- 2 "Winter guest" (Schauinsland, 1899: 101). oh Sept. 1911 24 Apr.- ? (Dill and Bryan, 1912). 5 June 1912- 22 Dec. & Several noted 29 and 30 January; 1 col- 1973, 12 Meir’: lected from a flock of 5 (Bailey, 1956: 97). 1923 8-13 Apr. 2 Maximum of a dozen at one time (Wetmore, THSey ie 29 Apr.- 2 A few along lagoon (Wetmore, Mic) is 14 May 1951 12 May ? (POFT). 1955 10 Feb. - Presence noted but context of notes suggests that Ruddy Turnstones were meant (POFI). 1959 28 Apr.- i (Kramer, ms.). 1 May 1961 7-8 Mar. 30 (Woodside and Kramer, 1961). 4-10 Sept. 30 Count; all along lagoon (Walker, ms. a). 1963 11-13 Feb. 6 Along lagoon (BSFW, POBSP). 3-10 Dec. 8 Count of 7 on outer beach; 1 along lagoon (Walker, ms. b). 1964 10-11 Mar. 30-35 Count of 27: 1 (4%) on outer beach, 26 (96%) along lagoon (BSFW, POBSP). 16-20 Sept. 8 With Golden Plovers (BSFW, POBSP). 213 Table S-l. (continued) Population Date of Survey Estimate Remarks and References 1965 6-11 Mar. 15+ 15 on 7 March, increased thereafter (POBSP). 17-21 July O (POBSP). 5-12 Aug. 3 Seen on 9 August; west side of island (POBSP). 1966 10-16, 20-21 June 0 (POBSP). 20-23 Oct. 2 In winter plumage (POBSP). 1967 18-19 Mar. 9-12 Around lagoon (BSFW, POBSP). 7-12 June 0) (POBSP). 5-11 Sept. 5 (POBSP). 1968 17-19 Mar. 15 Around lagoon; none seen on beaches (BSFW, POBSP). 1969 26-29 Mar. al Along lagoon (BSFW). RED PHALAROPE Phalaropus fulicarius Status Rare visitor; four records: 1896 or 1897, January-February 1913, February 1963, October 1966. - Observations The Red Phalarope was first reported from Laysan by Schauinsland (1899: 101) who listed the species as a "winter guest." It is not known whether this record is based on a sighting or a specimen. Bailey (1956: 97) saw several of these phalaropes along the shallow waters of the lagoon on 6 January 1913; he saw two on 9 January, and several the next day when a female was collected (USNM 240986). A second female was collected by Willett on 13 January (CMNH 188914). All were in winter plumage. Willett (ms.) states that these phalaropes were frequently noted in flocks of plovers and turnstones until the middle of February. Recently POBSP personnel twice saw Red Phalaropes on Laysan in the central lagoon; two on 11 February 1963 and one on 22 October 1966. 214 NORTHERN PHALAROPE Lobipes lobatus Status Rare visitor; two records: March 1965, December 1967. Observations Northern Phalaropes were first recorded from Laysan when Clapp shot a male and a female (USNM 494118, 494119) on 7 March 1965 as they fed near large flocks of Golden Plovers, Ruddy Turnstones, and Wandering Tattlers on the south shore of the lagoon (Clapp and Woodward, 1968: 25). Both were very fat and in winter plumage. On 13 December 1967 Northern Phalaropes were again recorded from Laysan when Kridler (BSFW) saw two, both in winter plumage, "in excellent light for several minutes with field glasses at a distance of 20 to 25 feet." [GLAUCOUS-WINGED GULL Larus glaucescens ] Status Hypothetical; one unconfirmed record. Observations A gull collected by one of Schauinsland's correspondents (in 1896 or 1897) and initially identified as a Glaucous Gull (Larus hyperboreus) by Schauinsland (1899: 101) was reidentified as probably a young female glaucescens by Rothschild (1893-1900: 307). Three additional sight records are attributed to this species or to Herring or Glaucous Gulls. On 6 December 1963, Kramer (ms.) saw a gull, almost as large as an albatross, that was "gray all over with black and white flecks.” It was feeding in flooded Cyperus mats near the south end of the lagoon. He found the same gull feeding along the lagoon shore the following day and recorded it as either a Glaucous-winged or a Herring Gull. A second gull, tentatively identified as an immature Glaucous-winged Gull by POBSP personnel, was seen off the west beach on 6 March 1965. It was first seen when it alighted on the water near a group of Black-footed Albatrosses that had gathered to feed on garbage thrown over the stern of the support vessel. It flew away before any attempt to collect it could be made and was not seen on Laysan during the subsequent survey of the island. Two other gulls, believed to be adult Glaucous (Larus hyperboreus ) or Glaucous-winged Gulls, were seen near the south point of the island 2i15) on 10 March 1965 but were so wary that they could not be approached closely enough either for accurate identification or for collection. Since we have not seen Schauinsland's specimen and since all sight records are inadequate for certain identification, we prefer to regard the occurrence of Glaucous-winged Gulls on Laysan as hypothetical. GLAUCOUS GULL Larus hyperboreus Status Accidental; one record: January 1906. Observations Bailey (1956: 98) reported that a specimen (BPBM 4546) collected by Bompke on 29 January 1906 was in the Bernice P. Bishop Museum, Honolulu. Bailey thought the specimen might prove to belong to the race L. h. burrovianus. (See also Glaucous-winged Gull above.) HERRING GULL Larus argentatus Status Rare visitor; three records: April 1906, January 1913, February 1963. Observations Bailey (1956: 97) reported two specimens of Herring Gulls taken on Laysan prior to POBSP investigations. A female, collected by Bompke on 20 April 1906 and deposited in the Bishop Museum (BPBM 4539), was sub- sequently identified as Larus argentatus smithsonianus. Bailey remarked, however, that E.H. Bryan, Jr. had informed him that this bird might prove to be L. a. vegae upon further critical comparison. The second specimen (USNM 240940), an immature female, was collected along the shore of the lagoon by Bailey on 25 January 1913. It was later identified by Dr. Herbert Friedmann as L. a. vegae. The third specimen (USNM 493352) was collected on 12 February 1963 by POBSP personnel (Clapp and Woodward, 1968: 26). This bird, a female in first nuptial plumage, was subséquently identified as L. a. vegae by Mrs. Roxie C. Laybourne and Dr. Lester L. Short, Jr. of the Bureau of Sport Fisheries and Wildlife. BONAPARTE'S GULL Larus philadelphia Status Accidental; one record: December 1912. 216 Observations Bailey (1956: 98) collected an immature female (USNM 240984) on 27 December 1912 as it hovered low over the water along the shore of the lagoon. | . a This is the only confirmed occurrence of Bonaparte's Gull in the Northwestern Hawaiian Islands. A single individual was recorded (with- out comment) from Sand Island, Midway Atoll, 29 January 1963 by POBSP personnel (Sibley and McFarlane, 1968: 318). BLACK-LEGGED KITTIWAKE Rissa tridactylus Status Accidental; one record: winter 1906. Observations Bryan and Greenway (1944: 118) reported this species from Laysan on the basis of fragments in the Bishop Museum. Wetmore, in his field notes of 1923, stated that the remains (BPBM 4537), consisting of wings, feet, tarsi, and bill, had been found on the beach by Bompke during the winter of 1906. He also noted that the wings were in partial molt. SOOTY TERN Sterna fuscata Status Abundant breeder; maximum recent estimate: 2,000,000. Usually present from mid-February through mid-October. Absent except for occa- sional wandering birds during remainder of year. Most nesting is from early April to early September. Nests in shallow scrapes in the soil over most of the island, exclusive of the beaches and lagoon shore, but chiefly in the Eragrostis association of the island interior. Populations The Sooty Tern is presently the most abundant bird nesting on Laysan. Peak populations occur in summer but are difficult to estimate because of the immense numbers and the large area covered. Most, if not all, esti- mates (Table ST-3) are conservative. Recent estimates are generally higher than figures given earlier in the century for probably two reasons-- (1) these earlier visits were usually made early in the season before population maximums had been reached; (2) recent estimates usually in- clude birds using the island but which may be absent at the actual time of the count (e.g., absent mates of incubating birds). Schauinsland's and Dill and Bryan's estimates for 1896 and 1911 indicate huge populations, probably not greatly different from present real numbers. The latter estimated 333,900 birds, with the colony still increasing in size. Fisher's description of the area covered by the breeding colony in 1902, which is similar to that used today, implies a population of at least several hundred thousand birds. Estimates by Willett and Bailey for 1913, Munter for 1915, and Wetmore for 1923 were too early in the breeding season to be directly comparable with either the earlier or the more recent figures. Diggs, who made the September 1918 estimate which seems very large for that time of year, was not a biologist and his estimate is probably little more than a guess. Therefore, it is not possible to determine accurately the effects on this species caused by denudation of the island by rabbits or by the depredations of the feather poachers who took very large numbers of Sooty Terns. Annual Cycle The timing of the Sooty Tern breeding cycle on Laysan may vary more than a month from year to year. A new breeding cycle begins with the appearance of a swirl, or swirls, of birds offshore at night. Later these swirls move over the island at night but remain offshore during the day. As the swirls grow larger, birds begin landing on the island at night and stay over and around the island longer during the day. At about this time a few birds lay, but usually abandon their eggs during the day when the flock moves off. Not until large numbers lay do the birds remain on their eggs. The first groups to lay and incubate form loose nuclei around and among which later arrivals expand the colony. Members of sub-groups within the population usually lay in close syn- chrony, but the sub-groups found on different parts of the island may vary by several weeks in their stage of breeding. These sub-groups are not clearly defined but form a nearly continuous colony, both in space and time. On Laysan the first birds probably return to the vicinity of the island in January (one observation in 1913), or possibly even in December (1963), and begin building up substantial numbers by the end of February. Some birds were settling on the ground in February 1963. All March ob- servers found birds swirling over the island, and most of them reported at least a few birds on the ground. The earliest known egg laying occurred in 1959, when Kramer (ms.) found small chicks 28 April to 1 May. The eggs from which these chicks hatched must have been laid during the last days of March or the first week of April. Munter's (1915) observations indicate egg laying by at least the first week of April. Eggs may be laid as late as late June (1964, 1967) or even mid-July (1965), but in most years the peak period of egg laying falls between mid- or late May and mid-June. Hatching begins as early as late April (1959) and continues to at least mid-August (1965) with the peak period usually from mid- or late June to early July. Young may begin to fly by late June (1959) or more often July, and most of them have fledged by the end of August or the 218 first week of September. The last chicks probably fledge in late September or October when nearly all the adults and young have left the island, leaving behind a few crippled and emaciated chicks which probably die within a few weeks. No November observations are available but Sooty Terns were reported on two of three December visits. Between 3 and 10 December 1963 they were common at night, possibly indicating the beginning of the pre-breeding swirls. A single adult seen in December 1967 was probably a traveling or feeding bird that was attracted to the island briefly as it passed through the area. Eeology Breeding: Sooty Terns nest over nearly the entire surface of the island except the open sand beaches, the open lagoon shore, and dense shrubby vegetation. Fisher (1903a: 779) described "a great colony which extends along the upper half of the interior slope completely around the island, with only a few interruptions, and are thus found almost entirely among the bushy grass; on the west side the community extends nearly to the low bluff overlooking the sea." In 1923 Wetmore mentioned colonies "in the Sesuvium" and "at the tobacco patch." These birds probably were attracted to any vegetation available at this time when most of the island was barren sand. The nesting area used in recent years resembled that described by Fisher. In June 1966 nesting occurred continuously around the island except for a small gap between the north end of the lagoon and the broad north beach. On the west and south sides nearly all birds were in the Eragrostis association, while some of those on the east side were in Ipomoea cover, and a few were found under Pluchea. Heaviest concentra- tions of birds occurred on the northwest, northeast, and southwest corners of the island, and a huge swirl of birds not yet nesting was present over the southeast corner when the party left the island on 21 June. Eggs are simply deposited on the sand, usually near or under a grass clump or other protective vegetation. Non-breeding: Birds not occupied with eggs or chicks are usually found with the breeders. Late in the nesting cycle large numbers of these unemployed birds come to the island in company of those returning to feed the remaining chicks. Birds without eggs roosted on open sand on the east side of the island in June 1966 and September 1967 (flocks of 500+), but they were never more than several yards from nesting birds. Specimens Ninety Sooty Tern skins from Laysan are currently distributed in various museums as indicated in Table ST-l1. Seventeen additional mounted 219 specimens are distributed as follows: BPBM (1 male); DMNH (2 in Bahamas exhibit); SUI (14 in Laysan exhibit). Also preserved are at least 2 skeletons (USNM); 1 alcoholic (BPBM); and 72 eggs (BPBM, 37; USNM, 35). Banding and Movements The POBSP banded 130,425 Sooty Terns on Laysan (Table ST-2). Forty- eight birds banded on Laysan were recovered elsewhere as follows: Johnston Atoll, 12; French Frigate Shoals, 4; Lisianski, 20; Midway, 1; Kure, 4; Japan, 1; Philippines, 1; Marshal Islands, 1; Phoenix Islands, 1; Line Islands, 1; Atafu, 1; at sea in Western Pacific, 1 (Appendix Table 4-8a). Forty birds banded elsewhere were recaptured on Laysan as follows: Johnston Atoll, 23; French Frigate Shoals, 3; Lisianski, 10; Midway, 2; Kure, 2 (Appendix Table 4-8b). The high number of recaptures involving Johnston Atoll is undoubtedly because most of them were color-marked individuals. Table ST-1. Locations of Sooty Tern skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 5 5 7 iy BPBM als 1 3 5) CMNH Al il O 2 DMNH 4 4 2 10 MCZ 2 2 O y SUL 2 alas O 13 UMMZ, db 2 O 3 USNM (non-POBSP) ay 13 2 32 (POBSP) 3 1 0 4 Totals 36 LO 1h 90 Table ST-2. Sooty Terns banded on Laysan by the POBSP. Period of Survey Adults Young Totals 1964 Sept. 900 1,000 1,900 1965 July 9,186 2,814 12, 000 Aug. 30, 900 9,100 40, 000 1966 June 69, 900 0 69, 900 1967 June 5,000 0 5,000 Sept. 1,425 200 1,625 Totals pb ese 13, 114 130,425 220 Table ST-3. Observations of Sooty Terns on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1890 16 July ? Dense population with eggs and small chicks (Lyons, 1890: 90-91). 1891 16-27 June % Nesting (Rothschild, 1893-1900: 40). Eggs present (Munro, 1930: 687). 1895 Sept. 2 4 adults collected by Hall (AMNH, BPBM). 1896 2h June- 10, 000- "On the verge of selecting their nesting 24 Sept. 100, 000 places [24 June]" (Schauinsland, 1899: 48). 1902 16-23 May ? Evidently very abundant; fresh eggs (Fisher, 1903a: 780). 22 May ? 5 fresh eggs collected (USNM). 1903 Apr. ? 1 specimen and 36 eggs collected by Bryan (BPBM). 1907 16 May 2 4 collected by Schlemmer (MCZ). 1911 24 Apr.- 333,900 Laying dates not given, but up to 7 5 June to 9 eggs per square yard present by 4 June; on 23 April 2 colonies of ca. 500 each; thousands appeared ca. 6 May and increased daily; final estimate made from area x density measurements 4 June (Dill and Bryan, 1912: 13). 1912 22 Dec.- % Several seen first on 7 January; ar- 1913 11 Mar. riving in great numbers by 28 February; no colonies established by 11 March (Bailey, 1956: 102). 1915 3 Apr. 10, 000 Beginning to lay in east-central portion of the island (Munter, 1915: 139). 1918 8-10 Sept. 500, 000 Nesting; most common species (Diggs, ms.). 1923 8-13 Apr. rg Large flock circling east of lagoon (Wetmore, ms.) 2/7 Apr. ? Laying begun; perhaps 20 eggs laid by this date (Dickey, ms.). * 221 Table ST-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1923 28 Apr. 2 Terns flooded out of colony (Dickey, TMS ie 29 Apr.- 12, 000 Egg-laying and fresh eggs (Wetmore, 14 May Msi) fe 4-8 May ? 30 fresh eggs collected (USNM). 1930 2-18 Aug. most abundant Young about 3/4 grown (Wilder, ms.). species 1936 7-8 Mar. 2 Evidently no nests found (Trempe, ms.). 1950 23 June abundant Eggs and young (POFT). 1951 12 May 2 Eggs (POFI). late June- 115, 800 Based on count of birds present on early July island (Brock, 1951b: 18). 1955 10 Feb. 2 (POFL). 1957 25 June- 400, 000 (Woodside, ms. b). 3 July 8-12 July ? Heavy concentrations; hatching eggs and some older young (Labrecque, 1957: Ie 1958 27 May- t Thousands of eggs; those near the center 4 June of the colony about 1 week incubated, those near the periphery fresh; more being laid (Warner, ms.). 1959 28 Apr.- Egg laying to small downy young; most 1 May with eggs (Kramer, ms.). 1961 7-8 Mar. ? Not nesting; "very numerous" in air, a few on the ground (Woodside and Kramer, msie)!< 4-10 Sept. ? Most colonies with flying immatures; no eggs or small young observed; great in- crease in numbers at night (Woodside, Msis, Cyl. 222 Table ST-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1962 14-19 June 1963 11-13 Feb. 3-10 Bee. 1964 10-11 Mar. 16-20 Sept. 1965 6-11 Mar. 17-21 July 5-12 Aug. 1966 26-31 Mar. 10-16, 20-21 June abundant Some colonies almost entirely on eggs; others almost entirely with downy young; others not nesting (Kramer and Beardsley, ms.). 1, 000 Not nesting; mostly overhead; some birds settling (BSFW, POBSP). ? Not nesting; common in air at night, but none seen on ground; none seen during the day (Walker, ms. b). 300, 000- Pre-breeding; 3,000 to 4,000 on the 600, 000 ground by day (BSFW, POBSP). 50, 000- Ca. 25,000 flying immatures; a small 75,000 number of still unfledged young (BSFW, POBSP). 7, 000= 2 separate breeding colonies forming; 12, 000 largest (5,000-10,000) on ground for short periods during day; smaller group of a few thousand circling and spending little time on ground (POBSP). 450, 000 Ca. 50,000 young; stage of sub-colonies varied from heavily incubated eggs to a few fully feathered chicks, none of which could fly; many areas had no eggs or chicks (POBSP). 1, 000, 000 Also 200,000 young from downy to flying immatures. Ca. 200 heavily incubated eggs near north end of lagoon; many areas with adults only; restless mil- ling behavior at dusk (POBSP). 250, 000 Mostly over the island; several hundred on ground at night; no eggs or young (BSFW). 2, 000, 000 Also 500,000 nests with eggs; stages of cycle ranged from birds just settling on the ground to slightly incubated eggs (POBSP). 223 Table ST-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1966 17-18 Sept. 20, 000- Primarily present at dusk; some emaci- 30, 000 ated (apparently deserted) almost full- grown chicks and several hundred dead chicks (BSFW). 20-23 Oct. 100 Few crippled immatures and a few others flying with adults (POBSP). 1967 18-19 Mar. 4,000 Small swirls; ca. 200 on the ground at night (BSFW, POBSP). 7-12 June 1,543, 000 All stages from birds settling only at night to newly hatched young; estimated 257,000 eggs, thought to be about 1/4 of total to be laid (POBSP). 5-11 Sept. 60, 000 Several thousand young from 3/4 grown to fully fledged; most birds evidently post-breeding (POBSP). 13 Dec. a 1 adult heard flying overhead (BSFW). 1968 17-19 Mar. 7,500 Not more than several hundred present in 3 mornings; several thousands swirling over island in late afternoon and night; one flock of ca. 200 on ground at night (BSFW, POBSP). 1969 26-29 Mar. 50,000 Birds swirling; no signs of nesting (BSFW). 9 Sept. thousands Only adults seen; evidently post-nesting (BSFW). GRAY-BACKED TERN Sterna lunata Status Common breeder; maximum recent estimate: [40,000]. Present from late December or early January through mid-September; absent remainder of year. Most nesting is from March through early August. Nests on ground in semi-open areas near the outer beaches and near the central lagoon. Populations No population estimates were made until 1911, when Dill and Bryan estimated 50,000 birds from 24 April to about 5 June 1911 (Table GBT-2). 22h On 3 April 1915, Munter estimated 5,000, and Wetmore found only about 1,000 between 29 April and 14 May 1923. These figures indicate that the number of breeding birds declined from 1911 to 1923. In 1923 Wetmore noted heavy egg predation by Bristle-thighed Curlews and Ruddy Turnstones, perhaps as a result of almost complete destruction of nesting cover by rabbits. No further numerical estimates were made until after the mid-1950's. The maximum figure of 40,000 given by POBSP in June 1967 (if not ex- cessive) approximates the 50,000 estimate of 1911 and suggests a population recovery from the low point reached when the island was nearly devoid of vegetation. Annual Cycle Birds arrive in late December or January with numbers increasing to their maximum about April. Egg laying may begin by late February (1913) or early March (1961, 1965) or may not begin until after mid-March (1964). The peak laying usually occurs from mid- through late April. Laying may continue through early June (1958, 1967), but birds laying this late have probably lost eggs earlier in the season. Wetmore found fresh eggs in mid-April, but by mid-May most of these had been destroyed by curlew and turnstone preda- tion or by flooding, and some birds were re-nesting. Hatching may begin as early as the last week of March and may continue through early July. Most young, however, probably hatch in mid- to late May. Fledging begins as early as late May but most young fledge in July or early August. Num- bers decrease with the fledging of young and by early September the popu- lation is reduced to a few immatures, adults, and weak non-fledged young that probably have been abandoned. No birds were found during the single October visit (1966), and no visits were made in Noyember. It is unlikely that more than a few wandering birds visit the island from late September until the beginning of the next cycle. Ecology Breeding: In mid-May 1902, Fisher (1903a: 780) found Gray-backed Terns in two interrupted bands around the island. Most were scattered in a narrow strip close to the beach on the seaward slope, entirely out- side the Sooty Tern colony. A second set of separated, smaller colonies ringed the lagoon, inside the Sooty colony. Dill and Bryan (1912: 15) found these terns on the rocks at the south end of the island and in small rookeries on the east side in 1911. In 1913, the birds nested on open sand at the south end of the island and apparently lost many eggs to the surf (Bailey, 1956: 98-100). Wetmore (ms.) described nesting in broken and eroded coral and sandy areas on and above the beaches on the northwest shore, and on the crest of the highest point at the south end of the island. Many birds laid too far out on the beach and their eggs were destroyed by surf. 225 The colonies observed during POBSP visits in June 1966 and 1967 were distributed in a pattern similar to that found by Fisher in 1902. Birds were scattered over most of the island wherever Ipomoea was present, es- pecially along the central lagoon and along the ocean beach. Considerable numbers nested also among the Scaevola on the beach crest, especially along the southern half of the west shore. Gray-backed Tern colonies were more open and less crowded than Sooty Tern colonies, and the birds seemed to prefer a vegetated edge bordering open areas. Eggs were laid on open sand, in the lee of coral rocks and drift, on exposed rocks, and sometimes under bushes. Non-breeding: Pre-nesting birds observed in mid-February 1963 ar- rived after dark and milled about over the island. Small groups rested briefly on open sand beaches, but they flushed readily if approached. Throughout the breeding season, roosting flocks, probably of non- nesters, form at night in open areas, particularly on the beaches at the outer edges of the Scaevola. From July to September most of these birds are immatures. Specimens One-hundred and ten Gray-backed Tern skins from Laysan are currently distributed in museums as indicated in Table GBT-l1. An additional 11 mounted specimens are in the Laysan exhibit at SUI. Also preserved are at least 3 skeletons (BPBM, 1; USNM, 2); and 24 eggs (BPBM, 11; USNM, 13). Banding and Movements The POBSP banded 45 adult Gray-backed Terns in February 1963. No interisland movements were recorded. Table GBT-1. Locations of Gray-backed Tern skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 1 7 9 23 BPBM 5 2 9 14 CMNH al il (0) 2 DMNH 5 3 iL 9 MCZ 3 2 6 11 SUL 3 2 O 5) UMMZ 3 5 O 8 USNM (non-POBSP) 22 13 1 36 Other* iL IL O 2 Totals 3} 36 26 110 *Law Coll. (1d, 1 9). 226 Table GBT-2. Observations of Gray-backed Terns on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1828 24 Mar. 2 Possibly recorded by Isenbeck (see Rothschild, 1893: v). 1891 16-27 June great On eggs (Rothschild, 1893-1900: 38). numbers 1896 24 June- 2 Nesting (Schauinsland, 1899: 101). 2h Sept. 1902 16-23 May ? Heavily incubated and hatching eggs (Fisher, 1903a: 780-781); 3 eggs of advanced incubation collected (USNM). 1903 20-30 Apr. ? At least 6 specimens and 10 eggs col- lected by Bryan (AMNH, BPBM). 1905 spring ? 6 collected by Schlemmer (MCZ). 1907 16-18 May 2 5 collected by Schlemmer (MCZ). 1911 24 Apr.- 50, 000 Fresh eggs and young in all stages of 5 June development (Dill and Bryan, 1912: 15). 1912 22 Dec.- 4 Many present on 22 December; numbers 1913 11 Mar. increasing by 24 January. Several thousand present by 24 February when first egg was found; no young by 11 March (Bailey, 1956: 98-100; Willett, ms. ) ies Sy None, 5,000 Only eggs, no young (Munter, 1915: 139). 1916 9 Feb. a few (Munter, ms.). 1923 8-13 Apr. common (Wetmore, ms.). 13 Apr. common 10 freshly incubated eggs collected (USNM). Ca. 50 nesting at south end of island (Dickey, ms.). 29 Apr.- 1, 000 Fresh to slightly incubated eggs 14 May (Wetmore, ms.). 1936 7-8 Mar. ? (Trempe, ms.). 227 Table GBT-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1950 23 June common Eggs (POFI). 1951 12 May ? Eggs (POFI). 1957 25 June- 4,000 Adults (Woodside, ms. b). 3 July 8-12 July ? (Labrecque, 1957: 18). 1958 27 May- large 4 June numbers Arriving; beginning to lay on 1 June (Warner, ms.). 1961 7-8 Mar. iy Sparse colonies; beginning to lay (Woodside and Kramer, ms.). 4-10 Sept. very few Mostly immatures (Woodside, ms. c). 1963 11-13 Feb. 200 Pre-breeding (POBSP). 3-10 Dec. Z (Walker, ms. b). 1964 10-11 Mar. 200 Not nesting (BSFW, POBSP). 16-20 Sept. 9 All unfledged, very weak young (BSFW, POBSP). 1965 6-11 Mar. 3, 000- Fresh eggs, birds laying (BSFW, POBSP). 5,000 17-21 July 5,000 Ca. 2,000 young, 1/2-grown to fledged (POBSP). 5-12 Aug. 1,600 1,500 young, mostly fledged; ca. 50 adults (POBSP). 1966 26-31 Mar. many Few nests with eggs (BSFW). thousands 10-16, 12,000 Mostly heavily incubated eggs and 20-21 June small chicks; also many flying imma- tures; ca. 1,000 eggs and 4,000 chicks (POBSP). 17-18 Sept. very few 3 or 4 near-fledging young seen (BSFW). 20-23 Oct. O (POBSP). 228 Table GBT-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1967 18-19 Mar. 1, 000 15 eggs found (BSFW, POBSP). 7-12 June [40, 000] Egg laying to flying immatures; estimated 10,000 nests (POBSP). 5-11 Sept. 40 Ca. 10 large young; some starving, evidently abandoned; very few adults seen (POBSP). 13 Dec. 0 (BSFW). 1968 17-19 Mar. 5,000 Scattered, small groups along west beach. Most pre-nesting; 2 nests with eggs found (BSFW, POBSP). 1969 26-29 Mar. 2,000 Eggs (BSFW). 9 Sept. 1 An immature bird (BSFW). BLUE-GRAY NODDY Procelsterna cerulea Status Rare visitor; two records: September 1964, June 1967. Observations On 19 September 1964 Walker (BSFW) saw one of these terns on the southwest side of the island at a distance of about 18 inches. Stadel (POBSP) saw another on the night of 11 June 1967. This species was not reported previously from Laysan but its appearance there is not surpris- ing since it is common on some of the inner islands of the leeward chain. BROWN NODDY Anous stolidus Status Common breeder; maximum recent estimate: 20,000 to [40,000]. Present year round but populations much smaller from mid-fall through early spring. Most birds breed from March through September but some also breed during all other months of the year. Usually nests on the ground under or near vegetation throughout the island. Often builds a nest of grass, leaves, and plant stems. 229 Populations Recent estimates, with the exception of the June 1967 estimate which seems excessively large, indicate populations of 10,000 to 20,000 birds during the early summer nesting peak (Table BrN-2). Munter's estimate of 6,000 in April 1915 and Dill and Bryan's figure of 5,500 in May 1911 indicate the same order of magnitude for populations then as in the 1960's. These figures are probably lower than maximum figures from recent visits because they were made earlier in the breeding cycle. Brown Noddies were less popular with feather hunters than some other species, so probably were not destroyed in great numbers. The low number found by Wetmore in 1923 (500 birds) probably was the result of the destruction of island vegetation by rabbits. Annual cle The Brown Noddy is one of the most, if not the most, irregular breeding species on Laysan. Eggs may be laid or may hatch and young may fledge in any month of the year. However, like other species that breed irregularly on Laysan, most of the birds nest in what appears to be a regular annual cycle. The breeding season begins with increased egg laying in April or early May, with a peak laying period from mid-May through early June. The peak hatching period occurs from mid-June through mid-July and the largest numbers of young probably fledge during August or early September. During the latter part of the fledging period, numbers of adults decrease and flying young make up a large proportion of the popu- lation. A large proportion of the adults and immatures apparently leave the island from late September or October through February or March. A gradual build-up in numbers of adults probably begins in late Winter, culminating in the onset of a new peak breeding period in the spring. Ecology Breeding: Brown Noddies nest over most of the island but appear to be more common in the Scaevola zone around its periphery, particularly on the west and northwest sides. Smaller numbers nest in the Eragrostis association, and in the Ipomoea zone around the lagoon, especially at the north end. Wetmore found them preparing to nest only near the north end of the lagoon in 1923, but did not describe the nest sites in detail. Although small colonies may be formed, these noddies are not charac- teristically colonial but build scattered nests throughout their habitat. Most nests are merely depressions in the sand under Scaevola or Eragrostis, lined loosely with grass, leaves, and sometimes bones, and on rare occasions unlined. Occasionally more substantial nests are built On procumbent limbs of shrubs, or a platform may be built of sticks and grass in low Scaevola, completely off the ground. The tops of Eragrostis clumps are also used. 230 Non-breeding: Fisher (1903a: 783) aptly described the activities of non-breeding Brown Noddies as follows: "Noddies like to gather in little companies on the beach, or on rocks near the shore, where they sit for hours dozing away or preening their feathers." Similar behavior has been noted on many subsequent visits, especially those made late in the breed- ing season when many fledged immatures join these diurnal resting flocks. At night most non=-nesting birds roost on Scaevola bushes. Specimens Eighty-six Brown Noddy skins from Laysan are currently distributed in museums as indicated in Table BrN-1l. Six additional mounted specimens are distributed as follows: BPBM (1 male); CMNH (1 female in Laysan group); DMNH (1 in Bahamas group); SUL (3 birds in Laysan group). Also preserved are at least 2 skeletons (USNM) and 10 eggs (USNM). Banding and Movements The POBSP banded 1,225 Brown Noddies: 1 young in September 1964; 35 young in October 1966, and 953 adults and 236 young in September 1967. No interisland movements were recorded. Table BrN-1. Locations of Brown Noddy skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 6 3 5 14 BPBM 1 iL 2 y CMNH 1 il (0) 2 DMNH 3 3 e) 6 MCZ 5 iL al i SUI 4 6 0) 10 UMMZ 3 3 O 6 USNM (non-POBSP) 19 16 iL 36 Other* AL O 0) 1 Totals 13 3h g 86 *Law.Coll. (1c). Table BrN-2. Observations of Brown Noddies on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1891 16-27 June plentiful Nesting (Rothschild, 1893-1900: 42). 231 Table BrN=2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1895 Sept. 2 3 or 4 adults collected by Hall (BPBM). 1896 2h June- ? Nesting (Schauinsland, 1899: 101). oh Sept. 1902 16-23 May ? Laying recently begun (Fisher, 1903a: 783). 1903 Apr. 3 Juvenile collected by W.A. Bryan (BPBM). 1904 2h May 2 3 adults collected by Schlemmer (MCZ). 1907 2 May 2 4 adults collected by Schlemmer (MCZ). 1911 23 Apr.- 5,500 Fresh eggs present during last week of 5 June May (Dill and Bryan, 1912: 15). 1912 22 Dec.- 400 Incubated eggs to 1/2-grown young on 1913 11 Mar. (nesting) 23 December. Nesting in colonies of 50 to 100 pairs (Bailey, 1956: 106); young birds fairly plentiful by 1 Janu- ary and fledged young present on 1 March (Willett, ms.). 3 Apr. 6, 000 Eggs and young (Munter, 1915: 139). 9 Feb. very Nesting on ground (Munter, ms.). common 8-13 Apr. ? Not nesting (Wetmore, ms.). 29 Apr.- 500 Selecting nest sites and beginning to 14 May build nests (Wetmore, ms.). 7-9 May ? 8 fresh eggs collected (USNM). 2-18 Aug. plentiful Eggs and young (Wilder, ms. b). 7-8 Mar. z (Trempe, ms.). 12 Dec. ? 3 or 4 at south end of island (Coultas, ms.). 23 June common Eggs and young (POFI). 232 Table BrN-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1951 12 May 2 Eggs (POFI). late June- ? Combined count of this species and the early July Black Noddy of 9,521 (Brock, 1951b: 18). 1957 25 June- 10, 000 Adults (Woodside, ms. b). 3 July 1958 27 May- ? Moderately to heavily incubated eggs; 4 June no young seen (Warner, ms. ) 1959 28 Apr.- ? None found nesting; groups of 25 to 50 1 May along beaches (Kramer, ms.) 1961 7-8 Mar. very few No eggs noted; only 4-5 immatures seen (Woodside and Kramer, ms.). 4-10 Sept. zh Eggs to flying immatures; mostly imma- tures. Some colonies with 1/2- to 3/4-grown young; a few birds on eggs (Woodside, ms. c). 1962 14-19 June ? Some downy chicks (Kramer and Beardsley, ms.). 1963 11-13 Feb. 100 1 heavily incubated egg (POBSP). 3-10 Dec. ? Low numbers; eggs to fledglings (Walker, ms. b). 1964 10-11 Mar. 100 Ca. 10 nests with eggs; 2 nearly fledged young observed (BSFW, POBSP). 16-20 Sept. 2, 000 A few eggs and young found; most non- breeding (BSFW, POBSP). 1965 6-11 Mar. 500-1, 000 Eggs to flying immatures (POBSP). 17-21 July 15, 000 Half-incubated eggs to fledged young; ca. 5,000 young (POBSP). 5-12 Aug. 20, 000 Egg laying to fledged young; ca. 1,000 nests with eggs and ca. 5,000 young (POBSP). 1966 26-31 Mar. ? At least several thousand; eggs to recently hatched young (BSFW). 235 Table BrN-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1966 10-16, 20, 000 Fresh eggs to nearly fledged young; 20-21 June ca. 5,000 nests with eggs, 5,000 with young (most recently hatched) (POBSP). 17-18 Sept. ? No chicks seen (BSFW). 20-23 Oct. 800 Only flying immatures (POBSP). 1967 18-19 Mar. several 1 egg found (BSFW, POBSP). hundreds 7-12 June [40, 000]* Eggs to fledglings; most nests with eggs. Estimated 10,000 nests (POBSP). 5-11 Sept. 10, 000 Eggs to fledged immatures; estimated 10 nests with eggs; 50 small young and 200 large young; at least 1/5 of the flying population comprised of imma- tures (POBSP). 13 Dec. ? Several flocks of 75-100 birds seen; no nests found (BSFW). 1968 17-19 Mar. 500-1,000 Most not nesting; few scattered large young and several nests with eggs seen (BSFW, POBSP). 1969 26-29 Mar. 100 Several found incubating eggs; another brooding a rather large downy chick (BSFW). 9 Sept. ? Several found incubating eggs (BSFW). *Hstimate considered excessive. BLACK NODDY Anous tenuirostris Status Common breeder; maximum recent estimate: 5,000. Most breeding occurs from November through July; birds present in varying numbers throughout remainder of year. Builds bulky nest in woody vegetation. 234 _ Populations ‘ Spring populations in 1911 and 1913 were at least 3,000 to 4,000 birds (Table BIN-3). Munter's 1915 estimate of 20,000 seems unbeliev- ably high; he may have confused this species with the Brown Noddy. Wetmore's estimate of 600 in 1923, the lowest breeding season estimate, suggests that the lack of vegetation may have reduced the population by 75 percent or more from the 1911 to 1913 period. There is little evidence that the population size has changed since 1911 to 1913. Bailey and Willett counted 1,151 active nests in January 1913, indicating a minimum nesting population of 2,302 birds. This is the highest number of nests ever recorded on the island, but there are no recent counts during the same time of year. Neither are there any 20th century population estimates for either summer or fall, when popu- lations may be larger than in the spring. Annual Cycle Black Noddies are present on Laysan throughout the year, but are apparently most common near the end, or after completion, of the nesting eycle. Egg laying may begin in October (1967), November (1963), or December (1912, 1915, 1964) and continue at least into May (1951, 1962, 1965, 1966). Observations of an egg and a young bird in September 1964 and young birds in September 1969 indicate that at least a few eggs may occasionally be laid through August. Observations from five other September visits (1918, 1961, 1965-67) indicate that in most years nesting has ended prior to September. Bailey's observation of many new eggs being laid in March after an earlier egg peak in January, as well as several recent sets of observations, indicate that there may be several nesting cycles within one breeding season. Within the usual breeding period, hatching occurs from as early as late October through late June or perhaps early July. Similarly, fledging may occur from late December (1967) through early August. Evidently the young do not leave the island immediately after fledging. A non-breeding period usually extends from mid- or late August to at least mid-October. Ecology Breeding: Palmer in 1891 (Rothschild, 1893-1900: 43) recorded Black Noddies nesting "in some numbers on the north side of the island, sitting around in clusters." Fisher (1903a: 774, 784) found them nest- ing in Chenopodium and another bush (probably Scaevola) in scattered communities over the island, either near the sea or in the interior. [In 1912 and 1913, Bailey (1956: 108) reported nesting in the few remaining low bushes circling the lagoon. Munter (1915: 139) found four or five colonies in low bushes and a small colony nesting "on the tops of lime- stone or phosphate rocks at the south end of the island." 289) In 1923 Wetmore (ms.) found them nesting on the corrugated iron roof of an old building, inside the building (one chick), and in the few remaining trees, and on rock ledges around the beaches. Nesting on artificial structures and rocks was undoubtedly because of lack of vegetation in areas where the birds normally nested. Evidence of com- petition for nest sites was shown by the nesting of Black Noddies within a few feet of nesting Red-footed Boobies--much closer than under present conditions. In recent years Black Noddies have nested most densely in the Casuarina tree near the landing on the northwest side of the island where at least 100 active nests were counted in December 1967. In June 1966 and March 1968 this tree contained 48 and ca. 205 nests, respectively. Other nests have been found in widespread locations in Scaevola, Cocos, Pluchea, and Eragrostis (Fig.41). Scaevola, particu- larly the taller growth along the northwest rim of the island, seems to be the next most favored nest site after Casuarina. Fisher (1903a: 784) described the nests of this species as follows: "from 18 inches to 3 feet up...constructed of twigs, usually morning-glory stems and leaves, and are from 10-12 inches in diameter. Usually the nests are built flat on top of bushes, or sometimes below in a crotch. There is scarcely any hollow, and occasionally a few feathers enter into the lining of dried leaves. The nests are in a large number of cases com- pletely plastered over with droppings and are used year after year." Figure 41. Black Noddy at nest with downy young, March 1965. Photo by R.B. Clapp. 236 Non-breeding: These noddies, especially recently fledged young, congregate during the day in flocks on the beaches, particularly the western ones. Adults and immatures gather at dusk and roost on the tops of Scaevola. These congregations are often much larger than the breeding population and include birds from other islands. Specimens One-hundred twenty-one Black Noddy skins from Laysan are currently distributed in museums as indicated in Table BIN-1. Ten additional mounted specimens are distributed as follows: AMNH (1 in Laysan group); BPBM (1 juvenal); CMNH (1 adult); SUL (7 birds in Laysan exhibit). Also preserved are at least 3 skeletons (BPBM, 2; USNM, 1); 2 alcoholics (BPBM, USNM); and 4 eggs (BPBM, 1; USNM, 3). Banding and Movements The POBSP and BSFW banded 325 Black Noddies on Laysan Island (Table BIN-2). Five Black Noddies banded on Laysan were recaptured as follows: French Frigate Shoals, 3; Lisianski, 1; Pearl and Hermes Reef, 1 (Appen- dix Table 4-9a). Seven banded on French Frigate Shoals and 1 banded on Midway were recaptured on Laysan (Appendix Table 4-9b). Table BIN-1. Locations of Black Noddy skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH nal 6 4 21 BPBM G 3 3 13 CMNH 2 1 iL ye DMNH 6 2 O 8 MCZ 4 iL aL 6 SUL i 3 2 12 UMMZ 1 y 0) 3) USNM (non-POBSP) 23 io D 5 (POBSP) 5 6 2 13 Other* 2 2 @) 4 Totals 68 35 iu 121 *Moseley (1 9); Law Coll. (16, 19); Phelps Goll. (1¢). Table BIN-2. Black Noddies banded on Laysan. Period of Survey Bander Adults Young Totals 1963 Feb. POBSP Sif ) Sif PGS aie > 237 Table BIN-2. (continued) Period of Survey Bander Adults Young Totals 1964 Mar. BSFW 4 ©) 4 Sept. POBSP 19 1 20 1966 Oct. POBSP 36 (65 )* 101 1967 Sept. POBSP iLL 9 126 1968 Mar. POBSP 27 10 ay Totals 20 85 325 *Age not reported; all were at least locals. Table BIN-3. Observations of Black Noddies on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1891 16-27 June 2 Eggs present (Rothschild, 1893-1900: 43). 1896 24 June- ? Nesting (Schauinsland, 1899: 101). 2h Sept. 1902 16-23 May ig Nests in considerable numbers; per- haps third in relative abundance among the terns. All eggs were more or less advanced in incubation; fledged young were common (Fisher, 1903a: 784). 1903 April ? 16 specimens and 1 egg collected by Bryan (BPBM). 1904 24 May ? 1 collected (by Schlemmer) (MCZ). 1905 spring ? 1 young collected (by Schlemmer) (MCZ). 1907 2 May i 4 specimens collected (by Schlemmer) (MCZ). 1911 24 Apr.- 3,000 Fresh eggs to fledged young (Dill and 5 June Bryan, 1912: 15). 1912 22 Dec.- 3, 000- Numerous on 22 December and beginning 1913 11 Mar. 4,000 to nest. Many nests with eggs on 24 December. 1,151 nests censused on 17 January and eggs present; young hatched 1913 11 Mar. (cont'd.) 1915 1916 1918 1923 1923 1930 1936 1950 1951 3 Apr. 9 Feb. 8-10 Sept. 8 Apr. 8-13 Apr. 14 Apr. 29 Apr.- 14 May 2-18 Aug. 7-8 Mar. 12 Dec. 23 June 12 May late June- early July 20, 000 2,000 ? numerous ? ? 238 Table BIN-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References gi 22 Dec.- shortly afterward and nearly full- grown young were present by 10 Feb- ruary; fully one-half of young lost subsequently; eggs seen again in March (Bailey, 1956: 108); first young seen 17 January; re-nesting by 20 February; eggs plentiful again by 1 March (Willett, ms.). Eggs and young in nests (Munter, 1915: 139). Thousands nesting; eggs to more than 1/2 grown young (Munter, ms.). No nests or young (Diggs, ms.). Colony on buildings near camp, ca. 9% with nests under construction, 90% with eggs, only a few with young; colony at south end of island (about 100 pairs), about half building nests and half with eggs (Dickey, ms.). (Wetmore, ms.). Colony in tobacco patch building nests (Dickey, ms.). Fresh eggs and young (Wetmore, ms.). A few pairs seen (Wilder, ms. b). Nesting in Casuarina trees, eggs present but not clear from ms. whether young were present (Trempe, ms.). About 40 in Casuarina trees (Coultas, ms.). Eggs and chicks (POFT). Eggs (POFI). Not distinguished from Brown Noddy in diurnal count of 9,521 birds (Brock, WeeulaR ws}) 239 Table BIN-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1957 2 June- 1, 000 (Woodside, ms. b). 3 July 1958 27 May- ? Few eggs; young from about one week 4 June to fledged; aggregations of fledged birds (Warner, ms.). 1959 28 Apr.- ? All stages of young (Kramer, ms.). 1 May 1961 7-8 Mar. ? From eggs to newly hatched young to 2/3-grown young (Woodside and Kramer, Msieye 4-10 Sept. many Immatures; no nests found (Woodside, mishpe)ie 1962 14-19 June common Eggs and young (Kramer and Beardsley, ms. e 1963 11-13 Feb. 200 Eggs; one examined was well-developed (POBSP). 3-10 Dec. ? Most in Casuarina tree had newly hatched young; some eggs present (Walker, ms. b). 1964 10-11 Mar. 1, 000 Estimated 150-200 nests with eggs and same number with young (BSFW, POBSP). 16-20 Sept. 300 1 nest with egg and 1 chick noted; most of population non-nesting (POBSP). 1965 6-11 Mar. 1, 000- Most nests with-eggs, but young from 2,000 recently hatched to fledged also ob- served (POBSP). 17-21 July 3,500 Estimated 1,500 young present, from half-grown young to fledged birds (POBSP). 5-12 Aug. 4,000 Estimated 1,000 flying young present (POBSP). 26-31 Mar. ? Ca. 300-400 in the Casuarina tree; at night many hundreds more; from eggs to nearly fledged and flying young (BSFW). ako Table BIN-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1966 10-16, 4500 A few half-grown young to flying im- 20-21 June matures. Ca. 5 nests with eggs; also ca. 500 young present (POBSP). 17-18 Sept. thousands No nesting noted (BSFW). 20-23 Oct. 4 000 None nesting; flying immatures seen (POBSP). 1967 18-19 Mar. 350 Eggs and nearly fledged young present; some seen nest-building. In the Casua- rina tree 8 of 10 eggs checked were fresh and 6 nearly fledged young were counted (BSFW, POBSP). 7-12 June 5,000 From small to large downy young in nests; flying immatures observed; most nests had large or medium-sized downy young. Sample count of 38 nests in the Casuarina tree: 6 (16%) with small downy young; 13 (34%) with medium-sized downy young and 19 (50%) with large downy young (POBSP). 5-11 Sept. 2, 000 None nesting (POBSP). ig} WAGs 2 100 nests checked in Casuarina tree: 42 were being constructed or had been constructed recently. Of those with contents, 53 (91%) contained eggs and 5 (9%) small downy chicks; also several very large chicks, almost capable of flight seen (BSFW). 1968 17-19 Mar. 3, 000 Very slightly incubated eggs to fledged immatures. Of 79 nests with contents counted, 69 (88%) held eggs; 1 (1%) a small downy chick; 1 (1%) a medium chick; and 8 (10%) large young. No less than 250 active nests present (BSFW, POBSP). 1969 26-29 Mar. 1,400 + In sample count of 100 nests, 5% were empty but active, 71% contained eggs, most of them well incubated, and 24% contained small downy young (BSFW). 9 Sept. ? Nests contained downy to near-fledging young. Most nests held large downy young (BSFW). aa a 2k1 WHITE TERN Gygis alba Status Common breeder; maximum recent estimate: 1,500. Breeds throughout the year but with a definite peak from about May through August; present in smaller numbers during remainder of year. Lays single egg chiefly on rocky outcrops and ledges, less frequently on limbs or vegetation, rarely on the ground. Populations Not since mid-June 1891 has the White Tern been considered abundant (Table WhT-3). In 1902 Fisher considered it "one of the least abundant of the breeding seabirds.” It was one of the species most prized by feather hunters and undoubtedly suffered heavily during the feather raids of 1909-1910 and 1915. Maximum populations recorded during that period were about 75 in May and June 1911; 80 in early March 1913 and about 400 in April 1915. These estimates (except the one for 1915) and those for September 1918, and April and May 1923 are lower than recent estimates for the same periods and suggest that present populations are larger than those during the early part of the century. Annual Cycle Eggs or young may occur during any month of the year but there is a distinct breeding peak during spring and summer. Eggs may be laid as early as late November or early December (1912, possibly 1963) and as late as early October (1963) but most laying probably occurs from late April through mid- or late May. Peak hatching and fledging periods are respectively from early June through late June and from mid-July through mid-August. That no nesting birds were reported by the February 1963 survey party probably reflects inadequacies in the survey rather than the actual ab- sence of breeding birds. Bailey's observations of eggs and young in February 1913 as well as recent observations of young in March (1961, 1964, 1967 to 1969) and eggs in December (1963, 1967) indicate that some White Terns usually breed in January. March estimates, ranging from 100 to 500, suggest that numbers begin to increase during this month, and a gradual build-up in numbers probably continues during the succeeding month or months. Ecology Breeding: Early observers found White Terns nesting in a wide variety of sites on Laysan. In 1899, Schauinsland (1899: 57) reported finding eggs on the bare sand, on the rim of the lagoon's salt crust, on bare rock cliffs near the edge of the surf, and in the forks of branches in the bushes. Fisher's (1903a: 785) observations suggest that in 1902 these birds nested more commonly in the interior than they do at present. He ake found small colonies scattered over the island interior with the largest near the fresh water pond near the south end. Here the White Terns laid their eggs "on lumps of phosphate rock, among bunch grass, or under the overhanging shelter of some shrub or clump of vines..." Munter (1915: 139) also noted nesting on the scattered phosphate rocks in the southern part of the island. Wetmore (ms.) reported nests on rocks and rocky ledges of the south, southeastern, and north beaches, and also on the framework of the buildings and on piles of guano. More recent observers found White Terns nesting primarily around the periphery of the island with a relatively small proportion in the interior. Eggs were found chiefly on the rocks and rock ledges of the southwestern and south beaches and on the rocks along the north beach, many of which are nearly hidden from view by encroaching vegetation. Small colonies occur on the rock piles in the interior south of the lagoon. Nests were also reported in the Casuarina tree, in Cocos trees (July 1965) and in Scaevola bushes (September 1966, June 1967). Probably the paucity of nests reported from Scaevola indicates that such nests are more easily overlooked than those on the rocks, but may indicate a genuine preference for rock or ledge nesting sites. Nests occur from ground level to 8 feet or more above the substrate on the larger rock ledges. Nests on the ground, however, are apparently far less common than those which are slightly elevated, even if no more than one or two inches. Only Schauinsland and Fisher of the earlier observers stated or implied that nests were found on the ground and only three such nests were reported during recent POBSP surveys. In August 1965 an adult was found incubating an egg in a slight depression in the sand under a Scaevola bush and two eggs were found under clumps of Era- grostis within the Sooty Tern colony. Many nests were found less than a foot from the ground, usually in a niche on the side of a boulder or on a flat stone ledge. Non-breeding: Some non-breeding birds roost on the rockpiles and ledges mentioned above and are often abundant in the taller vegetation, particularly Scaevola along the west beach, and the Casuarina and Cocos trees. Most non-breeding birds, however, are absent from the island. Specimens Seventy White Tern skins from Laysan are currently distributed in museums as indicated in Table WhT-l1. Seven additional mounted specimens are distributed as follows: BPBM (4, including 2 juvenals); SUI (3 birds in Laysan exhibit). Also preserved are at least 3 skeletons (BPBM, ale USNM, 2); 3 alcoholics (BPBM, 2; USNM, 1 young); and 21 eggs (BPBM, 4; USNM, 17). Banding and Movements The BSFW and POBSP banded 480 White Terns on Laysan through 1969 (Table WhT-2) but no interisland movements were recorded. 243 Table WhT-1. Locations of White Tern skins from Laysan. Adult Adult Museum Males Females Other Totals AMNH 8 7 7 ee BPBM 4 y y 12 CMNH 2 O () 2 DMNH 2 O I 3 MCZ 2 aL 1 4 SUL 2 1 il 4 UMMZ O 1 2 3 USNM (non-POBSP) uf 6 6 19 Other* 1 O O 1 Totals 2 20 22 70 *Hachisuki (1c). Table WhT-2. White Terns banded on Laysan. Period of Survey Bander Adults Young Totals 1963 Feb. POBSP 31 O Syl 1964 Sept. POBSP 155 y 159 1965 Mar. POBSP 84 a 85 1966 Mar. BSFW 6 O 6 1967 Sept. POBSP 189 9 198 1968 Mar. POBSP 1 fo) 1 Totals 1 80 Table WhT-3. Observations of White Terns on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1890 16 July ? Small flock near lagoon (Lyons, 1890: gl). 1891 16-27 June great Nests with eggs (Rothschild, 1893- abundance 1900: 36). 1895 Sept. ? At least 4 adults collected by Hall (BPBM, MCZ). 1896 24 June- 2 Eggs and young (Schauinsland, 1899: 2h Sept. 5O))s aby Table WhT-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1902 16-23 May ? One of the least abundant of the breeding seabirds; fresh eggs to fully fledged young (Fisher, 1903a: 785). 23 May ? 3 heavily incubated eggs collected (USNM) . 1903 7-15 Apr. f 11 specimens (including 2 juveniles on 7 April) and 3 eggs collected by Bryan (AMNH, BPBM). 1904 2k May ? 1 collected by Schlemmer (MCZ). 1907 8 May 2 2 collected by Schlemmer (MCZ). 1911 24 Apr.- 75 Only 4 seen during the first week of 5 June survey. Found nesting on 15 May (Dill and Bryan, 1912: 15). 1912 22 Dec.- 1913 11 Mar. 80 2-3 pairs present 22 December; an egg found 29 December hatched on 6 January and the young was able to fly by 10 March. Eggs seen in January and Febru- ary. Numbers increased to more than 50 on 11 February and about 40 pairs were present on 1 March; on 11 March not more than six pairs present (Bailey, 1O56: TOs Willette mss IUSHB) S) fake 400 Eggs seen (Munter, 1915: 139). 1916 9 Feb. small No eggs (Munter, ms.). number 1918 8-10 Sept. 5 (Diggs, ms.). 1923 8-13 Apr. common Mostly eggs; a few hatched young pres- ent on 8 April (Dickey, ms., Wetmore, ms.). 18 Apr. 100 (Dickey, ms.). 29 Apr.- 80 Count; eggs and young destroyed by 14 May heavy surf between visits (Wetmore, ms.); 14 fresh eggs collected (USNM). ahs Table WhT-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1930 2-18 Aug. scarce A few pairs nesting in buildings and 3 pairs seen on rocks; some with eggs, some with young (Wilder, ms. b). 1936 7-8 Mar. ? (Trempe, ms.). 12 Dec. 2 4 or 5 around house (Coultas, ms.). 1950 23 June a few (POFT). 1951 12 May 2 Eggs. noted (POFT). late June- 150 Based on diurnal census (Brock, 1951b: early July iN3})) 5 1955 10 Feb. 4 (POFI). 1957 2 June- 500 Adults (Woodside, ms. b). 3 July 8-12 July ? Eggs to fully grown young (Labrecque, 1957: 18). 1958 27 May- ? Eggs and young; all eggs examined mod- 4 June erately or heavily incubated; most chicks less than 2 week old (Warner, ms). 1959 28 Apr.- ? Eggs through fully feathered immatures 1 May (Kramer, ms.). 1961 7-8 Mar. ? Eggs or small chicks; only 1 flying immature (Woodside and Kramer, ms.). 4-10 Sept. ? Eggs to immatures but mostly flying immatures (Woodside, ms. c). 1962 14-19 June common With eggs, downy chicks, and fully feathered young (Kramer and Beardsley, ms.). 1963 11-13 Feb. 100 Paired but no eggs seen (BSFW, POBSP). 3-10 Dec. ? Eggs and half-grown young (Walker, ms. b). 1964 10-11 Mar. 400 Eggs to flying immatures; ca. 18 eggs and 12 young seen (BSFW, POBSP). 246 Table WhT-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1964 16-20 Sept. 400 Estimated 15 eggs and 25 young; 1 egg hatched during visit (POBSP). 1965 6-11 Mar. 200-300 Eggs and a few chicks (POBSP). 17-21 July 1,500 Also ca. 500 nestlings from downy chicks to flying immatures; 1 egg seen (POBSP). 5-12 Aug. IL Feo) Also ca. 200 eggs and 500 young; fresh eggs to flying immatures (POBSP). 1966 26-31 Mar. 4OO+ Some incubating eggs (BSFW). 10-16, 900 Eggs to flying immatures; also ca. 20-21 June 100 eggs and 100 young (POBSP). 17-18 Sept. 200-300 Hundred or so present during the day and at least several hundred at night; eggs found (BSFW). 20-23 Oct. 100 Several eggs and a few fledged young (POBSP). 1967 18-19 Mar. 100 1 egg and 3 half-grown young (BSFW, POBSP). 7-12 June 1, 000 Eggs through flying immatures; estimated 200 nests (POBSP). 5-11 Sept. 800 Fresh eggs through flying immatures; many nests with medium-sized or large young. Of 15 nests in a sample count: 5 (33%) contained eggs, 1 (7%) had a small downy chick, 7 (47%) had medium- sized or large chicks, and 2 (13%) con- tained near fledging young (POBSP). 13 Dec. % Fairly common along beaches; 3 on eggs (BSFW). 1968 17-19 Mar. 300-350 Most not nesting; 3 nests with eggs and 1 large chick seen (BSFW, POBSP). 1969 26-29 Mar. 250 Eggs to young in all stages (BSFW). ALT HORNED PUFFIN Fratercula corniculata Status Accidental; one record: February 1963. Observations The only record is a skeleton (USNM 497918) collected 12 February 1963 by POBSP personnel. The bird was evidently one of many washed up on the Northwestern Hawaiian Islands in the winter of 1962 to 1963. At least 15 other individuals were found by POBSP personnel on Kure, Midway, and Pearl and Hermes Reef (Clapp and Woodward, 1968: 30). Still other puffins, some of which may have been the same birds reported by POBSP field workers, were reported from Kure by Robbins (1966: 53) and from Midway by Fisher (1965: 357). LAYSAN MILLER-BIRD Acrocephalus f. familiaris Status Endemic. Formerly a common to abundant permanent resident; now extinct. Nested during early summer in bunch grass near the lagoon. Populations The Laysan Miller-bird was first collected for science by Palmer and Munro in 1891 and was named by Rothschild the next year (1892b: 109). Earliest accounts considered it "plentiful" or "abundant" but few numeri- cal population estimates were ever published (Table LM-2). The Laysan Miller-bird became extinct sometime between 1915 and 1923 and only sketchy details of its life history were reported by the few biologists to observe the species in life. The most useful informa- tion available is a summary by Bailey (1956: 117-118) and the original report by Fisher (1903a: 805-806) from which most of the following account is taken. Its life history was probably very similar to that of its only relative in the central Pacific--the Nihoa Miller-bird (Acro- cephalus f. kingi) of Nihoa, some 563 miles to the southeast. Habits The Miller-bird was fearless and its confiding ways were described by all biologists who encountered it. Birds visited tables at mealtime, alighted on and very near people and searched inside occupied buildings for insect food. Munro (1942b: 2) described feeding inside lighted buildings at night and commented that they became a pest in the laboratory by perching on, tipping and breaking test tubes. Birds were always described as very active or "busy." Much of this activity was devoted to the pursuit and capture of several species of moths 248 locally called "millers," hence the common name "miller-bird." Accord- ing to Munro the moths were swallowed entire. Feeding took place in and around buildings and in various types of vegetation, particularly the mat-like Portulaca surrounding the lagoon where caterpillars were a major food item. Activity, including singing, reached peaks in morning and late afternoon and birds retired to the shelter of bushes or grass during the warmer parts of the day. The song was described as "liebliches,” (lovely) (Schauinsland, 1899: 44) and "musical" (Fisher, 1903a: 806); and its call was "a harsh deep note much resembling that of a thrush" (Rothschild, 1893-1900: 2). Nesting Nests were usually placed about two feet from the ground in the middle of Large clumps of bunch grass. Fisher stated that largest numbers were found along the inner edge of the bush grass area near the lagoon. The nest was best described by Fisher (1903a: 806) as follows: "The structure itself is composed of dried grass stems and blades, fine rootlets, white albatross feathers. The bowl is 1 3/4 inches wide by the same depth, and the diameter of the mouth is somewhat less than that of the interior, so that the edges of the cup overhang a little. It is lined with fine rootlets, shredded grass, and white albatross feathers, the last being a very characteristic feature of all nests,...Occasionally a trace of down was found on the inside. The outer portion of the nest is rather loosely held together, and forms a globose mass 3 1/2 inches in diameter." Nesting was just getting underway in mid-May 1902 (1 clutch of 3 eggs; 1 incomplete clutch of 2 eggs and many nests "apparently just ready for eggs"). Eggs were taken by Palmer in late June 1891 and Dill and Bryan (1912: 22-23) found nests with eggs and young birds in late spring or early summer. The clutch size was two or three. Eggs varied in size (Fisher) from 22 x 15 mm to 19 x 14 mm. The ground color varied from very pale olive buff through greenish white, to almost pure white. Most eggs were blotched and spotted with olive chiefly at the larger end and often with tiny white lines and specks scattered over the entire egg. Although its eggs were known to be sometimes eaten by the Laysan Finch (Dill and Bryan, 1912: 23), extinction was almost certainly due to destruction of habitat by rabbits. An excellent photograph of this species is found in Fisher (1903a: Fig. 43) and Bailey (1956: 118). Specimens Seventy-seven Laysan Miller-bird skins are currently distributed in museums as indicated in Table LM-1. Five additional mounted specimens are distributed as follows: SUI (2 in Laysan exhibit; 1 in extinct bird case); DMNH (2 birds in Laysan exhibit). Also preserved are at least 2 skeletons (BPBM); 4 nests (BPBM, 2; USNM, 2); and 4 eggs (BPBM, 1, USNM, 3). - Table IM-l. Museum Totals 249 Locations of Laysan Miller-bird skins. Adult Adult Males Females Other Totals 13 9 0) 22 12 9 3 24 1 alt 0) 2 6 3 0 9 O 0 4 4 y 3 3 10 y 1 i}: 6 0 a 11 T1* *Law Coll. (1c, 1); Hachusiki (1); Carnegie Mus. (10); Acad. Nat. Sci. Phila. (2 ct). **According to Greenway (1958: 393), at least one specimen is at Bremen and at Denver. Table LM-2. Observations of Laysan Miller-bird on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1891 16-27 June 1896 1902 1903 1904 1907 1911 24 June- 24 Sept. 16-23 May 22-23 May 20-28 Apr. 12 May 17-19 May 24 Apr.- 5 June "plentiful" Two nests with eggs, 24 June (Palmer in Rothschild, 1893-1900: xi, 2). 2 Present (Schauinsland, 1899: 100). “abundant” "One of the most abundant of the four strictly land birds; many empty nests and two with eggs" (Fisher, 1903a: 806). 2 3 eggs and 2 nests collected (USNM). ? At least 26 specimens and 2 nests and eggs collected by W.A. Bryan (AMNH, BPBM). z 1 collected by Schlemmer (MCZ). ? 8 collected by Schlemmer (MCZ, PANS). (under 300) "least abundant of the indigenous [land] species." A few nests with eggs and young (Dill and Bryan, 1912: 22-23). 250 Table IM-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1912 22 Dec.- "abundant" (Bailey, 1956: 117). 1913 11 Mar. “at least "Least plentiful of land birds on 600" Laysan, but...in no present danger of extinction" (Willett, ms.). Tigh) 8h sgore perhaps "fairly common" (Munter, 1915: 140). 1,500 1916 9 Feb. % A few seen around buildings (Munter, jis) 1918 8-10 Sept. extinct? Not seen (Diggs, ms.). 1923 8-13 Apr. extinct (Wetmore, ms.). LAYSAN HONEY-EATER Himatione sanguinea fraithii Status Endemic. Formerly common permanent resident; extinct since 1923. Nested in bunch grass and Chenopodium bushes near the lagoon during spring and early summer. Populations The Laysan Honey-eater was first reported by Isenbeck (Kittlitz, 1834: 125) in 1828 and was formally described by Rothschild (1892b: 109-110) in 1892 from specimens taken by Palmer and Munro the previous year. Although all subsequent workers reported the species, the only population estimates (Table LH-2) are those made in 1911 (about 300), 1915 (about 1,000), and just prior to its extinction in 1923 (3). The few published observations suggest that honey-eaters were never very abundant on Laysan. Isenbeck considered it "not very common" in 1828 and other workers through the early part of the 20th century stated or implied that though it occurred in fair numbers it was nevertheless the least common of the endemic species. Any major reduction in numbers apparently occurred after 1902. Dill and Bryan estimated the population at 300 birds in 1911 and in 1913 Bailey reported "a few" confined to the few remaining patches of rapidly disappearing vegetation. Munter's 1915 estimate of "about 1,000 birds" and "fairly common" is perhaps too generous since he considered them not very numerous the following year. At any rate only three birds remained by 1923 and the last of these perished in a sandstorm on 20 April (Wetmore, ms.). 251 Habits The few biologists who observed the Laysan Honey-eater left few detailed accounts of its life history. The most useful information is contained in the original observations of Fisher (1903a: 803-804) and in the compilation by Bailey (1956: 119, 122). The honey-eater, like the miller-bird, was very active, constantly flitting about in vegetation or near the buildings. Most accounts sug- gest that it was less fearless and confiding than the miller-bird although birds did occasionally enter buildings for moths and Dill and Bryan (1912: 22) reported four birds roosting inside a building at night. Birds originally fed regularly on nectar from the large flowers of Capparus sandwicheana and changed to Portulaca and Sesuvium when the Capparus became extinct. When feeding in bushes, birds flitted rapidly about reminding Schauinsland (1899: 44) of hummingbirds. Fisher (1903a: 804) made a similar comment on birds feeding on the low mat flowers except that birds were then walking from flower to flower rather than hovering before them. Insects were a very important part of the diet and probably more important than nectar at some seasons. Fisher (1903a: 804) reported their feeding on small, green caterpillars taken from Chenopodium bushes and all observers saw them feeding on the abundant moths ("millers") which occurred throughout the island. The moths were held in one foot while the wings were removed and only the soft parts were eaten. Palmer (in Rothschild, 1893-1900: 4) described the song as low, sweet, and consisting of several notes. One bird, captured and hand-held, sang and answered Palmer's whistle apparently unafraid. Birds were said to be usually silent except during the breeding season. Singing was re- ported between December and mid-June. Honey-eaters occurred over the entire island (Fisher, 1903a: 804) but were more abundant in the interior near the lagoon, probably (in 1902) attracted by the flowering vegetation as well as by nesting habitat. Nesting was usually in the bunch grass near the lagoon but Schauinsland (1899: 101-102) found some nests in Chenopodium. The nest was a cup-shaped affair of rootlets and grass, lined with fine rootlets and usually albatross down. Fisher (1903a: 804) commented that nests could be distinguished from the very similar nests of the miller-bird by the absence of white feathers but apparently this was not always true. Schauinsland (1899: 101-102) further distinguished honey-eater nests by their tighter construction and more shallow cup. Details on the breeding cycle are unavailable. A few nests with eggs and/or young were recorded by various observers but without specific dates. A nest with a single egg was taken by Fisher (1903a: 804) some- time in mid-May and at least one egg was taken by W.A. Bryan on 10 May (BPBM). Bailey (1956: 122) gave the clutch size as "four to five” and several sets of three were taken by early collectors. The eggs were 252 white, spotted with grayish or reddish-brown on the larger end. More detailed descriptions of nests and eggs are given by Schauinsland (1899: 101-102) and Fisher (1903a: 804). A photograph of the nest is found in Fisher (1903a: Fig. 51) and Bailey (1956: 121). Specimens Eighty-five Laysan Honey-eater skins are currently distributed in museums as indicated in Table LH-1. Six additional mounted specimens are distributed in museums as follows: BPBM (1); DMNH (2 co’ in Laysan exhibit); SUI (2 birds in Laysan exhibit, 1 in extinct bird case). Also preserved are at least 2 skeletons (BPBM, USNM); 3 nests (BPBM) and 1 egg (BPBM). Table LH-1. Locations of Laysan Honey-eater skins. Adult Adult Museum Males Females Other Totals AMNH 7 y 5 16 BPBM 8 y at 23 CMNH 1 alt ©) 2 MCZ 6 3 2 11 SUL O i O 1 UMMZ 0 0 8 8 USNM 10 Hi 3 20 Other* 3 iL 0) 4 Totals 35 21 29 B5%** *¥Law Coll. (1c, 19, both missing); Acad. Nat. Sci. Phila. (1c); Carnegie Mus. (1°). x*\dditional specimens are said (Greenway, 1958: 409) to be at Berlin, Bremen, Denver, London, Stanford University, Stockholm. Table LH-2. Observations of Laysan Honey-eaters on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1828 2h Mar. not very Isenbeck in Rothschild (1893-1900: v). common 1891 16-27 June fair Rarest of island birds; full grown young number present (Palmer in Rothschild 1893-1900: 255) Table LH-2. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1896 24 June- 2 Present; one nest found (Schauinsland, 24 Sept. 1899: 101-102). 1902 16-23 May q Least abundant of the endemic land birds; by no means rare; only 1 nest (with eggs) found (Fisher, 1903a: 803-804). 1903 19-25 Apr. ? At least 22 birds, 3 nests and 1 egg collected by W.A. Bryan (AMNH, BPBM). 1904 12-13 May ? 2 collected by Schlemmer (MCZ). 1907 23-24 May ? 6 collected by Schlemmer (MCZ). 10, 15 June ? 2 juveniles collected by Schlemmer (MCZ). 1911 24 Apr.- 300 Not common; a few nests with well- 5 June incubated eggs or young (Dill and Bryan, 1912: 22). 1912 22 Dec.- few Less common than miller-birds and 1913 11 Mar. finches; confined to vanishing vegeta- tion (Bailey, 1956: 122). 1915 3 Apr. 1, 000 Fairly common (Munter, 1915: 140). 1916 9Q Feb. ? In pairs; not very numerous (Munter, ms.). 1918 8-10 Sept. ? A red bird seen (Diggs, ms.) was almost certainly this species. 1923 8-20 Apr. 3 Three remained around rocks at south end of island; none survived the storm of 20 20 Apr. extinct April (Wetmore, ms.). LAYSAN FINCH Psittirostra c. cantans Status Endemic. Abundant; most recent estimates: about 10,000 birds. Occurs throughout the island in all habitats. Most nesting is from early May through July. Nest is a cup of grass and plant stems placed in a grass tussock or (formerly) a bush. 25) Populations The Laysan Finch, always a conspicuous part of the island avifauna, was mentioned by nearly all visitors to Laysan. Its bright colors, song and hardy nature made it a popular cage bird and ships visiting Laysan frequently transported birds to Midway, Honolulu and perhaps elsewhere. The species was formally named (Wilson, 1890: 341) from a captive bird purchased in Honolulu from a shipment of 60 imported in 1889. A common term used by various observers through 1915 in describing the finch population, "everywhere in abundance," is understandable. The finches were tame, constantly in evidence and entered occupied buildings where they explored everything in sight, roosted, sang and (in 1923) even built nests. Few observers, however, attempted the difficult task of estimating the finch population (Table LF-3). Dill and Bryan estimated 2,700 birds in 1911 and Munter estimated 4,000 in 1915. Finches were still "very common" in 1916. The population dropped to 100 birds in 1923 but had increased to at least 1,000 in 1936. This drastic fluctua- tion in numbers seems to have paralleled the degradation and subsequent recovery of the vegetation. Birds were again considered "quite abundant” in 1950 by Brock who estimated about 5,000 birds the following year. Woodside estimated about 5,000 birds in 1957. Warner estimated a con- servative 10,000 birds as the result of a transect census in 1959. Numbers are believed to have remained at about this level to the present time (Table LF-3). Habits Fisher's observations of 1902 (1903a: 804) are equally appropriate today: "quite fearless and unsuspicious. It is also saucy to a marked degree, and ignores the presence of man when he is peaceably disposed. One can not walk ariywhere without encountering them singly or in little flocks, diligently searching for food among the bushes, or out in the open. When disturbed they eye the intruder with interest or half in doubt and utter their low, mellow linnet-like call. They do not fly far, but prefer to alight soon, and run along the ground, or elude pursuit by suddenly crouching under a grass tussock." Schauinsland (1899: 44) stated that at meal time finches would "sit on the edge of our plates and share our rice and bacon." Finches today are only slightly less confiding. Food Fisher (1903a: 804), the first to report on this species at length, noted that these finches were "fond of the soft parts of grass stems, tender shoots of bushes, seeds, and especially of eggs." More recent observers have been more specific about vegetation eaten. Apparently Tribulus seeds and seeds of Eragrostis are a major staple, more observers having noted these plants being eaten more often than any other (Labrecque, 1957: 17; Wilder, ms. b; Kramer, ms.; Warner, ms., Crossin, POBSP). Other items apparently frequently eaten are inflorescences of coconuts (Walker, ms. b; Crossin) and the seeds and buds of Boerhavia and a es ae ae 2D Portulaca (Crossin; Kramer, ms.). They have also been noted feeding on the centers of flowers of Ipomoea (Kramer, ms.) and Nicotiana (Crossin). Their passion for birds’ eggs has been described by numerous ob- servers and any eggs left unattended for a few moments are soon broken and their contents eaten by the finches. Finches also concentrate at the edges of tern colonies and take advantage of any disturbance (such as human entry) by flying in and attacking unattended eggs. Species whose eggs are known to have been eaten by the finch include Bulwer's Petrel and the Wedge-tailed Shearwater, Sooty, Gray-backed and White Terns, and Black and Brown Noddies (Fisher, 1903a: 801; Bailey, 1956: 123; Wetmore, ms.; Crossin, Stadel, POBSP). Bryan and Dill (1912: 22) indicated that the finches also ate finch eggs but no more recent observer has confirmed this. The contents of albatross eggs, most probably old and rotten, are also avidly eaten (Fig.42) but it is doubtful if the finches themselves break the eggs. Probably most albatross eggs eaten are those broken by other animals (man, seals, Bristle-thighed Curlews) or those that may have burst due to a combination of high internal pressure from egg-decomposition and rough contact caused by storms (as in the in- stance of eggs wind-rowed along the lagoon edge). Figure42, Laysan Finches feeding on remains of albatross egg, March 1965. Photo by R.B. Clapp. 256 Crossin (POBSP), who watched these birds feeding in Sooty Tern colonies during June 1966, gives the most detailed notes on the egg- eating process: ; When [POBSP] personnel moved through the tern colony, finches were observed on numerous occasions to approach an exposed egg and peck through the shell with...forceful blows. Pieces of the shell were then chipped off and flipped aside until a...large hole...usually [a ragged circle] of about 3/4 inch diameter [was made]. The birds then sipped the contents, lifting their heads back as if drinking water. Crossin's observations also suggest that egg-eating is a learned trait. In one instance a number of immature finches "were given every opportunity to attack freshly uncovered eggs, but ignored them and con- tinued feeding on seeds of Boerhavia and other plants." On the other. hand "as soon as a tern was flushed from its egg, nearby adult finches quickly proceeded to the egg and began feeding." Observations by Crossin and by Stadel (POBSP) in June 1967 clearly indicate that human disturbance coupled with the finches' propensity for eating eggs can result in considerable Sooty Tern egg mortality. Crossin believed that at least several thousand eggs were destroyed by finches in June 1966 and Stadel estimated that more than 10 percent of the eggs then present were destroyed in June 1967. Other species of terns may also have a large proportion of their eggs destroyed under the effect of human disturbance. In 1923 Wetmore (ms.) noted that most, if not all, White Tern and Black Noddy eggs in the area of the Tanager Expedition campsite were destroyed within a few hours of the field party's arrival. Bailey (1956: 123) found the brunt of preda- tion to be on Black and Brown Noddies and on Gray-backed Terns but comments by Dill and Bryan (1912: 22) also suggest that considerable egg destruction may occur as the result of human disturbance. Being somewhat opportunistic feeders, Laysan Finches have been ob- served feeding on meat. Munro (1942b: 2) reported them feeding on maggots and flesh of dead birds, and more recently, in March 1965, POBSP personnel noted them feeding on the flesh of dead albatross chicks. Reproduction Annual Cycle: Although no one visit could obtain sufficient data to delimit the breeding cycle, the various sets of observations (Table LT-3) seem clearly to indicate spring and summer nesting. Bailey reported eggs being laid in February and March but no nests were found recently during those months despite fairly intensive checks of potential nest sites during two March visits (1968 and 1969, BSFW). However, Bailey was on Laysan continuously for nearly three months and made much more thorough observations than could recent observers, who seldom spent more than a few days. Be 257 Thus although eggs may occasionally be laid in February and March, other observations and collections suggest that most egg laying occurs from late April through May and June and that most young have fledged by the end of July or early August. Young have been seen in the nest as late as mid-September (1964) but evidently few nests are active in this month, or in following months until the beginning of another breeding season. Nest Sites: On Laysan, clumps of Eragrostis are used almost ex- clusively for nest sites. In both 1913 and 1923 however, when far less Eragrostis was available than earlier or at present, birds were seen building nests in holes in piles of phosphate rock. In 1923, one bird built a nest in one of the old buildings, inside one of the windows against a board (Wetmore, ms.) and in 1902 Fisher found one or more nests in Chenopodium bushes. Apparently only Eragrostis is now used for nest sites. Crossin (POBSP) has given the best recent survey of nest sites. He noted that: in the vast majority of cases the sites chosen were the dense portions of the grass clumps where the old blades had fallen down and [where] new green blades overhung. The dense mats of dead grass blades usually formed the foundations for the nest bottoms. All nests were...par- tially to completely hidden [from] view by overhanging grass blades. A few nests were built entirely within the mass of dead overhanging leaves within a clump. Grass clumps growing adjacent to old Scaevola bushes seemed to be especially favored; one such clump contained three old nests and an active one with one egg. In open grassy areas away from Scaevola the birds choose either very dense single clumps, or much more often, the area between two immediately adjacent clumps of grass where leaves from both plants form a dense canopy and an abundance of fallen dead blades between the two. Seven nests found by Crossin varied from 4 to 17 inches (X = 13.1) above the ground but he noted that the nest height was invariably dictated by the size and form of the grass clump with all nests found in the most secluded portion of any respective clump. Fisher (1903a: 805), the only observer to report nest sites in any detail, noted that nests in grass clumps were "in each case...wedged in the center of the tussock, well hidden by the tall grass stems." The Nest: The only observer who published a description of the nest was Fisher (1903a: 85) who reported that "it is made of rootlets, twigs, and coarse grass, and the whole structure is rather loosely put together. The sna Tow cup is 2-3/4 inches in diameter and is tied with shredded grass. Crossin, who has observed more nests closely than any other observer, noted that "nest dimensions were markedly uniform as was the nature of the 258 nest material.” Outside heights of the seven nests ranged from 2-1/4 to 3 inches (X = 2.7), and outside widths ranged from 4-1/2 to 6 inches (x= Bel)he The ranges and mean bowl depth and width for the seven nests were, respectively, 1-3/4 to 2-1/4 (x = 2.1) and 2-1/2 to 3 (x = 2.9) inches. Crossin added the following remarks on the composition of the nest: In the few nests found which were just begun, long grass rootlets formed the basis for the bottom and side walls. In all finished nests dead grass blades and stems [were] interwoven among the rootlets and the entire structure -ee| was] composed of these plant portions. There [was] essentially no cup lining, the entire cup portion being constructed of smaller and finer grass blades and root- lets. The finished structure is...compact and the sur- rounding grass blades allow the nest to remain in place for long periods. Eggs: Fisher (1903a: 805) described the eggs at some length. Clutch size, despite Fisher's (op. cit.) pronouncement that "three eggs are laid” ranges from 2 to 4 eggs, with 3 most often found. Introductions Laysan Finches were successfully introduced to Midway in 1891 and following years and survived until 1944 after the accidental introduction of rats. Recently (in March 1967) birds were introduced to Southeast Island, Pearl and Hermes Reef, by BSFW personnel and are now well estab- lished there. Laysan Finches breed successfully in captivity and the survival of the species seems assured. A statement by Dill in 1911 (1912: 22) seems appropriate: "One of the last birds to disappear from the island will be the Laysan Finch....Laysan Island is an ideal place for this bird, but should anyone be rash enough to introduce it to a civilized community it would be a pest that would rival the English Sparrow." However, several importations to Honolulu were unsuccessful, perhaps due to some insect=borne bird disease (cf. Warner, 1968: 109-110). Specimens One hundred ninety specimens of Laysan Finches are currently distribu- ted in museums as indicated in Table LF-l1. Eight additional mounted specimens are distributed as follows: AMNH (1 male and 1 female in Laysan exhibit); BPBM (2 females); DMNH (1 male and 1 female in Laysan exhibit); SUI (2 birds in Laysan exhibit). Also preserved are at least 26 skeletons (BPBM, 1; UMMZ, 20 aviary birds; USNM, 5); 2 alcoholics (USNM); 4 nests (BPBM, 1; USNM, 3); and 5 clutches of eggs (BPBM, 2; USNM, 3). Banding Five hundred sixty-six Laysan Finches were banded by BSFW personnel (Table LF-2). Table LF-1. Locations of Laysan Finch skins. Museum Males Females Other Totals AMNH 16 23 y 43 BPBM 16 15 6 37 CMNH 1 a ) 2 MCZ t 2 0 9 SUL iL 0 2 3 UMMZ 0 0 14 Vyxx USNM (non-POBSP) 34 33 6 73 (POBSP) 2 2 0 4 Other* 3 2 0 5 Totals 80 oh 32 190 *Yale Univ. (1 9); Hachisuki (1); Moseley (10°); Law Coll. (1d, 19). **Includes 2 aviary skins. Table LF-2. lLaysan Finches banded on Laysan by the BSFW. Number Banded Immature and Period of Survey Males Females Unknown Totals 1964 March iL 0 0 a Sept. 16 11 (0) 27 1966 March 96 139 9 aby Sept. O @) 1 iL 1967 Sept. 20 0 @) 20 1968 March 33 29 2 64 Sept. 38 27 44 209 Totals 2 2 15 5 Table LF-3. Observations of Laysan Finches on Laysan. Population Date of Survey Estimate Breeding Status, Remarks, References 1828 24 Mar. rather common Vague description by Isenbeck (in Rothschild, 1893-1900: vi). 1891 16-27 June common Eggs and small young (Rothschild, 1893-1900: 5). 260 Table LF-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1894 26 Nov., 2 Single specimen collected (MCZ). 8 Dec. 1896 24 June- qi Breeding (Schauinsland, 1899: 101). 24 Sept. 1902 16-23 May ? Everywhere; several nests found, all with fresh eggs (Fisher, 1903a: 804- 805). 22-23 May ? 2 nests with fresh eggs (1 with 2 eggs, 1 with 3 collected) (USNM). 1903 17 Apr.- % At least 28 skins and specimens col- May lected by W.A. Bryan or his associates (AMNH, BPBM), including a nest and egg on 10 May. 1904 10 May ee 1 collected by Schlemmer (MCZ). 1905 19 Sept. z Present (Wilder, 1905: 393). 1907 16-17 May ? 5 collected by Schlemmer (MCZ). 1911 24 Apr.- 2,700 Many nests with fresh eggs found in 5 June May (Dill and Bryan, 1912: 22). 1912 12 Dec.- ? "Abundant;" first egg on 11 February; 1913 11 Mar. another on 3 March and a set of 3 on 10 March (Bailey, 1956: 123). TESS) strona 4, 000 "All parts of island" (Munter, 1915: 140). 1916 9 Feb. very common Many singing (Munter, ms.). 1918 8-10 Sept. ? Found everywhere; most common around old buildings and at southwest end of island (Diggs, ms.). 1923 8-13 Apr. few One building nest inside window of house on 8th (Wetmore, ms.). 20-23 Apr. numerous Some seen nest building (Ball, ms.; Dickey, ms.). 261 abundant Table LF-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1923 29 Apr.- 100 Four introduced from Midway on 30 14 May April (Wetmore, ms.); some nest build- ing by 20 April (Ball, ms.); complete clutch (3 eggs) 10 May; several pairs nest building 12 May (Wetmore, ms.) and nest with fresh eggs collected (USM) . 1924 6 May only a few (Wilder, ms.). 1930 2=18 Aug. many (Wilder, ms. b). hundreds 1936 7-8 Mar. many No nests found (Trempe, ms.). ; hundreds 12 Dec. 1, 000 (Coultas, ms.). 1950 23 June quite (Brock, 1951a: 372). abundant 1951 late June- 5,059 Estimated from transect censuses early July (Brock, 1951b: 18). 1955 10 Feb. thousands (POFT). 1957 25 June- 5,000 (Woodside, ms. b). 3 July 8-12 July everywhere (Labrecque, 1957: 17). 1958 27 May- 10, 100 Estimate based on transect census and 4 June believed to be conservative. Eggs in all stages of incubation to week old young (Warner, ms.). 1959 28 Apr.- ? Most paired but no nests found (Kramer, 1 May ms.). 1961 7-8 Mar. very More than in 1959; no nests found abundant (Woodside and Kramer, ms.). 4-10 Sept. 10,000 No active nests; several recently fledged immatures (Woodside, ms. c). 1962 14-19 June very (Kramer and Beardsley, ms.). 262 Table LF-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1963 11-13 Feb. 10, 000 Wo nests found (POBSP). 3-10 Dec. ? (Walker, ms. b). 1964 10-11 Mar. 10, 000 No signs of nest-building noted (BSFW, POBSP). 16-20 Sept. thousands Very abundant; one near-fledged young seen being fed by parent (BSFW, POBSP). 1965 6-11 Mar. 3, 000- No nests found (POBSP). 5,000 17-21 July 10, 000 Many flying young still begging for food, about 5,000 fledged immatures (POBSP). 5-12 Aug. (40,000)* About 10,000* fledged immatures. One young seen being fed by parent on 6 August (POBSP). 1966 26-31 Mar. 7,400 Estimate based on transect census; no breeding noted; series captured (BSFW). 10-16, 20-21 June 10, 000 New nests through fully fledged young; estimated (15,000)* young; 1,000 eggs (POBSP). 17-18 Sept. very (BSFW). abundant 20-23 Oct. 10, 000 No nests found (POBSP). 1967 18-19 Mar. 4, 000- No nests found (BSFW, POBSP). 5,000 7-12 June 10, 000 Series captured for transplantation. Eggs to flying immatures (POBSP). 5-11 Sept. several Abundant; post-breeding; several young(?) thousands seen begging for food (POBSP). 13 Dec. very (BSFW). common *Estimate probably excessive. 263 Table LF-3. (continued) Population Date of Survey Estimate Breeding Status, Remarks, References 1968 17-19 Mar. several No nests found despite investigation thousand of several hundred potential nest sites (BSFW, POBSP). 1969 26-29 Mar. 11,882 Estimate based on transect census. Over 500 potential nest sites checked but no nests found (BSFW). 2-3 June % 365 Eragrostis clumps investigated: 9 nests found: 6 apparently used pre- viously, 1 with a pre-laying bird; 1 with 1 egg and 1 with 1 young (BSFW). 9 Sept. abundant (BSFW). Mammals RABBIT Orytylagus cuniculus (EUROPEAN HARE) Status Introduced in 1903 and subsequently. Extirpated in 1923. Observations Dill and Bryan (1912: 9) reported that rabbits were introduced to Laysan about 1903 but Bryan later (1915: 293) indicated that these animals were introduced in 1903 and 1904. Most secondary sources repeat the former statement but Tomich (1969: 30) recently stated that the rabbits were intro- duced in 1902 and 1903. Presumably this conclusion is based on Dill and Bryan's statement that Max Schlemmer "could not give the exact dates {of introduction] but thinks the first were imported eight or nine years ago, or about 1903." However, W.A. Bryan (in Dill and Bryan, 1912: 26) clearly indicated that the rabbits were introduced "shortly after my former visit" (April-May 1903). In any case, these animals were brought to Laysan on more than one occasion (Dill and Bryan, op. ns ic The rabbits, which were said to have been imported for the purpose of starting a rabbit-canning business, consisted of domestic rabbits, Belgian and English Hares. They prospered greatly and began to cause serious damage to the vegetation (Table R-1). By 1911 the rabbits had eliminated several species of plants that had been present in 1903. - 264 Dill and Bryan's report of their effect eventually resulted in two expeditions which were sent to destroy the rabbits. The former, which spent three months on Laysan in the winter of 1912-1913, killed many thousands but failed to exterminate them, largely due to a lack of ammu- nition (Salisbury, ms.). The second expedition, the Tanager Expedition of 1923, arrived too late to do little more than apply the coup-de-grace as the animals had nearly extirpated themselves by having eaten almost all of their potential food saurces. Table R-1l. Observations of rabbits on Laysan. Period of Observation Observations 1910 16-18 Jan. "A large number of rabbits was found there, and if they should increase to a much greater number, I am of the opinion that they would destroy the vegeta- tion and then attack the birds' eggs. ‘The rabbits should be exterminated." (Jacobs, ms.). 2 Sept. "The rabbits...are increasing very rapidly and will no doubt disturb the birds in time if they do not do so now." (Cochran, ms. c). 1911 24 Apr.- Many bushes killed. Often seen feeding in the green 5 June juncus (=Sesuvium portulacastrum) near the lagoon where at times "...there are so many ears protruding that they resemble a vegetable garden." Stated to have been captured by frigate birds (Dill and Bryan, 1912: 10). Oe 22 Apr. ",.eapparently no material increase in the number of rabbits." (Cochran, ms. a). 1912 22 Dec.- Found everywhere and often utilizing petrel burrows NOS ees Max. for shelter. "Each chenopodium bush would shelter half a dozen, and even the foundations of the build- ings were undermined; each clump of grass contained a warren, and out on the open flats could be seen dozens of bunnies feeding on grasses pushing their way through cracks in the phosphate rock." (Bailey, 1942: 154). 100 killed first day on island. 5,024 killed during entire visit but several thousand re- mained unkilled (Salisbury, ms.). 1914 11 Sept. "found the number of rabbits greatly decreased" (Log of the Thetis). "There seems to have been a decrease in the number of rabbits..." (Brown, ms. c). Elschner (1915: 30), however, indicated that the earlier at- tempt to destroy them "seems to have been rather un- successful. Table R-1. Period of Observation 1915 1916 1918 1923 3 Apr. 6 Feb. 8-10 Sept. 7 Apr.- 14 May (continued ) 265 Observations "The rabbits were found to be very plentiful. They were seen wherever green patches existed. Twenty of them were caught and taken off to the ship for food. About 15 of them were found dead near one of the buildings. They are rapidly eat- ing off the vegetation of the island." (Munter, 1915: 140). "The rabbits...are multiplying very fast and the vegetation is disappearing rapidly....The men were set to work catching rabbits. Twenty only were captured as they were very active and easily es- caped pursuits." (Munter, ms.). "The domestic breed, reddish brown, gray, and white were found in number in and around their burrows among the stones, shrubberies and green juncus growing near the lagoon. ‘They could be run down easily and caught in the hands or could be chased into small caverns among the stones or rocks and caught as they seemed to have little endurance when on the run....Only in one spot, and that near the Southwest end of the lagoon, was there any no- ticeable amount of dead shrubbery and no rabbits at all were to be found at any time during our stay feeding upon this plant....The green juncus growing lower on the ground was quite fresh and seemed to cover an immense area around the lagoon. Here and [there] upon this the rabbits were seen feeding at times. These rabbits,...while numerous in many parts of the island, were not by any means found to be as plentiful as might be expected when consider- ing the length of time left unmolested in which to breed. Only in one case was a young rabbit seen and he was caught and taken aboard....we attempted to estimate the number of rabbits on the island, which we figured to be not more than a hundred at the most liberal estimate, twenty five of these which we caught ourselves for food...." (Diggs, ms.). A few hundred rabbits present when the expedition arrived (Wetmore, 1925: 103). "Low areas here [near the lagoon]...were covered with a mat of Sesuvium...that was making a brave struggle against the depredations of the rabbits. The latter hopped or squatted about among the albatross and 266 Table R-l1. (continued) Period of Observation Observations 1923 7 Apr.- shearwaters occasionally taking alarm and dashing 14 May away to run down a hole. About 150 to 200 have been killed here since our arrival. All are of large 'Belgian Hare' size and vary- ing color. Grizzled grays and grayish browns pre- dominate with numbers of reddish brown, dull black, and blackish brown. All had the abdomen well dis- tended two or three that were examined were not at all fat though in good flesh. I noted scattered carcasses of animals that had died before our ayn- rival....All rabbits seen were adult but Reno found embryos in several females." (Wetmore, ms., 9 April). "When we left May 14 no rabbits had been seen for the past 8 days."* 1924 5 May No rabbits noted (Wilder, ms. a). *Extract from Major Reno's report on the success of the rabbit extermination. Ms. material in the files of the Bureau of Sport Fisheries and Wildlife, Honolulu. GUINEA PIG Cavia porcellus Status Introduced in 1903 and/or 1904 and once common. Extirpated during the period from December 1911 through March 1912. Observations According to W.A. Bryan (1915: 293), guinea pigs were introduced to Laysan at the same time as the rabbits. Subsequent information on their habits on Laysan is scant. Dill and Bryan eee 10) reported that guinea pigs were rather abundant in the thick juncus (=Cyperus) at the south end of the island during the spring of 1911. The guinea pigs evidently fared poorly in competition with the rabbits for only four were seen, all of which were killed, during the visit by the Biological Survey party in 1911-1912 (Salisbury, ms.). EE 267 BOTTLE-NOSED DOLPHIN Tursiops truncatus Status One sighting offshore: May 1958. Observations Rice (1960a: 407) reported that a half-dozen were seen 27 May 1958 from shipboard about three kilometers southeast of Laysan. Bottle-nosed Dolphins are predominantly an inshore species (Rice, op. feast) and probably occur more commonly offshore Laysan than the single record might indicate. HAWAIIAN MONK SEAL Monachus schauinslandi Status Common resident present throughout the year. Maximum recent esti- mates: 326 for an aerial count; 314 for a terrestrial count. Observations Tables HMS-1 and HMS-2 briefly summarize earlier and more recent observations of monk seal populations on Laysan. Kenyon and Rice (1959) present a thorough summary of much of what is known of the life history. The paucity of observations in the late 1800's and early 1900's as well as the scant numbers seen indicate that seals had been nearly extir- pated on Laysan, primarily by sealers, by about the turn of the century (Kenyon and Rice, 1959: 215). Not until the latter half of the 20th century did populations show much recovery. Recent population counts, while variable, seem to indicate a relatively stable population with a breeding population on the order of 140 to 160 animals. Counts from 1966 through 1969 tend to be somewhat lower than for preceding years but we think it likely that this change can be largely at- tributed to the effect on counts of the greater incidence of disturbance during the latter period. Variations in age-class terminologies used by different observers make it difficult to assess yearly production of offspring but the data suggest that some 50 to 80 pups are born yearly. These young are born from at least as early as the first week of January through at least the end of June with most being born in the period from mid-February through May. A considerable number of monk seals has been tagged on Laysan (Appendix Table 2). One of these animals, tagged as a pup on Laysan on 18 March 1968, was later reported present on Johnston Atoll from late July through early December 1968 (Schreiber and Kridler, 1969: 842). Details of the tagging program as well as more detailed recent information on the life history of the seal is to be presented at a later date by the BSFW. 268 Table HMS-1. Observations of Hawaiian Monk Seals on Laysan prior to 1951. Date of Survey Remarks and References 1828 2h Mar. "On the beach several small Seals...were found" (Isen- beck in Rothschild, 1893-1900: vi). 1857 1 May Seals numerous (Paty, 1857: 40). 1896 24 June- A skull, skin, and parts of two other skulls and skins 24 Sept. were given to Schauinsland by Schlemmer (Bailey, 1952b: 4), this material later serving as the basis of Matchie's (1905) deseription of the species. 1911 24 Apr.- No seals seen (Dill and Bryan, 1912: 9). 5 June 1912 22 Dec.- A single seal found on the north end of the island on 1913 11 Mar. 30 December was collected by Willett. No others were seen (Bailey, 1952b: 6). 1915 July-Sept. 5 July: single seal seen at south end of island was shot for oil. 15 August: seal seen on beach. 22 August: one seal seen in trip around island. 24 August: a seal killed. 14 September: another seal killed (Schlemmer and Schlemmer, ms.) 1923 7 Apr.- A male and a female collected and another pair seen 14 May (Dickey, ms.). 1936 7-8 Mar. "Right or ten were seen in the water or on the sand.” (Trempe, ms.). ca. 12 Dec. 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UT Semstq_ °sdnd ynq sesseto ase TayjO [Te seumsqns sanSTF stuL I “ssuUTTZeek pue sa—npeqns yI0q Sepn[ouT [e074 4IMpeqns *(MASd) Wig 9 = - oO Gig LCO= BOgL Ji = 408s 16 *peaiunoo S[TeuLue JO Taqumu [e309 9a. go ale Saseyueoteg °*sTeas paseum 6g Sepntout qunop “*(MaSd) (6°ST) 62 (ee) et (6°T8) 641 TI2 anf g 696T Seouetejey pue sHreuey “ON = Paqtmoo yuMoD Fo eyed sdng SSUT [Leon syrnpeqns SvINpYy sTeeas gO °ON (penutquod) *¢g-SWH eTAeL eT DOMESTIC HORSE Equus caballus Status A hypothetical introduction; see accounts of Donkey and Mule, below. DONKEY Equus asinus Status Hypothetical; may have been present from about 1905 through 1910. Observations Wilder (1905: 392) noted that "one old donkey” was present in September 1905. Cochran* implied that this or another donkey was removed from Laysan by Schlemmer in 1910. Possibly this animal is the same as the mule removed from the island in July 1910 (see below). MULE Equus asinus X Equus caballus Status Introduced ca. 1891 by the guano company. Last mule removed from island ca. July 1910. Observations The guano company brought mules to Laysan to pull carts of guano to the loading wharf. Munro (1946: 60) indicated that mules were present in June 1891. Since the tramway had not been completed by the preceding November, presumably the mules were introduced early in 1891. Their occurrence on Laysan was noted on several subsequent occasions. Thomas (ms.) saw "a small herd" of mules during his visit in May 1902. Jacobs (ms.) recorded that two "almost wild" horses (probably mules) were present in January 1910. The Honolulu Evening Bulletin for 6 August 1910 reported that the single surviving mule was removed from the island on or about July 1910 by the crew of the schooner Concord. Whether this mule was the donkey referred to above and whether Schlemmer was along during this visit is not known but seems at least likely. ¥Letter from C.S. Cochran to the Secretary of the U.S. Treasury, dated 8 September 1910. Record Group 26, U.S. National Archives. ae 275 PIG Sus scrofa Status Introduced by the guano company about 1890; evidently not present for more than a few years. Observations In June 1891 Munro (1946: 60) noted that "A few hogs roamed around, feeding on the dead albatross..." F.D. Walker (1909: 30) also noted that pigs were present during this visit and that they were known to feed on the tubers of a "false yam" which Tomich (1969: 79) has identified as Boerhavia. Two pigs were still present a little more than two years later (Farrell, 1928: 399) but none was noted as present in May 1902 (Thomas, ms.). DOMESTIC CATTLE Bos taurus Status Introduced in the early 1900's; subsequently died, or were removed from Laysan. Observations Only two observers mention the presence of cattle. In May 1902, Thomas (ms.) noted that several cows, kept for the use of Schlemmer, were present. Subsequently Wilder (1905: 392) reported that "a few milch cows" were present in September 1905. Probably these animals were removed from the island during the next few years as none was present in January 1910. Reptiles GREEN TURTLE Chelonia mydas Turtles, presumably this species, were first recorded on Laysan co- incident with the first known report of the island by Europeans in 1828 (von Kittlitz in Rothschild, 1893). Other early observers (e.g. Brooks, 1859) usually reported their presence but indications of the number present were rarely given. Numbers of turtles seen on Laysan by different observers are summarized in tabular form below. Only those observations which give some idea of their abundance or breeding status are included here. Laysan Green Sea Turtle populations have clearly decreased gradually and steadily throughout the last two centuries, certainly as a direct result 276 of predation and disturbance by man. From a formerly abundant resident of the island, these turtles have become uncommon to rare and apparently are in distinct danger of becoming extinct at this breeding station. No recent observers, either POBSP survey teams or those of the U.S. Fish and Wildlife Service, have found any direct evidence that the species still breeds on Laysan although Woodside (1961) found some "nests" in September 1961, none of which, when examined, contained eggs. Table GT-1. Observations of Green Turtles on Laysan.* Number Date of Survey seen Remarks and References 1828 24 Mar. # Some very large turtles seen (Rothschild, 1893-1900: vi). 1857 1 May numerous (Paty, 1857: 40). 1858 14 Jan. B 6 small turtles killed (Log of the U.S.S. Fenimore Cooper). 1882 26-30 Jan. ? 104 turtles taken by crew of fishing schooner Ada (Hornell, 1934: 432). 3 May i 26 turtles taken by crew of Ada (Hornell, 1934: 432-433). 1886 ca. late % Some turtles killed (Farrell, 1928: 253- Sept. 254) by crew of schooner General Siegal. 1896 2h June- ? Numerous on Laysan's coasts; often in 24 Sept. whole schools (Schauinsland, 1899: 64). One female captured contained several hundred eggs. 1905 19 Sept. ? A few turtles shot (Wilder, 1905: 392). 1911 24 Apr.- ? 1 turtle killed for food (Dill and Bryan, 5 June 1912: 421). 1912 22 Dec.- 2 "Turtle appeared occasionally;" [1 or more 1913 11 Mar. killed for food]. (Salisbury, ms.). 1915 3 Apr. ? Decaying turtle meat found in building (Munter, 1915: 138). [Turtles evidently killed for food by Japanese]. *Many early reports do not specifically identify turtles seen on Laysan as Green Turtles. However, as the Hawksbill and Ridley are not known to breed in the Northwestern Hawaiian Islands, we feel it a safe assumption that early records referred to the Green Turtle. 277 Table GI-1. (continued) Number Date of Survey seen Remarks and References 1915 12-31 July 4 Turtles captured for food on 14 July (1 small), 15 July (2), 27 July (1) (Schlemmer and Schlemmer, ms.). 1-31 Aug. 2 Turtles killed or captured on 7 August (1) and 31 August (1) (Schlemmer and Schlemmer, ms.). 1-31 Oct. 3 Turtles killed or captured 11th (2 small, 1 large) (Schlemmer and Schlemmer, ms.). 1-30 Nov. 5 Turtles "turned over" and presumably captured on 8th (2 large); others cap- tured 9th (2) and 22nd (1 small) (Schlem- mer and Schlemmer, ms.). 1918 8-10 Sept. in Many caught by crew of Hermes (Diggs, ms.). abundance 1923 8 Apr. at least 5 seen on west shore; largest thought to 50 weigh over 200 lbs. (Dickey, ms.; Ball, ms.). 10 Apr. 5) Seen on west shore; 1 ca. 50 lb..turtle cap- tured for food (Dickey, ms.). 3 May 2 Seen along beach (Wetmore, ms.). 8 May 3 Small, 1 captured for food (Wetmore, ms.). 1934 26 June 2 A few large turtles seen (Baylis, ms.). 1936 7-8 Mar. 10-12 Seen along eastern beach (Trempe, ms.). 12 Dec. a (Goultas, msi). 1950 23 June Gap 10) (POFI). Some presumably tagged (cf. Brock, 1951a: 371). 1951 12 May % Turtles numerous along northeast, north, and west side of island. No tagged turtles seen (POFI). 1954 3 Nov. at Seen; medium-sized (POFI). 1961 7 Mar. 6 Seen along beaches. All 40 lbs. or more but no evidence of egg laying (Woodside and Kramer, ms.). 278 Table GI-1. (continued) Number Date of Survey seen Remarks and References 1961 4-10 Sept. 3-4 2 females tagged. 5 nest sites examined but no eggs found (Woodside, ms. c). 1963 11-13 Feb. 6 All seen were tagged by Kramer who saw no small turtles and no signs of egg laying (POBSP). 3-10 Dec. 3 Counted 3 December. A male and 2 females tagged 4 December (Walker, ms. b). 1964 10-11 Mar. aE seen on southeast beach 10 March. About 2 feet long (BSFW, POBSP). 16-20 Sept. 2 Seen close to shore on 16 September (POBSP). 1965 6-11 Mar. 2 Seen on southwest beach 7 March; one, a female, had been tagged in December 1963 (POBSP). 5-12 Aug. 2 Seen offshore west beach (POBSP). 1966 26-31 Mar. 2 Females (BSFW). 10-16, Females seen along beaches; 1 ca. 18". 20-21 June 3 2 ca. 36" (POBSP). 17-18 Sept. 2 Females; 1 tagged, the other had been tagged February 1963 (BSFW). 20-23 Oct. 2 Large females counted 21 October. Neither previously tagged (POBSP). 1967 18-19 Mar. 2 1 male tagged (BSFW). 7-12 June nh 3 medium-sized, 1 large (POBSP). 21-24 Sept. iL 1 male tagged (BSFW). 13 Dec. 6 All females, 2 of which were returns. 1 had been tagged in February 1963, and was the same turtle that returned in September 1966 (BSFW). 1968 17-19 Mar. 5 4 (3 females and a male) newly tagged and 1 returned (the male tagged in September 1967) (BSFW). 279 Table GI-1. (continued) Number Date of Survey Seen Remarks and References 1969 26-29 Mar. 2 Counted 26 March. Both small, tagged. 6 others seen swimming offshore (BSFW). 2-3 June 3 Counted 2 June (BSFW). 9 Sept. O 9 nest sites found, 1 along south shore and 2 groups of 4 on northwest corner. 1 site investigated contained no eggs (BSFW). FOX GECKO Hemidactylus garnotii The first report that the Fox Gecko occurred on Laysan was Werner's (1901: 382) statement that this species occurred there, apparently on the basis of collections made by Schauinsland in 1896. Snyder (1917) later reported that eggs had been found on Laysan, presumably by himself in May 1902. Subsequently, Willett (ms.) indicated that this species was oeca- Sionally seen during the visit of the Biological Survey party from December 1912 to March 1913. No more recent observers have reported this large gecko (or, for that matter, any gecko) from the island. It seems likely that the Fox Gecko was accidentally introduced by early visitors to the island and has since become extirpated. SNAKE-EYED SKINK Ablepharus boutonii The earliest report of the occurrence of this lizard on Laysan was by Werner (1901: 385), who examined Schauinsland's reptile collections. Snyder subsequently observed that, of 10 specimens collected there (pre- sumably in May 1902), none had uninjured tails. He attributed these deformities to predation on the skinks by the birds (Snyder, 1917). Judging from field notes taken by the POBSP, skink populations on this island periodically exhibit great variation in size. On surveys from February 1963 through August 1965, none or very few skirkswere seen, most being found in local concentrations in some of the rock formations. In mid-June 1966 the skinks were evidently somewhat more abundant Since some 22 individuals were collected in various locations around the island. During this survey a nest containing 23 eggs was found in a shallow depression beneath an old lumber pile. By the following October these liz- ards were abundant enough so that POBSP observers reported them as "very numerous." POBSP specimens: 3, September 18-19, 1964, USNM 157670-157672; 21, June 13, 1966, USNM field numbers 10897-10917. 280 ACKNOWLEDGMENTS Field work on Laysan was made possible by a co-operative agreement between the Department of the Interior, Bureau of Sport Fisheries and Wildlife, and the Smithsonian Institution. Special appreciation is par- ticularly due Mr. Eugene Kridler, Refuge Manager of the Hawaiian Islands National Wildlife Refuge, who not only granted us access to the islands but also allowed us the use of a massive amount of reports and notes largely compiled by him during his many visits to Laysan from 1964 through 1969. He also allowed many POBSP personnel to accompany him on the regular inspections of the refuge conducted under his leadership by the Bureau of Sport Fisheries and Wildlife. Mr. Michio Takata, Hawaii Division of Fish and Game, kindly made available reports and data collected by his personnel. We are also grateful to Mr. Ronald L. Walker, Mr. David H. Woodside, and Mr. Raymond J. Kramer for the generous use of their field notes and data. Mr. Edwin H. Bryan, Jr., Manager of the Pacific Science Information Center, Bernice P. Bishop Museum, made an invaluable contribution by granting us access to his voluminous reference files and spent a con- siderable amount of time answering queries by POBSP personnel. Margaret Titcomb, Librarian of the same institution, was helpful in uncovering various manuscript materials pertaining to the 1923 visit of the Tanager Expedition. Others who aided in finding relevant historical material and to whom we owe thanks include: Thomas R. Howell, who loaned as a copy of the field notes kept by Donald R. Dickey during the Tanager Expedition; Eric Laysan Schlemmer, who loaned us a diary kept by his father and himself during an extended stay on Laysan in 1915, as well as other historical materials; Jean Dabagh of the Hawaii State Archives, Honolulu. A great many other individuals, particularly those at the U.S. National Archives, were of considerable help in uncovering previously little known manuscript material. Dr. Alexander Wetmore very courteously allowed us full use of the i valuable and detailed field notes that he made as leader of the 1923 . Tanager Expedition. These notes contained a wealth of unpublished informa- j tion that has contributed much to both the biological and historical sections of this account. Dr. Wetmore loaned us photographs used in this account as well, and both he and E.H,. Bryan, Jr., were kind enough to comment on the history section. Special acknowledgment is due Dr. Philip S. Humphrey, principal investigator of the POBSP, whose foresight and optimism made the POBSP a reality. The following POBSP personnel participated in field work on Laysan: Kenneth E. Amerman, A. Binion Amerson, Jr., Allen H. Anderson, Kenneth C. Balcomb, F. Allen Blagden, David L. Burkhalter, Richard D. Chandler, Roger B. Clapp, Richard S. Crossin, Robert L. DeLong, Charles A. Ely, Robert R. Fleet, Patrick J. Gould, C. Douglas Hackman, Brian A. Har- ‘ rington, J. Vincent Hoeman, David I. Hoff, Dayle N. Husted, T. James Lewis, C. Robert Long, Robert W. McFarlane, Richard W. Merrill, David L. Pearson, 281 Philip C. Shelton, Fred C. Sibley, Frank H. Smith, Jr., Dennis L. Stadel, Jeffrey P. Tordoff, F. Christian Thompson, Robert W. Tuxson, J. Douglas Whitman, William 0. Wirtz II, George S. Wislocki, and Paul W. Woodward. Among our co-workers, we should especially like to single out for thanks A. Binion Amerson, Jr., Philip C. Shelton, and Paul W. Woodward, each of whom has recently had to struggle with long and sometimes re- calcitrant manuscripts, and whose comments on various sections of the manuscript were both enlightening and helpful. Jane P. Church and Mae H. Esterline aided in editing and proofing various stages of the manuscript. Figure 5 was redrawn by Tina Clapp. Transportation to Laysan was provided by the U.S. Military Sea Trans- port Service, the U.S. Navy and the U.S. Coast Guard. We wish to thank the various individuals who provided this support and especially the officers and men of the vessels involved. The following U.S. Navy personnel assisted in field work on Laysan: Ronald R. Amerson, Edward King, and Charles Williams, Jr. The examination of Laysan specimens was made possible through the co- operation of the curators of the foilowing museums: American Museum of Natural History (Dean Amadon); Bernice P. Bishop Museum (Edwin H. Bryan, Jr.); Chicago Natural History Museum (Austin L. Rand); British Museum (Natural History) (Ian Galbraith); Denver Museum of Natural History (Alfred M. Bailey); J.E. Law Collection (John P. Hubbard); Museum of Comparative Zoology (Raymond A. Paynter, Jr.); Museum of Natural History, State Univer- sity of Iowa (George Schrimper and Walter C. Thietje); Academy of Natural Sciences of Philadelphia (James Bond); University of Michigan Museum of Zoology (Robert W. Storer); United States National Museum (Richard Zusi). For collection and compilation of botanical data we acknowledge the efforts of C. Robert Long and Charles Lamoureux (University of Hawaii). The camera copy of the manuscript 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, 1971). 282 LITERATURE CITED Agassiz, A. and H.L. Clark. 1907-1912. Hawaiian and other Pacific Echini. Mus. Comp. Zool. Harvard, Mem. 34: vii and 383 pp. Alfken, J.D. 1903. Beitrag zur Insecten fauna der Hawaiischen und Neuseeldndischen Inseln...Schauinsland 1896-97. Zool. Jahrb., Syst., Jena 19: 561-628. American Ornithologists' Union. 1957. Check-list of North American birds. 5th ed. Lord Baltimore Press, Baltimore. xii and 691 pp. Amerson, A.B., Jr. 1966. Ornithodoros capensis (Acarina: Argasidae) infesting Sooty Tern (Sterna fuscata) nasal cavities. J.. Parasit. 52: 1220-1221. ---1968. Tick distribution in the central Pacific as influenced by seabird movement. J. Med. Ent. 5 (3): 332-339. Anonymous. 1905. An island schooner's bunch of disasters. Paradise of the Pacific 18 (9): 17-18. ---1939. Manure. The Sales Builder 12 (1): 2-22. ---195la. Hawaiian Monk Seal - marine rarity - Zoonooz 24 (7): 3. ---1951b. The Laysan Island Cyclorama and late news from Laysan. Elepaio 12: 1-2. Aoki, J. 1964. Some oribatid mites (Acarina) from Laysan Island. Pac. Insects 6 (4): 649-664. ---1965. Notes on the species of the genus Epihohmannia from the Hawaiian Islands (Acarina: Oribatei). Pac. Insects 7 (2): 309-315. Bailey, A.M. 1918. The monk seal of the southern Pacific. Nat. Hist. 18: 396-399. ---1934. Wanderers of the seas: albatrosses which nest on coral and volcanic rocks in the Pacific Ocean. Nat. Hist. 34: 273-281. ---1942,. The portulaca flats of Laysan. Aud. Mag. 44: 150-161. ---1952a. lLaysan and Black-footed Albatrosses. Denver Mus. Nat. Hist., Muse Pict 6.930) pp. ---1952b. The Hawaiian monk seal. Denver Mus. Nat. Hist., Mus. Pict. 7. 32 pp. ---1956. Birds of Midway and Laysan Islands. Denver Mus. Nat. Hist., Mus. Rigging 12, IO ii 283 Baldwin, P.H. 1945. Fate of the Laysan Rail. Aud. Mag. 47: 343-348. ---1947. The life history of the Laysan Rail. Condor 49: 14-21. Ball, S.C. (ms.). Field notebook kept during the 1923 Tanager Expedition. B.P. Bishop Museum, Honolulu. Barr, R. 1903. Hornschwdmme aus dem Pacific...Schauinsland, 1896-97. Zool. Jahrb., Syst., Jena 19: 27-36. Bassham, B.L. (ms.). Cruise report of the Reliance for March 1936. Rec. Group 26, U.S. Nat. Arch. 2 pp. Baylis, J.S. (ms.). Cruise report for the Itasca for the month of June 1934. Rec. Group 26, U.S. Nat. Arch. 6 pp. Beardsley, J.W. 1966. Insects and other terrestrial arthropods from the Leeward Hawaiian Islands. Proc. Haw. Ent. Soc. 19: 157-185. Bergh, R. 1900. Ergebnisse einer Reise nach dem Pacific (Schauinsland, 1896-97). Die Opisthobranchier. Zool. Jahrb., Syst., Jena 13: 207-246. Berry, 5.S. 1910. Diagnoses of new cephalopods from the Hawaiian Islands. Proc. U.S. Nat. Mus. (No. 1713): 407-419. Bitter, G. 1900. Die phanerogamische Pflanzenwelt der Insel Laysan. Abh. Nat. Ver., Bremen.16: 430-439. Brock, V.E. 195la. Some observations on the Laysan Duck, Anas wyvilliana laysanensis. Auk 68: 371-372. ---1951b. lLaysan Island bird census. Elepaio 12: 16-18. ee N.C. 1859. Cruise of the Gambia. Pac. Comm. Advertiser, 11 Aug. 1 59. ---1860. Islands and reefs west-north-west of the Sandwich Islands, Pacific. WNaut. Mag. 29: 499-504. Brown, J.H. (ms. a). Report to the Captain Commandant of the Coast Guard of observations made during a cruise to Laysan, Lisianski, etc., in March 1915. Rec. Group 26, U.S. Nat. Arch. 14 pp. ---(ms. b). Report to the Captain Commandant of the Coast Guard of the Cruise made to the Hawaiian Bird Reservation in February 1916. Rec. Group 26, U.S. Nat. Arch. 12 pp. ---(ms. c). Report to the Captain Commandant of the Revenue Cutter Service of a cruise to the "Bird Islands" in September 1914. Unpublished ms. material, Rec. Group 26, U.S. Nat. Arch. 4 pp. 284 Bryan, E.H., Jr. 1932. Notes and exhibitions. Proc. Haw. Ent. Soc. Crier ---1942. American Polynesia and the Hawaiian Chain. Tongg Publ. Co., Honolulu, 253 pp. ---1954. The Hawaiian Chain. B.P. Bishop Mus. Press, Honolulu. iii and 71 pp. ---1958. Check list and summary of Hawaiian birds. Books about Hawaii, Honolulu. 28 pp. Bryan, E.H., Jr. and J.C. Greenway, Jr. 1944. Contribution to the orni- thology of the Hawaiian Islands. Bull. Mus. Comp. Zool. Harvard 9}: 79-142. Bryan, E.H., Jr. et al. 1926. Insects of Hawaii, Johnston Island and Wake Island. B.P. Bishop Mus. Bull. 31: 94 pp. Bryan, W.A. 1901. Key to the birds of the Hawaiian group. B.P. Bishop Mus. Mem. 1: 259-332. ---1911. Laysan Island, a visit to Hawaii's bird reservation. Mid-Pac. Mag. 2: 303-315. ---1912. The introduction and acclimization of the Yellow Canary on Midway Island. Auk 29: 339-342. ---1915. Natural history of Hawaii. Hawaiian Gazette Co., Ltd., Honolulu. 596 pp. Bureau of Sport Fisheries and Wildlife, U.S. Fish and Wildlife Service, Kailua, Hawaii (unpublished reports and notes): Kridler, E. personal correspondence, 1968-70. ~ ---1964. [Report on] Hawaiian Islands National Wildlife Refuge... [survey]...September 16 through 27, 1964. 31 pp. ---1966. Hawaiian Islands National Wildlife Refuge trip report. 2 pp. ---1966. Hawaiian Islands National Wildlife Refuge trip - September 8-28, 1966. 34 pp. ---1967. Refuge log book for 6 March - 1 April 1967. 39 pp. ---1967. Refuge log book for 19-29 September 1967. 12 pp. ---1967. Hawaiian Islands National Wildlife Refuge...Preliminary trip report - December 7 through 18, 1967. 13 pp. ---1969. Hawaiian Islands National Wildlife Refuge spring trip. March 19 - April 6, 1969. 32 pp. 285 Olsen, D.L. 1969. Hawaiian Islands National Wildlife Refuge Field trip, May 28 - June 11, 1969. 18 pp. Kridler, E. 1969. Hawaiian Islands National Wildlife Refuge Field trip report, August 19 - September 23, 1969. 32 pp. Butler, G.D., Jr. 1961la. Insects and other arthropods from Laysan Island. Proc. Haw. Ent. Soc. 17: 379-387. ---1961b. Stratiomyiid fly associated with dead albatrosses on Laysan Island. Proc. Haw. Ent. Soc, 17: 331-332. Butler, G.D., Jr. and M.D.F. Udvardy. 1966. Basking behavior of the Hawaiian monk seal on Laysan Island. J. Wildlife Manage. 30: 627-628. Butler, G.D., Jr., and R.L. Usinger. 1963. Insects and other inverte- brates from Laysan Island. Atoll Res. Bull. 98: 1-30. Caspers, H. 1968. Biology of a hypersaline lagoon in a tropical atoll island (Laysan). Recent Adv. Trop. Ecol. 1: 326-333. Chapin, E.A. 1925. Descriptions of new internal parasites. Proc. U.S. Nat. Mus. 68 (No. 2603): 1-4. Christophersen, E. and E.L. Caum. 1931. Vascular plants of the Leeward Islands, Hawaii. B.P. Bishop Mus. Bull. 81. 41 pp. Clapp, R.B. 1968. Three unusual shorebirds from Midway Atoll, Pacific Ocean. Elepaio 28: 76-77. 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 F.C. Sibley. 1967. New records of birds from the Phoenix and Line Islands. Ibis 109: 122-125. 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. Clark, A.H. 1908. Descriptions of new species of crinoids, chiefly from the collections made by the U.S. Fisheries steamer"Albatross" at the Hawaiian Islands in 1902....Proc. U.S. Nat. Mus. 34 (No. 1608): 209-2k4., ---1912. Notes on the Laysan Finch. Auk 29: 166-168. Cochran, C.S. (ms. a). Report to the Captain Commandant of the Revenue Cutter Service of an inspection of the Hawaiian Bird Reservation in April and May 1912. Rec. Group 26, U.S. Nat. Arch. 2 pp. ---(ms. b). Report to the Captain Commandant of the Revenue Cutter Service of a cruise to the Hawaiian Bird Reservation in December 1912. Rec. Group 26, U.S. Nat. Arch. 4 pp. 286 ---(ms. c). Report to the Secretary of the Treasury of a survey of the Hawaiian Bird Reservation in August and September 1910. Unpublished ms., Rec. Group 26, U.S. Nat. Arch. 2 pp. ---1949. Ophiuroidea of the Hawaiian Islands. B.P, Bishop Mus. Bull. 195. 133 pp. Coultas, W.F. (ms.). Notes taken on Laysan Island, December 1936. Extracts from a letter in the files of the Bureau of Sport Fisheries and Wildlife, Honolulu. 1 p. Crossin, R.S. and L.N. Huber. 1970. Sooty Tern egg predation by Ruddy Turnstones. Condor 72: 372-373. Dall, W.H., P. Bartsch, and H.A. Rehder. 1938. A manual of the recent and fossil pelecypod mollusks of the Hawaiian Islands. B.P. Bishop Mus. Bw. 1539 233) joo. Dickey, D.R. (ms.). Notes taken during the Tanager Expedition. Diggs, J.T. (ms.). Report of the cruise of the U.S.S. Hermes among the islands of the Hawaiian Group, 1918. Rec. Group 45, U.S. Nat. Arch. 2h pp. Dill, H.R. 1913. The albatross of Laysan. Amer. Mus. J. 13: 185-192. ---196la. The mating and nesting habits of Fregata aquila. Wilson Bull. 28: 153-157. ---1961b. The albatross of Laysan. Wilson Bull. 28: 172-175. Dill, H.R. and W.A. Bryan. 1912. Report of an expedition to Laysan Island in 1911...U.S. Dept. Agr. Biol. Surv. Bull. 42: 30 pp. Donaghho, W.R. 1953-1954. Ornithological notes. Elepaio, 14: 9-11, 18-21, 30-33, 41-43, 46-49, 57-59. Dutcher, W. 1905. [Bird protection in the Pacific]. Bird-lore 7: 301-306. 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. Edwards, R.L. 1961. Studies of the Philopteridae (Mallophaga) from birds of the order Procellariformes. 1. The genus Halipeurus Thompson. J. Parasit. 47: 125-157. Elschner, C. 1913. Corallogene Phosphat-Inseln Austral-Oceaniens und ihre Produkte. Max Schmidt,°Lllbeck. 120 pp. Translation of Chapters I and IV in the files of the Pacific Ocean Biological Survey Program, Smithsonian Institution, Washington, D.C. ---1915. The leeward islands of the Hawaiian Group [Reprint from Honolulu Advertiser]. Honolulu, 69 pp. 287 Emery, C. 1899. Ergebnisse einer Reise nach dem Pacific (Schauinsland 1896-1897). Formiciden. Zool. Jahrb., Syst., Jena 12: 438-440. Farrell, A. [ed.]. 1928. John Cameron's Odyssey. The MacMillan (Oss New York. 461 pp. Ferris, G.F. 1927. Fourth report upon Diptera Pupana from the Philippine Islands. Philippine J. Sci. 34: 207-233. Fisher, H.I. 1960. Records of the Bar-tailed Godwit and Tufted Duck on Midway Atoll. Condor 62: 480. ---1965. Bird records from Midway Atoll, Pacific Ocean. Condor 67: 355-357. Fisher, W.K. 1903a. Birds of Laysan and the leeward islands, Hawaiian Group. U.S. Fish. Comm. Bull. 23 (pt. 3): 767-807. ---1903b. Notes on birds peculiar to Laysan Island, Hawaiian Group. Auk 20: 384-397. ---1904a. On the habits of the Laysan Albatross. Auk 21: 8-30. ---1904b. Three boobies interviewed. Condor 6: 89-9}. ---1906. The Starfishes of the Hawaiian Islands. U.S. Fish. Comm. Bull. septs 3): .9o7=1131. ---1907. The holothurians of the Hawaiian Islands. Proc. U.S. Nat. Mus. 32 (No. 1555): 637-744. Fosberg, F.R. 1962. Miscellaneous notes on Hawaiian plants -- 3. B.P. Bishop Mus. Occ. Papers 23 (2): 29-44, Fowler, H.W. 1927. Fishes of the tropical central Pacific. B.P. Bishop Mus. Bull. 38: 32 pp. ---1934. The fishes of Oceania-Supplement 2. B.P. Bishop Mus. Memoir 11 (6): 385-466. Fowler, H.W. and S.C, Ball. 1924. Descriptions of new fishes obtained by the Tanager Expedition of 1923 in the Pacific Islands west of Hawaii. Proc. Acad. Nat. Sci. Phila. 76: 269-274. ---1925. Fishes of Hawaii, Johnston Island, and Wake Island. B.P. Bishop Mus." BULLS. 262931. pps Frohawk, F.W. 1892. Description of a new species of rail from Laysan Island (North Pacific). Ann. Mag. Nat. Hist. III, 9: 247-ah9. Fullaway, D.T. l1914a. A new species of Oodemas from Laysan Island. Proc.) Hawes hdt. Sec.3s) 1On ---1914b. A list of Laysan Island insects. Proc. Haw. Ent. Soc. 3: 20-22. 288 Garrett, L.E. and F.H. Haramoto. 1967. to December 2, 1915. Rec. Group 22, U.S. Nat. Arch. (original in possession of E.L. Schlemmer). 32 pp. Schreiber, R.W. and E. Kridler. 1969. Occurrence of an Hawaiian monk seal on Johnston Atoll, Pacific Ocean. J. Mammal. 50: 841-842, Sibley, F.C. and R.W. McFarlane. 1968. Gulls in the Central Pacific. Pac. Sci. 22: 314-321. 295 Simon E. 1899. Ergebnisse einer Reise nach dem Pacific (Schauinsland 1896-97). Arachnoideen. Zool. Jahrb., Syst., Jena 12: 414-437. Sluiter, C.P. 1900. Tunicuten aus dem Stillen Ocean. Ergebnisse einer Reise nach dem Pacific, (Schauinsland 1896-97). Zool. Jahrb., Syst., Jena 13: 1-35. Smythe, W.R. 1960. Monk seals on Laysan Island. Elepaio 20: 78-79. Snyder, J.0. 1904. A catalogue of the shore fishes collected by the steamer Albatross about the Hawaiian Islands in 1902. U.S. Fish. Comm. Bull. 22: 513-538. ---1917. Notes on Hawaiian lizards. Proc. U.S. Nat. Mus. 54 (No. 2224): 19-25. Speiser, P. 1902. Studien Uber Diptera pupipara. Zeitschr. Syst. Hym. Dipt. 2: 145-180. Stackpole, E.A. 1953. The sea-hunters. J.D. Lippincott Co., New York. 510 pp. Starbuck, A. 1878. History of the American whale fishery from its earliest inception to the year 1876. p. 1-779 in Report of the Commissioner of Fish and Fisheries for 1875-1876. Washington Government Printing Off. Steindacher, F. 1900. Fische aus dem Stillen Ocean. Ergebnisse einer Reise nach dem Pacific, Schauinsland 1896-97. Denks. Math.-Nat. K.K. Wiss. Wien 70: 483-521. Stoops, L. 1958. Life on Laysan. Paradise of the Pacific 70 (9): 14-15. Strasberg, D.W. 1956. Notes on the blennoid fishes of Hawaii with descrip- tions of two new species. Pac. Sci. 10: 241-267. Studer, T. 1901. Madreporarien von Samoa, der Sandwich-Inseln und Laysan. Zool. Jahrb., Syst., Jena 14: 388-428, Suman, T.W. 1964. Spiders of the Hawaiian Islands: catalog and biblio- graphy. Pac. Insects 6 (4): 665-687. Svihla, A. 1959. Notes on the Hawaiian monk seal. J. Mammal. 40: 226-229, Swezey, O.H. 1914. Two new species of moths from Laysan Island. Proc. Haw. Ent. Soc. 3: 18-19. Thomas, C. (ms.). Report of explorations made by the United States Fisheries Commission steamer Albatross in and about the Hawaiian Islands during 1902. Rec. Group 22, U.S. Nat. Arch. 136 pp. 296 Thompson, G.B. 1948. Mallophaga collected by the Tanager Expedition. B.P. Bishop Mus. Occ. Papers 19: 195-200. Thrum, T.G. [comp.]. 1902. Hawaiian Almanac and Annual for 1903. T.G. Thrum, Honolulu. 202 pp. ---1905. Hawaiian Almanac and Annual for TSO). Ee Thrum, Honolulu. 255 pp. Timberlake, P.H. 1919. Descriptions of new genera and species of Hawaiian Encyrtidae (Hymenoptera). Proc. Haw. Ent. Soc. 4: 197-231. Tomich, P.Q. 1969. Mammals of Hawaii. A synopsis and notational biblio- graphy. B.P. Bishop Mus. Spec. Pub. 57: 1-238. Treadwell, A.L. 1906. Polychaetous annelids of the Hawaiian Islands collected by the steamer Albatross in 1902. U.S. Fish. Comm. Bull. 23 (pt. 3): 1245-1181. Trempe, A.D. (ms.). Report on the bird life observed on the cruise of the Reliance in March 1936. Rec. Group 26, U.S. Nat. Arch. 5 pp. Tsuda, R.T. 1965. Marine algae from Laysan Island with additional notes on the vascular flora. Atoll Res. Bull. 110: 31 pp. ---1966. Marine benthic algae from the Leeward Hawaiian group. Atoll Res. aye, IN ALS) yoyo) Udvardy, M.D.F. 196la. Additions to the check list of Hawaiian birds. Elepaio 21: 83-90. ---196lb. The Harold J. Coolidge Expedition to Laysan Island, 1961. Elepaio 22: 43-47. ---1963. Data on the body temperatures of tropical sea and water birds. Auk 80: 191-194. Vaughan, T.W. 1907. Recent Madreporia of the Hawaiian Islands and Laysan. U.S. Nat. Mus. Bull. 59: ix and 222 pp. Walker, F.D. 1909. Log of the Kaalokai. The Hawaiian Gazette Co., Ltd., Honolulu. 64 pp. Walker. R.L. (ms. a). Excerpts from Leeward Island journal, September 2nd to 12th, 1961. Hawaii Dept. of Fish and Game, Honolulu. 4 pp. ---(ms. b). Notes on a visit to Laysan Island, December 3-10, 1963. Hawaii Dept. of Fish and Game, Honolulu. 25 pp. Warner, R.E, 1958. Wildlife of Laysan Island. Elepaio 19: 8-10, 20-23. 297 ---1963. Recent history and ecology of the Laysan Duck. Condor 65: 3-23. _=---1968. The role of introduced diseases in the extinction of the endemic Hawaiian avifauna. Condor 70: 101-120. ---(ms.). Completion report-Midway and Laysan Islands bird studies. Hawaii Dept. of Fish and Game, Honolulu. 11 pp. Weins, H.J. 1962. Atoll environment and ecology. Yale Univ. Press, New Haven. xxii and 532 pp. Werner, F. 1901. Ergebnisse einer Reise nach dem Pacific, Schauinsland 1896-97. Zool. Jahrb., Syst., Jena 14: 380-387. 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). Wheeler, W.M. 1934. Revised list of Hawaiian ants. B.P. Bishop Mus. Occ. Papers 10 (21): 1-21. | Wilder, G.P. 1905. A short trip to the Midway Islands with Capt. A.P. Niblack in the U.S.S. "Iroquois." Hawaiian For. and Agr. 2: 390-396. ---(ms. a). Extracts from a letter received by the Bureau of Biology Survey on 12 May 1924. Bureau of Sport Fisheries and Wildlife, Kailua. ---(ms. b). Report of a trip to Laysan in the summer of 1930. Rec. Group 22, U.S. Nat. Arch. 4 pp. Willett, G. 1919. Notes on the nesting of two little-known species of petrel. Condor 21: 60-61. ---(ms.). [Extracts from a report made to the Bureau of Biological Survey], Bureau of Sport Fisheries and Wildlife, Kailua, Hawaii. Wilson, C.B. 1950. Copepods gathered by the United States Fisheries Steamer Albatross from 1887 to 1909...U.S. Nat. Mus. Bull. 14 (3): 141-441. Wilson, N. 1964. Ixodes laysanensis, a new species of tick from birds on Laysan Island (Metastigmata: Ixodidae). J. Med. Ent. 1: 165-168. Wilson, S.B. 1890. On a new finch from Midway Island, north Pacific. Ibis 7: 339-341. Woodside, D.H. (ms. a). Report on the population of Laysan Teal and the transport of teal to [the] Honolulu Zoo. [1957]. Hawaii Dept. of Fish and Game, Honolulu. 4 pp. ---(ms. b). Notes on a survey of Laysan Island, July 1957. Hawaii Dept. of Fish and Game, Honolulu. 298 ---(ms. c). Report on a survey of Laysan Island, September 1961. Hawaii Dept. of Fish and Game, Honolulu. i and 32 pp. Woodside, D.H. and R.J. Kramer. (ms.). A report on a survey trip to the Hawaiian Islands National Wildlife Refuge, March 1961. Hawaii Dept. of Fish and Game, Honolulu. 32 pp. Woodward, P.W. and R.B. Clapp. 1969. First records of Baird's Sandpiper from the central Pacific. Elepaio 30: 25. Zimmerman, E.C. 1948a. Insects of Hawaii. Vol. 2. Apterygota to Thysanoptera. Univ. of Hawaii Press, Honolulu. vii and 475 pp. ---1948b. Insects of Hawaii. Vol. 3. Heteroptera. Univ. of Hawaii Press, Honolulu. v and 255 pp. ---1948c. Insects of Hawaii. Vol. 5. Homoptera: Sternorhyncha. Univ. of Hawaii Press, Honolulu. vii and 464 pp. ---1948d. Insects of Hawaii. Vol. 4. Homoptera: Anchenorhyncha. Univ. of Hawaii Press, Honolulu. vii and 268 pp. ---1957. Insects of Hawaii. Vol. 6. Ephemeroptera-Neuoptera-Tridroptera and supplement to Vols. 1 to 5. Univ. of Hawaii Press, Honolulu. ix and 209 pp. ---1958a. Insects of Hawaii. Vol. 7. Macrolepidoptera. Univ. of Hawaii Press, Honolulu. ix and 542 pp. ---1958b. Insects of Hawaii. Vol. 8. Lepidoptera: Pyraloidea. Univ. of Hawaii Press, Honolulu. ix and 456 pp. 299 Appendix Table 1. Scientific visits to Laysan Island, 1828-1969. Date Personnel Vessel 1828 24 Mar. C. Isenbeck Moller 1859 May N.C. Brooks Gambia 1890 16 July A.B. Lyons Kaalokai 1891 16-27 June Rothschild Expedition Kaalokai Henry C. Palmer George C. Munro 1892 ca. 18 Nov.- J.J. Williams : Liholiho 1893 ca. 22 Jan. 1895 ca. Sept. W.H. Hall (BPBM)* C.D. Bryant (2) 1896 24 June- H.H, Schauinsland (BM) H. Hackfeld 2h Sept. 1902 16-23 May Charles H. Gilbert (SU) Albatross Walter K. Fisher (SU) Charles C. Nutting (SUI) John O. Snyder (SU) 1903 Apr.-May William A. Bryan (BPBM) z ca. 1903-1908 Max Schlemmer Various vessels 1905 19 Sept. Gerrit P. Wilder (HDAF) Iroquois 1906 26 Jan.- Paul E.H. Bompke 2 6 June 1910 16-18, W.V.E. Jacobs Thetis 26 Jan. 1911 24 Apr.- State University of Iowa Thetis 5 June Expedition Homer R. Dill (SUI) Clarence J. Albrecht Charles A. Corwin Horace C. Young (SUT) William A. Bryan (BBS) *A glossary of abbreviations indicating professional affiliation is appended at the end of this table. 300 Appendix Table 1. (continued) Date Personnel Vessel 1912 22 Dec.- G.R. Salisbury (USN) Thetis 1913 11 Mar. Alfred M. Bailey (BBS) David T. Fullaway George Willett (BBS) William S. Wallace (SU) 1914 11 Sept. Carl Elschner Thetis 1915 3 Apr. William H. Munter (USCG) Thetis Members of crew 1916 9 Feb. William H. Munter (USCG) Thetis Members of crew 1918 8-10 Sept. Crew of the Hermes Hermes 1923 7 Apr.- Tanager Expedition* Tanager 14 May 8-14 Apr., Alexander Wetmore (BBS) 29 Apr.-14+ May 8 Apr.-14 May 8-14 Apr. 8-14 Apr., 29 Apr.-14 May 8-14 Apr. 8-14 Apr., 29 Apr.-14 May 8-29 Apr. (ornithologist) E.C. Reno (BBS) (small mammal control expert) David T. Fullaway (BPBM) (entomologist) Ditlev Thaanum (BPBM) (conchologist ) Edward L. Caum (BPBM) (botanist) Chapman Grant (naturalist) Donald R. Dickey (naturalist, photographer) *Individual itineraries given under entire period during which members of the expedition were encamped on the island. Two other persons, not official members of the survey party, visited the island briefly on 14 May. These were Theodore Dranga, who was collecting shells for L.A. Thursten, and Austin Jones, a friend of the ship's surgeon. Appendix Table 1. (continued) Date Personnel 1923 +7 Apr.- Tanager Expedition (continued) 14 May 8 Apr.-14 May Eric L. Schlemmer (assistant to Wetmore) 8 Apr.-l4+ May Stanley C. Ball (BPBM) (biologist) 8-29 Apr. J.W. Thompson (preparator) 14 May John Baker (collector) 14 May Gerrit P. Wilder (botanist) 8 Apr.-14 May George Higgs (USN) (cook, camp assistant) 1924 6 May Gerrit P. Wilder 1928 1 Mar. Victor Pietschmann (VM) 6 May William G. Anderson (Capt.) 1930 2-18 Aug. Gerrit P. Wilder (BBS) 1936 7-8 Mar. Alfred D. Trempe (BBS) B.L. Bassham (USCG) Members of crew 12(-15?) Dec. C. Templeton Crocker Expedition William F. Coultas (AMNH) Toshio Asaeda (AMNH) 1950 23 June Vernon E. Brock (HDFG) POFI personnel 1951 12 May POFI personnel late June- George Vanderbilt Pacific early July Equatorial Expedition George Vanderbilt Robert R. Harry (SU) 301 Vessel Tanager Pelican Lanikai Lanikai Pioneer Reliance Zaca Hugh M. Smith Hugh M. Smith Pioneer 302 Appendix Table 1. Date 1951 late June- early July 1954 1 Nov.* 3 Nov. 1955 10 Feb. 1957 7 Jan. 15 Apr. 2 June- 3 July 8-12 July 28 Dec. 1958 27 May- 4 June 28 June 1959 28 Apr.- 1 May *Another derial survey may have been made on 5 December. (continued) Personnel Vanderbilt Expedition (continued) Vernon E. Brock (HDFG) Anita Vanderbilt Lucille Vanderbilt B. Green T,. Ivar Vatland Johnson A. Neff (BSFW) Philip A. Dumont (BSFW) POFI personnel Donald L. McKernan POFI personnel Karl W. Kenyon (BSFW) Dale W. Rice (BSFW) Karl W. Kenyon (BSFW) Dale W. Rice (BSFW) David H. Woodside (HDFG) Richard E. Warner (HDFG) Alphonse Labrecque Al Stoops Dale W. Rice (BSFW) Richard E. Warner (HDFG, BSFW) Prentis Burtis Frederick W. Landers Dale W. Rice (BSFW) Richard Takahashi (USN) Dale W. Rice (BSFW) Raymond J. Kramer (HDFG) George D. Butler, Jr. (UA) Hubert Caspers (HU) William R. Smythe (HSPA) Vessel Pioneer Aerial survey Charles H. Gilbert Hugh M. Smith Aerial survey Aerial survey Coast Guard ship Koyu Maru Aerial survey Matagorda and Chautauqua Aerial survey Matagorda 303 Appendix Table 1. (continued) Date Personnel Vessel 1959 20-27 July Miklos D.F. Udvardy (UBC) - George D. Butler, Jr. (UA) C.H. Danforth Charles W. Daniel Richard E. Warner (HDFG) *1960 23 Aug. POFI personnel Charles H. Gilbert 1961 7-8 Mar. David H. Woodside (HDFG) Planetree Raymond J. Kramer (HDFG) 4-10 Sept. Harold J. Coolidge Expedition Ironwood (0800-0800) Richard E. Warner (UC) George D. Butler, Jr. (UA) Edward C. Jestes (UH) Charles Lamoureux (UH) A. Starker Leopold (UC) Miklos D.F. Udvardy (UBC) Robert L. Usinger (UC) Martin Vitousek (UH) Ronald L. Walker (HDFG) David H. Woodside (HDFG) 1962 14-19 June Raymond J. Kramer (HDFG) Stone County John W. Beardsley (HSPA) (0930-1645 ) David H. Woodside (HDFG) David B. Marshall (BSFW) 1963 11-13 Feb. Raymond J. Kramer (HDFG) Moctobi A. Binion Amerson, Jr. (POBSP) (1600-1400) F. Allen Blagden (POBSP) Robert W. McFarlane (POBSP) Fred C. Sibley (POBSP) William 0. Wirtz, II (POBSP) 3-10 Dec. Ronald L. Walker (HDFG) Matagorda Roy T. Tsuda (UH) and Nixon Wilson (BPBM) Chautauqua 1964 10-11 Mar. Eugene Kridler (BSFW) Planetree A. Binion Amerson, Jr. (POBSP) (0900-0730) Loren Kroenke (UH) Edward O'Neill (BSFW) Ronald L. Walker (HDFG) George S. Wislocki (POBSP) *Time of arrival and departure, where known, are listed under the dates of visit for surveys made during the 1960's. 304 Appendix Table 1. Date 1964 1965 1966 16-20 Sept. (0900-0600) 19-20 Sept. (0900-2000) 6-11 Mar. (1100-1500) 17-21 July (2040-1000) 5-12 Aug. (1530-2100) 26-31 Mar. (1345-0730) 15 May Senaanes (1345-1530 10-16 June (0945-0330) 20-21 June (1300-225 ) (continued ) Personnel Kenneth E. Amerman (POBSP) Alan H. Anderson (POBSP) Robert Banner (UH) Richard W. Merrill (POBSP) J. Douglas Whitman (POBSP) Paul W. Woodward (POBSP) Alan Lee Young (UH) Eugene Kridler (BSFW) John W. Beardsley (UH) Robert R. Fleet (POBSP) C. Robert Long (POBSP) Ronald J. Walker (HDFG) William O. Wirtz, II (POBSP) Kenneth E. Amerman (POBSP) Roger B. Clapp (POBSP) J. Vincent Hoeman (POBSP) Dennis L. Stadel (POBSP) Charles Williams, Jr. (USN) Richard S. Crossin (POBSP) Brian A. Harrington (POBSP) Dayle N. Husted (POBSP) Jeffrey P. Tordoff (POBSP) Richard §. Crossin (POBSP) Kenneth E. Amerman (POBSP) Brian A. Harrington (POBSP) Dayle N. Husted (POBSP) Jeffrey P. Tordoff (POBSP) Eugene Kridler (BSFW) Andrew J. Berger (UH) Nelson Rice (HDFG) Ronald L. Walker (HDFG) Members of the crew Richard S. Crossin (POBSP) Kenneth C. Balcomb (POBSP) Richard D. Chandler (POBSP) David I. Hoff (POBSP) David L. Pearson (POBSP) Philip C. Shelton (POBSP) Frank H. Smith, Jr. (POBSP) Vessel Shearwater Basswood Shearwater Shearwater Shearwater Buttonwood Charles H. Shearwater Gilbert 305 Appendix Table 1. (continued) Date Personnel Vessel 1966 17-18 Sept. Eugene Kridler (BSFW) Ironwood Sherwin Carlquist (CC) (1340-1450) Karl W. Kenyon (BSFW) Warren Roll (HSB) Ronald L. Walker (HDFG) 20-23 Oct. Kenneth C. Balcomb (POBSP) Tawakoni Patrick J. Gould (POBSP) (1945-1210) Brian A. Harrington (POBSP) T. James Lewis (POBSP) 1967 18-19 Mar. Eugene Kridler (BSFW) Basswood C. Douglas Hackman (POBSP) (1330-1430) Ernest Kosaka (HDFG) John Maciolek (BSFW) Richard Wass (UH) 7-12 June Robert L. DeLong (POBSP) imc OOns David L. Burckhalter (POBSP) LT 2086 (0900-0900) Dennis L. Stadel (POBSP) LT 2087 F. Christian Thompson (POBSP) Robert Tuxson (POBSP) 5-11 Sept. Charles A. Ely (POBSP) LT 2081 Ronald Amerson (USN) LT 2086 (1930-0730) Roger B. Clapp (POBSP) LT 2087 David I. Hoff (POBSP) Edward King (USN) 21-24 Sept. Eugene Kridler (BSFW) Buttonwood Robert Ballou (BSFW) (1645-1600) John L. Sincock (BSFW) Ronald L. Walker (HDFG) 13 Dec. Eugene Kridler (BSFW) Ironwood (1100-1600) 1968 17-19 Mar. Eugene Kridler (BSFW) Ironwood Roger B. Clapp (POBSP) (1700-1600) Karl W. Kenyon (BSFW) Ernest Kosaka (HDFG) John L. Sincock (BSFW) Sept. Eugene Kridler (BSFW) LTronwood Derral Herbst (UH) Robert Eddinger ae John L. Sincock (BSFW) 306 Appendix Table 1. (continued) Date Personnel Vessel 1969 26-29 Mar. Eugene Kridler (BSFW) Buttonwood Karl W. Kenyon (BSFW) (0730-1530) George Laycock (NAS) David L. Olsen (BSFW) John L. Sincock (BSFW) 2-3 June Eugene Kridler (BSFW) Mahi Karl Bathen (UH) (0630-1600) Thomas Clark (UH) Ronald Kent Ernest Kosaka (HDFG) James McVay (UH) David L. Olsen (BSFW) William Patzert (UH) John L. Sincock (BSFW) Douglas Yen (BPBM) 9 Sept. Eugene Kridler (BSFW) Buttonwood John Maciolek (BSFW) (0730-1430) David L. Olsen (BSFW) John L. Sincock (BSFW) Glossary of Abbreviations AMNH American Museum of Natural History BBS Bureau of Biological Survey BM Bremen Museum BPBM Bernice P. Bishop Museum BSFW Bureau of Sport Fisheries and Wildlife CC Claremont College, Claremont, California HDAF Hawaiian Department of Agriculture and Forestry HDFG HawaiiDivision of Fish and Game HSB Honolulu Star Bulletin HSPA Hawaiian Sugar Planters Association HU Hamburg University NAS National Audubon Society POBSP Pacific Ocean Biological Survey Program POFI Pacific Ocean Fisheries Investigations, Bureau of Commerical Fisheries SU Stanford University SUL State University of Iowa UA University of Arizona UBC University of British Columbia UC University of California UH University of Hawaii USCG United States Coast Guard USN United States Navy VM Vienna Museum Appendix Table 2. Date 1828 1859 1890 1891 1892 1893 1895 1896 1902 24 Mar. ca. 18 Nov.- ca. 22 Jan. ca. Sept. 2k June- 24 Sept. 16-23 May 307 Results of scientific visits to Laysan Island, 1828-1969, with emphasis on bird observations.* Results Observations of birds (Kittlitz in Rothschild, 1893- 1900). 2> varieties of plants collected were subsequently lost. Potatoes, onions, and pumpkins planted. Observations of birds (Lyons, 1890); plants collected. Observations of birds (Rothschild, 1893-1900; Munro, 1930, 1941a, 1941b, 1941c, 1942a, 1942b, 1944, 1945, 1946, 1947, 1953) and many bird specimens collected (no less than 153 skins of 19 species). The follow- ing birds were described: Laysan Rail by Frohawk (1892), Laysan Teal by Rothschild (1892a), Laysan Miller-bird, Laysan Honeyeater, and Laysan Finch (the latter subsequently synonomized) by Rothschild (1892b), and Laysan Albatross by Rothschild (1893). Two new moth species described. Photographs and a collection of birds made. Two Black-footed Albatross captured and transported to Honolulu. Collections of birds (at least 21 skins of 9 species). Observations and collections of birds (Schauinsland, 1899; Rothschild, 1893-1900). Hartert (1919-1927) lists a number of Schauinsland's and Rothschild's specimens (at least 271 skins of 25 species, and others sent to them by correspondents) in the Tring Museum and describes a new subspecies of White Tern from one of Schauinsland's specimens. Collections of: crus- tacea, vascular plants, foraminifera, ectoparasites, shrimp, insects, sea squirts, fish, corals, hydroid, turbellarium, oligochaete, molluscs. Observations and collections of birds (78 skins of 20 species) (Nutting, 1903; Fisher, 1903a, 1903b, 1904a, 1904b). Notes on the anatomy and pterylosis of a Laysan Finch were reported by Clark (1912). Collec- tion of: plants and reptiles; mallophaga, corals, brachyuian and macruian crabs, hydroids from offshore, erustacea, starfishes and other echinoderms, medusae, polychaetous annelids, fish, lizards. *Papers by Fosberg (1962), Caspers (1968), and St. John (1970) were obtained too late in the preparation of the manuscript for inclusion in this and the following appendix table. 308 Appendix Table 2. (continued) Date Results 1903 Apr. -May Plants, insects, and birds (331 specimens of 25 species--inyolving 189 skins, 6 mounts, 24 skeletons, 8 alcoholics, 7 nests, and 92 clutches of eggs) col- lected by W.A. Bryan. ca. 1903-1907 Collections of birds (at least 134 skins, 9 mounts, and 30 eggs) and fish by Schlemmer. 1905 19 Sept. Insects collected; a coconut tree planted; Casuarina, Paritium, Thespesia, Chrysopphyllum, grasses and lily bulbs left to be planted; Laysan Finches captured and transferred to Midway (Bryan, 1912). 1906 26 Jan.- Birds collected, some of which were new distributional 6 June records. 1910 16-18, Several bird skins prepared by Japanese and seized by 26 Jan. the Thetis later found their way into various museum collections. 1911 24 Apr.- Observations of birds (Dill and Bryan, 1912; Dill, 5 June 1913, 1916a, 1916b; W.A. Bryan, 1911)*; collection of insects, plants; 398 birds of 25 species collected. 1912 22 Dec.- Observations of birds (Bailey, 1934, 1942, 1952a, 1913 11] Mar. 1956; Willett, 1919) and at least 401 specimens of 31 species collected; collections of insects, plants, and a monk sealg 100 coconut sprouts planted; 128 Laysan Rail captured for transport to other areas, a number of Laysan Finches captured and transported to Midway Atoll; 5,024 rabbits killed. 1914 11 Sept. Study of geology; analysis of lagoon water and phos- phates. 1915 3 Apr. Observations of birds, a few notes on rabbits (Munter, 1915). 1916 Q Feb. Brief observations of birds and rabbits; samples of ‘beach sand, shells, and shellfish collected for W.A. Bryan. 1918 8-9 Sept. Observations of birds, turtles, seals, and rabbits. *Several of these papers are primarily reprints or barely different versions of one another, ee 309 Appendix Table 2. (continued) Date Results 1923 7 Apr.- Observations of birds (Wetmore, 1925); extermination 14 May of rabbits; many photographs taken; a number of vas- cular plants sowed; 8 Laysan Rails introduced from Midway. Collected: arachnids, insects, crustacea, searstars and other echinoderms, polychaetous annelids, foraminifera, vascular plants, sponges, fish, nematode, cestode, many birds (at least 319 skins of 21 species), 2 seals, algae, molluscs. Observations by Wetmore of turnstone predation on eggs later reported by Crossin and Huber (1970). 1924 6 May Evidently only a few sketchy observations of animals made (Wilder, ms. b; Wetmore, 1925). 1928 1 Mar. Collected: 8 species of Cypraeacea and 1 polychaete. 6 May Larval hemirhamphids collected offshore by Anderson. 1930 2-18 Aug. Collected: plants, insects, bird eggs, marine organisms (crabs, fish), some artifacts, 4 species of flies. Various plants introduced and some notes taken on bird life. 1936 2-8 Mar. Brief observations of birds, seals, and turtles. 12 DEC. Very brief observations of birds, seals, turtles, and vegetation; small collection of birds made. 1950 23 June Laysan Teals counted, unspecified number of turtles tagged, seals reported (Brock, 1951a). 1951 12 May Census of seals. late June- Census of seals and turtles; extensive fish collections; early July bird census (Brock, 1951b). 1954 1 Nov. Aerial seal count. #955 10 Feb. Observations of teal, brief notes on other birds. 1957 7 Jan. Aerial seal census; albatross census (Rice and Kenyon, 1962). UG) Ayah Aerial seal census. 25 June- Nine teal captured and 8 taken to Honolulu Zoo (1 3 July died in transit); seals censused and 25 seals tagged; seabird populations estimated; 400 albatrosses banded; observations of teal (Warner, 1963). 310 Appendix Table 2. Date 1957 8-12 July 28 Dec. 1958 27 May- k June 28 June 1959 28 Apr.- 1 May 20-27 July 1960 23 Aug. 1961 7-8 Mar. 4-10 Sept. (continued ) Results Amateur observations of birds. Aerial albatross and seal censuses (Rice and Kenyon, 1962). 3,302 birds of 9 species banded; census of fauna with emphasis on seals, young albatrosses, Laysan Teal, and Laysan Finches (Warner, 1958). Observations of birds; emphasis on Laysan Teal* (Warner, 1963). 36 Laysan Teal captured and transported to Honolulu Zoo; young seal collected for British Museum; seals tagged; Laysan Finches captured for transport to Honolulu Zoo. Aerial seal census. Census of: seals, teal (Warner, 1963), Laysan Finch, albatross. Detailed observations of seals, Laysan Finch, Laysan Teal; notes made on other birds. Col- lection of plants, crustacea, and insects; weather observations, mapping of vegetation. Collected: insects, crustacea, and other arthropods; plants by Butler and Daniel. Observations on Laysan Teal (Warner, 1963) and body temperature of birds (Udvardy, 1963). Seal census. Count of seals; survey of island for fish bait. Census of seals and turtles; Laysan Teal observations (Warner, 1963). Census of Laysan Teal, Laysan Finch, seals, and turtles. 275 birds of 2 species banded; 2 turtles tagged; previous photographic stations refurbished and new ones established. Collected: 8 Laysan Teal by Leopold and Warner; plants; crustacea; insects; arachnids; marine algae by Lamoureux. Geological ob- servations and specimens by Jestes; geophysical measurements by Vitousek; rectal temperatures of seals taken by Udvardy; body temperatures taken for 8 species of birds (Udvardy, 1963); Laysan Teal ob- servations (Warner, 1963). *Notes on teal in captivity captured during this visit were reported by Ripley (1960) and Lint (1960). Appendix Table 2. Date 1962 14-19 June 1963 1964 1965 1966 11-13 Feb. 3-10 Dec. 10-11 Mar. 16-20 Sept. 19-20 Sept. 7-11 Mar. 17-21 July 5-12 Aug. 26-31 Mar. 15 May Syl (continued) Results Census of seals; observations on vegetation and avi- fauna; collection of ectoparasites, crustacea, insects by Beardsley. 540 birds of 11 species banded; 10 birds of 8 species collected by POBSP; 2/ seals tagged, 1 seal collected; 6 turtles tagged; distribution and condition of vege- tation noted. Ectoparasites, including a tick, collected by Wilson; plants introduced; seals, turtles, albatrosses, shore- birds, and teal censused; 3 turtles tagged and 17 Laysan Teal banded by Walker; marine algae and vas- cular plants collected by Tsuda; island water salinity tested. Census of seals and turtles; geophysical measurements by Kroenke; 516 birds of 2 species banded by POBSP; 442 birds of 9 species banded by BSFW; ticks collected. 3,792 (plus any Brown Boobies) birds of 15 (or 16) species banded by POBSP. Collected: 9 Bonin Petrels, some insects, spider, and plants; turtles noted. Birds observed, seals censused, no turtles seen, teal counted, insects and plants collected; 40 birds of 3 species banded by BSFW. 18,273 birds of 12 species banded by POBSP; turtles noted. 15,470 birds of 5 species banded by POBSP; 4 birds, ectoparasites, Berlese samples collected. 44,442 birds of 8 species banded by POBSP, bird ob- servations made; 1 turtle tagged. Observations of birds; census of seals, turtles, Laysan Teal; estimates of Laysan Finch population from transect counts; 333 birds of 8 species banded; 4} Laysan Finches captured for transport to Honolulu Zoo; refuge sign erected; photographs of vegetation made; Chenopodium seeds planted. Pictures taken of wildlife. 312 Appendix Table 2. Date 1966 1967 1968 1969 10-16, 20-21 June ali -18 sept e 20-23 Oct. 18-19 Mar. 7-12 June 5-11 sept ° 21-24 Sept. 13 Dec. 17-19 Mar. Sept. 26-29 Mar. 2-3 June 9 Sept. (continued) Results Observations of birds; census of seals and turtles, 72,454 birds of 9 species banded by POBSP; 58 bird specimens of 9 species collected; 17 ectoparasite collections made from 7 species of birds; 30 Berlese samples taken from nests of 4 species of birds. Transect census of finches and teal; observations of birds; census of seals and turtles; 1 turtle and 11 seals tagged; 1 turtle recaptured; 14+ birds of 4 species banded. Observations of birds; census of seals, turtles, shore- birds; 536 birds of 6 species banded by POBSP; 6 bird specimens of 4 species collected. Observations of birds, census of seals, turtles; 4 Golden Plovers banded by BSFW; 33 seals and 1 turtle tagged. Observations of birds and turtles; census of seals; 8,092 birds of 11 species banded; 2 seal skulls col- lected and 12 seal pups tagged. Observations of birds; 4,085 birds of 15 species banded by POBSP. Collected: 7 bird specimens of 4 species, hippoboscid flies, ticks from various birds. 34 birds of 3 species banded; 1 turtle tagged. Observations on vegetation and wildlife; census of seals and turtles; 2 turtles tagged. Census of seals and turtles; observations of birds; 115 birds of 8 species banded by POBSP; 64 Laysan Finch banded by wildlife personnel; 5 bird specimens of 3 species collected by POBSP; 1 Laysan Teal carcass collected by BSFW; 4 turtles and 36 seal tagged, 1 turtle recaptured. Data are not available at present. Observations of birds, censuses of seals, turtles, teal, albatrosses, and Laysan Finch. 38 seals and 2 turtles tagged. Teal, turtles, and seals censuses; 25 seals tagged. Brief observations on birds; seals, and teal censused; 2 seals tagged; a few observations made of marine life. 313 Appendix Table 3. Publications on collections and studies (with the exception of birds) made on Laysan Island.* Protozoa Rhumbler, 1906. Records foraminifera collected by Schauinsland. Cushman in Edmundson Reports collection of 24 species of foramini- et al., 1925. fera by the Tanager Expedition. Porifera Barr, 1903. Records 2 species of demonspongiid sponges collected by Schauinsland. Preiwisch, 1903. Records 2 species of calcarean sponges col- lected by Schauinsland. Coelenterata Hartlaub, 1901. Reports a hydroid collected by Schauinsland. Studer, 1901. Reports 18 species of Madreporia collected by Schauinsland (cf. Schauinsland, 1899). Nutting, 1905. Reports hydroid collections made offshore May 1902. Mayer, 1906. Reports hydromedusa from collections made offshore by the Albatross Expedition. Vaughan, 1907. Reports corals (Madreporia) collected by Albatross Expedition; revises some of Studer's taxonomy, and describes several new species. Nutting, 1908. Reports 2 species of coral (Alcyonaria) from collections made offshore by the Albatross Expedition. Platyhelminthes Plehn, 1899. Reports a turbellarian collected by Schauinsland. Chapin, 1925. Describes a new species of cestode from an Hawaiian monk seal collected by the Tanager Expedition. *Authors are in chronological order. 314 Appendix Table 3. (continued) Rschelminthes Chapin, 1925. Mollusca Schauinsland, 1899. Bergh, 1900. Berry, 1910. Pilsbury, 1917. Pilsbury, 1920. Pilsbury, 1927. Schilder, 1933. Dall, Bartsch, and Rehder, 1938. Butler and Usinger, 1963 « Annelida Michaelsen, 1899. Treadwell, 1906. Treadwell in Edmundson et al., 1925 Holly, 1935. Describes a new species of nematode from a Hawaiian monk seal collected by the Tanager Expedition. Mentions occurrence of 2 cephalopods and several gastropods. Reports 3 species of opbsthobranchs collected by Schauinsland. Lists 2 cephalopods recorded by Schauinsland. Describes 3 species of molluscs collected by W.A. Bryan. Lists 2 species of molluscs, one collected by W.A. Bryan, the other by Capt. Brown of the USCG. Lists 2 species of barnacles collected by the Tanager Expedition. Reports 8 species of Cypraeacea collected in March 1928. Lists 10 species of pelecypods collected at or offshore Laysan, most of them by the Albatross Expedition. Records 2 species of land snails. Records an oligochaete collected by Schau- insland (also mentioned by Schauinsland, 1899). Reports collections of 5 polychaetes in May 1902. Reports 5 species of polychaetes collected by the Tanager Expedition. Reports a single polychaete collected in March 1928. Bly Appendix Table 3. (continued) Annelida (continued) Hartman, 1966. Summarizes earlier records and gives current taxonomy. Arthropods Butler and Usinger, Records plant-arthropod, bird-arthropod, 1963. and lake-arthropod associations. Arachnomorpha (Arachnida) Schauinsland, 1899. Lists species also reported by Simon (below). Simon, 1899. Reports 6 species collected by Schauinsland. Butler, 1961a. Reports 3 species of Acarina and 5 Araneida collected in April and July 1959. Butler and Usinger, Lists new records of 3 species of Acarina 1963. and 2 Araneida from collections made in September 1961 and June 1962 and summarizes earlier records. Aoki, 1964. Records 8 species of oribatid mites (Acarina) from collections made in December 1963. Suman, 1964. Gives recent summary of spiders known from Laysan. Lists 9 species. Wilson, 1964. Describes new tick (Ixodes laysanensis) (Acarina, Ixodidae) collected December 1963. Aoki, 1965. Records 2 species of oribatid mites (Acarina) from collections made in December 1963. Kohls, Sonenshine, and Records occurrence of Ornithodorus capensis Clifford, 1965. group (Acarina: Argasidae). Amerson, 1966. Reports infestation of Sooty Tern nasal cavities by ticks (Acarina: Argasidae) from specimens collected 8 August 1965. Beardsley, 1966. Reports a new distribution record for a spider from collection made September 1964, Garrett and Harimoto, Summarizes earlier records of Acarina. 1967. 316 Appendix Table 3. (eontinued) Arachnomorpha (Arachnida) (continued) Amerson, 1968. Crustacea Schauinsland, 1899. Lenz, 1901. Sars, 1903. Ortman, 1905. Rathbun, 1906. Edmundson in Edmundson et al., 1925. Wilson, 1950. Butler, 1961la. Butler and Usinger, 1963. Chilopoda Bryan et al., 1926. Insecta (Hexopoda) Rothschild, 1894. Schauinsland, 1899. Emery, 1899. Meyrick, 1900. Reports distribution and hosts of ticks (Acarina) from collections made by the POBSP. Mentions some of crustaceans he collected. Reports 25 species of crustacea collected by Schauinsland. Records a brine shrimp collected by Schau- insland. Reports a single species of schizopod from the vicinity of Laysan that was collected by the Albatross Expedition. Reports brachyuran and macruran crabs col- lected by the Albatross Expedition. Reports 59 species of decapods, and a phyllopod, collected by the Tanager Expedition. Reports a copepod collected north of Laysan by the Albatross Expedition. Reports an isopod from April, July 1959 collections. Reports an isopod from collections made in September 1961 and June 1962. Reports a chilopod from collections made by the Tanager Expedition. Describes 2 moths from Laysan. Lists some insects he collected. Reports 4 species of ants collected by Schau- insland. Describes 2 noctuid moths collected by Schau- insland. 317 Appendix Table 3. (continued) Insecta (Hexopoda) (continued) Speiser, 1902. Alfken, 1903. Perkins, 1906. Kellogg and Paine, 1910. Fullaway, 19l4a. Fullaway, 1914b. Swezey, 1914. Perkins, 1919. Timberlake, 1919. Bryan et al., 1926. Ferris, 1927. Bryan, 1932. Wheeler, 1934. Thompson, 1948. Reports a hippoboscid fly from Schauinsland's collection. Records 17 species of insects collected by Schauinsland. Reports 7 species of insects collected by Wilder in 1905. Reports 10 species of mallophaga collected in May 1902. Reports 60 species of insects from collections made in 1905 by Wilder (6 species), 1911 by Bryan (3 species), and 1912 by Fullaway (55 species). Describes a new beetle from material col- lected in April 1911 and December 1912. Describes 2 new species of moths from col- lections made in 191e. Describes a new species of Otiorrhynchine beetle from specimens collected by W.A. Bryan. Describes a new species of encyrtid (Hymenop- tera) from specimens collected in December 1912. Records ca. 34 species of insects collected by the Tanager Expedition. Reports a hippoboscid fly. Reports 4 species of flies collected in August 1930. Gives a summary listing 7 species of ants from early collections. Reports 10 species of mallophaga collected by the Tanager Expedition. Appendix Table 3. 318 Zimmerman,* 1948a Zimmerman, 1948b. Zimmerman, 1948c. Zimmerman, 1948d. Ross, 1951. Zimmerman, 1957. Zimmerman, 1958a. Zimmerman, 1958b. Hardy, 1960. Butler, 196la. Butler, 1961b. Edwards, 1961. Maa, 1962. Butler and Usinger, 1963 (continued) Insecta (Hexopoda) (continued) Lists 3 cockroaches, an embiopterid, an earwig and 10 mallophaga. Lists a nabid and a mirid. Lists a coccid. Lists a delphacid. Reports that an embiopterid collected by the Tanager Expedition as Oligotoma insularis by Bryan et al., 1926, is actually Oligotoma (Apothonia) oceania Ross sp. nov. Supplement lists an embiopterid. Lists 6 noctuid moths. Lists 3 pyralids and a pterophorid. Describes a stratiomyid fly from specimens collected by the Tanager Expedition. Summarizes insects collected by Wilder, Bryan, Fullaway, and the Tanager Expedition and adds collections of 68 species made in April and July 1959. Reports association of stratiomyid flies with albatross carcasses from observations in July 1959. Describes a new species of mallophage collected from Christmas Shearwaters. Collected ca. 1904-1907. Reports specimens of hippoboscid flies col- lected in August 1930 and April-May 1959. Adds new insect records from collections of September 1961 and June 1962 and summarizes old records. Lists ca. 189 species from Laysan. *Zimmerman, and Hardy, in the Insects of Hawaii series, present distribu- tional records derived primarily from the Tanager collections, but ex- tensively revise taxonomy, reidentify specimens, and identify to species hitherto unidentified specimens. 349 Appendix Table 3. (continued) Insecta (Hexopoda) (continued) Hardy, 1964. Hardwick, 1965. Beardsley, 1966. Maa, 1968. Echinodermata Fisher, 1906. Fisher, 1907. Clark, 1908. Agassiz and Clark, 1907-1912. Clark in Edmundson et al., 1925. Fisher in Edmundson et al., 1925. Clark, 1949, Chordata Urochordata Sluiter, 1900. Lists 2 dolichopodid flies. Discusses specimens of Heliocoverpa (Noctuidae) collected in September 1961. Reports 5 species of insects collected in September 1964. Reports hippoboscid flies from collections made 1963-1967 by the POBSP. Reports 6 species of starfish, most newly described, from collections made offshore by the Albatross Expedition. Reports 6 species of holothurians collected by the Albatross Expedition. Reports 2 newly described species of Crinoidea that were collected offshore by the Albatross Expedition. Reports echinoderms collected by the Alba- tross Expedition. Reports 8 Ophiuroidea, 9 Echinoidea, and 9 Holothuroidea collected by the Tanager Ex- pedition. Reports 2 starfish (Asteroidea) collected by the Tanager Expedition. Reports 6 species of brittle stars (Ophiur- oidea) from collections made by the Albatross Expedition and gives summary lists of other echinoderms previously collected from Laysan. Records 6 species of sea squirts collected by Schauinsland. 320 Appendix Table 3. Chordata Vertebrata Pisces Steindacher, 1900. Jordan and Snyder, 1904. Snyder, 1904. Gilbert, 1905. Fowler and Ball, 1924. Fowler and Ball, 1925. Fowler, 1927. Fowler, 1934. Strasberg, 1956. Reptilia Patysel Ooi. Schauinsland, 1899. Rothschild, 1893- 1900. Werner, 1901. (continued) Reports 51 species of fish collected by Schauinsland. Reports fish collected at Oahu, Hawaii, and Laysan, the latter collection by Max Schlemmer, but does not always give collection localities. Lists 7 species from Laysan. Reports 33 species of fish collected by the Albatross Expedition. Records 24 species of deep sea fishes col- lected in the vicinity of Laysan by the Albatross Expedition and one species collected at Laysan by Max Schlemmer. Describes a new family, genus and species for a fish collected by the Tanager Expedition. Reports 71 species of fish collected by the Tanager Expedition. Lists 7 species, 6 from the Tanager collec- tions of which 1 was not reported in Fowler 1925, and 1 from a collection in May 1893. Lists 2 additional species of fish from Laysan from collection made in August 1930. Revises taxonomy of Hawaiian blennioid fishes recording 3 species from Laysan. Mentions presence of turtles in 1857. Gives observations of turtles from 1896. Indicates turtles were seen in 1828. Reports 2 lizards from Schauinsland's visit. ByZIL Appendix Table 3. (continued) Reptilia (continued) Wilder, 1905. Mentions that turtles were shot in September 1905 . Dill and Bryan, 1912. Mentions occurrence of turtles and their eggs, and the killing of one turtle for food. Snyder, 1917. Reports collections and observations of 2 lizards in May 1902. Munro, 1946. Reports that "one or two species" of lizards were present in 1891. Brock, 195la. Mentions field party went ashore to tag turtles. Anon., 1951b.; Reports that 15 turtles were seen in December Coultas, ms. 1936. Udvardy, 1961b. Gives turtle observations made September 1961. Beardsley, 1966. Reports collection of a skink in September 1964; erroneously records it as new record. Mammalia Paty, 1857. Mentions presence of seals in 1857. Rothschild, 1893- Mentions seals were seen in 1828. 1900. Matschie, 1905. Describes Hawaiian monk seal from skull collected by Schauinsland. Wilder, 1905. Mentions presence of donkey and a few cows in September 1905. Dill and Bryan, 1912. Gives information on introduction and habits of rabbits; presence of guinea pigs and ab- sence of seals. Munter, 1915. Mentions presence and capture of rabbits in April 1915. Bailey, 1918. Notes collection of a seal specimen by the Biological Survey party 1912-1913. Wetmore, 1925. Reports that only a few hundred rabbits were present in 1923. Bea Appendix Table 3. Mammalia (continued) Munro, 1946. Anon., 1951b, Coultas, ms. Bailey, 1952b. King, 1956. Labrecque, 1957. Warner, 1958. Kenyon and Rice, 1959. Svihla, 1959. Rice, 1960a. Rice, 1960b. Smythe, 1960. King and Harrison, 1961. Udvardy, 1961b. King, 1964, Butler and Udvardy, 1966. (continued) Reports that mules and a few hogs were present in 1891. Reports that 5 seals were seen in December 1936. Summarizes earlier information on monk seal (with liberal quotes). Original information includes specimen data on seal collected in December 1912 and on 2 specimens collected by the Tanager Expedition; observations made in December 1936, seal count made in June 1951. Detailed summary of previous information on Hawaiian monk seal. Gives a few notes on seals made in July 1957. Gives a few observations of seals from May-= June 1958. Results of Hawaiian monk seal aerial survey January and April 1957, and ground survey June-July 1957. Summarizes earlier population data and gives results of counts or estimates made in May 1951, November 1954, and February 1955. Records offshore observations of bottle- nosed dolphins on 27 May 1958. Gives Hawaiian monk seal observations from aerial surveys in December 1957 and June 1958, and from ground survey May-June 1958. Reports seal observations made April-May 1959. Gives detailed notes on a young seal col- lected 4 June 1958. Gives results of seal counts made September 1961. Presents historical summary 1824-1954. Reports observations made July 1959 on the basking behavior of the monk seal. 328 _ Appendix Table 3. (continued) Mammalia (continued) Schreiber and Reports movement of a monk seal tagged on Kridler, 1969. Laysan in March 1968 to Johnston Atoll, 547 nautical miles to the south-southeast. Flora Reinbold, 1899. Reports algae collected by Schauinsland. Schauinsland, 1899. Lists 2/7 species of vascular plants with notes on them from observations and col- lections made in 1896; lists more algae collected; and mentions two species of algae found in the lagoon. Bitter, 1900. Reports 26 species of vascular plants col- lected by Schauinsland. Fisher, 1903a. Reports list of vascular plants collected in May 1902. Lemmermann, 1905. Reports 45 species of algae collected by Schauinsland. Wilder, 1905. Lists plants introduced in September 1905. Elschner, 1915. Gives observations of vascular plants and an algae made in September 191. Rock, 1916. Describes sandalwood on Laysan as a new variety from collections by Schauinsland, Bryan, and Fullaway. MacCaughey, 1918. Brings together Lemmermann's records and others. Christophersen and Reports on 4 vascular plants collected by Caum, 1931. the Tanager Expedition and summarizes records and observations from visits in 1896, 1902, 1903, and 1911. Lamoureux, 1963. Reports 24 species of vascular plants col- lected in September 1961 and gives historical summary; reports plants colleeted in August 1930 and April and July 1959. Tsuda, 1965. Summarizes earlier records and reports algal collections made in April 1923 (43 species or 32h Appendix Table 3. (continued) Flora (continued) Tsuda, 1965. (cont'd.) varieties), September 1961 (20) and December Tsuda, 1966. Saito, 1969. Geophysical Schauinsland, 1899. Muhle, 1902. Elschner, 1915. Kroenke and Woollard, 1965. Tsuda, 1965. 1963 (57). A total of 106 species or vari- eties listed of which 72 are new records. Also gives observations of 22 species of vascular plants collected in December 1963. Corrects specific epithets of algae (3 species of Liagora; 1 Halimeda) that were listed inaccurately in Tsuda, 1965. Reports 4 species of algae (Laurencia) identifying and reidentifying collections made by Tsuda in December 1963. Gives extensive descriptive notes on the island including analyses of guano and lagoon water; occurrence of pumice, notes on cli- matic conditions. Reports studies of a few rock samples col- lected by Schauinsland. Gives various descriptive material on the geology as well as analyses of lagoon water, bird excrements, and guano. Gives gravity observations made in March 1964, Reports salinity data for water collected at the ocean edge and in the central lagoon. »-1. 325 STe10OL odo 89OT og Sie eke} Q6T 9 &8 6ST TE 9eT TOT 42 LE 6einaie eG T 629°9 006°69 000'ZS 006‘T St ie D )), Lt it 9 €T € OT val Z cI LE + 12 oe wh €T ge Ge OTT LT 40% HOE €9T 3g GT OT 16 get lzn 00g € We € 9 rf 61 TS Q6T G6T a GOE J TOT OOT 6 r{o) 612 g6 TLT‘T =: 96h G20*S 002 Ege jis OOT HnS*Z 259 66€ L96T 996T G96T + €96T 00¢ 00¢ STeZ0], youtd ueskeyT use, 89 TUM AppoN ¥oeta Appon umorg uzay, Ayoos Ute], peyoeq-Aery auoysuany, Appny Z2T932] SUTTapuem MeTIND peystyy-eTIsTId ZJ2ACTd uapTtoy Tee, uesfeyT PiTgezestag Yeory &qoog pezoos-pey kqoog umorg Aqoog pedej-en Tg patgotdorzl, pettey-ped Teryed wr0yis AWo0o0g TIVeMTB9yG seuysTaAyO TaYeMIeVsyS peTTej-espom Terjed §,7eaTng Terzeq uTuog ssoreqrty uesfeT ssozqZeqry peyoos-Hoela *soTouese snotTzea fq uesfeT uo papueq spxtq Jo s[¥}04 Sutpueq ATreaR “*T-+ eTqQeL xppueddy *SZUSMOAOCU UT POATOAUT SPITq JO Tequmu [e104 3} ST Seseujueted ut TequmNx, *SpueTST OM} Ue. eTOM UO peangdeoer atom Spitq fo Jequmu e sodUTS SuTAom spatq jo Zequmu [e404 ueY. 10780878 9q AeW syUSMAeACM Jo saequmuU TeIOL,. 326 (y2T)92T (g)Q@ Hon (E)E (T)T (T)T (SE)BE (T)T (92)9@ (IT (T)T (h)h ex(Z)e (07) TeI0L B29 4V € eysely ueder spuetTs] eutddtrttud TIOVW nsgervy pueTs{[ xtusoud pueTsT seuqstazyo TIo9Vy upeTefeny TL99,V nwen T T pueTST oyem on €?g +1 G TlIo1y uoysuyor 9 o ali T 1 T TL0O7V eany S T Z T T TIo7vy ASMPTW 8 T T € T ro) joeay Souzey pue [ree O€ OT I tT 1 1 FASUCTSTT O€ vE € +1 + Z STeoyg e1eSTigq youezq7 T T seToeUuUuTg Teupzey a wy ta rs Q cd @ a w bo ta = w EB ha Ro swory ueskeyT oF ct ie) (e) Qu KR @) jor fe} i= Oo Qa 5 S< of i) @) ct Qu Le 2) it) i] = (9) @ 0a Fee a) » 0 oo q 4 @ ct ct Hb 5 I Pi) 5 i) ct n Ze (e) La) ! 3 BOI = tc) re; w) ty ° ow » © ct ty O {e) @ I ky ct (o) Q ct © @ > no a 8 8 ct bs © ° 0) © FS ct = n O a 5 fas S 0g a o ro) RH 8 a ct cq n bh iy) < ® o) © io) ct {0je} co ow w Qu (= ct jon co) io © fo) o} 8 5 () toy" {e) (e} e) (p) joy) [- lo” lox n BK “ < a Q *g-h eTaeL xppueddy x° DUCTS] ueske] SUTATOAUT SPATq pepueq Fo 4JUEeWeAOM pUeTST194UT 327 Eze L6t (S)S (neh (t/t (mt (ne) 6g (OT eee (Tt (orm (EE (worg) TeqoL ST ST T T OT € eeS 4V q T T exSeLly G Gg WU T ueder S S iL + spueTsr eutddrtttud T il T TIOVV NFSIV T T T puels] xp~useoud T T U puels]l sewmzstzyoO T U T TI0vV uteteleny T I t TLOFV nueN € ro) 2 pue[sI ayem +6 2S rll 6€ Tl [TIo1y uorsuyor te gill 4 G yh g TIOVY e1ny , Z tl it TIoyy AeMpTW €L ig 1 aif T o jeaoy Soutoy pue [zee ie) SS iL 0g +1 0g THSUCTSTT S9 GE € + 2 S2 i STeoyg eqesTag youer7 T seToeUuUuTd Tapurey me oe ee es eS ee Gh "pepe wea 9 ct ©) a 0) ro ° i om «4 o # 5 © a ct a © i = o) Coin ni 90 Ou bon C em a ch 3389 Appendix Table 4-5b. (continued) Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex Lisianski I. (continued) 1517-27725 10-19-66 I U Laysan I. 09-06-67 S U 1757-30113 06-02-67 A U Laysan I. 09-06-67 A F (in roost- ing club) 1757-30151 06-03-67 A M Laysan I. 09-06-67 A U | 1757-30181 06-03-67 A F Laysan I. 09-06-67 A F 587-90052 09-01-67 A M Laysan I. 09-10-67 A M 587 -90068 09-01-67 A M Laysan I. 03-18-68 A M Midway Atoll, Eastern I. ~508-51758%* 08-15-57 U U Laysan I. 03-10-64 A U Pearl and Hermes Reef, Kittery I. 1737-26549 06-26-63 N U Laysanire 09-16-64 A F Johnston Atoll 05 -20-65* e Ss Pearl and Hermes Reef, Little North I. 5558-83563 06-23-63 N U Laysan I. 03-09-65 A F Pearl and Hermes Reef, Seal I. 1737-26562 06-26-63 N U Laysan I. 09-16-64 A F *Marked with orange leg streamer. **Banded by BSFW (DR). 334 Appendix Table 4-6a. Movements of Red-footed Boobies from Laysan. Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex 587-805 03 09-17-64 I U Johnston Atoll 05-26-65 I U 587-805 07 09-17-64 A U Laysan I. 08-09-65 A U Lisianski I. 09-02-67 A U 5587-80544 09-17-64 A U Johnston Atoll 02-15-66 A F 587-80553 09-17-64 A U East I., French 06-11-66 A U Frigate Shoals (with small chick) East I., French 03-11-67 A U Frigate Shoals East I., French 05-28-67 A U Frigate Shoals Trig I., French 06-24-67 A U Frigate Shoals (with large downy chick) q 587-80557 09-17-64 I U Laysan I. 09-07-67 A U . Johnston Atoll 01-06-69 A U 587-80561 09-17-64 A U Johnston Atoll 12=13=65.u Namen 587 -80564 09-17-64 A U Johnston Atoll 02-22-66 A F Laysan I. 10-20-66 A U Laysan I. 06-09-67 A U 587 -80570 09-17-64 Johnston Atoll 04-23-66 A U Johnston Atoll 01-06-69 U U 587 -80575 09-17-64 I U East I., French 08-13-66 S U Frigate Shoals Trig ie, French 08-23-66 A F Frigate Shoals Whale-Skate I., 06-05-67 A U French Frigate Shoals 335 Appendix Table 4-6a. (continued) Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex 587 -80580 09-17-64 I U Whale-Skate I., 08-31-65 S U French Frigate Shoals 587-80595 09-17-64 I U Lisianski I. 08-31-67 A U 587-80598 09-17-64 I U Johnston Atoll 02-22-66 S U 587 -80607 09-17-64 A U Johnston Atoll 02-22-66 A F 587 -80620 09-17-64 I U Johnston Atoll 06-24-65 A U 587-80625 (ole en ly to) a Johnston Atoll 02-22-66 S U Lisianski I. 10-19-66* S U 587-80643 09-17-64 I U Whale-Skate I., 08-16-65 S U French Frigate Shoals 587 -80646 09-17-64 I U Johnston Atoll 03-30-65 I U 587-80663 09-17-64 I U Lisianski I. 10-19-66 A U 587-80697 09-17-64 A U Johnston Atoll 02-15-69 A U (roosting) 587 -80700 09-17-64 A U East I., French 06-12-66 S$ U Frigate Shoals 767-41051 03-07-65 A U Laysan I. 10-20-66 A U Laysan I. 09-07-67 A U Johnston Atoll 02-15-69 A U (roosting) Johnston Atoll 06-25-69 A U 767 -41069 03-07-65 A U Lisianski I. 10-19-66 A U 767-41072 03-07-65 A U Johnston Atoll 02-18-66 A M 767 -41098 03-07-65 A U Johnston Atoll 02-18-69 A U 336 Appendix Table 4-6a. (continued) Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured . Date Age Sex 767-41204 03-07-65 AU Laysan I. 10-20-66 A U Johnston Atoll 01-06-69 A U 767 -41231 03-08-65 A U haysant i ' 10-20-66 A U Johnston Atoll 01-05-69 U U 767-41236 03-08-65 A U Johnston Atoll 02-22-66 A F 767 -41241 03-08-65 A U East I., French 06-10-66 A U Frigate Shoals East I., French 05-26-67 A U Frigate Shoals (on egg) 767-4142 03-08-65 A U Johnston Atoll 01-04-69 A U Johnston Atoll 01-07-69 A U (dead?) 767 -41401 03-10-65 A U Trig I., French 06-24 -68 Frigate Shoals 767-4404 03-10-65 A U_ Kure Atoll 06-30-68 A 767 -41411 03-10-65 A U East I., French 06-12-66 AP Frigate Shoals Laysan I. 10-20-66 A U 767-4436 03-10-65 A U Laysan I. 10-20-66 A U Johnston Atoll 01-05-69 A U 767 -41469 03-10-65 A U Laysan I. 10-20-66 A U Trig I., French 06-19-67 A U Frigate Shoals 757 -25639 08-09-65 I U Johnston Atoll O4-23-66 I U 1757-25643 08-09-65 I U_ Lisianski I. 06-04-67 S$ 1757-25650 08-09-65 I U East I., French 06-10-66 S U Frigate Shoals 337 Appendix Table 4-6a. (continued) Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex 1757-25650 08-09-65 I U_ Whale-Skate I., 07-02-66 S U French Frigate Shoals Trig I., French 08-13-66 S U Frigate Shoals 7157-25652 08-09-65 -I U Lisianski I. 09-03-67 S U 7157-25656 08-09-65 I U Tisiansks 1. 10-19-66 S U Lisianski I. 09-03-67 S U 1757-25658 08-09-65 I U Namu Atoll, Mar- 03-24-66 UU shall I., ca. 08°00'N, 168°10'E 7157-25662 08-09-65 I U Wake Island 06-18-66 S U 7157-25667 08-09-65 I U Johnston Atoll 12-28-65 I U Whale-Skate I., 08-15-66* S U French Frigate Shoals 1757-25668 08-09-65 I U Johnston Atoll O41-03-66 I U 7157-25693 08-10-65 I U Trig I., French 06-08-67 S U Frigate Shoals 157-5742 08-09-65 I U_ Whale-Skate I., 06-26-66 S U French Frigate Shoals 157-25 744 08-09-65 I U Kure Atoll 08-07-66 S U 157-5751 08-09-65 I U Lisianski I. 10-19-66 S U 7157-25754 08-09-65 I U East I., French 06-10-66 S U Frigate Shoals Lisianski I. 10-19-66 S U Johnston Atoll O4-03-67 S U 338 Appendix Table 4-6a. (continued) Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex | 7157-25759 08-09-65 I U East I., French 05-26-67 S U | Frigate Shoals . 7157-25769 08-09-65 I U Whale-Skate I., 06-25-66 Ss U French Frigate Shoals 7157-25779 08-09-65 I U Johnston Atoll 03-31-66 i Oi 7157-25832 08-09-65 I U Wake I. 06-19-66 S U Wake I. 12-30-66 S U 7157-25835 08-09-65 I U Johnston Atoll 01-01-69 A U 7157-25837 08-09-65 I U Johnston Atoll 03-31-66 I U 157-25 O41 08-09-65 I U Lisianski I. 10-19-66 Ss U 157 -25843 08-09-65 I U Whale-Skate I., 06-04-67 S U French Frigate Shoals 7157-25850 08-09-65 I U Johnston Atoll 05-27-66 I U 7157-25853 08-09-65 I U East I., French 06-10-67 Ss U Frigate Shoals Whale I., French 06-17-67 S U Frigate Shoals 157-25 854 08-09-65 I U Johnston Atoll 02-22-66 I U 157-25 856 08-09-65 I U Johnston Atoll 02-18-69 A U (roosting) | 757-25896 08-09-65 I U Johnston Atoll 02-23-66) Eamau 71757-25898 08-09-65 I U Johnston Atoll O3=31-65,. ent Laysan I. 10-20-66 S U ; Johnston Atoll 02-18-69 Au : (roosting) 1757-25921 08-09-65 I U Johnston Atoll 02-22-66 I U 339 Appendix Table 4-6a. (continued) Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex 1757-25931 08-09-65 I U Kure Atoll 09-16-66 I U 7157-25932 08-09-65 I U Johnston Atoll 01-06-66 S U (dead) 7157-25953 08-09-65 I U Johnston Atoll 06-12-66 Ss U 7157-25986 08-09-65 I U Laysan I. 10-20-66 S U Kure Atoll 05-30-67 S U 7157-25897 08-09-65 I U Whale-Skate I., 06-03-67 S U French Frigate Shoals 1757-25682 08-10-65 I U Kure Atoll 06-04-67 S U 1757-25693 08-10-65 I U Trig I., French 06-08-67 S U Frigate Shoals 1757-25791 08-10-65 I U Whale-Skate I., 06-06-67 S U French Frigate Shoals 1757-25798 08-10-65 I U Lisianski I. 10-19-66 S U 1757-28125 10-21-65 A U Johnston Atoll 01-06-69 U U 1757-28129 10-21-66 A U Southeast I., Pearl 08-28-67 A U and Hermes Reef 1757-28513 10-22-66 A U Trig I., French 06-08-67 A U Frigate Shoals 1757-28517 10-22-66 A U Lisianski I. 09-04-67 A U 157 -28533 10-22-66 A U Trig I., French 06-19-67 A U Frigate Shoals 587-85218 06-09-67 S U_ kure Atoll 07-04-67 S U 587-85 223 06-09-67 A U Johnston Atoll 02-18-69 A U (with egg) (roosting) 587-85 237 06-09-67 A U Johnston Atoll 01-05-69 A U 310 Appendix Table 4-6a. (continued) Original Banding Data Recapture Data ! Band Number Date Age Sex Where Recaptured Date Age Sex | 587-85 244 06-09-67 A U Lisianski I. 09-03-67 A U *Marked on leg with orange streamer. Appendix Table 4-6b. Movements of Red-footed Boobies to Laysan. Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex East I., French Frigate Shoals 767 -43301 08-07-65 S U East I. 06-11-66 A U Laysan I. 10-20-66 A U Laysan I. 09-07-67 A U 767 -43851 08-25-65 S U Laysan I. 10-22-66 Ss .U Trig I., French 06-19-67 S U Frigate Shoals 1757-26094 06-10-66 A U Laysan I. 09-10-67 A U 7157-26175 06-11-66 Ss U Laysan I. 09-07-67 A U 1757-26187 06-11-66 A U Laysan I. 10-20-66 A U 1757-29055 05-26-67 I U Laysan I. 09-06-67 S U 757-36081 06-10-67 (I) U Laysan I. 09-10-67 (A?) U Trig I., French Frigate Shoals 1517-27495 07-03-66 Ss U Laysan I. 09-08-67 A U 1757-27615 08-13-66 S U_ Laysan I. 09-10-67 S U 157-35 854 06-08-67 S U Laysan I. 09-06-67 S U 341 Appendix Table 4-6b. (continued) Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex Original Banding Data Whale-Skate Island, French Frigate Shoals 1737-38033 06-14-63 S U Whale-Skate I., 06-23-66 A U French Frigate Shoals Laysan I. 10-20-66 A U 1597-27251 06-24-66 A U_ Laysan I. 10-21-66 A U Johnston Atoll | 1737-44156 02-26-64 I U East I., French 06-10-66 A U 3 Frigate Shoals Laysan I. 10-20-66 A U East I., French 05-27-67 A U Frigate Shoals (with small chick) 737-44.958 12-12-65 A U Laysan I. 09-10-67 A U 737 -44-960 12-12-65 A U Laysan I. 10-20-66* A U Laysan I. 09-07-67 A U 1737-44963 12-13-65 A U Laysan I. 09-06-67 A U 7737-44965 12-13-65 °9°A °U- ‘ Laysan: 2. 10-20-66 A U 737-44989 12-28-65 A U Laysan I. 06-12-66 U U Laysan I. 10-20-66 A U 737 -43532 02-19-66 S U Laysan I. 10-22-66 S U 1737-43562 02-22-66 I U Laysan I. 10-20-66 S U 737 -43569 02-22-66 A F Laysan I. 09-08-67 A U 737 -44686 02-23-66 A M _ ULisianski I. 06-18-66 UU Laysan ©. 10-20-66 A U 737 -44693 02-23-66 A M Laysan I. 10-22-66 A U 1737-43648 03-24-66 S U Laysan I. 06-29-67 S U 342 Appendix Table 4-6b. Original Banding Data (continued) Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex Kure Atoll 737-95987 12-30-64 A U_ Laysan I. 03-10-65 A U Lisianski I. 7157-27939 10-19-66 S U Laysan I. 09-07-67 S U 1757-27962 10-19-66 A JU Laysan I. 09-10-67 A U 1757-27984 10-19-66 S U Laysan I. 09-08-67 S U 7157-28457 10-19-66 A U Laysan I. 09-10-67 A U 1757-28499 10-19-66 U U Laysan I. 09-10-67 A U 7157-28705 09-02-67 s(7)U Laysan I. 09-06-67 A(?)U Midway Atoll, Eastern I. 767 -4.0297 07-22-65 N U Laysan I. 10-20-66 S U Wake I. 767 -48159 06-18-66 I U Laysan I. 10-20-66 S U Kure Atoll 07-04-67 S U *Marked with orange streamer on leg. Appendix Table 4-7a. Original Banding Data Movements of Great Frigatebirds from Laysan. Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex 587 -80701 09-20-64 A #F(?) Kure Atoll 05-07-66 S U 587-80708 09-20-64 A F Trig I., French 06-24-69 A F Frigate Shoals 757-25 564 06-12-66 A M(?) East I., French 05-30-67 A F(?) Frigate Shoals 343 Appendix Table 4-7a. (continued) Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex 757 -28563 10-22-66 U U_ kure Atoll 05-09-67 S F 7157-28576 10-22-66 L U Kure Atoll 07-20-68 St well 1757-28622 10-22-66 L U In Adanan River, 01-12-68 U U Bayugan, Agusan Province, Philippine Ws Gas CPIOVN 125°HO'E 757 -28634 10-22-66 L U Balusan, Sorsogon, 02-77-68 UU Philippine Is. ca. 12°40'N, 124°00'E 7157-28653 10-22-66 L U Kure Atoll 05-30-68 S M 1757-28697 10-22-66 L U In the Sea of Bawang, 03-13-68 UU Secogon I., (Carlos Mun.) Philippine Is. Cael 20 "Ne | 123 110. 'b 587-85 247 06-09-67 S U Near Virac, Philippine03-77-68 U U IES CEs aS ea OUINS 124°10'E 587-85364 06-09-67 S U Kure Atoll 06-30-67 S F *Found floating on river. Bird subsequently died. Appendix Table 4-7b. Movements of Great Frigatebirds to Laysan. Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex Kure Atoll 767 -45160 05-16-66 S U Laysan I. 10-20-66 Ss U 3h) Appendix Table 4-8a. Movements of Sooty Terns from Laysan. Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex 1793-82159 09-17-64 A U Lisianski I. 06-16-66 A U 793 -82184 09-17-64 A U Lisianski I. 06-04-67 A U 823-01100 09-17-64 A U Lisianski I. 06-18-66 A U 823 -18811 07-17-65 A U_ Lisianski I. 06-04-67 A U 843 -63403 07-18-65 I U Johnston Atoll 06-15-69 A U 843 -65340 07-18-65 A U East I., French 06-15-68 A U Frigate Shoals 843 -67038 07-19-65 A U Johnston Atoll 08-10-67 A U 893 -00856 08-05-65 A U Johnston Atoll 08-19-65 A U 893-01179 08-05-65 A U Lisianski I. 06-17-66 A U 893-01681 08-05-65 A U Lisianski I. 06-17-66 A U 893 -05046 08-05-65 A U Eastern I., Midway 06-21-66 A U Atoll 893 -07386 08-05-65 A U Lisianski I. 06-04-67 893 -094.02 08-06-65 A U Lisianski I. 06-18-66 A U 893 -09661 08-06-65 A U East I., French 05-28-67 A U Frigate Shoals 893-10752 08-06-65 A U Honshu, Japan, O4-26-68 U U 35°20'N, 139°35'E 893-11955 08-06-65 A U ULisianski I. 09-03-67 A U 893 -12201 08-05-65 A U Kure Atoll 05-13-67 A U 893 -16895 08-06-65 I U Johnston Atoll 06-16-69 A U 893-17133 08-06-65 I U Johnston Atoll 09-23-68 A U 863 -62311 08-07-65 A U lMisianski I. 06-19-66 A JU 893 -12998 08-07-65 A U Johnston Atoll 08-07-67 A U 3a) Appendix Table 4-8a. (continued) Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex 893-16158 08-07-65 I U Atafu Atoll 01-25-67%* AU 863 -65847 08-08-65 A U Johnston Atoll 08-15-67 A U 863-67412 08-08-65 A U Lisianski I. 06-18-66 A U 863-67615 08-08-65 A U_ Lisianski I. 06-18-66 A U 863-69621 08-09-65 I U At sea, 11°09'N, Before ie 101 140°53'W 12-06-65 863-70245 08-09-65 A U Eee ie, bine 12-29-67 A U ce 863-70414 08-09-65 A U_ Kure Atoll 05-16-66 A U Kure Atoll 06-07-67 A U (breeding) 863 -70890 08-09-65 A U_ Awayon, Silago, 12-17-65 Ui U Southern Leyte, Philippine Is. 863 -71956 08-09-65 A U Lisianski I. 06-04-67 A U 863 -72007 08-09-65 A U Paocais I., Phoenix 01-28-67 A U Se. 863-74962 08-10-65 AU Johnston Atoll 07-11-66 A U 903 -33483 06-11-66 A U East I., French 08-23-66 A U Frigate Shoals 903 -41890 06-11-66 A U Lisianski I. 06-04-67 A U 903 -34.664. 06-12-66 A U Ebjedik I., E. Kwa- 06-21-67 A U jalein, Marshall Is. 903 -43528 06-12-66 A U Lisianski I. 06-04-67 A U 903 -43923 06-12-66 A U Lisianski I. 09-03-67 A U 903 -45621 06-12-66 A U Kure Atoll 06-14-67 A U (nesting) 903-50101 06-13-66 A U Kure Atoll 06-12-67 A U (nesting) 346 Appendix Table 4-8a. (continued) Original Banding Data Recapture Data Band Number Date | Age Sex Where Recaptured Date Age Sex 903 -60540 06-13-66 A U Lisianski I. 06-04-67 A U 903 -61808 06-13-66 A U Lisianski I. 06-17-66 A U 903 -61956 06-13-66 A U Johnston Atoll 07-10-66 A U 903 -63810 06-13-66 A U Lisianski I. 06-03-67 A U 903 -5 2322 06-14-66 A U Lisianski I. 06-04-67 A U 903 -64.636 06-14-66 A U East I., French 05-28-67 A U Frigate Shoals 903 -65555 06-14-66 A U Johnston Atoll 07-25-67 A U 903 -82859 06-15-66 A U Johnston Atoll 05-23-69 A U (on egg) 993 -45954 06-07-67 A U Johnston Atoll 08-07-67 A U *Blood sampled. **Caught by a native hunter. Appendix Table 4-8b. Movements of Sooty Terns to Laysan. Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex French Frigate Shoals 723 -62369 06-09-63 A U Laysan I. 08-05-65 A U 863 -26728 08-25-65 A U Laysan I. 06-10-67 A U (with downy chick) 863 -26796 08-25-65 A U Laysan I. 06-10-67 A U (on nest) 347 Appendix Table 4-8b. (continued) Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex Johnston Atoll 7153-14254 08-02-63 A U Laysan I. 06-10-67 A U 1753-20707 08-26-63 A U Laysan I. 07-19-65* A U 7153-20853 08-26-63 A U Laysan I. 06-10-67 A U 753 -20889 08-26-63 A U Laysan I. 06-15-66 A U 1753-21404 08-31-63 A U aysam ele 09-17-64 A U 753-21699 09-01-63 A U lLaysan I.¥* 06-14-66 A U 7153-23285 99-06-63 A U Laysan I. 07-19-65 A U 1753-23432 09-09-63 A U Laysan I. 06-15-66* A U 1753-23733 09-10-63 A U Laysan I. 08-08-65 A U 71753-24138 09-15-63 A U Laysan I. 06-10-67 A U Laysan I. 06-08-67 A U 7153-24227 09-15-63 A U Laysan I. 06-10-67 A U 753 -bh23 09-17-63 A U_ Laysan I. 06-15-66 A U 7153-82465 05-14-64 L wU Laysan 1. 07-18-65 A U 843 -78332 07-17-65 A U Laysan I. 08-10-65 A U 84.3 -78822 07-18-65 A U Laysan 1. 08-08-65 A U 843 -78978 07-18-65 A U Laysan I. 06-10-66 A U 843 -82173 07-22-65 A U Laysan I. 06-07-67 A U 843 -82822 07-22-65 A U Laysan I. 08-08-65* A U 84.3 -83669 07-24-65 A U Laysan I. 06-11-66* A U 843 -83987 07-24-65 A U Laysan I.** 06-10-66 A U 843 -85955 07-26-65 A U Laysan I. 08-05-65 A U 348 Appendix Table 4-8b. Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex Johnston Atoll (continued) 843 -91342 843 -99861 Kure Atoll 1773-19689 813 -91288 Lisianski 823 -19014 823 -19043 823 -19068 823 -19072 903 -904.22 903 -96566 903 -98068 943-14501 943 -17848 943 -18114 08-06-65 08-22-65 05-20-64 08-27 -64 07-16-65 07-16-65 07-16-65 07-16-65 06-16-66 06-16-66 06-16-66 06-18-66 06-18-66 06-77-66 A A (continued ) U U U U U Laysan Laysan Laysan Laysan Laysan Laysan Laysan Laysan Laysan Laysan Laysan Laysan Laysan Laysan Laysan 08-10-65* A U 06-11-67 06-20-66 06-20-66 06-12-66 06-07 -67 06-10-67 06-12-66 06-11-66 06-13-66 06-13-66 06-10-67 06-10-67 06-10-67 06-11-67 06-11-67 06-11-67 A VeU A U (breeding) AD A U A aU) AU A U A U A U AU A U (nesting) A U (nesting) A Ui (nesting) A U (breeding) A OU (breeding) A U 349 Appendix Table 4-8b. (continued) Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex Midway Atoll, Eastern I. 903 -00280 08-28-65 A U Laysan I. 06-10-67 A U 903 -02280 08-29-65 A U Laysan I. 06-11-67 U U *Marked with orange leg streamer. **Bird collected. Appendix Table 4-9a. Movements of Black Noddies from Laysan. Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex 723 -60413 02-12-63 A U Trig I., French 06-24-68 A U Frigate Shoals 723-6042} 02-12-63 AU Whale-Skate I., 08-30-65 A U French Frigate Shoals 723 -60435 02-12-63 A U Southeast I., Pearl 05-30-67 A U and Hermes Reef 723 -60446 02-12-63 A U Whale-Skate I., 08-15-65 A U French Frigate Shoals 712-00697 10-22-66 U U Lisianski I. 09-02-67 A U Appendix Table 4-9b. Movements of Black Noddies to Laysan. Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex Whale-Skate I., French Frigate Shoals 863 -23696 08-15-65 A U Laysan I. 03-18-68 A U 350 Appendix Table 4-9b. (continued) Original Banding Data Recapture Data Band Number Date Age Sex Where Recaptured Date Age Sex Whale-Skate I., French Frigate Shoals (continued) ———— 712-01740 08-16-65 A U Laysan I. 09-10-67 A U 712-01782 08-16-65 A U _ Laysan I. 03-17-68 A U 712-01362 08-29-65 A U Laysan I. 10-22-66 A U 712-01477 08-29-65 A U Laysan I. 10-20-66 A U 863 -27548 08-29-65 A U Laysan I. 03-18-68 A U (with egg) 7112-58324 06-26-66 A U Laysan I. 03-18-68 A U (with egg) Midway Atoll, Sand I. 542-77233 08-10-62 U U Laysan I. 09-07-67 A U *Banded by BSFW. Appendix Table 5. Distribution of Laysan bird skins and mounts. More specimens have been taken at Laysan over a longer period of time than at any other locality in the Northwestern Hawaiian Islands. Most of the many specimens collected at Laysan are now in major American museums; others were exchanged to foreign museums and an unknown number are in private collections or small museums. The following listing was completed through an examination of museum. catalogues and (usually) skins present in the following museums: AMNH American Museum of Natural History, New York BMNH British Museum (Natural History), London BPBM Bernice P. Bishop Museum, Honolulu CMNH Chicago Natural History Museum, Chicago DMNH Denver Museum of Natural History, Denver JEL Law and Bailey collections, presently at Rockbridge Alum Springs, Virginia MCZ Museum of Comparative Zoology, Cambridge PANS Academy of Natural Sciences of Philadelphia SUL State University of Iowa, Iowa City =——_ 351 Appendix Table 5. (continued) UMMZ University of Michigan Museum of Zoology, Ann Arbor USNM United States National Museum, Washington D.C. (Also included are a few specimens from the D.R. Dickey (DRD) collection, now at the University of California, Los Angeles (7) ). This listing includes all major Laysan collections but time did not permit the checking of all individual Laysan specimens. Whenever possible, museum catalogues were checked against specimens present in the collections. In appendix tables specimens are listed under the museum where catalogued. Coverage of various museums is uneven: AMNH was covered in much better depth than was MCZ; data from AMNH and USNM are more accurate than those at BPBM where duplications and errors are known. Despite errors, this listing should be of value to anyone working with the Hawaiian area and the central Pacific. Hopefully, others using this listing will make ap- propriate corrections when errors are found. Appendix Table 5-1. lLaysan specimens in major museums. Species AMNH BPBM CMNH DMNH MCZ Misc. SUI UMMZ USNM Totals Black-footed Albatross ie ae 6 i 2 3 ey me ak 82 Laysan Albatross 26 15 8 7 4 OP wh hr Be 136 Hybrid Albatross 1 dL iL 3 Bonin Petrel a 420 6 8 2 ron 2 121 Bulwer's Petrel 12 3 i Gy 1 8 29 valle Sooty Shearwater 1 iL Wedge-tailed Shearwater 12 18 2 iL Sh wes Suess 72 Christmas Shearwater De ly) dell 4 4 5 1 Mh Te hi) 88 Sooty Storm Petrel 7 6 3 2 pe LON Seay 46 Red-tailed Tropicbird a4 12 6 8 1 6 18 16 92 Blue-faced Booby ee tO) 3 2 O55 Ty lS 54 Brown Booby ale 2 3 Red-footed Booby ast t 3 2 y Oh ee mel 53 Great Frigatebird 14 9 6 y 4 hen Ee BS) 87 Mallard 2 2 Laysan Teal PPG 2 6 2 13 61 Pintail al 1 Bufflehead 1 al) Harlequin Duck 1 1 Laysan Rail 46 = 28 4 3 3 2 4 1G | BS 140 Semipalmated Plover 1 iL Golden Plover 10 3 2 a 2 Do Sine NS) 53 Black-bellied Plover dl dL Ruddy Turnstone 8 5 2 6 2 3 "6." 25 57 Bristle-thighed Curlew 12 8 iG 6G AS) 2 Bh soi Tec AUG) 77 Wandering Tattler 8 2 rials 4 21 Greater Yellowlegs 1 ik 352 Appendix Table 5-1. (continued) Lesser Yellowlegs Sharp-tailed Sandpiper Pectoral Sandpiper Baird's Sandpiper Dunlin Marbled Godwit Bar-tailed Godwit Sanderling Red Phalarope Northern Phalarope Glaucous Gull Herring Gull Bonaparte's Gull Black-legged Kittiwake Sooty Tern Gray-backed Tern Brown Noddy Black Noddy White Tern Laysan Miller-bird Laysan Honey-eater Species AMNH BPBM CMNH DMNH MCZ Misc. SUI UMMZ USNM Totals 2 2 al i 2 2 2 2 a il il it a a 1 i i. i 2 2 2 1 i al 2 3 a a 1 a 17 6 22 4 eT 3 786 107 23. «4k 2 OP aaeeliste 2. 16" Ota S6 121 14 5 3 iG 7 leeclS iG aa 92 Be 115) 5 8 6 Pe eG.) 8S) iam 22: i KG 2 3 4 C. Bouag CG 22° 25 2 2 9 4 eee 82 16 7) 23 2 2 32 2 SES Roy 8 2a) 90 45 =e 2 2 9 2 5 i 198 Laysan Finch Totals Appendix Table 5-2. Laysan specimens at the American Museum of Natural History (includes mounts). Bailey Coultas Misc. Rothschild Schauinsland Schlemmer Bryan ID)ah aL Hall Species Black=-footed Albatross Laysan Albatross Bonin Petrel Bulwer's Petrel Wedge-tailed Shearwater Christmas Shearwater Sooty Storm Petrel Red-tailed Tropicbird Blue-faced Booby Brown Booby Red-footed Booby Great Frigatebird W OVO’ PP ine} bw | el MMM wN hk ee) ne = HW fD 5B) Appendix Table 5-2. (continued) o iS ao das& ed n © cal 3 S S | nv o Ss P . 1) =) @) o a ee Se SS teem o Aig Seals oat ES ia ~ @ > el w& -« 5 © fs io) Species A a o aA & SF 8 aA a = Laysan Teal ORs Guy lon al 22 Laysan Rail 4 a 29 12 46 Golden Plover 10 10 Ruddy Turnstone 8 8 Bristle-thighed Curlew ape de a cae TRS 12 Wandering Tattler 8 8 Bar-tailed Godwit aL aL Sooty Tern 1 1 Cols} ALY Gray-backed Tern ee wile i) dik 23 Brown Noddy 1 13 14 Black Noddy 2 5 a 22 White Tern 2 oy 2 22 Laysan Miller-bird 3 1 fer lO 22 Laysan Honey-eater 3 1 Cane 16 Laysan Finch Laie To een Hy Gh SIg) 45 Totals 12 29 39 6 5). iO) 1s; 252 3 459 Appendix Table 5-3. lLaysan specimens at the Bernice P. Bishop Museum (includes mounts). Species Bryan Hall Misc. Rothschild Schauinsland Wetmore Total Black-footed Alba- tross Tl 5 12 Laysan Albatross slay aL ale 2 15 Hybrid Albatross i 1 Bonin Petrel 9 2 1 5 3 20 Bulwer's Petrel 2 HE 3 Sooty Shearwater al i Wedge-tailed Shear- water 6 aL 5 2 2 2 18 Christmas Shearwater 1 2 i i 11 Sooty Storm Petrel 6 6 Red-tailed Tropic- bird Tf 1 4 12 Blue-faced Booby 2 al i 10 Brown Booby ab 1 2 Red-footed Booby 2 L us if Great Frigatebird iL 3 3 2 9 354 Appendix Table 5-3. (continued) Golden Plover Ruddy Turnstone Bristle-thighed Curlew Red Phalarope Sooty Tern Species Bryan Hall Misc. Rothschild Schauinsland Wetmore Total Laysan Teal 12 i. a 2 16 Pintail iL i Harlequin Duck 1 all Laysan Rail 15 8 5 28 Golden Plover 2 1 3 Ruddy Turnstone al 4 5 Bristle-thighed Curlew 2 2 2 2 8 Wandering Tattler ill aL 2 Glaucous Gull aL als Herring Gull dL at Black-legged Kitti- wake il aL Sooty Tern 3 2 a 6 Gray-backed Tern 4 2 ell ve 4 Brown Noddy it al 3 5) Black Noddy 13 a. alt i155 White Tern 6 3 5 2 al ay Laysan Miller-bird 21 } 5 Laysan Honey-eater 19 1 2 all 23 Laysan Finch oh 12 3 3 he Totals 166 16 68 3h aby 2 341 Appendix Table 5-4. lLaysan specimens at the Chicago Museum of Natural History (includes mounts). Species Bailey Dill Schlemmer _ Misc. Totals Black-footed Albatross 2 4 6 Laysan Albatross 4 4 8 Hybrid Albatross all i Bonin Petrel 4 2 6 Bulwer's Petrel iL aL Christmas Shearwater 2 y Sooty Storm Petrel 3 Red-tailed Tropicbird 1 3% 6 Blue-faced Booby il 3 Red-footed Booby 1 3 Great Frigatebird 2 1 6 Laysan Rail a 1** 4 2 2 16 i 2 MURPANNNMW~ANNNMWD 355 Appendix Table 5-4. (continued) Species BawleyDLLE Schlemmer Misc. Totals Gray-backed Tern Brown Noddy Black Noddy White Tern Laysan Miller-bird Laysan Honey-eater Laysan Finch Totals 2D 23 1 4 83 MMM MW WM wy NM WM MAW fo *Japanese hunters, 1909-1910. **Townsend (captive in Honolulu). Appendix Table 5-5. lLaysan specimens at the Denver Museum of Natural History (includes mounts). Species Bailey BSFW Dill Misc. Wetmore Totals Black-footed Albatross 6 Laysan Albatross Bulwer's Petrel Wedge-tailed Shearwater Christmas Shearwater Red-tailed Tropicbird 4 Blue-faced Booby 2 Red-footed Booby Great Frigatebird Laysan Teal 2 Laysan Rail Golden Plover Bristle-thighed Curlew Sooty Tern Gray-backed Tern Brown Noddy Black Noddy White Tern Laysan Miller-bird Laysan Honey-eater aL Laysan Finch Mr FAH nN fh -h 1* 1* kh WNMNMWONONMArPWhwWFMNM OFM ANA UMMA NYH 1* MVRPNMNMFNMUAFTF W Totals 26 2 V4 it ine) Went Xe) No) *Japanese hunters, 1909-1910. 356 Appendix Table 5-6. Laysan specimens at the Museum of Comparative Zoology. Species Schlemmer Bailey Misc. Totals Black-footed Albatross Laysan Albatross Bonin Petrel Bulwer's Petrel Wedge-tailed Shearwater Christmas Shearwater Red-tailed Tropicbird Red-footed Booby Great Frigatebird Laysan Teal Laysan Rail Ruddy Turnstone Bristle-thighed Curlew Sooty Tern Gray-backed Tern Brown Noddy Black Noddy White Tern Laysan Miller-bird Laysan Honey-eater Laysan Finch Totals igh mn q 139 he FONE ENA AFENME DOF hk | ae hk i Fob ER OAWAFENUE ND OF HH NOON PRE bh * Oo MO *From Flood brothers. Appendix Table 5-7. Laysan specimens at the State University of Iowa (includes 106 mounts). Species Albatross Bailey Dill Totals Black-footed Albatross 12 Laysan Albatross 34 34 Bonin Petrel Tt Bulwer's Petrel Wedge-tailed Shearwater Christmas Shearwater 2 Sooty Storm Petrel Red-tailed Tropicbird iL Blue-faced Booby Red-footed Booby aL Great Frigatebird dL 2 Laysan Teal Laysan Rail Golden Plover Ruddy Turnstone Bristle-thighed Curlew Wandering Tattler | ee MW MWFEWDYEF OWOWF WN | ee MWMWEWY FOO DNF Fuofl - Bonin Petrel Sil Appendix Table 5-7. (continued) Species Albatross Bailey Dill Totals Sooty Tern 2 2 27 Gray-backed Tern i 15 16 Brown Noddy 13 13 Black Noddy ale 18 19 White Tern 3 h 7 Laysan Miller-bird 3 3 Laysan Honey-eater it 3 y Laysan Finch 5 5 Totals als) 2 201 eal Appendix Table 5-8. Laysan specimens at the University of Michigan Museum of Zoology. Species Bailey Dill Misc. Totals Black-footed Albatross Laysan Albatross Wedge-tailed Shearwater Christmas Shearwater Sooty Storm Petrel Red-tailed Tropicbird Blue-faced Booby Red-footed Booby Great Frigatebird Laysan Rail Golden Plover Ruddy Turnstone Bristle-thighed Curlew Wandering Tattler sooty Tern Gray-backed Tern Brown Noddy Black Noddy White Tern Laysan Miller-bird Laysan Honey-eater Laysan Finch kh PR He FOFWNADOWNADOAWYHWO FH ON & 11* PH MAW EPMADWANHDOFMOHPHENMOWH ODS Derr bh hr Totals 133 19 Gy *Japanese hunters, 1909-1910. 358 Appendix Table 5-9. Laysan specimens at the U.S. National Museum (includes mounts). Species Albatross Bailey BSFW Dill Misc. POBSP Wetmore Totals Black-footed Albatross 2 6 y 12 ak Laysan Albatross 4 8 6 1 32 Hybrid Albatross 1 1 Bonin Petrel 3 26 2 4 16 al 52 Bulwer's Petrel 6 3 20 29 Wedge-tailed Shear- water ale 2 30 33 Christmas Shearwater 2 aly il 4 il iby 4O Sooty Storm Petrel aL iLL 1 4 17 Red-tailed Tropicbird 1 3 5 "i 16 Blue-faced Booby in 9 13 Red-footed Booby 4 11 15) Great Frigatebird 5 5 19 29 Mallard 2 2 Laysan Teal 3 3 iL 6 13 Bufflehead all BL Laysan Rail 7 26 2 35 Semipalmated Plover al! 1 Golden Plover 2 iL 1 5 4 13 Black-bellied Plover IL ill Ruddy Turnstone il 1 2 11 10 2 Bristle-thighed Curlew 3 5 10 18 Wandering Tattler iL iL 2 4 Greater Yellowlegs I aL Lesser Yellowlegs 2 2 Sharp-tailed Sandpiper alt 1 Pectoral Sandpiper 2 2 Baird's Sandpiper 2 2 Dunlin dl al Marbled Godwit 1 alk Sanderling 1 alt Red Phalarope all 1 Northern Phalarope 2 2 Herring Gull il 1 2 Bonaparte's Gull il 1 Sooty Tern 2 9 1 2 y 18 36 Gray-backed Tern 3 9 2 2 20 36 Brown Noddy aL 15 6 14 36 Black Noddy 3 14 a 2 13 16 4g White Tern 6 1 12 19 Laysan Miller-bird 6 2 3 11 Laysan Honey-eater 6 10 2 3 21 Laysan Finch 12 13 35 2 4 sla al Totals 65 183 L875 6 76 263 716 SBS) Appendix Table 5-10. Minor Laysan holdings. British Museum (Natural History) Species Rothschild Schauinsland Wetmore Totals Laysan Albatross aL IL Bonin Petrel 1 dl. Great Frigatebird y M Totals al! A 1 6 D.R. Dickey Collection Black-footed Albatross i 2 2 J.—~. Law Collection Species Bailey Total Black-footed Albatross 1 il Laysan Albatross 1 1 Bonin Petrel all all Christmas Shearwater 1 1 Sooty Storm Petrel 2 2 Laysan Rail 2 2 Golden Plover 2 2 Ruddy Turnstone 2 2 Bristle-thighed Curlew 2 2 Gray-backed Tern 2 2 Brown Noddy 1 dL Black Noddy 2 2 Laysan Miller-bird 2 2 Laysan Honey-eater 2 2 Laysan Finch 2 2 Totals 25 25 Academy of Natural Sciences 6f Philadelphia Species Dill Rothschild Schlemmer Totals Laysan Albatross als 1 Laysan Miller-bird 1 1 2 Totals 1 1 il 3 360 Appendix Table 5-11. Present deposition of specimens from major expeditions. Expedition AMNH BPBM BMNH CMNH DMNH DRD JEL MCZ SUI PANS UMMZ USNM Totals Albatross 13 65 78 Bailey 12 be 25) 25 My 133 183 LOX Bryan 29 166 195 Coultas 39 39 DTLILTL 6 2 52 201 Ee 5 362 Hall Cali 21 Misc. IO 6&8 4 5 4 19 6 116 Rothschild 103 «34 i il 139%* Schauinsland 252 15 4 27 1** Schlemmer 3 1 131 aL 136 Wetmore ke 1 wi 2 263 322 POBSP 76 76 BSFW 2 48 50 Totals 459 341 6 63 99 2 2 139216 3 bongo *A number of specimens were scattered among private collections. **An unknown number are in other European museums. Appendix Table 5-12. lLaysan specimens by expedition. ce) q ue} ay n P= | u un “o n {d) ae g 4 5 p © is} p . Ay a 3 G) fe) qt @ qo @ = qo q qo 2) icp) a ou o =| yj Q A E> Ss) fal co n faa) yp ect ict ~» » . fo | a Lop) [e) a [a0 ed Oo {e) oO Oo (0) {e) Species 4 aa (sats W2e se 69 | biiol4 dsiwoe To shovel J sysoboye 8 6 ; 2ravi nt spb Fring: A PRELIMINARY CORAL REEF ECOSYSTEM MODEL Edited by ee edition ben GoredistChan SeeaWe Mien ane J... Ge ZaleMmans Wi. A major goal of the CITRE planning effort was to develop a concep- tual framework for a systems model of a coral reef ecosystem suitable for computerization. Both the process of development and the prelimin- ary results may be of some value to future comprehensive studies of complex reef systems. The preliminary model was conceived not only to outline a possible mathematical model, but also to integrate the proposed research of many specialists into a coherent framework with defined interactions. While the specialists generated the model structure, they also had to modify their research plans to incorporate all major aspects of the system as identified in the model. No one model can deal with all aspects of an ecosystem. Its proper- ties, however, should ultimately resemble those of the system being modeled and its Jevel of abstraction should suit its intended purpose. Thus no model is static; it must evolve as available knowledge of the ecosystem increases. The preliminary model described here is one fixed point in such an evolving process. It represents an intermediate level of ecological abstraction, which, because of the diversity of the reef system, is nevertheless highly complex. It is based on the great diver- sity of viewpoints and professional experiences of the investigators involved. Yet, while its details are open to question, it represents a useful benchmark from which to proceed. The model was developed during a two-week workshop at Glover's Reef, British Honduras, in November 1971. Forty-one participants (see Intro- duction) representing a wide variety of reef-related disciplines and with experience in many coral reef areas attended this workshop. The principal activity was the development of a conceptual model sufficiently comprehensive that it could be applied to most reef areas. However, the shallow reef flat areas of Glover's Reef served as the prime focus for the initial effort and this may be reflected in some of the details presented here. Overall, the result represents a unique synthesis of current information about the tropical reef ecosystem. Process of Model Development CITRE participants were initially chosen to be members of one or more of the nine working groups (Introduction) with responsibility for a major aspect of coral reef ecology. Most working groups met before the workshop for initial discussion of their subject area and an intro- duction to modeling concepts. The workshop itself included lectures introducing the scientists to some aspects of modeling theory, and presentations by working groups as they fitted their subject matter nto the general modeling framework. The working groups proceeded by internal discussions, consultations with the modelers and with other working groups, and field observations to check on the general applicability of the model elements developed. The minor inconveniences of working at a remote site were far outweighed by the freedom from distractions and the ability to "consult the environment." The model was designed to depict the flow of carbon through the coral reef ecosystem. Each working group first defined the "compartments," or functional units between which the flows were to be measured. An approximate limit of twenty compartments per working group was set to fix the overall model complexity within present computer capabilities. Inter-group discussions resolved problems of overlapping or overlooked functional units. Diagrams were then developed for each compartment relating it to other compartments and flows in the system. The compart- ments were listed in a matrix, and the interactions between compartments determined jointly by the working groups involved. Some quantification of these interactions was attempted (at least in terms of orders of magnitude), but this was not completed and is omitted here. Revisions and adjustments continued throughout the workshop. The terrestrial section was kept simplified as the primary concern was its interactions with the reef system. The model developed through a cyclical progression. A statement of the purpose of the model and the definition of compartments was followed by the conceptualization of the initial model and field observations to check the mode] validity, leading to changes in the conceptualization and (more rarely) in compartment definitions. With each permutation the mode] became more refined and (hopefully) accurate. The Model Compartments Three basic elements characterize the model: compartments, flows (fluxes) and external driving forces (forcing functions). The compart- ments are the carbon or carbon-equivalent storage units of the system and may be defined as plant or animal types that act as processing units for the food which they ingest or produce, cr as material pools such as detritus or carbon dioxide through which substances pass as they cycle in the system. Compartments in the CITRE model may be considered "functional groups" in that they are abstract condensations of groups of organisms or substances that apparently have the same or similar func- tions inthe ecosystem. In this sense they are “ecological species" as opposed to conventional taxonomic species. For example, Halimeda, Penicillus, Rhipocephalus, Udotea, Jania, and Corallina are distinct genera of algae (even belonging to two phyla); yet in the model they are all considered part of the same compartment--the noncrustose, calcium carbonate-producing macroalgae designated XCBALG* [6] (see Table I). Likewise two compartments are distinguished on whether motile organisms remove solid substratum as they feed ("scrapers": designated XSCRAP [35] ), or eat only the overlying organisms ("browsers": XBROWS [362 ). Al] dissolved organic compounds containing nitrogen, and not associated with some other compartment, are similarly grouped (XDON [67] ). Compartment boundaries not only combine dissimilar species on functional grounds, in some cases they even separate parts of organisms with distinct roles or locations. Marine grass blades (XBLADE [9] ) were separated from marine grass roots (XROOT [10] ) because of the functional differences inherent in the location of these parts above versus below the sediment surface. Transport from one part of the plant to another thus becomes a flow between compartments. When it appeared that the inorganic carbon cycle could be most easily treated as a dis- tinct system, separate compartments were established for the calcium carbonate in the walls of living organisms (XORG [84] ) and the remaining (organic) carbon in the organisms. The organisms were therefore defined in the mode] as controlling the flow of carbon to their skeletons rather than flowing that carbon through the organism compartments. Table I enumerates the preliminary working compartments identified for the CITRE model, with their acronyms and numbers. Flows In order that compartments may function within the model, they must be linked or coupled; flows (or fluxes) between them are measured in whatever is the currency of the model--i.e., the material with which exchanges are made. Flows, like compartments, are based on the general objectives of the modeling program. In the case of the CITRE model the following materials were considered significant to the model of a coral reef ecosystem: carbon, inorganic carbon, organic carbon, nitrogen, phosphorus, calcium, biomass, and energy. It was finally agreed to flow carbon through this model because of its mutual importance to the biochemical and geochemical cycles, even though other flows are also important. For example, the nutrients (P0,, NO, etc.) which are important in feed-back loops controlled by photosynthésis,~are flowed in the model, but for mathematical consistency, are converted to carbon equivalents. * Although there is no standard convention for naming compartments, flows, or other components, certain consistencies make bookkeeping easier and more accurate, such as the use of acronyms. Here X---- stands for com- partment, thus XCBALG for CarBonate producing ALGae and XDON for Dissolved Organic Nitrogen. For convenience, all compartments were numbered consecutively as they appeared in the matrix (Fig. 9). 10 Forcing functions Forcing functions are driving forces or variables which originate outside the system of reference but which influence the behavior of the system. These variables include light, temperature, inputs of materials, and other influences not under the control of the system. The diagrams used in this model to illustrate compartments and flows are feedback dynamics or Forrester diagrams (Forrester, 1961). The units of compartments gues flows in the CITRE model are grams of carbon per sayaye Uren (gC m-2) and grams of carbon per square meter per day (gC day-!) respectively. Figure 1 shows the symbols used in the dia- arane: and briefly describes meanings attached to these symbols. Although these diagrams may become very complex, they are useful for a graphic representation of the model and as a means to facilitate discussion. The general characteristics of the total CITRE ecosystem model as described by these diagrams is outlined below. The forcing functions (external driving forces), compartments and flows between compartments, together delineate a preliminary total ecosystem model for a coral reef. Both the large number of compartments (104) and the number of flows between them form a very complex model. Figure 2 graphically illustrates the couplings and ecological rela- tionships of one compartment, the fleshy algae (XFLALG [5] ). C02 input from XDISOL (80), a compartment of common interest to the Geology and Nutrients-Detritus groups, is controlled by four integrating functions, three of which (#1 through #3) are abiotic functions that regulate poten- tial photosynthesis. Integrating function #4, coupled with the other three, gives realized photosynthesis. Dashed arrows from the integrating functions to the valves on the flows show that the integrating functions influence flow without contributing material to it, that is they are informational or control couplings rather than material flows or fluxes. Six feedback loops or cyclical flows, of three basic types, affect this compartment. Two similar loops (diagrammed as one) indicate the cycling of carbon, or carbon equivalents of oxygen, respectively, between the dissolved organic carbon (XDOC [62] ) or oxygen (X02 [70] ) compart- ments and XFLALG (5). These loops are controlled by the amount of the "upstream" or donor compartment present as well as by temperature, salinity, and exposure (integrating function #1). Another type of loop in the diagram shows the cycling of carbon to XTURF (4), XPHPL2 (12), and XPHPL3 (13). Flows to these three compartments from XFLALG (5) are reproduction, and flows back from them are recruitment from settling and growth. The third loop type is that diagramming CO? uptake from XDISOL (80) during photosynthesis, and its return during respiration. Outputs to XDON (67) and XDOP (69) are excretion, as is in part the flow to XDOC (62). Floating macroalgae, XDTPLT (8) break off from XFLALG (5) and continue living; detritus (XDETR3 [65] and XDETR4 [66] ) is similarly derived. XFLALG (5) also furnishes carbon to XBROWS (36), XGRAZV (48) and XBROWV (49), because these animals eat algae. The dotted arrows on this last set of flows show that these flows are controlled in part by the amount of algal material available, and in part by the biomass of the grazer. Six more "single compartment" models are also shown in a similar diagram format (Figures 3-8). These diagrams all resemble Figure 2 in their complexity and mode of working. Space does not permit the inclusion of diagrams for all 104 compartments, although these were developed during the two-week workshop.* Connectivity matrix While feedback dynamics diagrams are a convenient method for showing individual compartments in detail, they cannot be combined for a model of this scale without becoming unmanageable mazes. An alternative graphical representation that has desirable properties is the connectivity matrix (Figure 9). This matrix is a square binary matrix showing the presence or absence of flows from each compartment to each other one. The size of the matrix is determined by the number of compartments (104 in the case of the CITRE model). Each compartment, from X1 through X104, is listed both across the vertical axis and down the horizontal axis. The direction of flow is from compartments on the horizontal to those on the vertical axis. Flows are indicated by a dot " " located at the intersection square of two components of the matrix. Thus, in Figure 9 for example, the single dot on the bottom line (XORGC 104 ) indicates a flow from XORGC (104) to XDOC (62) but not the reverse. The connectivity matrix shows other information of interest. As its name implies, it illustrates the total "connectivity" or percentage of possible interaction. The CITRE matrix with 104 compartments has (104)2 or 10,816 potential interactions. In this coral reef ecosystem model there are about 2,000 interactions or 20% connectivity. Since connectivity is highly dependent on compartment definition and at this tevel of resolution concerns only material flows, it is not possible to distinguish at this time between properties resulting from the modeling approach, and those inherent in the ecosystem. Complex ecosystems may be characterized by a much higher percentage of information "flows" or non-material interactions. However, these interactions are even more dependent than material flows on the level of model resolution. * Copies of the working drafts may be obtained from A. L. Dahl at Department of Botany, Smithsonian Institution, Washington, D. C. 20560. Some working groups are continuing to develop their parts of the model. 12 There are two disadvantages to the connectivity matrix: it does not show external forcing functions (temperature, currents, etc.) or information processing (which are shown by dashed arrows on the diagrams), and it does not indicate the magnitude of the flows. A more complex form of matrix, a coefficient matrix, substitutes numerical values for the binary indicators of the coe Seema matrix. Turnover rates then appear in the principal diagonal (al], a 2° A33 seus) of the matrix, and flux or transfer rates appear in the off- RR esuE elements. The CITRE matrix of turnovers and transfers is incomplete and is therefore omitted here. While there is no immediate prospect of continuing to develop the CITRE model in its present form, it should now be possible to begin to piece together the major elements, and to quantify the essential relationships of the coral reef ecosystem. Only in this way will an overall picture of this most fascinating biological community ultimately be assembled. Literature Cited Forrester, J. W. 1961. Industrial dynamics. 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Figure 9. CITRE preliminary ecosystem matrix; reef-flat submodel. The dots indicate carbon or carbon-equivalent flows from compartments on the left to compartments along the top. Blanks indicate no connectivity, and the question marks indicate possible carbon flow between various Organic compounds and benthic plants. A COMPARATIVE SURVEY OF CORAL REEF RESEARCH SITES* Arthur L. Dahl, Ian G. Macintyre, and Arnfried Antonius The complexity of coral reef environments has long attracted scientific interest, but as yet few if any research programs have compared reefs from different areas or oceans with a view to understand- ing the overall functioning of a reef ecosystem. Plans for such programs recently were initiated at the Smithsonian Institution, and it was found that in the undertaking of an interdisciplinary investigation of reef ecosystems, the first critical step is to choose research sites that can meet stringent scientific criteria. The Smithsonian's comparative information on potential reef research sites in both the Caribbean and Pacific is being presented here in the nope that other investigators will find it useful in planning future reef studies. The data were originally compiled as a report on the research site selection process for the Smithsonian programs, and this has largely determined the present format. The observations are primarily qualitative in nature, presenting a broad view of comparative reef structure and composition. Their value lies in their common perspective, having been compiled by a closely integrated site selection team whose members, within a short time, visited many coral reef areas throughout the Caribbean and much of the Pacific, applying common criteria to achieve a common goal. The site search was conducted for two programs of the Smithsonian Institution. The first, Investigations of Marine Shallow Water Ecosystems (IMSWE) , was organized within the National Museum of Natural History with the support of the Smithsonian Environmental Sciences program to conduct an integrated biological and geological analysis of coral reef communities. The second was to be a much larger, multi-institutional program, the Comparative Investigations of Tropical Reef Ecosystems (CITRE), developed under the auspices of the Smithsonian Office of Environmental Sciences with a planning grant from the International Decade of Ocean Exploration Office at the National Science Foundation. The plans for the CITRE program included the development of a complete systems analysis model of the reef ecosystem based on energy and material flows through various reef components. For both programs, the scientific advantages and greater research potential of a scientifically "ideal" reef site were given first priority, with purely logistical considerations * Tnvestigations of Marine Shallow Water Ecosystems (IMSWE) contribution No. 3; supported in part by the Comparative Investigations of Tropica] Reef Ecosystems project under NSF Grant No. GX-28676 as part of the International Decade of Ocean Exploration program, 38 : : | considered secondarily, and thus a fresh look was taken at many potential reef research areas with the context of the programs in mind. An "ideal" reef was considered to be one that is subject to little external stress, and that is characterized by as many of the scientific criteria as possible. Since the criteria by which sites were judged have a crucial bearing on the conclusions of the site analysis, they are outlined below in some detail: 1. Ample development of all characteristic reef zones, from the "reef flat" to deep water communities in depths of 50 to 100m. & 2. A steep fore-slope, to facilitate field observations by telescoping the zonation. 3. Vigorous reef growth where interactions with the surrounding water mass can be measured and with a good geological record of past development. 4, Unidirectional current flow for periods long enough to permit cross- reef metabolic studies of the type successfully used at Eniwetok (Johannes et al. 1972), 5. No overriding unique characteristics with respect to the reefs in the same area. 6. Freedom from major terrestrial, human, or natural catastrophic influences in the present or recent past. Some practical criteria could not be ignored: 1. Sufficient accessibility to meet the needs of a large program. 2. A harbor and some accommodations and facilities, or the possibility of developing these at reasonable cost. 3. Availability of research vessel support. 4. Probability of a stable government with a favorable attitude towards the program. 5. Suitability for research needs such as multiple sampling, monitoring with shore-based instrumentation, drilling for geological samples, etc. Since the published literature has only scattered information on many of these features, a questionnaire (Fig. 1) was developed and sent to numerous scientists with field experience in the Caribbean and the tropical Pacific. The response was generally good, particularly for the Caribbean, and provided many suggestions for potential coral reef research sites. Following preliminary evaluation of the questionnaires, survey teams were sent to the most promising reef areas to investigate 39 their appropriateness in terms of the established criteria. The standardized reports of the survey teams, based primarily on their first hand observations, make up the body of the present paper. For the sake of brevity, no attempt has been made to cite the many earlier published reports on the areas described. Reef descriptions were compiled from field notes and photographs. At least one, and generally two or more of the authors participated on all survey teams in order to provide a basis for valid comparative judgements. In all instances the surveys focused on the set criteria although additional details included in the reports may reflect the special interests of individual team members rather than the unusual prominence of a particular group. The conclusions in terms of site preference, were conditioned by the criteria for the proposed ecosystem programs, and would differ if other factors were to be included. Charges for facilities or transportation, where given, are those at the time of the surveys (generally 1971) and are included only where such information is difficult to obtain to give a rough measure of practicality. For convenience and clarity, and Caribbean and Pacific sites are treated separately. The Pacific survey was inevitably less thorough, and many other areas warrant detailed examination. The Indian Ocean and more distant Pacific areas such as Australia were excluded as being impractical for a U.S.-based program. CARIBBEAN AREA Until recently, Caribbean coral reefs were generally considered inferior to their Indo-Pacific counterparts, mainly because previous studies concentrated on the southern Florida and northern Bahama area (e.g. A. G. Mayor; T. W. Vaughan), which is noted for its marginal coral-reef development. Recent investigations, however, indicate that this earlier assessment of Caribbean reefs is not altogether accurate. Although reef growth in the Caribbean is accomplished by considerably fewer coral species than in the Indo-Pacific, the reef construction processes are comparable. In other words, the same variety of reef types occur in both oceans, as well as an overall similarity in zonation with depth. Data on potential research sites were compiled for the Caribbean Sea and areas to the north. Regions of coral growth in the southwestern Atlantic, south of the barren area of heavy sedimentation off the Orinoco and Amazon rivers, were excluded from consideration owing to logistical problems and lack of scientific justification (see Laborel, 1967). Three sources of data were used: (a) the literature; (b) standard questionnaires sent to experienced workers in the Caribbean region; and (c) report from the site-survey teams. Information from the questionnaires is condensed in Table 1. 40 From accumulated information and the personal reports of program members, a number of sites were selected for detailed field examination, including Acklins Island (Bahamas), St. Croix (U.S. Virgin Islands), Discovery Bay (Jamaica), Glover's Reef (British Honduras) and the San Blas Islands (Panama). The data in Table 1 incorporate the conclusions of the survey team reports, which follow. Acklins Island, Bahamas* The reefs off the north and east coast of Acklins Island (22°30'N F 74-00'W) in the eastern Caribbean were surveyed during two visits; the first on Oct. 20-24, 1970 by a two-man team, and the second from April 13-23, 1971 by an eight-man team. (0) Acklins Island is part of the Crooked Island District of the Bahamas, but in fact is separated from Crooked Island by "The Going Through", a tidal channel (Fig. 2). The nearest populated center is Nassau, and from there access to Acklins is by small plane or boat. An old airport at Pinefield Point will soon be replaced by the new one under construc- tion at Pompey Bay. At present, charter airlines land on a road near this bay. Apart from this service, two weekly flights are available from Nassau to Crooked Island. The trip to Acklins is completed by taxi and ferry (small outboard). A 27 m vessel based in Chester on northern Acklins is used primarily for shipping and ferrying from Nassau to Chester, but would be available for charter at $100/day, and it could feasibly be considered for a floating hotel or ferry for reef diving. Small boats are readily available, but outboards are not. The population of Acklins is sparse and scattered along the western and northern coasts. Only a few settlements can be called communities. The cost of living is very high because nearly all commodities are imported. Local facilities for visitors are few and also expensive (e.g. a small cottage with barest essentials costs $11(US)/day/person, and a half-ton truck $24/day with driver). A research team would probably have to bring “bed and board" with it. The island is composed of Pleistocene calcarenite, apparently predominantly oolitic material. Its topography is relatively flat but a series of striking "fossil dune ridges" (up to 30 m relief) parallel the present coastline. The water table is close to the surface (2-3 m) so that fresh water is readily available. The soil is very thin except in some depressions. Most of the vegetation is a heavy growth of low brush, generally about six to ten feet high, but there are a few scattered trees (coconut, mahogany, casuarina and tamarind). Mangrove swamps are well developed on the northern end of the Bight of Acklins. *Field survey: October 20-24, 1970, by W. H. Adey, and I. G. Macintyre. Second field survey: April 13-23, 1971, by W. H. Adey, A. Antonius, A. L. Dahl, P. M. Kier, I. G. Macintyre, M. E. Rice, and T. R. Waller. at First Visit: October 20-24, 1970 The eastern and northern areas of Acklins were investigated as well as the central lagoon area (Bight of Acklins). Heavy swells created rough weather conditions and turbidity for two days, a condition common in the area from October to April. The following sites were examined: 1. West side of Atwood Harbour, north coast of Acklins Island. The patch reef here is no more than 15 m from shore, and it abounds with fish; a hawk's bill turtle also was observed in the area. Much of the reef surface is covered with crustose coralline algae, especially of the genera Porolithon and Neogoniolithon but large growths of Montastrea annularis, Acropora palmata, Millepora, Porites astreoides, and Porites porites were also present. The sandy bottom of the harbor, where examined, had well-developed Thalassia beds with large heads of Neogoniolithon strictum. In the southwest where the current was strong only a thin sediment cover was present over the rock ledge and smal] heads of Siderastrea radians and Favia fragum occurred. 2. Patch reef directly shoreward of bank barrier reef off east coast of Acklins Island, opposite Golden Grove. The reef here is about one and a half kilometers offshore. The abundant patch reefs in this area are surrounded by a sandy bottom and have about 6 m of relief. A very high percentage of the reef surface is covered by Porolithon, Neogoniolithon and other corallines. No large caverns were observed as described elsewhere on the Bahama Banks (Storr, 1964), but there were “overhangs” here and there. Large colonies of Acropora palmata and Millepora were observed along with abundant but small colonies of Agaricia agaricites. Additional common corals include three species of Diploria, Dichocoenia stokesii, Porites astreoides, and Montastrea annularis. A great abundance of alcyonarians was also noted. 3. Inner edge of bank barrier reef, northeast coast of Acklins Island off Pinefield Point. Going seaward from the patch reef there is a zone of abundant Montastrea annularis and Diploria coral heads leading to the turbulent inner edge of the reef flat, where there are large overturned growths of Acropora palmata (some were live growths) and large growths of Millepora. Rock surfaces and coral debris are heavily coated with coralline algae. Fish were abundant, including a number of different types of parrot fish, some quite large, which seem to inhabit an open network of smal] tunnels under the coral-rock floor. Be on Spring Point on the west coast of Acklins Island in Acklins ight. For a distance of about 150 m from shore a very thin layer of sediment covers the smooth rock pavement. Rich algal growths of Halimeda, Penicillus, Batophora and Acetabularia were observed, along with two species of corals: Siderastrea radians and Flavia fragum. A few colonies of Manicina areolata were also noted. 42 i=) t+etet +SN}C7S [POLL 10d i=) oO+ + SOldL [Loe 2907 -_ + - + 0 - + - - + + + 0 + = + + + + + + + 5 + + 0 + + + + 0 a 0 5 oP + - + - - + - + 0 + + 0 - 0 + = ILGLSsSID0V Ppequnys Lpuf) ¥S9ANZINATS F994 eau UvaqqlLue) - asuodsay euLeuUOLysant SPST -I 41avl soqo7] ap sels} OD] XH epnuwuag LULWLG pue|s] soupuy pue[sy] sul 4s seweueg uewhe) puedy joay uauueg yueg Oupadg *SAe) Oupad ‘ske) pue jaoy ualuueg Aeg AAVAO0IS LQ POL ewer eaue oObuLwog “45S 2 OLuedSLH XLOU9 *35S Spue|S] ULDALA pue[s] euoW OLY OJUaNd eo LULWOG Sd, [Ljuy uasse] Sa[Aeuj4 SO] Sep Luebury *9yo09 *enbeqng sanboy soq eB anzaua/j oroeun) eJUPW ePUeS OLUeSOY LAP SPLST eLquio| oo eSNG 43 + + + + 5nqe}S LPOL4L 10d "4a[Zjany *y pue “AqhLy “> “Cf SJJesd “Y SOUUNW “7 “P SUeWLLLEW “d *f SedAQULoRW “9 *T Sbue] °9 “Pp ‘puey “S$ *7 Suud[5 “mM “d ‘493S189 “fC “Lita “4 “uy fLued *7 “VW ‘deneg “M *y Ssnluqjuy ‘y ‘Aepy “HM :BULMO| LO} a43 Aq pal{ddns auam sazLs ueaqqlue) UO UOlZeUNOJUL ZUa|eALNba pue SatLeuUolysant Le21zoeUdWL SOltt [Loe [e007 wevboud papun} *s*p abue, e& spuemoz 9apnz14ze quawUseAOH peyedioique ayz AL |edLseq + JO alqeuorejun + + + + dLGLSSad0y a{qissod wo umMouxXUN = Q ++t+te+e+ + +e te tit paqung si pun +14 Oot 4 + 4+) fb ze Sadnzonu4s $994 *d fadoge eluaqLud Uol{.alasS a—as + + Kay a{qeLuea quasaud uO a[qeuorrs spue|s]T seg ues ‘sXke) sapue|oy eweurd pue|S~T uso) 2RaV9 skey anbuanbng|y ske) um0 UN) pue|syT sMaupuly “4S pue|S~T aduapLAOud Plo yueg wopeouoy yueg eueutas (enBevedin J}0) Spue| Sy ueLquio|o9 $994 S,49AO|Y jaay asnoyry61y pue|sy ajpsouaunl faay AdLUUeg SPUNPUOH YSlLzLug 391S 44 : | Visit: April 13-23. 1971 5. Atwood Harbor, North Coast of Acklins Island (see Fig. 3) Back Reef This area is characterized by small patch reefs scattered on a rock platform which generally has a thin sand cover. These protected patch reefs have a relatively dense coral growth consisting ee of Acropora palmata, Montastrea annularis, Porites astreoides, Agaricia agaricites, Dichocoenia stokesii, Dichocoenia stellaris and Porites porites. Reef Crest Coral composition on the lee side of the reef crest is almost identical to that of the patch reefs. However, the remainder of the reef crest is dominantly composed of Acropora palmata and Montastrea annularis which form an open framework marked by the common occurrence ; of dead overturned coral communities. oy Upper Fore-Reef Slope The dominant characteristic of this area of the reef is a gently dipping rock pavement. The two distinct zones observed are a) barren 7 zone 1.5-6 m and b) grooved zone 6-18 m. Barren Zone: Predominantly a knobby rock bottom with a thin cover- ing (2 cm) of benthic algae and fine to coarse carbonate sand. Crustose corallines generally are thriving under this algal-sediment cover. hh Smal] coral colonies scattered over the rock surface include Diploria Fi strigosa, Siderastrea siderea, Siderastrea. radians, Dichocoenia stokesii, Montastrea cavernosa, Montastrea annularis, Agaricta _agaricites » Porites astreoides, Porites porites. Colphyllia natans, Isophyllia sinuosa, __ Meandrina meandrites. Sargassum sp., gorgonians, and Millepora sp. increase noticeably in depths less than 4 m and large patches of boring sponges are common. Scattered shallow (1 m) depressions in the rock pavement have a well-rippled coarse sand or rubble bottom. A few Diadema sp. occur under ledges in the walls of these depressions. Grooved Zone: The most striking feature of this depth zone is the grooves which are clearly visible in aerial photographs. Off Atwood Harbour they are generally less than 30 m apart and are commonly about 60 m long. The gently dipping rock pavement transected by these grooves is encrusted with a variety of tropical-reef corals, but the colonies are generally small, well-spaced and therefore do not form an inter- locking framework. Dominant corals include the following species: Diploria strigosa, Porites astreoides, Siderastrea siderea, Dichocoenia stokesii, pechpcoenta stellaris, Montastrea cavernosa, Montastrea annularis, Favia fragum, Meandrina meandrites, Diploria ~Tabyrinthiformis, Colpophyllia natans, Stephanocoenia michelinii. Gorgonians and sponges are also abundant in this depth zone. 45 The groove system begins in a general depth range of 5 to 10 m as an indistinct labyrinth of channels between the scattered coral growth. In our limited observations we did not find grooves originating as "spoon-shaped incisions" or "erosional pits," as reported for similar features off Andros Island (Newell et al., 1951, p. 25) and off Saipan Island (Cloud, 1959, p. 406). These indistinct channels coalesce to form a well-defined u-shaped groove which is relatively straight and has a smooth sediment-free surface covered with a thin algal growth and corals here and there. These u-shaped grooves, which are about 10 to 15 m long, widen from 15 cm wide and 15 cm deep to 1 m wide and 1 m deep where they open out in- to a still deeper and wider sediment-filled groove. In some places, up to three separate u-shaped grooves were noted to be feeding into the same broad sediment-filled one. The sediment-filled sections are generally about 50 to 60 m long with a maximum width of about 10 m. The lower sediment-filled sections tend to taper into a series of lobes at the outer limit of the grooves. The thickness of accumulated sediment in the troughs was not establish- ed; however, at the head of the sediment-filled sections the walls were 2 to 2.5 m high and gradually diminished in relief to about 1 m at the terminus. It is interesting to note that the grooves investigated were completely enclosed with no ready outlet into deeper water for the sediments. At the head of the sediment-filled sections there is a band 5 m wide of coral rubble which grades down-slope into a well-rippled sand (wave length, 60 to 150 cm, amplitude, 12 to 15 cm). This is an ex- tremely coarse to medium-grained skeletal calcarenite with gravel size material collecting in the ripple troughs. Texture gradation of the sediments continues down the groove so that about half-way down, the gravel is absent and there is simply a poorly sorted, extremely coarse to medium-grained skeletal calcarenite, which in turn grades into a well-sorted medium to coarse-grained skeletal calcarenite at the terminus of the groove. Without subsurface data it is difficult to establish whether these linear features have been formed by erosion or differential accumula- tion of organic limestone. Newell et. al. (1951) found that grooves off Long Cay in the Bahamas cut into oolite bedrock, which clearly in- dicated an erosional origin. Lower Fore-Reef Slope This area starts at depths ranging from 12 to 18 m, where the slope of the sea floor increases markedly, and continues to the depth limits of our observations, approximately 40 m. Large areas of well-developed and diverse deep-water coral commu- nities are predominant in the upper section of this area. Sediment chutes that traverse the coral communities here and there coalesce at 46 depths of 20 to 25 m to form an extensive sand flat. Below this sand flat the corals do not show any appreciable change in species composition but tend to construct large coral mounds covering areas of up to 220 sq. meters with approximately 3 m of relief above the surrounding sand slope. The dominant corals of the lower fore-reef slope consist of the following species, in order of their abundance: Montastrea annularis, Agaricia agaricites, Montastrea cavernosa, Mussa angulosa, Diploria labyrinthiformis, Meandrina meandrites, Fusmilia fastigiata, Wen lamarckiana, Colpophyllia natans, Porites astreoides, Stephanocoenia michelinii, Dichocoenia stokesii, ~Dichocoenia stellaris, Porites porites, >» Zsophyllia sinuosa, and Diplor: Diploria strigosa. strigosa. East Coast of Acklins Island 6. Off Pinefield Point The area observed begins at a depth of 20 m where there is a distinct sand flat. Coral heads are sparse and widely scattered, but gorgonians and benthic algae are very common. Below a depth of 20 m the slope of the sea floor increases and coral growths tend to be restricted to linear ridges which are separated by sand and gravel-filled grooves generally 3 m wide and 1 m deep. The ; coral assemblage is similar to that of the deep-water coral communities i noted on the lower fore-reef slope off Atwood Harbour. Corals give way i to a sand and gravel slope at depths of 35 to 40 m. r 7. Misconception Rock Misconception Rock consists of a small (100 sq. m), relatively flat rock pavement (well-sorted fine to medium calcarenite) in the reef tract that is exposed at low tide. It is approximately 1 kilometer north of { Creek Point off the east coast of Acklins Island. “ The rock surface is heavily pitted and etched, the characteristic erosional form of limestones in the splash zone (Newe11 et. al., 195i and the outer edges are covered with a heavy algal growth. Many large gastropods (Livona pica), Diadema sp., and chitons were observed in numerous shallow depressions. See ee ape As found in other areas of the Bahamas (Newell et. al., 1951) the bank barrier reefs off eastern Acklins Island appear to be a Holocene X reef veneer over pre-existing sand ridges that are exposed above sea level in places. 8. South of Golden Grove The broad (12 m) platform which is a characteristic feature of the insular shelf off the east coast of Acklins Island separates the reef crest and fore-slope communities by distances of over 1.5 km. 47 The bottom is a smooth rock pavement which has a thin (1 cm) sedi- ment-algal cover, consisting of a dense benthic algal layer with trapped sand- to silt-sized sediment. Small coral mounds, having relief to less than 1 m are widely spaced on the rock pavement. Coral heads, for the most part dead, are scattered over this rock surface. Dominant live corals are Diploria strigosa, Eusmilia fastigiata, Agaricia agaricites and Dichocoenia stokesii. Rich epifaunal and infaunal assemblages occur under the live and dead coral material, including the bivalve Barbatia domingensis, y the gastropod, Tegula fasciata, and the echinoids, Eucidaris tribuloides, Arbacia punctulata and Echinoneus cyclostomus. Gorgonians are common and some are encrusted by Millepora sp. Summary Atwood Harbour would be an excellent location for a base station as it is a good harbour for boats, and the Bahamian government is willing to cooperate with scientists working in their territory. The northeast corner of Acklins Island, east of Atwood Harbour and North of Pinefield Point, is Crown Land and arrangements could probably be made to set this area aside as a research site. The reefs have been rela- tively unaffected by human activity. Fishing appears to be a minor occupation carried out in a primitive fashion on an occasional basis. When the new airstrip is completed, accessibility to Acklins Island will be improved with regular weekly flights scheduled for the near future. Bi-weekly flights to Crooked Island are currently available from Nassau. A number of vacated small houses on the island might be used as accommodation facilities. They might be rented at reasonable rates, but it should be recognized that amenities are lacking. Apart from some of the small patch reefs, the reefs on the insular shelf are not well developed. The exposure of calcarenite in the reef tract off the east coast (Misconception Rock) indicates that many of the reefs may be veneers over Pleistocene sediment ridges. The shallow reef crests are commonly separated from the fore-reef slope coral communities by a very broad rock platform. Rough seas common during the months of October through April would prevent work on the reef crest and fore-reef slope during these months. St. Croix, U.S. Virgin Islands* St. Croix (17°45'N, 64°40'W) is the largest of the U.S. Virgin Islands, having a surface area of 22 sq. km (45 km long and up to 117 km wide; see Fig. 4). Its population is about 35,000, and apart from “field survey: January 26-28, 1971, by I. G. Macintyre, assisted by H. G. Multer. I ee er 48 tourism, the island supports a few other industries such as bauxite refining, oil refining, and the assembly of wrist-watches. This island is readily accessible as it is served by several major airlines. Transportation on the island is available through car rental at about $12/day or scooters at $7/day. Boats may be chartered, or rented less expensively through the Fairleigh Dickinson Laboratory. Hotel accommodations are costly, particularly during the winter season, but the marine laboratory has some rooms at reasonable rates for visit- ing scientists. Terrestrial Conditions The topography of St. Croix is "hilly" (maximum relief 355 m, Mt. Eagle). The island is composed of rocks having diverse origins and compositions, primarily Upper Cretaceous volcanic sandstones and mud- stones, tuffaceous sandstones, Upper Oligocene montmorillonitic mud- stones, and Lower Miocene argillaceous and sandy coral limestones. Despite the abundance of volcanic debris, no volcanoes are present on St. Croix; however, two small igneous bodies (a diorite and a gabbro) intrude the Cretaceous sedimentary rocks. Numerous small dikes are also present in the hilly ranges occurring on the eastern and northwestern sections of the island (Whetten, 1966). St. Croix Reefs Although St. Croix has a variety of reef types that would be excellent for scientific study, there appears to be no evidence of a reef marginal to deep oceanic waters, a serious drawback to the select- ion of St. Croix as a primary site. 1. Bank Barrier Reefs. As expected, the best developed reefs occur off the eastern or windward coast, and the most striking developments are the bank barrier reefs on the landward.edge of the insular shelf; these reefs parallel the northeastern and! southeastern coasts for distances of over 5 km. The bank barrier reef off Tague Bay (Fig. 2, Site 1) has about 5 m relief off the shoreward edge and is approximately 90 m wide. This reef is dominated by Acropora palmata, but an indistinct zonation was noted around the inner edge of the reef, where the base is characterized mainly by Montastrea annularis; the edge by Acropora palmata and large mounds of Porites porites (up to 2.5 m high and 4.5 m wide); and the top by Acropora palmata, Diploria sp., and Millepora sp. The seaward edge of this reef was not investigated during this brief visit; however the transect (Fig. 6) based on information received from Dr. Multer indicates that this seaward slope drops off to a depth of 14 m and gives a graphic representation of the dominant coral distribution across the reef. 49 2. Algal Cup Reefs There was not enough time to investigate the fringe reefs common around most of the promontories along the northeastern coast of St. Croix. However, the nearshore "algal cup reefs" occurring around Cottongarden Point were investigated (Fig. 5, Site 2). These reefs, which are characterized by pronounced rims and over hangs, have a relief of about 2 m. Their upper surfaces are covered by fleshy algae including Sargassum sp. and a few flat colonies of Porites astreoides. Chips from these algal reefs, broken off with great difficulty, revealed a dense agglomeration consisting mainly of crustose coralline algae and the distinctive red encrusting Fore Ye Homotrema sp. An unusual occurrence of Acropora prolifera in an area of high agitation was noted next to these algal reefs. Abundant algae are present on the sea floor surrounding the cup reefs, notably Penicillus sp. and Halimeda sp. (sediments are particularly rich in Halimeda- derived debris). 3. Buck Island Trail A rapid survey of the Buck Island National Trail (Fig. 5, Site 3) indicated that this is a well-developed "palmata reef." Although separated from shore by a narrow sandy belt, it can be considered a form of fringe reef. 4. Deep Patch Reef A deep patch reef directly east of Buck Island (Fig. 5, Site 4) consists of an area of large open frameworks, primarily Acropora palmata This patch reef has a relief of about 8-9 m above its base, which is covered with large heads of Montastrea annularis, Diploria sp., Montastrea cavernosa, and surrounded by a great abundance of alcyonariars. Millepora was found coating some of the dead alcyonarian branches. 5. Shelf Edge North of Buck Island This area was investigated in order to determine whether a shelf- edge reef exists at this location (Fig. 5, Site 5). The reconnaissance was carried out by a towed swimmer, and all observations were made from the surface of the water. Although no evidence of a shelf-edge reef was found, an interesting transition in bottom characteristics was noted: at depths of about 30 m a series of sediment chutes traverse a relatively steeply dipping (about 10-20°) rock surface that is covered by alcyonarians, sponges, algae and scattered coral heads. From the surface, there appeared to be little relief between the sediment chutes and intervening rock surfaces. The sparsity of corals suggests that reef framework construction was not occurring at this depth. Although a spur and groove system appeared 50 to be present at this location from the air survey, firsthand obser- vations showed that the "buttress systems” described off Jamaica by Goreau (1959) are not present in this area off St. Croix. Shoreward, the sea floor gradually grades into a rocky bottom having sand patches and scattered alcyonarians, sponges, algae and coral heads. Around depths of 15 m, large scattered colonies of Acropora palmata appear, and directly shoreward there is a zone of Millepora. Abundant colonies of Acropora palmata are present in depths of 3-6 m along with other corals such as Montastrea annularis and Diploria sp. 6. Other Reefs off St. Croix During the R/V Eastward cruise, two other reef types were investi- gated in this area: inactive and submerged reefs. The most prominent reef type off St. Croix is the inactive reef, of which Lang Bank is the best example. Inactive reefs are reefs that occur within depths of potential vigorous coral growth, but which are characterized by an absence of reef framework construction. Lang Bank, situated at the shelf edge east (windward) of St. Croix (Fig. 5) at depths of around 9-12 m, is ideally located for active barrier reef development. From limited bottom photographs, however, it is evident that an interlocking reef framework does not occur on this ridge which is covered with abundant alcyonarians and sponges and large sand patches, but only scattered coral heads. Dredge hauls from this area contained abundant alcyonarians, sponges, and small colonies of Porites porites, Porites astreoides, Diploria SP. Agaricia agaricites, Stephanocoenia michelinii, Acropora cervicornis, and Millepora sp. Coral debris heavily encrusted by coralline algae was also abundant. Meyerhoff (1926) suggested that the topography of the eastern St. Croix insular shelf is related to subaerial erosion of a Tertiary reef complex. Submerged ridges were noted on bathymetric profiles off the southern coast of St. Croix, established at depths of about 40 m. These ridges are interpreted to be submerged reefs that were establish- ed in relation to a pre-existing lower sea level, and that occur in depths greater than are commonly associated with vigorous growth of tropical reef corals. They appear to be common features off most eastern Caribbean islands (Macintyre, 1972). Pollution Extensive pollution of the marine environment is occurring off the southwestern coast of St. Croix, where a garbage dump, bauxite plant and oil refinery are located (see Fig. 4). It is expected that pollution levels will rise in the near future because sewage will be piped from Christiansted (the island capital) over to this side of the island. Despite the unfortunate aspects of this circumstance,: it would offer an opportunity for detailed comparison between polluted reefs off this coast and the non-polluted reefs to the east. Such studies might yield pele inate 5] valuable information on the degrees of stress imposed on reefs by pollution, and how well they survive it. West Indies Laboratory of Fairleigh Dickinson University The West Indies Laboratory is located on Tague Bay, on the north- eastern coast of St. Croix, and it overlooks Buck Island National Park (Fig. 5). This laboratory offers scientific facilities for researchers as well aS some accommodations, when available. There are 4 small laboratories equipped with aquaria, and an outdoor aquarium area Supplied by a dual saltwater system. The Laboratory also owns two 4.9 m (16') Boston whalers and three 5.8 m (19') Rabollo boats which may be rented by visiting scientists at a minimal cost. Owing to an extensive summer program, visiting scientists are advised to plan visits for less crowded periods of the year. An affiliate program is gradually being developed which will permit outside universities to have faculty and research space as well as teaching space privileges on the basis of a yearly charge. Summary Apart from the polluted reefs on the south coast of St. Croix, the coral reefs around this island have been relatively untouched by man because there is little need to exploit them for food. However, the absence of a reef marginal to deep water in the region is a drawback, as is the general lack of well-developed reef-flat zones on the shelf reefs (a distinct disadvantage in drilling operations). On the other hand, the numerous advantages of this location make St. Croix worthy of consideration as a research site, particularly for a smaller scale program. The cooperation and available facilities of W.I.L. would make operational costs on St. Croix considerably lower than elsewhere in the Caribbean; at the same time, W.I.L. would probably facilitate detailed and sophisticated experimentation which might be difficult to undertake and complete at less well-equipped sites. Following is a brief outline of the merits of St. Croix as a study site: 1. Facilities of the West Indies Laboratory, notably; a. inexpensive accommodations; b. fish pens; c. outdoor and indoor aquaria supplied by all P.V.C. non- - toxic dual saltwater systems and backed up by SUSE EEE emergency power unit; laboratory and office space; motor boats; e. reference library with microfilmed periodicals and print- out service; radiocommunication with field units; monitoring service which W.I.L. intends to establish off northwestern coast. Q. WO bh 52 2. The laboratory staff would be able to assist in servicing field equipment. Technicians working on Navy projects on western St. Croix might also be enlisted to aid with electronics problems. 3. Jet airport with daily flights to Washington, Miami, and New York costing less than $200 return. San Juan, P. R., is 45 minutes away by hourly scheduled flights. 4. A variety of reef types are present, including fringing, bank barrier, inactive, submerged, algal cup, polluted and non-polluted reefs. 5. Because St. Croix is an American territory there should be a minimum of red tape involved in having a section of reef tract set aside for scientific investigation; ground has already been broken with the Buck Island National Underwater Park. 6. There would be less difficulty in shipping equipment to a U.S. island, and financial advantages of a duty-free island in the purchase of equipment. Discovery Bay, North Coast of Jamaica* Jamaica, an independent member nation of the British Commonwealth, is a large tropical island located in the northwestern Caribbean Sea at the extreme northwest tip of the Nicaraguan Rise. It is 140 miles long and 60 miles wide and has mountain peaks in excess of 7,000 feet above sea level. Fringe reefs extend discontinuously along the north coast and are noted for their coral species diversity (Goreau and Wells, 1967). Discovery Bay (18°30'N, 77°25'W), the site of the Discovery Bay Marine Laboratory, is located about midway along the northern Jamaica coast, and it lies adjacent to a well developed section of the fringe reef (Fig. 7). The reefs in Discovery Bay (Fig. 8) generally may be divided into the following major zones (after Kinsey, 1970; J. Lang, personal communication, 1971). Lagoon Zone: This zone is about 400 to 500 m wide, and about | to 3 m deep. It has a submerged sinkhole 25 m deep, but the walls and bottom are mud covered and there are no rock exposures. Loose sand is the main constituent of the floor in the lagoon zone, and this precludes development of sessile benthic epifauna. Here and there, bare rock patches and coral heads occur with scattered gorgonians. Corals and rocks become more abundant seaweed, toward the rear zone. Rear Zone: The inshore face of the reef crest makes up this zone, which is about 0.3 to 2 m deep. Apart from the reef flat, this is the only reef area strongly affected by tidal fluctuations and characterized by strong water movement. Bottom types range from calcareous sand, and sand and coral fragments, to bare coral-rock exposures. Living coral is * Field survey 21 to 28 July 197] by I. G. Macintyre, assisted by P. Dustran, E. A. Graham, L. S. Land, and J. C, Lang. =~ a aa —e Ses ee a ieetieel Alto metederent a. 2 tie ons eae ig eters tai aoe 53 common but never covers the bottom entirely. Richest coral growth occurs in the area abutting the shoreward face of the reef flat. Reef Flat: The reef flat comprises the area just submerged at low tide. It is subjected to strong breaking wave action as well as occasional exposure to air and rainwater. The main component is a reticulate ridge-like structure 5 to 20 m wide, composed of dead colonies of Acropora palmata, which separates the lagoon from the sea. The unconsolidated skeletons of A. palmata form a lattice-like structure. Protected areas within the reef are locations for a restricted gorgonian fauna. Channels and pools in the reef flat may reach a depth of 2 m. Palmata Zone: This narrow zone, 5 to 10 m wide, abuts the seaward face of the reef flat and extends seaward to a depth of 4 to 6m. The full force of wave action occurs in this zone where Acropora palmata is predominant. The bottom consists of coarse sand and sand-scoured coral rock littered with dead and fallen A. palmata colonies. ~ Barren Zone: A band of reduced coral diversity occurs seaward, and it can be up to 20 m wide, ranging from 5 to 8m in depth. The most common organism, Diadema antillarum, which grazes on the bottom, may be responsible for the scarcity of sessile organisms. Isolated colonies of A. palmata occur along with small heads of Diploria strigosa, Montastrea annularis, and Millepora complanata. Buttress/Mixed Zone: In the Ocho Rios area studied by Goreau (1959) as well as other areas along the northern coast, the buttress zone merges abruptly with the barren zone. Although the component species remain the same, Discovery Bay differs slightly in architecture from the other areas in that the normal buttress zone becomes flatter, with a gradual increase in coral species and coral size. This "mixed" zone is 15 to 40 m wide, and slopes from 6 to 15 m in depth. Montastrea annularis, one of the dominant corals in the buttress formations, occurs in Discovery Bay in the form of huge rounded masses scattered in fields of A. cervicornis. Diploria spp. is also common, along with Dendrogyra cylindrus, Agaricia agaricites, Porites spp., and Colpophyllia natans. This is an area of rich gorgonian diversity. Cervicornis Zone: A. cervicornis becomes dominant seaward, and M. annularis reduced to small patches. This zone, which is 30 to 100 m wide and 20 to 25 m deep, is characterized by huge mounds of A. cervicornis 15 to 40 m wide with their long axis normal to the axis of the reef crest. These reefs (1 to 5 m relief) are separated by sand channels 3 to 50 m wide. Upper surfaces of these reefs dre generally level, and they gradually slope to about 10 m in depth at their seaward extremity, where they drop abruptly to sand level at about 20-25 m on their seaward face. Consolidated coral rubble covered with living A. cervicornis is typical (Land and Goreau, 1970). Other coral species appear on the steep seaward faces and lateral reef flanks adjacent to the sand areas, including Porites astreoides, Mycetophyllia sp., and Montastrea cavernosa as well as M. annularis. In Discovery Bay the A. cervicornis reefs drop steeply at the seaward edge to a zone of sand about 40 to 60 m wide. 54 Upper Sill Reefs: A series of elongate but discontinuous reefs are established at 40 to 50 m depths, and rise to about 25 m depths. Platy colonies of Montastrea annularis are dominant along with a mixed coral community composed primarily of Agaricia lamarcki, Mycetophyllia sp., other species of Agaricia, Scolymia sp., fleshy algae, sponges and antipatharians. Fore Reef Slope: The platy growths of Montastrea annularis along with Agaricia sp. and abundant sponges, gorgonians and antipatharians continue down the reef fore slope to a depth of 50 m, below which either Agaricia sp. dominate, or mixed coral growths occur consisting of Agaricia sp., Madracis sp., Montastrea cavernosa, and Mycetophyllia sp. Lower Sill Reefs: This zone, at depths of 60 to 70 m, is charact- erized by either elongate ridges having less than 10 m relief or isolated rocky outcrops. Corals occurring on these features include Agaricia sp., Montastrea cavernosa, Madracis sp., and Mycetophyllia sp. Sponges and antipatharians are also abundant. Deep Fore Reef Slope: Scleractinian corals become increasingly Sparse with depth, and the dominant benthic organisms are sclero- sponges and demosponges. The Discovery Bay Laboratory The Discovery Bay Laboratory is jointly operated by the Marine Sciences Research Center of the State University of New York, and the University of the West Indies. The Laboratory site consists of .06 sq. kilometers (15 acres) of coastal property. Facilities have been designed primarily for coral-reef research with emphasis on on-site investigations (SCUBA facilities) rather than for training programs. Therefore, only limited space is available to visiting scientists and there are no dormitory accommodations. The new central building consists of an air-conditioned research unit housing four small laboratories, a large "wet" laboratory, a dark room, instrument store, museum and two offices. The "wet" lab is sub- divided into six semi-enclosed research bays provided with all services including seawater and central tables for aquaria and sorting and hold- ing tanks. A separate reading and conference room is nearby. Support facilities include a machine shop, a boat and wood workshop, and a diving locker. Three boats are available a 6.7 meter (22-foot) twin outboard-motor vessel, a 4.6 meter (15-foot) work boat, and a 3.7 meter (12-foot) inflatable boat that can be carried by car to remote sites. A landrover and Volkswagen bus serve to transport equipment as well as personnel. The laboratory is equipped with a glass-distilled water supply, dissecting and compound microscopes, histological apparatus, drying oven, pH meter, top loading and analytical balances, photographic equip- ment and darkroom facilities, refrigerator, freezer, céntrifuge, by : 55 collecting gear, etc. The diving facility contains two high-capacity air compressors, a recompression chamber with air bank, SCUBA tanks, regulators and auxiliary diving equipment. Summary The reefs off Discovery Bay Laboratory probably are some of the most extensively studied reefs in the world, and the data already available would be a distinct asset to a research program. The Discovery Bay reefs are rich and well developed both in shallow and deep water. Since they occur very close to the coast, their location would facilitate close monitoring of various parameters. More- over, the relatively narrow horizontal extent of the reef biotope would be advantageous in instrument monitoring, as well as in on-site investigations. This site is readily accessible from the U.S. mainland, and accommodations are available locally at inexpensively priced guest houses (ca. $7/day single) for as many as 40 scientists. On the other hand, there is a significant but as yet undetermined terrestrial influence on the reef ecosystem in this area. The internal sediment found in association with the present reefs is characteristic- ally brown owing to the presence of bauxite or iron oxide that has been incorporated from terrestrial sources. From 1964 to 1967, a ship channel was dredged and blasted into Discovery Bay to permit docking of large ocean-going vessels in this bay. The channel is about 300 to 400 m from some of the key areas of potential research. The construction of this channel as well as frequent visits of large ships into the bay provide negative influences on the reef ecosystem in this area. The paucity of larger reef fish is notable in Discovery Bay. Although fish populations are sparse, fish traps are still used exten- sively, and therefore this location has limited potential for any proposed fish studies in association with the reef ecosystem. The poor development of a buttress zone in the area is an additional drawback. The Discovery Bay Laboratory is not eqippped to handle large coral- reef study programs because of its relatively small size. Research projects might also be hampered by customs delays and the need for import licenses owing to the problem of bringing onto the island scientific equipment and material from another. country. Glover's Reef, British Honduras* A first reconnaissance visit to British Honduras, 17-22 January 1971, included several diving stations along the barrier reef and *Preliminary survey: January 17-22, 1971 by A. Antonius and J. N. Weber. Field survey: June 20-27, 1971 by A. Antonius, A. L. Dahl, and K. Ruetzler. 56 Turneffe Island, and an air survey of the barrier reef, Turneffe Island, Lighthouse Reef and Glover's Reef. The information obtained strongly suggested that an extensive diving survey be focused on Glover's Reef, and this was conducted 20-27 June. Description of Reefs Glover's Reef (16°50'N, 87°50'W) is the southernmost of the three British Honduras atolls, about 75 km SE of Belize City, 45 km E of the mainland and 25 km E of the barrier reef. It is elongated in NNE-SSW direction, about 28 km long and 10 km wide. The atoll (Fig. 9) is surrounded by deep water (350-1000 m) within 1-2 km from the intertidal peripheral reef. The well developed surface-breaking atoll reef flat (mainly coral and coralline algae) is interrupted by two major openings: NE-entrance and SW-entrance (12 m deep). On its SE part, it supports a chain of six cays which are distributed over a distance of 10 km. The cays vary between 150 m and over 1 km in length and are more or less covered by coconut palms. The lagoon. in contrast to the other two British Honduras atolls, is rather deep, 8-15 m in most parts, with hundreds of patch reefs rising to the surface. Of the six cays on Glover's Reef, only Long Cay and adjacent Little Cay are permanently inhabited. Both islands are used as a resort for diving-oriented tourists. From the seaward margin of the peripheral reef a gradually sloping reef-front extends to the drop-off. The reef front varies in width from about 400 m (SW cays) to 1.5 km (Long Cay) and is about 500 m wide on the leeward side. The edge of the drop-off occurs at a depth of 15-25 m. A system of deep parallel grooves runs perpendicular to it. These grooves are particularly well-developed on the windward (E) side of the atoll. Sediments produced in the breaker zone are transported down the grooves permitting undisturbed coral growth in between. The leeward reefs are considerably more influenced by sediments which are driven out of the lagoon by wind generated currents. The windward side shows a dominating, lush, growing reef-coral community including sponges and gorgonians (Fig. 10). No attempt was made to gain a complete list of species during the survey, but the general appearance of the reef and the unparalleled size of most of the coral and sponge specimens observed strongly suggest that all of the known Caribbean species can be found there. On the sketches of the reef transects only dominant species are listed. Compared with the windward reef, the leeward side shows a Slight decrease in number of scleractinian species and is slightly less populated, probably due to sedimentation (Fig. 11). The coral cover ends around 45 m and gives way to a sand slope. The fore-reef slope is very steep at NE Cay, almost vertical at Long Cay and vertical to overhanging at SW Cay. It was explored at five different locations to a depth of 60 m. There, and as far down as 57 one could see in the clear water (horizontal visibility at least 50 m), lush coral, sponge and algal growth continues. The patch reefs rise to the surface from the lagoon floor, which is generally covered with sand and Thalassia. These reefs are mainly of Acropora palmata and A. cervicornis structure on top and Montastrea annularis below, with coral growth occurring to a depth of 3-4 m and occasionally 10 m. Every patch reef is a small individual reef-entity and ideal for a variety of experiments. The winds in the vicinity of the British Honduras atolls blow steadily from the East for most of the year, being replaced between November and February by occasional Northern winds, which occur only a few times every winter and last four or five days at a time. Thus, the wind-generated western currents in the atoll area are by far predominant over the main Caribbean current, which impinges on the British Honduras atolls in a northerly or southerly direction, and regularly changes its direction several times a year. Water temperatures at the surface vary between 25°C in winter and 31°C in the summer (open water); no significant temperature decrease was noticed during the dives (for detailed information on climatic, biologic and geologic features of Glover's Reef, see Stoddart, 1962). Summar. British Honduras is a politically stable country, and its official language is English. Relations between the government and resident North Americans or Europeans are cordial. The British Honduras govern- ment seems to look favorably upon educational and research projects, and duty exemption for all scientific and educational equipment and materials has already been granted. The remoteness of Glover's Reef makes it likely that it will remain undisturbed; still, there is good accessibility to Belize by air, and the atoll can be reached in any weather by boat because two-thirds of the trip takes place in a sheltered lagoon between the mainland and the barrier reef. There are no research facilities on the atoll at present, and it is not always possible to make satisfactory arrangements for an extended scientific program using tourist-oriented facilities. Glover's Reef is well suited for direct comparison with Indo- Pacific atolls as there are no land influences, no pollution, or other human disturbances; yet the large barrier reef, and two comparable but physiographically different atolls are readily accessible. While Glover's Reef is the only one of the three atolls which was not hit by the devastating hurricane Hattie in 1961, there is a hurricane potential; a small one passed over the atoll in November 1971 without causing major damage. In contrast to the other reefs surveyed in the Caribbean area, Glover's Reef atoll appears to offer the greatest variety of reef types Rey! ult) net i ee 58 and the optimum reef development in terms of population density and species diversity of reef corals and associated organisms. San Blas Islands, Panama* The San Blas Islands (9°39'N, 78°45'W) lie off the Caribbean coast of Panama between San Blas Point and Cape Tiburon (Fig. 12). During the Spanish conquest of Panama, the Cuna Indians were driven from the mainland to find refuge on these islands. Today, they have title to over 365 islands and a strip of land along the adjacent coast. They have jurisdiction over agreements to buy, settle or establish any forms of business on their islands. Standard access to the San Blas Islands is by means of light plane or small boat. The islands are about 110 km east of Colon, and it takes about 8 hours from there by boat. There are scheduled flights to several San Blas localities from Panama City, and a float plane service from Colon. One serious problem to be considered concerning travel by sea is that rough waters are common during the months of December to April, so the islands generally are not accessible by boat. Airstrips such as at Carti on the mainland (see Fig. 12) are accessible year round to light aircraft that can transport passengers to and from Panama City at a cost of $9.25 per one-way trip. Although this provides a rapid and in- expensive form of transportation to the area, and could be used by scientists when the islands are inaccessible by water, this air carrier cannot always operate on a specific schedule owing to the frequency of poor weather conditions. In addition, the short and uneven dirt air- strips preclude the use of larger aircraft so that only four or five passengers and light baggage loads can be accommodated on any one flight. The only housing facilities presently available in the area are located on Picofeo Island, which is the only privately owned San Blas island. These facilities consist of several sheds and some machinery (including a 6-volt generator) that are part of a disused copra- processing plant. One building serves as the Picofeo Hotel which is operated by Jose Garcia, one of two Panamanian brothers who own the island and its buildings. The hotel, which is an old wooden structure, offers minimal accommodations to visitors, who are expected to bring their own food supplies at $2.50/night. Extended stays can be arranged at cheaper rates. Outsiders are generally discouraged from setting up reisdence in the other islands. However, a few tourist resorts are now being developed in the San Blas Islands, which should provide for improved access and accommodations. A short initial visit confirmed the potential of the area, and *Preliminary survey: May 1-2, 1971 by A. L. Dahl and.A. Childs. Field survey: August 3-7, 1971 by A. Antonius, J. C. Land, and I. G. Macintyre (assisted by C. Birkeland, D. Meyer, M. McCosker, W. K. Sacco). 59 located some sites for further study. Ten locations were then visited during the four-day survey. These areas are marked on maps redrawn from sections of HO chart No. 2771, San Blas Point to Concepcion Bay (Figs. 13, 17, and 22). Following are brief descriptions of the areas in the order that they were visited: 1. Off Sail Rock, Profile A-A' Sail Rock is an exposed rock knob marking an isolated reef not far from Porvenir Island, where there is a very short airstrip (Fig. 13). The water was very turbid during the survey of the reefs in this area (less than 2 m visibility at a depth of 20 m) and there was a marked thermocline at a depth of about 3 m. The zonation across the profile is given in Fig. 14. Points worth noting in this area are the dominance of Agaricia sp. corals at the edge of the shallow platform and the lack of coral growth below about 20 m where the accumulation of fine sediments (rich in organic material) apparently prevents corals--other than the bladed Agaricia tenuifolia--from growing. 2. Picofeo Island The Picofeo Hotel is set on piles over the water at the north end of Picofeo Island (Fig. 13). This area is characterized by typical back-reef coral communities (Fig. 15) dominated by Porites porites. As was found at Sail Rock, the outer edge of the platform is marked by a rich Agaricia sp. coral community. The reef terminates in a shallow sand flat at a depth of about 6 m. 3. Off Sardingan Point, Profile B-B' Sardingan Point is on the mainland near San Blas Point and west of Picofeo Island (Fig. 13). The reef in this area drops off steeply to a depth of about 30 m (Fig. 16). An interesting aspect of this reef is that Agaricia sp. corals tend to dominate throughout the depth zones except in very shallow water, where Porites porites is the most abundant coral. 4. Off Salar, Profiles C-C', D-D', and E-E' The group of islands at Salar are among the inner San Blas Islands north of Macolla Point (Fig. 12). As names for the individual islands were not available, they have been numbered from southwest to northeast (Fig. 17). The survey concentrated primarily on the area between islands 2 and 3. Several dives were completed in this area of vigorous coral growth, spectacular drop-offs, and overhangs which are described in profiles C-C' (Fig. 18), D-D' (Fig. 19), and E-E' (Fig. 20). The general zonation of the reef flat of island no. 3 is also shown in a diagramatic sketch (Fig. 21). 5. Off Ogopuquip, Profile F-F' Three different sites were visited in the Holandes Cays (Fig. 22). 60 The first was at Ogopuquip on the southern side of the keys, where a profile (F-F') was prepared off the southwest point of Ogopuquip Island (Fig. 23). This area had the greatest diversity of coral species of any site visited in the San Blas Islands. 6. Holandes Cays Algal Ridge, Profile G-G' 4 $ The distinctive structure of the northern reef around the Holandes i} Cays is illustrated by profile G-G' (Fig. 24). This reef has a } prominent algal ridge that has been described in more detail elsewhere (Glynn, in press). : ; 7. Off Holandes Cays, Profile H-H' Beyond the algal ridge off the east end of Holandes Cays, there is ‘ a broad rock platform similar to that observed off many of the Bahamian Islands (profile H-H', Fig. 25). The survey did not extend to the drop- off which, according to the chart, probably occurs 200 m beyond the outer end of the profile. Summary Some reef areas, in particular the reefs off Salar Island, meet the scientific criteria established above for a research site including: well-developed reefs, steep fore-reef slope, lack of disturbance by human activity, and a close proximity to shore. The well-developed algal ridge off the Holandes Cays offers an opportunity to make compar- ative studies with similar reef structures in the Pacific. Buildings are available for conversion into research facilities. However, they are not located adjacent to well-developed reefs. Smithsonian Tropical Research Institute maintains marine laboratories and a research vessel elsewhere in Panama, and conducts research in the San Blas Island area. Two atypical characteristics of San Blas reefs, in comparison with | other Caribbean reefs, are the predominance of Agaricia sp. corals in all depth zones, and the presence of a well-developed algal ridge off Holandes Cays. The inaccessibility of this area by boat for five months of the year and the limited air transport service offered present a serious logistical problem in transporting personnel and material in and out of the San Blas area. As facilities on Picofeo Island are too distant from good reef development to be useful, it would become necessary to negotiate lease arrangements with the Cuna Indians for one of the un- inhabited islands. 61 PACIFIC AREA In the Pacific Ocean, coral reefs are scattered over a vast geo- graphic area. The only safe generalization about Pacific reefs is that no two reefs are identical. However, in spite of the great diversity in form and composition that characterizes these communities , a number of features demonstrate the underlying relationships within this assemblage. The coral fauna is remarkably uniform throughout the Indo- Pacific (Wells, 1969), with the highest diversity and the center of evolutionary radiation in the western tropical Pacific (Stehli and Wells, 1971). This fact frequently leads to the statement that reefs in the western Pacific are "rich" while those to the east are "impoverished. Whether this applies to all elements of the flora and fauna, however has yet to be demonstrated. The atoll reef form is another common Pacific feature, although involving many structural variations on the basic theme. The presence of a Porolithon algal ridge is often believed to be characteristically Pacific, and is certainly common there, while it rarely occurs on Atlantic reefs. The great number and diversity of tropical Pacific reefs has made the search for data on research sites very difficult. Information was compiled from many sources on reef structure and composition, and on logistic and practical arrangements. Most published descriptions of Pacific Islands include only terrestrial geography; specific local descriptions are rare, and only the shallow reef is mentioned. Charts from the U. S. Army Map Service and the Navy Hydrographic Office in- dicate with fair accuracy the presence or absence of shallow reef structure, and sometimes the nature of the offshore slope and the depth to which coral development might be expected. A few scientific papers include reef descriptions of some sort, but not for the deeper areas important to a complete reef program,and generally in localities that are otherwise unsuitable. Most of the past research on Pacific reefs. was conducted either by ship-borne expeditions to otherwise inaccessible islands, or in areas that have since been disturbed by development. Also, published information tends to be too dated to be reliable for current reef conditions; much of it predates World War II. While many questionnaires were returned by Pacific reef special- ists, their information did not always meet program requirements, perhaps because of the great size and complexity of the area. Personal experience tended to be limited to a few sites and to what could be seen from above the water. Areas of field experience and interpret- ations of reef quality depended on the respondant's special interest (an interesting reef to a coral specialist would not necessarily be selected by a bird or sea-snake expert, for example). Also, the information was often too generalized, referring to island groups rather than to specific reefs. The areas recommended and a summary ae ie Blot responses for the Pacific area are listed in able II. 62 L NNN Or NT rr SUOL} epuaUUodey 40 Jaquiny +++ tetteeteeett +SN}Ze9S [ROLL LOd - = + 0 " ¥ + + ¥ 0 = + 0 0 + + ¥ : = + 0 + ¥ 0 " - = + 0 + + 0 + + 0 i 7 0 + 5 - = + 0 0 + 0 + ¥ 0 + F - = + + + r Solptitoey = =alqissesoy = paquin}s L pur) 2907 Pau ILyLdeg - asuodsay auLeuUoLysant ++O+ +O+t44+04+4+000 ¥%59AN}ONAYS $994 “IT J1aVvl LLoIy asoy POUeS Spue|s] eoues NAa] LILA neq spue[s] ify pue|sy uojue) Spue|S] XLua0ud Langeun4 SpuelS] SLL Lg YOVOMLU YN LLY ouuy ountew Spuels] | LeysueyW Lbueveweburdey LLOFW ULAR LLOdy uy adeuod nal LLOFY ANLPLT Leo OM LLOFY LUFELN de, fay UaL>} need spue|s]T auLjour) AILS 63 *TTemey JO AJISIOATUN pue fa83u0, “W °D “Zeqem *N *¢ ‘Iteppoig "y “qd “yoTeIg “vy "Mm “YIINGeTS *¢ ‘teqreryoS *M “Y ‘Tepyoy ‘Vy ‘H ‘zoUEW *D “g *Ppel °S “H ‘souuevyor *y ‘eTomieey “YH ‘xoAouoBey *y ‘3zreqsoy ‘y “gy ‘A100 "Ss “W ‘TUR “I ‘VY ‘woOTg “71 “Y “sntuojuy “vy :SULMOTTOF ey Aq pouINjeA 919M SOITS ITFTOeg UO UOTIEWLOJUT JUSTeATNbe pue seaxTeUuUOT SS werso1d pepung *S°*f Inq TeUuOTeULEJUT e81eT e 02 Opnzti}e JUSUMLIEAOS poyedToTIUE 9Y2 AT Te oTseq + 8¢ °*d ‘anoge BTI9}1T1ID UOTJDEeTES 90S y OTqetIeA = F edTJOeIdWE 1O STqeOAeZUN = - o9TqQTSsod 10 uMoUyUN = Q quoseid Io oTqei1oae :4£9 ‘oseTedtys1y YOIeWsTg ‘UreIIAIg MON {[Tneqey {spueTs] uoWoTOS ‘eT[TTAUTeSnog “eyng ‘SpueTS] XTUeOYd ‘SpueTS]T J1eqTTO ‘(UTYRW) T1OWW TxeITIVINgG ‘spueTS] 1eqTTD ‘TTOIy eoI0UO ‘SpueTS] TTeysiew ‘oy,OM ‘spueTS] suTToIeD ‘o1onynN {spueTS] seuetiey ‘uedtes :ATUO oueu A gysessns soqTS T + + - + - uesd0 UeTpuy ‘eiqepTy pueTsussn) *N il + 0 + + 0 “SOTS] MOT i + + + + + "W'a'D “pueTs] uorey e9S IOUT] T + - - + 0 ‘yooy e1Ouysy eouTNy MeN T + + + co iT endeg ‘Aqseitow 310g 1k - + + a + BTUOpsTe) MON 19430 I = 0 + + + TITUPI, SPpueTST A9TIOS Z + + 0 + 0 pueTs] Suruuey T “P 0 0 cE 0 pues] seugstsy) SpueTS]T SUurT (aenuep) T + 0 S os 0 spueTs] Aaaray SpueTST oo) SUOT} BPUSUMUODOY +SN}21S SOTITTTOe{ sTqrsses.y peqinjstpuQ ,seimnjonI3¢ 31TS JO sequny TeOTIT Tog Jooy 64 It was obvious from the questionnaires and the other information available that data on the reef state and structure, and on current logistic arrangements, could only be obtained by actual site visits by scientists familiar with the basic criteria for a research site, aS was done in the Caribbean. As it would have been physically impossible to visit every potential site, it was necessary to select from those on which some data were available the areas with the best prospects for a satisfactory site. One important role of the questionnaires was to help pinpoint such areas for more detailed examination. Inevitably other areas which might be ideal have been overlooked because they were not sufficiently well known, often because they are more remote. Trans- portation in the Pacific however is improving rapidly, and will make more such areas logistically practical. A number of islands in the Pacific have recently been recommended for preservation as "Islands for Science" (Elliott, 1971). These were selected for their lack of human disturbance, and in many cases, have excellent coral reef development. A number of them were in fact considered at an early stage in the screening process. However, they are undisturbed precisely because they are inaccessible, and thus logistically impossible for a major research program. Site Screening An initial screening of possible sites in the Pacific reduced to a more manageable number the list of areas being considered although many excluded areas have characteristics that might be of great interest for more specialized programs. The following criteria were applied in the screening. Because of the need to find a reef with as few complicating external influences as possible, preference was given to atolls or barrier reef areas far from high island influences. For a large program, the need for regular air service or an immediately adjacent airstrip for possible charter service eliminated many less accessible sites. The possibility of acquiring or constructing and maintaining a facility for at least 20 people at reasonable cost was also considered important. All French territories were omitted from consideration because of un- certainties concerning long-term cooperation with government authorities. Great distance or travel time from the United States was also considered to be less desirable. The results of the screening are given below by island group, with the reasons for inclusion or exclusion from further consideration. Excluded from further consideration: a. Hawaiian Islands--Reef structure and diversity inadequate. Disturbed. b. Mariana Islands--High islands. Reefs marginal in quality. Serious disturbance by war and Acanthaster. c. Guam--Reefs seriously disturbed. d. Line Islands--Poor accessibility. Far from center of coral diversity. e. Phoenix Islands--Inaccessible. Canton disturbed. 65 Tokelau Islands--Inaccessible. Tonga--Poor accessibility. Cook Islands--Poor accessibility. Bismarck Archipelago, Solomon Islands, New Hebrides--High islands or inaccessible. New Caledonia, French Polynesia--French administration. Great Barrier Reef and other Australian reef areas--Great distance. 5a + n~u. A further breakdown was made of those areas not excluded. Those dis- cussed in more detail later in this paper are marked with an asterisk (*). Caroline Islands U. S. administration, good air service to district centers, many atolls, in center of coral diversity. 1. Palau*. Many recommendations. Greatest biological diversity in Pacific Islands. Regular air service. Some facilities. Considerable barrier reef area and one small atoll (Kayangl). 2. Helen Reef. Rich and undisturbed, but inaccessible. 3. Yap*. High island. Regular air service. Some rich reef areas. 4. Ulithi Atol1*. Airstrip and weekly Coast Guard flight, large atoll. Some disturbance during World War II. 5. Woleai Atoll*. Small, undisturbed atoll with abandoned air- strip. Excellent reefs. 6. Ifaluk Atoll. Very smal] atoll, inaccessible. 7. Truk. Accessible but reefs seriously disturbed. 8. Ponape*. Regular air service. High island with barrier reef. Some reef disturbance. 9. Ant Atoll*. Near Ponape, undisturbed except for recent Acanthaster damage. 10. Pakin Atoll*. Near Ponape, undisturbed. 11. Nukuoro. Inaccessible. 12. Kapingamarangi Atoll, Heavily populated, inaccessible. Marshall Islands U. S. administration. Many large atolls, mostly either disturbed Or inaccessible. Majuro*. Regular air service, disturbed. Arno*. Near Majuro, undisturbed. Wotho. Inaccessible. Ailuk. Undisturbed but inaccessible. Eniwetok. Accessible. Excellent facilities, disturbed. Clearance for foreign participants difficult to obtain. aArwnm— Gilbert and Ellice Islands Soon to have improved regular air service. 1. Funafuti. Serious World War II damage and heavily populated. 66 State of other reefs unknown. Fiji Islands Large high islands accessible. Rich reefs in outlying areas, no information on possible specific sites. Samoa Islands Accessible. Good fringing reef development. 1. Tutuila*. Reefs disturbed in more populated areas. 2. Manua Islands*. Undisturbed. 3. Rose Atoll. Small and totally undisturbed. Accessibility difficult. On the basis of the initial screening, a number of areas most likely to offer suitable sites were selected, including Palau, Yap, Ulithi, Woleai, Ponape (including adjacent Ant and Pakin), Arno, and Samoa (including Rose Atoll). A survey team was therefore sent to examine these areas. Fiji and the Gilbert and Ellice Islands also have great potential, but too little information was available on which to plan a site visit, and time did not permit the more lengthy survey that would therefore be required. U.S. Trust Territory* Of the enormous number of reef areas scattered throughout the Marianas, Caroline and Marshall Islands, most were too inaccessible to be considered, but even the accessible areas include vast reef tracts with innumerable potential sites. The experience and reports of the Acanthaster surveys conducted in Micronesia by the University of Guam (Tsuda, 1971) and the Acanthaster control teams operated by the U.S. Trust Territory Administration were of particular value in selecting promising areas for the survey team visits. Transportation in Micronesia is improving. Continental Airlines-- Air Micronesia now serves all the district centers (Koror, Yap, Saipan, Truk, Ponape, and Majuro) 2 to 3 times per week. In addition, Air Pacific has a charter service available for $180/hour (7-passenger air- planes), and there is a new charter service by Island Aviation, Inc., (rates around $120/hour). However, flying by other than Navy seaplanes is still restricted to those few islands with airstrips. Vessels service outlying islands at intervals of two weeks to several months. * Field survey: August 14-September 5, 1971 by A. Antonius and A. L. Dahl [assisted by R. Randall (Palau and Yap), R. T. Tsuda (Ponape) and B. Sablan (Majuro and Arno) ] 67 Local transportation is difficult, although small motor boats can usually be hired. The Acanthaster control teams have diving compressors and good motor boats at Palau, Truk, Ponape, and Majuro, and provided much of the logistic support for the survey team. There is an excellent marine laboratory and staff at the University of Guam, and the U.S. Trust Territory Government is highly cooperative. There is some resistance from the local populations to outside interference but in general they appeared favorably disposed towards scientific as opposed to commercial or tourist activities. There are small hotels in the district centers, but in outlying areas it is necessary to depend on the hospitality of the local people. Palau Palau (7°30'N, 134°30'E) is considered to have the richest reefs of any Pacific island area. The archipelago of volcanic and high and low limestone islands contains a complex of fringing and patch reefs partly surrounded by a barrier reef (Fig. 26). A small atoll occurs immediately north of the barrier reef. After a detailed examination of charts and consultations with local specialists, a number of sites ranging throughout the archipelago were chosen for detailed examination. This was a wholly inadequate sample of the diversity of sites available, so no generalizations should be made from the few descriptions presented here. 1. Ngeregong Island The first dive was at Ngeregong Island, on the windward (east) side of the barrier reef about 32 km south of Koror (Fig. 26). Ngeregong has an abandoned Japanese airstrip that, if restored, could simplify logistic arrangements. The outer reef slope, on the SE side of the island consists of a gentle sandy slope, with scattered coral growth, becoming even sandier with depth. The reef was poorly developed. 2. Angaur Angaur Island lies 40 km beyond Ngeregong, at the southern end of Palau, beyond the barrier reef (Fig. 26). It has an excellent Coast Guard airfield serviced weekly from Guam. The island, formerly an important source of phosphates, lacks major reef development except on the south and west sides. The team dived on the west (leeward) side of Angaur (Fig. 26), in the center of reef development. There is a smooth rocky flat with small scattered corals extending 700 m offshore to a depth of 5 m, followed by a gentle slope 200 m wide and going down to 8 m, with larger, more abundant corals. A steeper slope, 100 m wide with lush living coral coverage, extends down to 38 m, where it is interrupted by a sand flat 30 m wide. A slope with patches of coral interspersed with sand continues down from 40 m at 1 km offshore. The water was warm and clear. The lack of diverse coral habitats, the amount of sediment, and the extended zonation interrupted by terraces were considered undesirable for a research site. . The Ngemelis Islands are 40 km SW of Koror on the western barrier reef (Fig. 26). The islands are parts of a slightly elevated coral platform, and front on the outer reef margin on the west and a sheltered channel on the southeast, as well as the lagoon to the north. The islands can be reached from Koror by boat without going outside the barrier reef. Two dives were made, first on the SE side, off a vertical drop-off going down to 240 m, and then on the sloping west Side. 3. Ngemelis Islands, SE On the sheltered SE side of Ngemelis (Fig. 28), the shallow reef flat is 300 m wide, with few corals inshore, increasing to dense coverage of the reef edge at 0.5 m depth, with soft corals predominant. Beyond the edge is a vertical slope, mostly overhanging, dropping to 260 m. Alcyonarians dominate down to 30 m, with scleractinians and gorgonians equally represented. The slope begins to project outward at 40 m, collecting calcareous sediment; corals are scarce and appear not to grow beyond 30-60 m. The water temperature decreases with depth. The drop-off is spectacular, but soft corals are dominant, and there is apparently a constant flow of sediment. _ 4. Ngemelis Islands, W The west (leeward) side of the Ngemelis Islands (Fig. 29) has a broader reef flat 500 m wide with stony corals dominant at the seaward edge (0.5 m depth). There is a steep drop-off to 12 m with good coral coverage, with big buttresses dominated by Porites heads and rubble- filled chutes extending down to 20 m. A gentle slope extends down to 40 m with Pachyseris dominant, after which the slope becomes increasingly sandy and corals dwindle. Again water temperature decreased with depth. The site has high species diversity and a good drop-off, but sediment- ation limits coral growth at around 50 m. At both sites the drop-offs are very near the surface, and sponges are almost completely lacking. Current patterns in the area appeared quite complex, and this, together with the inadequacies in the reef structure, would rule this area out for certain types of reef research. Kayang] Atoll is several hours away by small boat, traveling up inside the western barrier reef, and then beyond the northern tip of Palau (Fig. 26). Kayangl would generally be accessible from Koror except in rough weather. There was some difficulty at first because of a recently instituted village policy of charging all visiting Americans $50, but the requirement was waived after the District Administrator explained the value of the research program to the islands. Three dives were made on Kayangl; in the shallow lagoon, and on the outer leeward and windward slopes. 5. Kayangl lagoon The first dive was on the west (leeward) side of the lagoon near the entrance channel (Fig. 30). The sandy bottom, 4-6 m deep, becoming 69 shallower toward the reef, separates the abundant patch reefs and coral heads. These are coral-covered from top to bottom, and show very great species diversity. The area is typical of shallow Pacific lagoons. 6. Kayangl leeward outer reef The leeward outer reef of Kayangl (Fig. 30) is topped by a 50 m wide flat of calcareous algae, leading to a reef edge with lush coral growth. There is a steep drop-off to 10 m depth, still with very good coral coverage and high species diversity, followed by a gentkeslope to 40 m with Pachyseris dominant. The slope flattens between 40 and 50 m, and becomes very sandy beyond 50 m. The temperature decreases markedly with depth, and the water becomes increasingly murky. The living reef ends between 50 and 60 m. 7. Kayangl windward outer reef The east side of Kayangl is very exposed, making it impossible in most weather to anchor a small boat. The windward outer reef (no diagram) commences off the islands with a broad algal-turf-covered flat, and then drops to a platform about 1000 m wide and 7-10 m deep with little coral growth. This was not followed to the drop-off. The lack of a developed windward coral community and the shallow lagoon prevent this from being as desirable a site as logistics and diversity might indicate. Yap Yap (9°30'N, 138°05'E), as a district center, is accessible (3 flights per week from Guam), and has excellent fringing reefs, espec- ially on the northwest side, with some lagoon development inside the reef (Fig. 31). It is a high island, so that there are considerable terrestrial influences, and the reefs have experienced some Acanthaster infestation. In four dives on the west side, a superficial count of corals by R. Randall yielded over 100 species. Profiles from the dives on different parts of the reef have been combined to yield a composite section (Fig. 32). 8. Off Gorror Two dives were made in the lagoon, on the SW side, between the reef and Yap Island. The first, off Gorror, was in a shallow sand flat area with turtle grass and occasional coral patches of high coral diversity (approximately 50 species). The water was rather murky. 9. Off Nif The second dive, off Nif, was in one of the deep lagoon areas. The Surrounding reef walls are covered with large corals down to the sandy bottom at 10.m. The reef top is exposed at low tide, with an algal turf cover and only small specimens of Favites. 70 10. Off Okau The leeward outer reef off Okau begins with a rocky fore reef flat, 1500 m wide, cut by long surge channels 2 m deep and 1-2 m wide. Coral coverage increases to the reef edge at a depth of 7 m, with only a few species less than in Palau. The slope then drops steeply to 25 m with good coral growth, beyond which a more gentle slope is dominated by Pachyseris. The water changes at around 20-25 m; above it is warm and clear, below cool and turbid. Coral growth seems to end at around 40 m. 11. Mil Entrance A final dive was made in the Mil Entrance (no diagram) on the west Side between Yap and Rumung Islands. The reef flat on the north side of the channel extends down to 2 m with good coral coverage. There is a drop-off with good coral growth down to 10 m, and a gentle slope with turtle grass deeper down. A strong current of turbid water was moving out of the channel, which contained many large sharks. Although the reef quality observed was high, the terrestrial influences, distance of the reef front from shore, and shallow limit to coral development are major disadvantages. There is also more chance of antagonistic feelings against Americans on the part of the local population in district centers. Ulithi and Woleai It was not possible during the short survey to arrange transportation to Ulithi or Woleai Atolls in the Yap district, but from other evidence and from conversations with chiefs from both areas, some useful inform- ation was collected. Ulithi (10°00'N, 139°45'E) is a large atoll northeast of Yap. It has an airstrip with weekly Coast Guard flights from Guam or the possibility of charter flights ($500-$650 for 5-7 passengers, one day round-trip). There is also ship service from Yap every two weeks. There are some sunken ships in the lagoon, and other remnants of war damage, as well as an increasing ciguatera problem, but with the large size of the atoll undisturbed areas must remain. This is one of very few relatively undisturbed atolls directly accessible by plane. Woleai (7°20'N, 143°45'E) is a small, totally undisturbed atoll presently inaccessible but with an abandoned Japanese airstrip. If the airstrip were ever repaired this could be an important site to consider, even though logistics would be somewhat more difficult. Earlier suggestions that the airstrip might be restored soon have not been confirmed. A.Antonius dived on the reefs during the 1969 Acanthaster.- survey, and they appeared to be well developed. Ponape Ponape (7°00'N, 158°00'E) is another district center, consisting of a 7] large volcanic island and several smaller islands surrounded by a barrier reef (Fig. 33). The reefs are generally subjected to consider- able terrestrial influences. There are three flights weekly from Guam to Ponape and two from Hawaii, making this the most accessible of the Caroline Islands. Not far from Ponape, however, are two small atolls, Ant and Pakin. Ant is an atoll of moderate size only 10 miles from Ponape, while Pakin, somewhat smaller, is 30 miles away. Both can be reached by small boat from Ponape within 3 hours. One dive was made on the outer slope of each atoll, and both had excellent reef development. 12. Ant The dive was on the leeward side just beyond the northern tip at the northernmost islet (Fig. 33). The reef crest at this point is 100 m wide, with a fore reef flat 200 m wide, extending down to the reef edge at 10 m (Fig. 34). Coral growth begins at 2 m depth, 30 m offshore, increasing to 90 percent coverage at 50 m offshore, and 100 percent at 100 m out. From the reef edge there is a steep slope down to 30 m depth with a coral cover of over 100 percent because of the overlapping table Acroporas which are dominant. The slope continues down to 50 m with interspersed sand cover increasing to 50 percent, although coral development continues much deeper. The water was warm and clear, with no change in temperature. Ant is an excellent reef, with a deep entrance, a good drop-off, and deep coral development. 13. Pakin The dive site at Pakin was near the center of the northern reef (Fig. 33), in an exposed though not windward area. The 80 m wide reef crest merges with a sloping fore reef 50 m wide and 7 m deep at the edge (Fig. 35). The first 20 m is bare of coral cover, but coral density increases to the rugged edge, with headshaped Porites dominant. From there an almost vertical slope descends as far as could be observed, with a good coral cover of large specimens and a considerable amount of algae. Sand patches begin at 50 m, but at 60 m the coral cover is still 30 percent and continues to the limit of visibility. Again there was clear warm water with no temperature change with depth. The reef is excellent for research, with a steep drop-off and coral development to beyond normal SCUBA range. Six gray sharks were observed. Ant has a single owner, an ardent conservationist, which might make arrangements for a long-term project difficult. Pakin has no channel into the lagoon at present, but there are government plans to open one and to build a pier in the lagoon. There would not appear to be any difficulty in locating a facility there, and logistical and construction Support on Ponape seems excellent. One dive was also made on the Ponape barrier reef (see below), but it was disappointing. 14. Mant Passage The brief dive in the Mant Passage, a northeast entrance through 72 the windward wide of the Ponape barrier reef revealed a smooth rocky flat at 4 m depth with 20-30 percent coral coverage and low species diversity. The water was rather turbid, and one gray shark 2.5 m long was seen. The frequent heavy rainfalls in Ponape might cause technical problems for a research program, as well as affecting the reef. Arno and Majuro Majuro (7°05'N, 171°10'E) and adjacent Arno (7°05'N, 171°45'E), two large atolls in the southern Marshall Islands, are the only accessible atolls in the area without government entry restrictions. Majuro has two weekly flights from Hawaii, and Arno can be reached easily from Majuro. Majuro has been reported as being disturbed, while Arno is relatively untouched but dives on both atolls revealed rich reefs with an excellent configuration. 15. Kinajon (Arno) The outer reef off Kinajon on Arno is on the sheltered south side (Fig. 36). The rocky fore reef flat 100 m wide develops good coral coverage and surge channels towards its edge at 10 m depth (Fig. 37). A steep slope with Porites heads dominant drops to 30 m where the angle of Slope lessens and sand patches appear. Pachyseris then becomes dominant. At 60 m the coral cover is still 20 percent and continues as far as can be seen. The water was warm and exceptionally clear (the water surface was visible from 60 m), with no change in temperature. The reef in general seemed excellent; on the other side where the deep entrance is, it is reportedly even better developed. 16. Laura (Majuro) On Majuro the reef was surveyed off Laura, a sheltered location on the southwest side (Fig. 36). The 60 m wide fore reef consisted of 20 m of bare surface, 20 m with a dense algal cover, and 20 m with corals, mainly Acropora, leading to the rugged reef edge at 5-8 m depth (Fig. 38). Deep surge channels cut into the reef. There is no real drop-off, but a gently rounded slope with valleys and ridges perpendic- ular to the shore, and also many shore-parallel steps. The coral cover decreases from 90 percent at the edge (5-8 m) to 60 percent at 12 m, to 50 percent at 20 m with increasing algal cover. The slope gets steeper with depth, becoming vertical at 40 m, where the coral cover is 20 percent. This site is accessible by road from Majuro, and has good reef features and clear water. Fishes were very abundant, but no sharks were observed. AMERICAN SAMOA* The Samoa Islands (14°15'S, 170°00'W) are a chain of volcanic *Field survey: September 7-12, 1971 by A. L. Dahl and A. Antonius (assisted by S. Ritterbush). 73 islands with fringing coral reefs in the South Pacific. While rather far from the center of coral diversity, their ready accessibility by direct flight from Hawaii and the strong interest of the local govern- ment warranted detailed examination of the reefs. The Manua Islands are remote from the main island of Tutuila and can be reached by government or commercial boat in about 8 hours. There has also been an intermittent float plane service. Because transportation is difficult it was only possible to dive on the reefs in the vicinity of the main anchorage of each island. Since the reefs in the Manua Islands were not transected no diagrams are given. 17. Tau (Manua) At Tau the dive was made off the northwest side (Fig. 39), a lee- ward but exposed area, about 300 m offshore. There is a rocky flat with huge boulders at 20-25 m depth. Few coral species are present, mainly Porites and Pocillopora, and specimens are small and scarce. 18. Olosega (Manua) The dive off Olosega was on the sheltered leeward (west) side about 200 m offshore, at 20-25 m depth (Fig. 39). The bottom is rocky, with huge blocks, forming valleys and ridges. Coral cover is more extensive and with more species than Tau, with huge coherent colonies of table- like Acropora and Porites (lobata?). Many alcyonarians are also present. 19. Ofu (Manua) The anchorage of Ofu is on the west side in a very sheltered location (Fig. 39), 200 m offshore from Alaufau, where the water is 10-20 m deep. The bottom has marked topographic relief, with smal] corals and red algae on top of the elevations, dense coral cover and good species diversity on the walls, and white sand in the bottoms of the troughs. The water was clear and warm. Fishes were abundant, but no sharks were observed. Two dives were made on the fringing reefs on the north and south sides of Tutuila, the main island of American Samoa. 20. Leone Bay (Tutuila) At Leone Bay on the windward south west side (Fig. 39), the reef was surveyed out to 350 m from the shore nearLogologo Point (Fig. 40). The reef structure is irregular, somewhat resembling a spur and groove system, with a shallow reef flat, and then large reef patches in deeper water, extending down to 25 m. The coral cover is very variable, some- times almost 100 percent, as at Ofu, but many helioporas are also present. A large flat of white completely detritus-free coarse sand occurs at 15 m depth. The water was warm and clear, with many fishes and no sharks. 74 21. Ogegasa Point (Tutuila) On the north side of Tutuila, at Ogegasa Point (Fig. 39), there is a vertical basaltic rock slope from the surface to 3 m, followed by a rocky flat with small scattered corals extending 50 m offshore to a depth of 7 m (Fig. 41). From here a slope with spur and groove configurations drops to 15 m and then a very steep slope down to 30 m, ending in an extensive sand flat. This slope has the best coral cover- age and richest species diversity of all the Samoan sites observed. The water was warm and slightly turbid, perhaps from a recent rainfall. Again the area was rich in fishes but lacked sharks. Apparently there are no good near-shore drop-offs around the islands of American Samoa. It was not possible to reach Rose Atoll, as the seaplane was damaged shortly before our arrival, and time and weather precluded boat transportation. Discussions with the Office of Marine Resources which recently surveyed the atoll, indicated that the land area was inadequate for any facility, so that it could only be used for short visits. 75 CONCLUSIONS The overall evaluation of the site information leads to the following conclusions for the Smithsonian-planned programs. In the Caribbean, the logistic problems and unique character of the San Blas Islands, the disturbance and political problems at Discovery Bay, and the poor reef structure of St. Croix and Acklins Island left Glover's Reef, British Honduras as the preferred site. In the Pacific no final decision was possible without further field surveys, but several areas showed good potential, including Pakin, Ulithi, and Arno. Ulithi was not visited by a survey team, and so requires further examination. A more detailed study of areas in Fiji and the Gilbert and Ellice Islands could also be productive. However, once the scientific suitability of a site was determined, it would still be necessary to negotiate with the local inhabitants for space and permission to work on their reefs. It is important to remember that the areas reported on here were selected and described in accordance with the specific program criteria listed in the introduction, not all of which would necessarily apply to other projects. We hope that others searching for a suitable location for a coral reef research program will be able to use this information, with due caution for its limitations, in selecting a site most appropriate to their needs. 76 ACKNOWLEDGEMENTS We should particularly like to acknowledge the support of the Environmental Sciences Program at the Smithsonian Institution and of the International Decade of Ocean Exploration office of the National Science Foundation (Grant GX-28676). In addition, the cooperation and support of the governments of the Bahamas, British Honduras, the U.S. Trust Territory and American Samoa were greatly appreciated, as was the assistance of the West Indies Laboratory of Fairleigh Dickenson University on St. Croix, the Discovery Bay Marine Laboratory on Jamaica, the Smithsonian Tropical Research Institute in Panama, and the University of Guam. The individuals who cooperated in this survey, either as participants on survey teams, contributors of questionnaires or site suggestions, members of the CITRE and IMSWE programs, or suppliers of local advice and support are too numerous to mention although many are acknowledged at appropriate places in the report. 77 REFERENCES CITEN Cloud, P. E. 1959. Geology of Saipan, Mariana Islands. 4. Submarine topography and shoal-water ecology. U.S. Geol. Survey Prof. Paper 280-K: 361-445. Elliot, H. 1971. Pacific oceanic islands recommended for designation as islands for science. South Pacific Commission. Regional Symposium on Conservation of Nature--Reef and lagoons. SPC/RSCN/ WPeetse o'26-July..1971.. 16 p. Glynn, P. W. In press. Aspects of the ecology of coral reefs in the western Atlantic region. In R. Endean and 0. A. Jones (eds.), Biology and Geology of Coral Reefs. 2 vols. New York, Academic Press. Goreau, T. F. 1959. The ecology of Jamaican coral reefs. 1. Species composition and zonation. Ecology 40: 67-90. Goreau, T. F., and J. W. Wells. 1967. The shallow water Scleractinia of Jamaica: revised list of species and their vertical dis- tribution range. Bull. Marine Sci. 17: 442-453. Johannes, R. E. et al. 1972. The metabolism of some coral reef communities: a team study of nutrient and energy flux at Eniwetok. BioScience 22: 541-543. Kinzie, R. A. 1970. The ecology of the gorgonians (Cnidaria, Octocorallia) of Discovery Bay, Jamaica. Unpublished Ph.D. thesis, Yale University. Laborel, J. 1967. Madréporaires des cétes du Brésil. Deuxiéme thése: l'Université d'Aix-Marseille. Land, L. S., and T. F. Goreau. 1970. Submarine lithification of Jamaican reefs. J. Sed. Petrology 40: 456-462. Macintyre, I. G. 1972. Submerged reefs. of Eastern Caribbean. Amer. Assoc. Petroleum Geologists Bull. 56: 720-738. Meyerhoff, H. A. 1926. Geology of the Virgin Islands, Culebra and Vieques: Physiography. N. Y. Acad. Sci. Scientific Survey of Puerto Rico and the Virgin Islands 4: 72-219. Newell, N. D., J. K. Rigby, A. J. Whiteman, and J. S. Bradley. 1951. Shoal-water geology and environments, eastern Andros Island, Bahamas. Bull. Amer. Museum Nat. History 97: 1-30. Stehli, F. G., and J. W. Wells. 1971. Diversity and age patterns in hermatypic corals. Syst. Zool. 20: 115-126. Stoddart, D. R. 1962. Three Caribbean atolls, Turneffe Island, Lighthouse Reef and Glover's Reef, British Honduras. Atoll Res. Buc NOW 87.5. 05/0: Storr, J. F. 1964. Ecology and oceanography of the coral-reef tract, Abaco Island, Bahamas. Spec. Pap. Geol. Soc. America 79. 98 p. Tsuda, R. T. (ed). 1971. Status of Acanthaster planci and coral reefs in the Mariana and Caroline Islands, June 1970 to May 1971. University of Guam, The Marine Laboratory, Technical Report No. 2, October 1971. 127 p. Wells, J. W. 1969. Aspects of Pacific coral reefs. Micronesica 5(2): 317-322. Whetten, J. T. 1966. Geology of St. Croix, U. S. Virgin Islands. Geol. Soc. Amer. Memoir 98: 177-239. a NN 4 ie sphere Aer 119 9 buol) Tae totew-Nsodd Hier edgod Baps [she ReNSSiedass che ghissderot HsptcOpPteh lo. SORE aaa }'> boat Wiwde orgsas o2 ONG h eige leh ae eae ) Horsayyseautche We eogme, P* OFTP RT Pao as Yi We : ‘ 4 15 FAA ‘ 1 3 Si} i , AOS ont ae yee FA A es a to (sit rota F ay Fal] ots ri Ll . fot itersv Fae : 15 ¢.0 Sa .L .2t997 989 hema 2 TO} ) .l ,swiaihoe : ai forts? .SOReamamm i / . i Aan aa \ 4 Poteet re Sele a los yporeyAS esupatl) am ays amven id avs AM ’ a on Site name Ocean area General location Longitude Latitude Charts General references //island; //Continental; //Atoll; //Fringing reef on //volcanic or //other base; {//Barrier reef. Description of site Notes K (0) a Features: Adequate comparable reef area for sampling (perhaps 1 km frontage) Well developed reef flat for drilling Reef development to at least 50 m depth Width across reef less than 300 m Reef near enough to shore to permit shore-based instrumentation Reef undergoing active construction Considerable species diversity Reef not obviously unique Major terrestrial influences absent Reasonably continuous reef accumulation Current and tidal flow patterns permitting cross-reef metabolic studies Variety of subsidiary site types in area Undisturbed by development, catastrophic storms, war, or pollution Probability of remaining undisturbed Weather permitting year-round work Accessible, within 1 day's travel of a Major airport and harbor facility Anchorage and landing for work boats Accommodations available Research space available Buildings available for conversion Electricity Fresh water Local suppliers (food, fuel, building materials) Research vessels available in general area Governmental authority Possible attitude toward project Previous scientific work (include references) SN ASsssss_ BSS WW SRAVE GAG 4 SN SEASsss B&H Be VEROass SW Ve Aerial Photographs (Source) Other notes Information supplied by Date A sketch map on the reverse would be helpful.. EEG DGsES SCRLs or approximate distances if possible. (Also idealized cross sections with reef zonations) Figure 1 - QUESTIONNAIRE FOR SITE SURVEY <= int orth East Po N 9 Pinefield ey "0 3) SH Atwood Harbo The Bight of Acklins Golden Grove 5 km Figure 2 — ACKLINS ISLAND 22930'N 74°00'W ; = HOPYVH GOOMLYV -S ALIS — € aunbi4 SI9OIN o00€ 0002 0001 uoljesabbex3 eae, x OG adojs mes iel4-pues< 29-3104 ro. Bena aa ey 4aMo7 & Ol Lun adojs jeay-a104 Jaddpy faay 9eq : gorse, < 19010 4094 Ap’ SE, ee ....Buck | f Ph a A — Garbage Dump B — Bauxite Plant ~~ pare ace 400 Fathoms — XN C — Oil Refinery 5 km Figure 4 — ST, CROIX 17945'N 64°40'W S3LIS ONIAIG ONIMOHS XIOY¥D “LS 4O GNA LSwa — g enbiy Ses; / —S=>S> gts es ee ~~ seg jeuoneny | yong (uapBO “a “N pue snuUar “q ‘Yyr!4 “7 Aq patiddns eyep) AV 3NOWL 44O 4535Y YAIYYVE ANVG- 1 ALIS- XIOUD “LS — 9 anbi4 S19}0N oxoss 00d esojnbue essnjj\ Byeoinj. Saqd0q “ds eijjAydoyaoA | ‘ds eulapueay sueqeu BI||Aydodjo5 SILO JIYUIWAge| eluoldiq BSOUJAACD PANSEIUOW\ SluloaIMao B1OdoIDY >= ‘ds eloieby Ge _za1apis Banserepis J-y Saeinuue eansequoyy sy esoblu1s e0jdiq esobiiujs elojdiq eyewjed e1odoiy auozZ eee SaplOedjse Sa}10qg eyeue|dwios esodaj|i/\| OO| ejeujed e1odoiy suejeu e1||Audodjo5 esoblujs elojdiq sluejnuue ead}Se]UO/\| Sa}l40d saqo0g auoZ Jeay pues ma, os oes sons Keg sae4 S| Ol O SIOLOWN —— N.SZoZZ N,OE081L CLs, te StL te rye ss eye ENS SSRIS Be Gas Od alixneg Jasiey P, a aes asia Yi < AV§ AYSAOISIC Z Asoyesoge WN; Aeg Asanoosiq auozZ uoobe| ee j : | 1e14 390u f a, i iu é ‘i fi pe Bu : Ne NIE DOINGS ty as Sie re see Sasa) Sa 00S ado|s jaay o104 watt tetses S}99Y ||!S 49M07 é ado|s jaa 2104 daaq auoz uoobe VOIVINVE ‘AVE ANAAOOSIG — 8 2Nb14 auoz eyewed auoz uadeg auoz 1eay yelf peo Shite = <— - (exe) ado|s 901-310} daap A Nw adojs jaaJ-a104 YS l { ea $4994 |IIS fe! ae ud auoz paxiw ea \ I) OO| 08 Og Ov O02 O 002 OO£ OOvV S12}2I\) Windward Reef Flat Leeward Peripheral : e a Reef . Peripheral ae Reef “es ne S ae se es Wis S LONG CAY; : omen ice /=:NORTH “NORTHEAST Se es «(CAN 2 Weg “"” LONG CAY NIN ie eee 5 km of 1 2 SOUTHWEST CAYS Figure 9 — GLOVER’S REEF (modified after Stoddart, 1962) 16°50'N 87°50’w Meters 500 Meters : G 20 ¢ Acropora palmata 2 @ Acropora cervicornis @Montastrea annularis 4 Dendrogyra cylindrus () 4 Diploria labyrinthiformis x 30 Diploria strigosa We Diploria clivosa ee @Agaricia agaricites @Agaricia tenuifolia 40 Madracis mirabilis Madracis decactis yy, Porites furcata lsophyllia sinuosa lsophyllia multiflora 50 Dichocoenia stokesi ( Dichocoenia stellaris Siderastrea siderea Favia fragum Meandrina meandrites Porites astreoides 60 \ AHelioseris cucullata is is D A Agaricia lamarcki A Agaricia undata 70 A Agaricia fragilis A Montastrea cavernosa Scolymia lacera is \\ \ \ 80 4 —= dominant on the fore-reef 90 @ — = dominant on the reef-edge A= dominant on the slope 10 x Vertical Exaggeration : Note GLOVER’S REEF 2 Figure 10 — TYPICAL WINDWARD REEF 500 Meters. Meters 20 @ Acropora palmata @ Acropora cervicornis 30 @Diploria labyrinthiformis #Montastrea annularis @Siderastrea siderea @Diploria strigosa ®@Diploria clivosa lsophyllia sinuosa Madracis decactis aviattraguinie> iy) Wiamer ie! NGS Lops A Asgaricia agaricites , 50 A Agaricia tenuifolia A Montastrea cavernosa Porites astreoides Meandrina meandrites Solenastrea bournoni 60 Dichocoenia stokesi Dichocoenia stellaris Porites porites __ Siderastrea siderea 70 Scolymia lacera Mycetophyllia lamarckiana 40 80 @ —= dominant on the fore-reef @—*_ dominant on the reef-edge A> dominant on the slope 90 10 x Vertical Exaggeration a GLOVER'S REEF Figure 11 — TYPICAL LEEWARD REEF Coco Bandero Cays i 58 Aes > aa f ox fs Ti! se ey ee) Ce £ x ae G a a ee ROS RCI APRS Soe oN Cie d) Lees 8. Ge cos, Mangles Point AOR ary Race. MAY FLOWER CHANNEL e Se SEES SS Sy. LA Sree aS RRC a fea es ZN oe © zee Se Aes: a7 wad Ct REARY AS SY RONG, io ti fo! = a 2 gf © “1 <{ ier’) i i A) a ) 2 i) 2 fo} a i == a” So eee The dominant coral below LANG 20 meters is Agaricia lamarcki Other significant corals include Montastrea cavernosa Madracis decactis _ Siderastrea siderea 30 Stephanocoenia michelinii Mussa angulosa Scolymia cubensis Scolymia lacera Mycetophyllia reesii Manicina areolata SAN BLAS ISLANDS, PANAMA — OFF SARDINGAN POINT Figure 16 — PROFILE B—B’ NM.8t08Z N,O€06 SONV1SI SV18 NVS — ZL eunbig Hels Bi AROS OE RUE Seatearenes Aniepxconyy C LO MC uy NEB aM RC, UN: én; ~ LAIST ey “s G Cooks 2 é ‘ ae wy | 3: se jens | NOYOW 7 . ~] fs ‘ : ‘ SABD SO}|NJID so 9-9 311dOud — BL enbi4 YV1VS 440 — SGNV1SI SV18 NVS ‘ds eimAjooS ‘ds eijjAydo 990A \ BSOUJAACD AAISe]UO|\| JO speay paia}je9S ‘ayeujwop “ds elosiueby 4o syIMmOosb Ajejq O¢ e490e| PIWA|OIS Badapis ea;}SesapiS ‘ds e1jj|Aydoyash | BSOUJAARO BadJSeEJUO|\] apnjou! sje4o9 1940 ‘ds Salldog pue |youewe| elolieby ‘siuejnuue earjse}UO/\| yO syymoiB Ajejd uaemjaq sindd0 pues YyolJ-epawijey V7 02 “jueUIWOp osje “ds eldiseby “sjauueyo pues pauljap [jan JOU aWOS “sue}eU el|j|AYdodjoD pue sjjejnuue eadjsej,UO//\ yo Ajuyews pasodwoo ‘sasseaz3nq padojanap Ajio0q Ol slueynuue ead]se}UO}/\ ‘ds eloeby SIUJODIAJ9D BIOdOJOY, SB}IOd se}Og apnjoul sjesOo 184109 "saluojoo eyewjed exodois\y abie| peia}jze9S O S1O1aIN 1 9 ,0—G 31140ud — 61 aunbi4 yvi1Vvs 440 — SGNV1S!I Sv1d NVS =_ smitiaigrnre ; Aon) Ott sees cae ~ bee ae Ss ; ‘] bss tye (8. 7 safuods ab1e7 . 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In deep areas there are abundant corals (Porites astreoides, Agaricia sp. and Diploria sp.) and Diadema sp. In shallow areas the bottom is covered with Padina sp. Middle Ridge. The surface, which is about 0.3 meters above sea level at low tide, is extensively pitted and etched. This is in marked contrast to the smooth rock floors of the troughs or tidal pools. Sampling indicated that this ridge is dominantly composed of crustose coralline algae. Inner Trough. Shallow rock bottom covered with algae, mainly Caulerpa sp. and Halimeda sp. over crustose corallines, Porites porites is common, Inner Ridge. As in the case of the Middle Ridge the surface is extensively etched and pitted. However, this ridge is composed, for the most part, of corals in a coarse calcarenite matrix. of the ridges. 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(0) . o¢ 3e|4 pues AjyissaAip pue oz afeianod je4109 ybiy Ol sje409 |jewWs yeseq 0 VIINLAL ‘LNIOd VSWD3DO - LZ ALIS — Ly aunbiy 02 Ol e1Odoljay YyYonw osje ‘jueUIWOp BiOdoIDY pue ‘Sa}1104g ‘e10do]|190q fe) slasaw VIINLAL ‘AVE JNO31 - 02 ALIS — OF BNb14 STATE OF KNOWLEDGE OF CORAL REEFS AS ECOSYSTEMS Marie-Héléne Sachet INTRODUCTION The following brief review of current knowledge about coral reefs as ecosystems was written early in 1972, mostly from material furnished by the CITRE working groups. It attempts to describe the status of reef ecology papers which are particularly relevant to the modeling approach. No information on the systematics of reef biota is included in this summary, although inventories are obviously fundamental to any bio-ecological work. This review has become out-of-date in certain respects and should, ideally, be thoroughly revised. Time for this is not available, and only a certain amount of updating has been done. The bibliography has been gone over very carefully, corrected and very much enlarged. I acknowledge with thanks the work of Dr. Bryce Decker in this matter, and the help of former CITRE participants, especially those working in the Smithsonian. In its present form, and with apologies for its shortcomings, I believe this review may still be of value, especially to readers not thoroughly familiar with the coral reef field, or those familiar with other aspects of coral reef research, e.g. systematics. Since the November 1971 workshop, and the January 1972 completion of the CITRE proposal, the results of a number of Symposia and Seminars have been published, as well as several special issues of journals, devoted largely to coral reef research. They demonstrate how active and varied such research is, and should be consulted for topics outside the scope of this review, for details of studies only briefly mentioned here, and for more recent developments not covered here. Among such volumes can be cited the Proceedings of the 1969 "Mandapam Symposium" issued in 1972 (Mukundan and Pillai, eds.), those of the 1972 International Helgoland Symposium, "Man in the sea," assembled in a whole volume (24:1973) of Helgolander wissenschaftliche Meeresuntersuchungen, the March and June 1973 issues of the Bulletin of Marine Science, containing 17 papers "In Memory of Dr. Thomas F. Goreau," and a recent issue of Atoll Research Bulletin (166-170) devoted to five papers on Acanthaster infestations. (Manuscript completed March 1974) 122 The results of an Acanthaster symposium at the Second Inter-Congress meeting in Guam, May 1973, have just been published in Micronesica (Dec. 1973). Also just received is the first volume of the long awaited treatise on Biology and geology of coral reefs (Jones and Endean, 1973). Several other symposia and review volumes are still in press, including the results of the "Floating Symposium" (2nd International Symposium on Coral Reefs, Great Barrier Reef), June 1973. Hot off the press at the time of the Glover's Reef workshop and the preparation of the CITRE proposal were Regional Variation in Indian Ocean Coral Reefs (Stoddart and Yonge, 1971) and the Supplement to Island Bibliographies (Sachet and Fosberg, 1971). BACKGROUND Following Charles Darwin's description and theory of atoll origin (1842, 1962), a growing controversy over the "coral reef problem" (i.e. origin of atolls and reefs) led to vigorous debates based mainly on information collected from published bathymetric charts and insufficient field data (e.g. Daly, 1910, 1915; Vaughan, 1916a, 1916b; Davis, 1928). An exception was the results of the Funafuti expeditions of the Royal Society (1904) and of the Australian Museum (Hedley, 1896-1900). Over the past 20 years, the "coral reef problem" controversy has abated, particularly since drilling techniques have made it possible to establish that atoll formation is indeed associated with subsidence--notably in studies by Ladd and co-investigators (Ladd et al., 1953; Ladd and Schlanger, 1960; Ladd et al., 1967). The enormous volume of work carried out on reefs and atolls in the earlier periods and in recent years is summarized in literature reviews such as those of Stoddart (1969a), Ladd (in press) and Glynn (in press). Major topics discussed in previous reports include reef formation (Ladd and Tracey, 1949), geology of reefs (Ladd, 1961; Fairbridge, 1950; Newell, 1959), biological zonation of reefs (Wells, 1954, 1957a, Goreau 1959), and coral physiology (Yonge, 1940, 1963, 1968) to list but a few. In addition, several bibliographies of coral-reef studies have been compiled recently (Pugh, 1950; Sachet and Fosberg, 1955, 1971; Wells, 1957b; Ranson, 1958; Milliman, 1965). As in the early expeditions, most of the work on specific atolls in the last 20 years has consisted of detailed descriptions and inventories (Ladd, ed. 1954-date; Sachet 1962b; Direction des Centres d'Expérimentations nucléaires, 1969). A qualitative description of the coral atoll ecosystem was prepared in 1957 by Fosberg (1961, 1963b). These descriptions furnish an excellent baseline for comparative studies and a starting point for the quantitative observations needed for model construction. It can be safely stated that neither the qualitative constitution nor the role of a single functional component has been completely elucidated in any one reef environment. Ecological processes on tropical reefs have been estimated on occasion by projecting data from long-term or detailed studies of temperate marine communities (Paine and Vadas, 1970; Newman, 1970). An aspect of reef ecology which has hardly been developed as yet b i 3 123 is any work permitting an estimation of carrying capacity for exploitation by human populations. GENERAL REEF SURVEYS Since Darwin's reef studies, there have been many important general investigations of the nature of these structures from various viewpoints. Some of these were one-man enterprises, such as those of Alexander Agassiz at the end of the 19th century aboard the ALBATROSS, in all the major reef areas of the world, those of J. Stanley Gardiner in the Indian and Pacific Oceans in the first part of the twentieth century, that of Wood-Jones on Cocos-Keeling, results published in 1909 and 1910, and that of Gibson-Hill, also on Cocos-Keeling just before World War II. David Stoddart made three detailed geomorphological studies of the British Honduras reefs and cays between 1959 and 1965. Notable were a substantial number of cooperative investigations involving people with different interests and backgrounds. Among these have been the Royal Society's and the Australian Museum's expeditions to Funafuti, Ellice Islands, reported on by David and Sweet in a volume published by the Royal Society of London in 1904 and in papers edited by Hedley (1896-1900); the Great Barrier Reef Expedition in 1928-1929, under the leadership of C.M.Yonge (1930a, 1930b;, 1931); the Carnegie Institution of Washington's Coral Reef Program, with work especially at the Dry Tortugas Laboratory and in American Samoa and Tahiti in the first third of this century. Involved in this were especially Alfred G. Mayor (Mayer), W.A. Setchell, and T. Wayland Vaughan; some of their publications are listed in the references. After World War II came a comprehensive program of work on Bikini and Eniwetok Atolls in connection with the atomic weapons tests there (Ladd, 1973). This work is published as an enormous series of papers collectively known as USGS Professional Paper 260, parts of which are still appearing. The USGS-Army Map Service Far East survey of the Northern Marshall Islands in 1951-1952 led to many papers, especially the Military Geography of the Northern Marshall Islands and Atol] Research Bulletin 113 (Fosberg et al., 1956, Fosberg and Carroll, 1965), and a review by MacNeil (1972). During the 1950's the Pacific Science Board's Coral Atoll Program sent multi-disciplinary expeditions to Arno Atoll, Onotoa Atoll, Raroia Atoll, Kapingamarangi Atoll, and Ifaluk. In 1958 and 1960, under this same program were two expeditions to Jaluit Atoll to study the effects of Typhoon Ophelia. Reports were published in the Atoll Research Bulletin. ORSTOM, the French overseas research organization, sponsors a multidisci- plinary study of coral reefs around Nosy-Bé, Malagasy. From 1967 to the present time the Royal Society has sponsored continuing work on Aldabra Atoll and other southwest Indian Ocean coral islands and reefs. A small research station is now maintained on Aldabra to continue this work. Preliminary results are presented in Philosophical Transactions of the Royal Society of London vol. B, 260, 1971. A comprehensive discussion of the geology of Aldabra has been published recently (Braithwaite et al. 124 1973). The Fondation Singer-Polignac during the 1960's sponsored a number of expeditions to New Caledonia and French Polynesia, and maintained a research vessel in the area for this purpose. Results are published in Cahiers du Pacifique and in a series of beautifully illustrated Mémoires. In connection with the nuclear tests on Mururoa, Tuamotus, the French government sponsored an extensive series of inves- tigations in the Tuamotus, especially Mururoa, and in Clipperton Island. Reports are published in Cahiers du Pacifique and in a series of duplicated papers on Clipperton Island. In 1971, a small research station ("Antenne de Tahiti") of the Muséum National d'Histoire Naturelle, Paris, was created in Moorea, near Tahiti, to continue such research, and an impressive list of papers is already available, several of which are cited below (Chevalier, Denizot, Salvat, etc.) The marine station of Endoume-Marseille has for some years sponsored coral reef investigations at the marine station at Tulear, Malagasy, comprehensive reports have been published in special issues of the Recueil des Travaux series and later of Téthys. A general description of the reefs of Madagascar was presented by Pichon (1972). The University of Hawaii recently acquired a station on Fanning Island in the Line Islands and during 1969 staged a broad-scale NSF- sponsored investigation of the Fanning Islands reefs. The results have been pubtished in Pacific Science (April 1971) and in a special report issued by the Hawaii Institute of Geophysics (Chave, ed., 1970); this is a continuing program, and further results can be expected. Since 1968, a University of Hawaii Sea Grant project to study various quantitative aspects of reefs and reef biota in Kaneohe Bay, Oahu, has been underway. The first results of that program have been published (Se seSmich) Gta vallls.) 1973): The work of several other surveys is mentioned throughout the following sections though the present discussion is by no means exhaustive. PHYSICAL ENVIRONMENT Much of the oceanographic work carried out in the tropical seas in the past, and especially in the last 25 years, is a source of environmental data on coral reefs. One example is work on surface currents, of Significance in the distribution of reef and island biota. Specific research on atolls and reefs, however, has been pursued only by a small number of investigators and with a limited scope. Work at Bikini Atoll (von Arx, 1954; Munk and Sargent 1954) has delineated the circulation pattern within the lagoon that is established in response to prevailing winds and influenced by tides, waves, and local ocean currents. The Bikini study, as well as the research at Fanning Island ( Gallagher et.al., 1971), provides volumetric estimates of exchanges of water between lagoon and ocean. Fanning Island investigations also produced measurements of salt and heat fluxes. The waters of the Great Barrier Reef have been studied by Brandon (1973). These studies and others devoted to specific physical parameters on reefs can contribute guidelines for an ecosystem model, but the majority of oceanographic projects on reefs have not included coordinated studies of interactions of the physical environment 125 with the biological reef communities; see for instance the work of Van Dorn, and Vastano and Reid, on Wake Island. Atoll meteorology was mostly limited, prior to World War II, to sporadic weather records (see for instance those used by Sachet, 1957), observations on storms and hurricanes, and the like. With World War II action, and the atomic tests in the Pacific, as well as the increase in geophysical research generally, much more information in meteorology and climatology has become available. Weather data are more complete and collected from more stations. Summaries such as those of R. C. Taylor (1973) and Zipser and Taylor (1968), and volumes of data such as those prepared for U.S. Joint Task Force Seven, are a few examples among many, which supply important data in the study of atoll ecosystems. See also studies by Montgomery (1973) and Quinn and Burt (1970), and the very detailed work of Blumenstock and Rex (1960) on Eniwetok. Even more closely related to the ecosystem approach are papers describing hurricanes, typhoons and other storms, and their effects on reefs and islands. The work of Blumenstock and others (1958, 1961) on Jaluit, that of Stoddart on the British Honduras reefs (1963, 1969d), and the Solomon Is. (1973), of Sachet relative to Clipperton (1962b) are a few examples. More general work on hurricanes and other meteorological phenomena is included in the work of Ramage and others, in the U.S. Navy Atlases (1956-68) and many other useful sources. Atolls have also been used as “observation platforms" (Lavoie, 1963) in atmospheric and other studies, notably by the Hawaii Institute of Geophysics and the U.S. Joint Task Force Seven (e.g. McCreary 1959). GEOLOGY The geological record of fossil reefs has been the subject of exhaustive studies, especially by petroleum geologists,and some of the reviews and bibliographies cited above are concerned with fossil reef studies, including paleoecology. Those references will not be detailed here (but cf. Ladd, 1957; Laporte 1974). Suffice it to say that a vast amount of knowledge of fossil reefs has accumulated, from which studies of modern reefs can derive data as well as ideas (Hedgepeth, 1957). To a certain extent, of course, the fossil and modern reefs cannot be separated. The modern sections of the reef are considered here to be material deposited in relation to present sea levels, approximately during the last 5,000 years. Since the late 19th century, modern coral reefs have been studied by geologists primarily to provide information on pre-existing environ- mental conditions documented in the lithological record of the geological column. With few exceptions these studies have concentrated on descriptions of reef components rather than on the processes responsible for component formation. However, much attention has been paid to the influence of sea level changes on the nature of present-day reefs (Fairbridge 1958; cf. Stoddart 1971). 126 Studies of reef morphology (reviewed by Stoddart, 1969a) generally have included mapping bottom topography at intermediate scales (1:50,000) and describing topographical zonation and size of features in such terms as mean reef width, knoll dimensions, etc. (e.g. Emery et al. 1954). There have been few quantitative studies of reef features, apart from the work in Kaneohe Bay, Oahu, by Roy (1970a), at Fanning Island by Roy and Smith (1971), and in the Maldives and elsewhere by Stoddart (unpublished) . Many studies concern the mineralogy, composition, and texture of reef and near-reef sediments with a view to understanding sequences in the geological record (e.g. Ginsburg, 1956; Neumann, 1965, Macintyre, 1970). Recent trends have been to examine CaCO. budgets in areas of carbonate sedimentation (Stockman et al., 1967;-Neumann and Land, 1969; Land, 1970; S.V. Smith et al., 1970, 19713; S.V. Smith 197la, 19733; Chave et al., 1971). Such studies have only considered some (or even only one) of the components in the budget; or they have integrated the components of the budget across a very restricted environment; some have not even dealt with tropical reef systems. What is needed now is the generation of a total carbonate budget through time, in order to describe the relationships among reef development, sediment generation, sediment transport, and subsequent deposition. Little information is available on former zonations in various stages of reef development, yet work on specific problems associated with coral reefs has indicated that significant data may result from exposing internal reef structures: Shinn (1963) investigated the origin of spurs in reefs off Florida and subsequently (Ginsburg et al., 1967) studied marine cementation and internal sedimentation within reefs off Bermuda. To date, rather little coring has been attempted (Mayor, 1924; Cary, 1931; Ladd and Schlanger, 1960; Stoddart 1971) on modern coral reefs, with the result that the internal structures and limestone facies of these reefs are largely unknown. Most of this work has involved only one or a few core holes per reef. More detailed drilling with a submersible drill that can penetrate 2 meters into the reef is underway on reefs of Jamaica (L. Land, personal communication, 1970). It has been suggested that living coral reefs consist of a thin veneer of modern reef growth over older foundations which generally dictate their present-day morphology (Newell, 1962; Stoddart, 1969a, 1973). However, modern reef construction has been shown to form massive wave-resistant frameworks, notably in Jamaica (Goreau, 1961b, Goreau and Land, 1974) and in British Honduras (Purdy, 1974). Numerous workers have investigated growth rates (e.g. Vaughan, 1915; Mayor, 1924; Shinn, 1966) or calcium carbonate deposition rates (e.g., Kawaguti and Sakumoto, 1948; Goreau 1959; Goreau and Goreau, 1959; Goreau, 1961a) or general reef Cal, production rates (e.g. Chave et al. 1971; S.V. Smith et al., 1971, S.V.>Smith, 1973, Glynn et al. 1971), as well as skeletal framework destruction by mechanical (Stoddart, 1963; Glynn et al., 1965; Ball et al., 1967; Perkins and Enos, 1968) and biological means (Duerden, 1902; Otter, 1937; Yonge, 1963; Goreau and 127 Hartman, 1963; Bakus 1964, 1967; Neumann, 1966). However, concurrent investigations of most of these processes have not been carried out at a single reef site in order to determine the relative importance of the various parameters controlling the net accumulation of modern reef framework. The success of framework construction depends largely on the growth rate of reef organisms. Other factors affecting construction include predation, competition between corals and other organisms, variation in distribution of coral and other species, biological erosion of skeletal framework and of frame-cementing agents, mechanical erosion, and the rate of sea level change. Cement infilling and lithification of internal sediment in various cavities tend to negate the destructive action of coral borers. For example, some coral heads collected from submerged reefs in the eastern Caribbean have been almost completely altered--through repeated boring, infilling, and lithification--to a dense micritic limestone (Macintyre, 1972). Biota have long been thought to be the major frame-cementing agents, but in recent years extensive Mg-calcite cementation of reef framework and inner framework has been recognized (summarized in Bricker, 1971). The relation, if any, of biological agents to this cementation is not altogether clear. The origin of magnesian calcite cement remains unexplained, but chemical analysis of reef interstitial waters may offer some clues to the processes responsible for its precipitation. Within the reef ecosystem, interactions between waters and solid CaCO. appear to be largely the result of biological calcification processes, or iftteractions between biogenic rock, detritus, and waters separated from open ocean waters--e.g., the waters within the interstices of skeletal fragments or the reef frame itself. Direct chemical precipitation and solution reactions of CaCO, in seawater appear to be inhibited or prevented by interactions of d?ssolved Organic compounds in sea water with CaCO. mineral surfaces (Chave, 1965; Chave and Suess, 1967, 1970; Suess, 1970). Chemical reactions between seawater and carbonate minerals may occur, for example, in the formation of ooliths, grapestone, whitings, and beachrock; but these reactions are, at best, minor in a coral-reef ecosystem, if they are inorganic at all. Two interactions between reef waters and biogenic carbonates appear to be important when considering the flow of carbon in a reef ecosystem. First, recent reports indicate that chemical or biochemical precipitation of CaCO. within the reef is as important, or even more important, than biogenié calcification with respect to binding the components to form a rigid, wave-resistant framework (see Bricker, 1971). The second interaction between reef waters and biogenic carbonates is the removal of dissolved carbonate from interstitial waters of sediments and deposition of CaCO. within the sediment components (see Bricker, 1971). The importance of this interaction is not only that it represents a flow of carbon from seawater into the reef ecosystem but that the result of this flow directly affects the character and hydraulic properties of the sediment. 128 In order to understand fully the processes responsible for the precipitation of CaC0., both in the framework and within sediment grains, it is necessary to investigate the whole reef CaCO. system-- namely both water and rock chemistry. By studying the whote chemical system, it should be possible to determine whether the cementation process involves only an internal cycling of CaC0O., or whether significant » | amounts of Caco, are derived from external source’. The C0, system in seawater provides a unique link between geochemical and biologiéal processes. The link is particularly important in coral reefs, where both calcification and organic carbon production-consumption greatly affect the system. The organic carbon transfer should be mentioned; it is sufficient for this paper to point out that (as discussed in some detail by Park, 1969) measurement of two parameters in the marine CO, system is sufficient to partition the system changes into inorganic a cato, precipitation-solution and organic C production-respiration. 7 Separating changes in deep ocean CO, into solution and oxidation has been undertaken by several authors (8.9. Park, 1968; S.V. Smith, 1971a). Both S.V. Smith (1973) and Kinsey (unpub. data and personal comm.) have studied diel variability in the CO, system in coral reefs. 1 S.V. Smith (1973) has related this variability to organic carbon pro- 4a duction and utilization and to calcification. The present major limitation of these short-term studies is the small amount of change in the marine CO, system caused by inorganic chemical processes and organic metabolic aétivities of the reef communities. Kinsey's studies were performed on water which "aged" over several hours and consequently experienced easily measurable changes in the CO, system. | Smith's studies dealt with water "aged" less than one hour therefore showing barely measurable perturbations in the CO, system. BIOTIC ENVIRONMENT The biotic components of the surrounding sea should not be ignored. Perhaps most important is the plankton, which is a significant food supply of the corals and many other reef animals. This has been studied with . | relation to reef animals by A.R. Emory (1968). Small pelagic fish and fry are also food for reef-dwelling carnivorous fish, as well as for sea-birds. Predaceous pelagic fish visit the reefs and prey upon the reef fishes. The same is true of porpoises, other small cetaceans and pinnipeds. Detritus of organic origin brought to the reefs by water movement is also an important input into the reef system (N. Marshall 1965). All these biotic components are discussed in more detail below. TERRESTRIAL ECOSYSTEM To the best of our knowledge, the only attempt at a generalization Wag) of the functioning of the terrestrial coral atoll ecosystem is the description written in 1957 by Fosberg (1961, 1963b). That description attempts to conceptualize the system in abstract, non-quantitative, non-mathematical terms, and to indicate functional groups of entities in the system which can be studied in terms of their relationships to the whole. This work was not followed up except for extending it to high islands. It resulted logically from the general program for atoll studies proposed at two coral atoll research symposia sponsored in 1951 by the Pacific Science Board (cf. papers in Atoll Research Bulletins 1, 2, 1951, Fosberg ed.). Five expeditions planned to provide comparable descriptive and inventory data on atolls of different types were carried out (to Arno, Onotoa, Raroia, Kapingamarangi and Ifaluk). Their results, as well as those of many other studies of land (and marine) aspects of atolls were published over the next 20 years in the Atoll Research Bulletin and elsewhere. An attempt at bringing together this great mass of data was made by Wiens (1962), but it was not entirely successful in representing the world atolls as an ecosystem. Other qualitative descriptions of the island ecosystem are included in Numata (1967) especially those papers by Jackson, and Sachet. See also Stoddart, 1969c. An idea of the enormous amount of basic information available on the terrestrial aspects of coral atolls may be gained by perusing the 261 pages devoted to this subject in Island Bibliographies and 236 in its supplement (Sachet and Fosberg 1955, 1971). From this wealth of data, and citing only a few specific references, we can derive a fair idea of components and processes involved in the terrestrial or cay ecosystem and influences on the adjacent submerged reef ecosystems. Some components and processes, such as nutrient content of ground water, salt spray, rain water, animal and plant bodies, and sediments, have never been quantified but can be measured by standard analytical procedures. Standing crops of the macroscopic organisms present no problem, but again, there are no quantitative data for coral islands. Estimates can be reasonably made by simple sampling, counting, and weighing procedures. Imports of organic matter (fish, squid, etc.) by seabirds and shore and wading birds into the terrestrial atoll ecosystem have never been estimated. Some observations have been made on the ground water of atolls, beginning with Charles Darwin (1839), but mostly within the last few years (Fosberg, 1959; Cox, 1951; Arnow, 1954, 1955; Tracey et al., 1961). The general nature of the atoll ground water lens has been established, but detailed studies of its behavior under varying conditions of geological structure and rainfall amounts and regimes, as well as of tidal ranges and regimes, are still required. The terrestrial geology of certain atolls has been studied by a number of expeditions and individuals (e.g. Royal Society, 1904; Wentworth, 1931; Fosberg et al., 1956, Newell, 1956; Fosberg, 1957a; McKee, 1958, 1959; Fosberg and Carroll, 1965; Sachet 1962b; Stoddart, 1962, 1969b). Very few quantitative measurements of sediments are available. Fairly detailed soil studies are available for a few Pacific atolls (Stone, 1951; Fosberg, 1954; Fosberg et al., 1956; Fosberg and Carroll, 130 1965; Tercinier 1956, 1969) as well as a general discussion of atoll soils (Stone, 1953). Most of the atoll soil types are very widespread and : the patterns rather simple. There has been no systematic investigation to determine if Caribbean atoll soil types correspond to those in the Pacific. The occurrence and origin of atoll phosphate rock in the central Pacific atolls were elucidated by Fosberg(1957b); and further observations have been reported by Stone (1953), Fosberg et al. (1956), Niering (1961, 1963), and Roy (1970b). Similar rock was described for the Indian Ocean atolls by Piggott (1968) and recently was found on Glover's Reef, British Honduras by Fosberg. Hutchinson (1950) deals in great detail with background information on phosphate accumulation. Substantial information is available on atoll floras and vegetation (Fosberg 1949; Hatheway, 1953, 1955; Fosberg, 1953; Fosberg et al. 1956; Fosberg, 1957; Sachet 1962a; Stoddart, 1962; etc.) Pure stands of certain trees and shrubs are frequently found, an unusual occurrence in the tropics. The land animals of coral islands are reasonably wel] known but information, except for that on birds, is mostly scattered in monographs and papers on the groups concerned. A few papers specifically on atoll faunas are found in. the Atoll Research Bulletin series and in the reports of the British Indian Ocean expeditions (Percy Sladen, etc.) of the early twentieth century. There is an extensive literature on atoll birds (listed and annotated by Sachet and Fosberg 1955, 1971). The atoll bird data have never been reviewed as a whole, but the Smithsonian field guides (Watson et al. 1963; King, 1967) and a number of regional papers (Amerson, 1959; Baker, 1951; Pelzl, ms.) are steps in this direction. Atoll insects, as well as the insects of high islands, are discussed in the Pacific Insects and Insects of Micronesia series (Gressitt, ed.). Papers that deal with the interactions of atoll faunas are available for Arno atoll (Marshall, 1951) and the Tokelaus (Hinckley, 1969). Data on the consequences to the rest of the atoll ecosystem of the presence of large numbers of seabirds are brought together by Hutchinson (1950). No work has been done on either terrestrial primary productivity or nitrogen fixation on atolls, nor is there much information on terrestrial food chains, predation or decomposition of organic matter under atoll conditions. Extrapolations may be made from other ecosystems, but it would be important to know just how well these extrapolations correspond to actual atoll patterns. HUMAN INFLUENCES The complex of human influences on and interactions with the cay ecosystem and the total reef ecosystem is quite apparent but difficult to characterize and summarize satisfactorily (Johannes 1970-71). The pesticide or, better, biocide, component is of general importance and is very probably pervasive in all parts of the system. This particular problem is dealt with specifically below. The nature of changes brought 131 about by human activity varies from obvious (Stoddart 1968a) and easily measured to very obscure and hard to estimate, but the importance of these changes to the system may not necessarily be proportional to their obviousness. Pollution in terms of addition of human wastes (sewage, etc.) can be estimated from standard figures and tables. The amounts and effects of solid-waste pollution are probably in proportion to amount of human activity on the cays and within the lagoon, with a substantial addition from drift materials washed in from the open ocean. An FAO conference on Marine Pollution (Ruivo, ed., 1972) included several accounts of pollution of reefs, atolls and lagoon (see especially Johannes, Chan, Bagnis). Increased sedimentation is another frequent result of man's activities, as exemplified by Kaneohe Bay, Oahu (Roy, 1970a). This latter problem is likely to be more severe on high islands than on atolls. One of the most significant effects of human activity is the destruction and/or alteration of habitats of other organisms (Sachet, 1963; Jackson, 1967). The replacement of natural forest by coconut plantation is a good example (cf. Stoddart, 1968). Except for simple measurement of area, we know of no way to measure such changes or their effects, or, in many cases, even to assign specific effects to particular causes. This does not in any way minimize their importance. A volume edited by Fosberg (1963a), deals with islands including atolls, with special reference to man's role in their ecology. The presence of a marine laboratory in Kaneohe Bay, Oahu, Hawaii, has led to a large number of studies concerning various aspects of human effects on the barrier and fringing reef complex there. Among the papers dealing with that bay are Bathen (1968); Roy (1970a); Smith et al. (1970, 1973); Smith (1971b); Caperon et al. (1971); Clutter (1972); and Johannes (1970,1972). These papers have dealt primarily with the high nutrient level and rapid deposition rate in the bay. Pollution of the biosphere by man-made chemical compounds has reached a level such that all faunal elements of the earth are contaminated (Risebrough et al., 1970). These compounds are of two types: 1) biocides (primarily chlorinated hydrocarbons) manufactured and distributed specifically to eradicate or control "pest" organisms; and 2) chemicals manufactured for and used in industrial processes which "escape" from their intended area of use (primarily polychlorinated biphenyls and heavy metals). The accumulated literature on the distribution of these com- pounds is extensive but only recently have their physiological effects On metabolic processes begun to be elucidated. New information on induction or inhibition of various enzyme systems and chemical reactions by man-made chemicals becomes available with each edition of the pertinent journals. These physiological effects are especially relevant to marine organisms, since accumulation and biological con- centration occur most readily in aquatic systems. While biocides have been detected by all studies thus far designed to investigate their presence or absence, few such studies have been carried out on coral atolls; and little investigation has been made of 132 biocide accumulation in any level of the food web of a coral atoll. To our knowledge, the only non-marine data available for a tropical island are in unpubl ished information on Sooty Tern eggs from the Dry Tortugas, Florida, in which DDE and PCBs were present in all samples (W. B. Robertson, Jr. pers. comm.). By an accident of geography, atolls have been the most numerous sites of atomic and nuclear atmospheric bomb-tests and, inevitably, the sites of pollution by radionuclides. Between 1946 and 1958, more than 59 tests took place at Eniwetok and Bikini atolls (Welander, 1969). Later Christmas and Johnston Islands were involved, and more recently, Mururoa and Fangataufa Atolls in the Tuamotus. In the Marshall Islands, a survey of the ecosystem had been carried out prior to any tests (Ladd, 1973), and periodic resurveys followed. They gave rise to an enormous literature much of it in classified or hard-to-get AEC reports. Only after many years did papers appear in scientific journals of more general distribution, as well as in proceedings of conferences and symposia (see for instance papers by Beasley, Beasley and others, Held, Held and others, Nelson and Evans, Templeton et al, and many others). Such information has not yet been integrated as a whole picture of radionuclide pollution on coral reefs and atolls, but much data continues to be accumulated. There is much less easily available information on Johnston Island, Christmas Island (Palumbo et al., 1966) or the Tuamotus. In the Marshalls, as the result of a catastrophic incident (Operation Castle, 1954) the effects of exposure to radiation on atoll human population became available for study. Detailed medical surveys of the exposed islanders have been carried out and repeated at regular intervals. See for example papers by Conard et al., Robbins et al. and Lisco and Conard. The sociological effects of the Castle disaster, as well as those of the displacement of the Eniwetok and Bikini populations are other aspects of human interference with the coral island ecosystem (Stoddart 1968a). Many others, the impact of war on atolls, of organized migration, of changes in economic patterns, could be mentioned, but cannot be detailed here. This type of information is beyond the scope of this review, but it is obvious that anthropology and history can furnish evidence relevant to the total picture of coral island ecosystems. MARINE BIOTA: BENTHIC PLANTS Benthic marine algae make major contributions to primary pro- ductivity, nitrogen fixation, community structure, organism distri- bution, carbonate production, and reef consolidation and destruction. Setchell (1926, 1928) was probably the first biologist to recognize fully the variety and importance of these roles of algae in coral reefs, and much work has followed his pioneer effort. Setchel] (1928) described zonation of algae across the reef, as have Kanda (1944), Doty and Morrison (1954), Gilmartin (1960), Doty 133 (1967, 1970) and Tsuda (1970). Seasonal occurrence has been described by Bernatowicz (1952) and Denizot (1969), and measured on Guam by Tsuda (1972), and Dahl (1972) has analyzed community structure of Samoan algae. Dahl (1971) has also demonstrated that certain benthic algae are useful as ecological indicators and can even provide a continuous record of environmental conditions. Studies on temperate and subtropical algae have demonstrated their importance in ecological investigations and have developed techniques for surveying field populations. Line transects can provide a useful basis for describing community structure and measuring seasonal variations, as in the work by Neushul (1967) on subtidal vegetation in western Washington, and the recent survey of subtidal ecology off southern California (Neushul et al., 1967; Clarke and Neushul, 1967). Experimental techniques can add considerably to the field data on community composition and ecology (Neushul and Dahl, 1967). Crustose coralline algae (Melobesioidae) may be at least as important as corals in the development of reef structures (Setchell, 1926; J.H. Johnson, 1961; Gross et al., 1969). These algae have been especially noted in the Pacific, where they have given their name to the 'Lithothamnion ridge," a striking but misnamed topographic feature of Pacific atolls; however, the importance of coralline algae to reef construction in general is probably greater than the extant reef literature would indicate (Denizot, 1972). In many areas of the Pacific (e.g. Ladd ét al., 1970; J.H. Johnson 1961) as well as in the Tethys Seaway (e.g. Lemoine, 1939; J.H. Johnson, 1965) and even in the Atlantic (Iams, 1969) encrusting coralline algae have been important or even the primary contributors to sedimentary formations. Considering the widespread occurrence of encrusting coralline algae, their importance in determining the nature of the substratum-water interface, and their presence in the Fossil record, they have been rather neglected. They are considered to be a difficult group (see for example W.R. Taylor 1960; Adey 1970) and have often been treated as part of the "dead" substrate. Adey has underway a detailed study of the role of coralline algae in the reef ecosystem. Calcareous green algae (mostly Halimeda spp.) contribute greatly to loose sediments in certain areas of the reef, especially certain zones of lagoons (Chapman, 1901; Emery et al., 1954; Hoskin, 1963). Stockman et al. (1967) and Neumann and Land (1969) have also shown the importance of the genus Penicillus in the formation of lime muds in Florida and the Bahamas. Land (1971) has studied the importance of sediment production by Melobesia, a red alga which commonly encrusts sea grasses. The roles of boring algae and microbenthic algae have yet to be delineated in respect to calcium carbonate breakdown though Nestéroff (1956) and others ascribe to them a major responsibility for intertidal erosion. 134 ’ Some attempts at including crustose coralline algae in regional studies of calcium carbonate budgets (S.V. Smith et_al., 1970, 1972a) and in investigations of primary productivity (Marsh, 1970; Littler, 1971 have been undertaken. The results are preliminary, however, as there is likely a variation by several orders of magnitude in growth and metabolic rates, as a function of light, temperature, species, and even part of an individual plant considered (Adey and McKibbin, 1970). Some preliminary productivity measurements have also been made on reef algae by Doty (1971) and Soegiarto (1972). Recent studies have 4 indeed shown that the alga-covered reef flat, a visually unimpressive y and often ignored component of the reef community (Dahl, 1972), was f in fact, twice as productive as areas of rich coral cover and supported large roving populations of herbivorous fishes (Johannes q et _all.,. 1972): f Symbiosis, or the occurrence of zooxanthellae within living coral tissues has long been known, but only recently has their role q as nutrient acceptors been demonstrated (Goreau and Goreau, 1960; Muscatine and Hand, 1958). The zooxanthellae also play a significant role in the calcification of some corals (Goreau, 1961a, Goreau and | Goreau, 1960). There is still considerable debate over the inter- action between plant and animal components. Recent work has only begun to explore the variety of marine symbiotic relationships (see for example, D.L. Taylor 1969a, 1969b, 1971). Information on grazing of algae by reef herbivores includes general observations on predation upon algae via studies of gut contents and results from caging; herbivore densities have not been reported beyond presence or absence, nor have the plant communities in which the experiments were carried out been rigorously defined (Randall, 1961, 1965; Mathiesen et al., 1972; Earle 1972). The slate-pencil urchin, Heterocentrotus mammillatus, has been observed to feed heavily on Porolithon sp. on the algal ridge of Bikar Atoll in the Marshall Islands, and to have a significant influence in the erosion of the reef edge (Fosberg, pers. comm.). aCe Ole NE ee Te Se "Sea grasses" are locally important in tropical nearshore environments. In his monograph, den Hartog (1970) summarizes under each species what is known of its biology, its role as food for herbivores (e.g. Randall, 1965), and its function as a substratum Stabilizer. Recently, the echinoid Diadema has been shown to graze heavily on Thalassia in certain situations in the Caribbean (Ogden et al., 1973). There have been rather few quantitative studies on the productivity of turtle grass (Thalassia testudinum), one of the more abundant and the best known species. Early studies were made % by Pomeroy (1960), Odum (1957), and Jones (1968) on productivity a measured by the 0, method. However, closer inspection by Zieman ‘ (1968) has shown ehat this method is suspect for Thalassia produc- tivity, as the leaves have the capacity to expand and store gasses in interstitial lacunae. This phenomenon has also been demonstrated by Hartman and Brown (1967) for the fresh-water species Elodea canadensis and Ceratophyllum dimersum. Pe Me sme yn) 135 INVERTEBRATES It is impossible here to discuss the individual roles of the numerous groups of invertebrates in the reef ecosystem, or the enormous amount of available work basic to an integrated study of the system. Many pertinent papers on invertebrates have appeared in Cahiers du Pacifique, which also include bibliographies of certain groups, and in recent Symposium volumes cited pp. 1-2. This review will only touch on a few highlights of recent research on invertebrates in their functions in the system. Since coral reefs are almost entirely the result of biological activity, an analysis of reef development may be approached by examining structural activities (construction, maintenance, and destruction) and the contribution of the various biota involved (calcification by hermatypes, binding and baffling properties of some soft-bodied species, and biodegradation by a variety of invertebrates). Work on growth rates or calcium carbonate deposition rates has been mentioned in previous sections. The minimal energy requirements of corals were investigated by Coles (1969), who concluded that three common reef species are capable of capturing and ingesting sufficient zooplankton to account for daily maintenance under laboratory conditions. Yet it was determined from a field study in Bermuda that the energy needs of corals were of an order of magnitude greater than could be met by the supply of drifting net zooplankton (Johannes et al., 1970). Obviously these metabolic Studies are not conclusive Further studies of coral metabolism should include examination of coral feeding and digestive mechanisms and the relation of these mechanisms to coral morphology and food type. Hartman and Goreau (1970) have called attention to the sclero- sponges aS an important constructional element besides scleractinian framebuilders in the sub-reef and deeper reef framework. It is also clear that many benthic reef invertebrates play an important role in maintenance processes. An example is that of the sponge Mycale laevis, which protects the lower surfaces of some massive corals weenie destructive effects of boring sponges (Goreau and Hartman, 966). Although corals are commonly the principal animal contributors of skeletal materials to the framework and loose sediments that make up the reef, several other invertebrate groups can make substantial additions (especially mollusks, echinoderms, and Foraminifera) . The latter, in addition to their commonly recognized role as stratigraphic markers, are frequently a prominent component of loose sediments. Some entire atoll beaches in the central Pacific are made up of the worn tests of Calcarina and Baculogypsina, two genera that inhabit windward reef flats. Hometrema forms conspicuous dark red crusts on the coral fragments seen on Glover's Reef and other atolls. 136 Skeletal framework destruction by biological means has been discussed above. Some processes are direct and obvious, such as the feeding of fishes, crabs, worms, snails, etc. on living corals. Notable also is the recent population explosion of Acanthaster planci which feeds upon and locally devastates reefs in Pacific areas (Antonius 197la, Chesher, 1969; Newman 1970; Randall 1972; Atoll Res. Bull. 166-170, 1973; Micronesica, 9(2), Dec. 1973; and numerous other recent papers). Feeding of fishes on invertebrates, e.g. corals and sponges, was studied by Randall (1967), Randall and Hartman (1968), and Bakus (1967). Many endolithic boring and burrowing forms are also known (e.g. sponges: Goreau and Hartman 4 1963, and Ruetzler, 1971; sipunculids: Rice, 1969; mollusks: R. Robertson, : 1970, and rates of destruction have been calculated for some of them (Neumann, 1966; Glynn et al. 1971). Interactions involving invertebrates have been summarized for Western Atlantic reefs by Glynn (in press). Lang (1970, 1971) obtained information on the interspecific aggressive behavior of : hermatypic corals, representing a feedback loop within a single § functional component. Research on the role of small interstitial animals in the coral reef ecosystem has lagged behind other areas of study. Preliminary sampling has indicated, for example, that groups such as Turbellaria, ‘ Nematoda, Polychaeta, and Crustacea are abundant in reef sediments, ‘ raising the possibility that they may provide a significant food resource for deposit feeders and possibly grazers and browsers (Renaud-Mornant, et al., 1971; Thomassin 1972; papers presented at the q "Floating Symposium" (see p.2) by Thomassin, J.F. Grassle and other } participants). FISH (AND OTHER VERTEBRATES) The place of fishes in the budget of the reef ecosystem seems to have been elucidated perhaps in more detail than that of most other vertebrate groups. Except for the green turtle, the biology of which is being studied in various parts of the tropics (e.g. Frazier ms.; Ehrenfeld 1974), little consideration has been given to such other vertebrates as seals or sea-snakes, which are only locally significant. Sea birds have already been mentioned with respect to the terrestrial part of the ecosystem. Estimates of ‘fish biomass have been made by Odum and Odum, 1955 (425 kg/hect.); Bardach, 1959 (450 kg/hect.); Randall, 1963b (1590 kg/hect.) Brock, 1954 (1850 kg/hect.); and Goldman and Talbot, in press (200- 2100 kg/hect. from different reef areas). These widely different estimates may be due in part to different sampling techniques and in part to the variation that is now known to occur between different parts of a single reef. As yet, there is no good information on differences between widely separated reefs, particularly from ocean to ocean. Some of the factors in fish biology which affect biomass may be detailed as follows: 137 Habitat selectivity is marked in reef fishes. Distribution is obviously affected by depth, shelter, food availability and competitive interactions. Various areas of the reef have different specificity in many zones. Up to 25 per cent of species may be restricted to single areas of the reef (Talbot, unpublished information) . From the work done on the movements of fishes on coral reefs (Springer and McErlean, 1962; Randall, 1961, 1962, 1963a; Bardach, 1958; C.L. Smith and Tyler, 1972; Moe, 1969; Winn and Bardach, 1960; Reese, 1964) it seems that the majority of reef fishes are restricted to a reef or even individual patch reefs, at least for long periods. Many species are territorial. Some few pelagic species such as some scombrids, carangids, and sharks, apparently move between reefs (Tester and Wass, personal communication). These species feed mostly on reef fishes. Within their territories, many fishes have different feeding and resting areas and may move up toamile or even more on daily feeding migrations (Winn and Bardach, 1960; Randall and Randall, 1963; Randall, 1963; Bardach and Menzel, 1957). Some fishes may have daily breeding migrations. Others may rest in caves down the deep reef front and feed on the reef flat or rest on patch reefs by day and feed over Thalassia beds at night. Some diel changes in distribution have been documented by Hobson (1965, 1968, 1972), Starck and Davis (1966), and Collette and Talbot (1972). The food habits of West Indian and Pacific coral fishes have been studied by a number of workers, and this information forms a solid qualitative basis for further study (Hiatt and Strasburg, 1960, Suyehiro, 1962, Randall, 1967; Plessis 1972; Starck, Emery, personal communications). Few data on feeding efficiency and conversion are available for coral reef fishes, but a fair amount is known about metabolic rates in temperate freshwater and marine fishes. It appears that conversion factors could be used to calculate the metabolism of coral reef fishes. Some further work is needed, however, on selected species of different sizes, activity patterns, and feeding habits for accurate ecosystem modeling. Most investigators who have worked on the problem of age and growth rates have found that conventional techniques are not always applicable to tropical fishes. Recent work (Moe, 1969) has shown, however, thatotoliths can be used for some species, and size-frequency techniques as well as aging population dynamics may yield data for representative species (Gulland, 1970). Relatively little is known of the reproductive biology of coral reef fishes (Breder and Rosen, 1966). Fishes exhibit a wide variety of reproductive mechanisms, from extreme parental care (e.g. mouth breeding) to random scattering of eggs. A few species have, been shown to undergo daily or annual breeding migrations (Randall and Randall, 1963; Reinboth, 1973; C.L. Smith and Tyler 1972). Sex ratios are greatly variable, and several types of hermaphroditism are common 138 Fecundity, recruitment, and egg and larvae losses are perhaps most poorly known. Reef fishes can also play a role in reef destruction. Bardach (1961) has documented the role of destruction by fishes on Bermudian reefs, while Glynn et al. (1971) have done likewise for fishes on the Pacific reefs of Panama. Grazing by parrot-fishes and some other reef fishes is commonly observed on reefs in all areas. PLANKTON At present, the quantitative contribution of phyto- and zooplankton to the reef ecosystem is unknown. Estimations of the standing crop of zooplankton around coral atolls and in the lagoon areas can be found in M.W. Johnson (1949, 1954), Gilmartin (1958), Mahnken (1966), Barnett (ms.), 0.A. Mathisen (1964), Odum and Odum (1955), and A.R. Emery (1968). Most investigators have found that the biomass of zooplankton in the vicinities of reefs and atolls and inside coral reef lagoons is higher than the biomass in the open ocean at the same latitudes, yet the reef zooplankton biomass has probably been underestimated due to the inade- quate sampling of epibenthic (hovering above the water-bottom interface or within the coral heads) and neustonic (near surface) forms. Present estimates of zooplankton biomass, however, are orders of magnitude (Odum and Odum, 1955) smaller than estimates of benthic organism and fish biomass. The standing stock of phytoplankton has been expressed by chlorophy11-A values (N. Marshall), but conversion of these values into carbon is unrealistic without further analysis. The literature does contain information on such necessary inputs to tropical phytoplankton as light, nutrients, and trace metals (Strickland, 1960; Jeffrey, 1968). One common component of the coral reef water column, Trichodesmium, has been studied extensively (Prabhu et al., 1966; Calef and Grice, 1966; Goering et al., 1966; Ramamurthy and Seshadri, 1966a, 1966b; Ramamurthy and Krishnamurthy, 1967). It is obvious that many reef organisms feed on zooplankton. Quantity of plankton utilized by one reef assemblage in Puerto Rico was determined by Glynn (ms). Glynn (in press and ms.) has also given quantitative data on feeding activity of fishes and invertebrates on zooplankton as well as data on zooplankton recruitment (Glynn, in press). NUTRIENTS AND DETRITUS Studies of total community metabolism on atoll reef flats situated in largely unidirectional currents were initially developed by Sargeant and Austin (1949, 1954). Their method has been applied in a number of later studies (e.g. Odum and Odum, 1955; Kohn and Helfrich, 1957; Gordon and Kelly, 1962; Milliman and Mahnken, 1961; Qasim and Sankaranarayanan, 1970; Odum et al., 1959). The method was first used in a comprehensive 139 manner at Eniwetok in 1971 during Project Symbios (Johannes et al., 1972). That project proved without doubt the efficacy of this approach; for instance, the symbios studies explained why reef communities are so productive biologically though bathed in waters very low in plant nutrients. Nitrogen fixation and unusually efficient recycling of phosphorus within the community are apparently responsible. These two phenomena, therefore, Should now be focused upon at the subcommunity and species levels. Random Searching, species by species, would probably not have revealed the great quantitative significance of these processes. Net reef photosynthesis and total nighttime respiration can be determined by measuring the increase in the oxygen concentration in the water as it crosses the reef during the day, and its decrease at night. Another method which can help monitor community productivity and respi- ration uses changes in the CO, system (S.V. Smith, 1972). The system was analyzed according to somé of the calculations outlined by Park (1969). This approach has a number of advantages (cf. S.V. Smith, 1973; Smith and Marsh, 1973; Kanwisher, 1963). Still another approach to community metabolism consists of measure- ments made in fenced enclosures with the water surface open to the atmos- phere; it has been successfully used to a depth of 1 m on the Great Barrier Reef by Kinsey (1972, and unpublished observations). The reef imports organic matter from upstream in the form of zoo- plankton and exports it downstream in the form of mucus aggregates and algal detritus (N. Marshall, 1965; Johannes, 1967, Johannes et al., 1972). This transfer is likely to be of great significance in lagoon ecology and potential aquaculture. A curious result of recent studies at Eniwetok (Pilson and Betzer, 1973) has been to demonstrate that levels of dissolved phosphorus change remarkably little across the reef. The mechanism by which this constancy is maintained is by no means clear as yet. It may be that organisms paving the reef constitute a significant phosphorus sink and that exchange between this sink and the overlying water constitutes a mechanism for buffering phosphorus levels in the water. Such a mechanism has been demonstrated between sediments and the overlying water in salt marshes by Pomeroy et al. (1965, 1967, 1971). ECOSYSTEM ANALYSIS Interest in a mathematical representation of ecological phenomena developed early in connection with the dynamics of populations and epidemics. Names such as Malthus, Pearl, Merhulst, Lotka, and Volterra will always be cited in any history of mathematical ecology. At the present time, much active research centers on extensions and elaborations of the original Lotka-Volterra competition equations. This approach is particularly popular in population ecology. In community and ecosystem ecology, the dominant models currently 140 used are compartment models. They deal with storages and flows of energy and materials in systems and have been usefully complementary to chemical and radiobiological techniques developed and utilized in ecology during the last twenty years. They will continue to be important, despite suggestions that ecologists expand their mathematical horizons (e.g. Clymer, 1972) precisely because of this compatibility with current and prospective experimental methodology. The current status of systems modeling in ecology is summarized by Patten (1971, 1972). A model from a geological view point is presented by Macintyre et al., 1974. The International Biological Program (IBP) has been instrumental in accelerating the pace of development of total ecosystem modeling (in particular with the Analysis of Ecosystems Project of the U.S. IBP). All the Biome Programs (Grassland, Deciduous Forest, Desert, Tundra, Coniferous Forest, etc.) are committed to some form of mathematical ecosystem analysis. Progress to date has been variable, and the degree of emphasis also differs from program to program. The Grassland Biome Program is explicitly engaged in total ecosystem modeling (cf. Bledsoe et al., 1971). Others (e.g. Deciduous Forest, Desert) are emphasizing process models as potential modules for eventual total ecosystem models. The general philosophy and methodology of the ecosystem analysis approach as developed for reefs at the Glover's Reef workshop are described elsewhere in this issue by Dahl et al. REFERENCES Adey, W. H. 1970. A revision of the Foslie crustose coralline herbarium, including a preliminary world phytogeographic survey. D. Klg. Norsk Vid. Selsk. Skr. 1970(1):1-46. Adey, W. H. and D. L. McKibbin. 1970. Studies of the Maerl species ... in the Ria de Vigo. Botanica Marina, 13:100-106. Agassiz, A. 1888. Three cruises of the ... Blake. 2 vols., Boston and New York. moon ----- 1892. Reports on the dredging operations ... by the U.S. Fish Commission Steamer "Albatross" ... II. General sketch of the expedition of the"Albatross", from February to May, 1891. Bull. Mus. Comp. Zool. Harvard, 23:1-89. ---------- 1894. A reconnaissance of the Bahamas and of the elevated reefs of Cuba. Bull. Mus. Comp. Zool. Harvard, 26(1):1-203. ---------- 1899. The islands and coral reefs of Fiji. Bull. Mus. Comp. Zool. Harvard, 33:1-67. <<-====ne 1903a. The coral reefs of the tropical Pacific. Mem. Mus. Comp. Zool. Harvard, 28:1-410. 141 Agassiz, A. 1903b. The coral reefs of the Maldives. Mem. Mus. Comp. Zool. Harvard, 29:1-168. ---------- 1905. Three letters from Alexander Agassiz to the Hon. George M. Bowers ... of the ... "Albatross." Bull. Mus. Comp. Zool. Harvard, 46:63-84. ---------- 1906a. Reports on the scientific results of the expedition ‘ by the U.S. Fish Commission steamer "Albatross" .... October 1904, to March 1905 ... 5. General report of the expedition. Mem. Mus. Comp. Zool. Harvard, 33:1-75. alata 1906b. The eastern tropical Pacific. Mem. Mus. Comp. Zool. Harvard, 33:1-75. Amerson, A. B., Jr. 1969. Ornithology of the Marshall and Gilbert Islands. Atoll Res. Bull., 127:1-348. Antonius, A. 1971la. The Acanthaster problem in the Pacific (Echinoder- mata). Int. Revue ges. Hydrobiol., 56:283-319. ---------- 1971b. Occurrence and distribution of stony corals (Anthozoa and Hydrozoa) in the vicinity of Santa Marta, Colombia. Mitt. Inst. Colombo-Aleman Invest. Cient. 6:89-103. Arnow, T. 1954. The hydrology of the northern Marshall Islands. Atoll Res. Bull., 30:1-7. wocecc---- 1955. The hydrology of Ifaluk Atoll, Western Caroline Islands. Atoll Res. Bull., 44:1-15. Bagnis, R. A. 1972. Ciguatera et intervention humaine sur les éco- systemes coralliens en Polynésie frangaise, in: M. Ruivo, ed., Marine pollution and sea life, pp. 597-600, London, Fishing News (Books) Ltd. Baker, R. H. 1951. The avifauna of Micronesia, its origin, evolution, and distribution. Univ. Kansas Publ. Mus. Nat. Hist., 3(1):1-359. Bakus, G. J. 1964. The effects of fish-grazing on invertebrate evolution in shallow tropical waters. Allan Hancock Found., Publ. Occas. Papers 27:1-29. SSSSSSSS55 1967. The feeding habits of fishes and primary production at Eniwetok, Marshall Islands. Micronesica, 3:135-149. Ball, M. M., E. A. Shinn and K. W. Stockman. 1967. The geologic effects of hurricane Donna in South Florida. J. Geol., 75:583-597. Banner, A. H. 1968. A fresh water "kill" on the coral reefs of Hawaii. Univ. of Hawaii Inst. of Marine Biol. Tech. Rept. No. 15, 29 pp. 142 Banner, A. H. and Bailey, J. H. 1970. Effects of urban pollution upon a coral reef system: a preliminary report. Univ. Hawaii Inst. Marine Biol., Tech. Rept., 25:1-66. Bardach, J. E. 1958. On the movements of certain Bermuda reef fishes. Ecology, 39(1):139-146. ---------- 1959. The summer standing crop of fish on a shallow Bermuda reef. Limnol. and Oceanogr., 4(1):77-85. ---------- 1961. Transport of calcareous fragments by reef fishes. Science, 133(3446) :98-99. Bardach, J. E. and D. W. Menzel. 1957. Field and laboratory observa- tions on the growth of some Bermuda reef fisheries. Proc. Gulf and Carib. Fisher. Inst. Ninth Ann. Sess., pp. 106-112. Bathen, K. H. 1968. A descriptive study of the physical oceanography of Kaneohe Bay, Oahu, Hawaii. Univ. of Hawaii--Inst. Mar. Biol. Tech. Rept. No. 14. Beasley, T. M. 1969. Lead-210 production by nuclear devices: 1946-1958. Nature, 224(5219):573. Beasley, T. M. and E. E. Held. 1969. Nickel-63 in marine and terrestrial biota, soil, and sediment. Science, 164:1161-1163. ---------- and ---------- 1971. Silver-108m in biota and sediments at Bikini and Eniwetok atolls. Nature, 230(5294) :450-451. Beasley, T. M., E. E. Held and R. A. Conard. 1972. Iron-55 in Rongelap people, fish, and soils. Health Physics, 22:245-250. Bernatowicz, A. J. 1952. Seasonal aspects of the Bermuda algal flora. Contr. Dept. Bot. Univ. Mich., 924:3-8. Bledsoe, L. J., R. C. Francis, G. L. Swartzman, and J. D. Gustafson. 1971. PWNEE: A grassland ecosystem model. U. S. IBP Grasslands Biome Tech. Rept. No. 64. Bloom, A. L. 1974. Geomorphology of reef complexes, in: Laporte, pp. 1-8. Blumberg, B. S. and R. A. Conard. 1961. A note on the vegetation of the northern islets of Rongelap Atoll, Marshall Islands, March 1959. Atoll Res. Bull., 84:4-5. Blumenstock, D. I. 1958. Typhoon effects at Jaluit Atoll in the Marshal] Islands. Nature, 182:1267-1269. wooo ----- » ed. 1961. A report on typhoon effects upon Jaluit Atoll. Atoll Res. Bull., 75:1-105. Blumenstock, D. I., F. R. Fosberg, and C. G. Johnson. 1961. The re-survey of typhoon effects on Jaluit Atoll in the Marshal] Islands. Nature, 189:618-620. 143 Blumenstock, D. I. and D. F. Rex. 1960. Microclimatic observations at Eniwetok. Atoll Res. Bull., 71:1-158. Braithwaite, C. J. R., J. D. Taylor and W. J. Kennedy. 1973. The evolution of an atoll: the depositional and erosional history of Aldabra. Phil. Trans. Roy. Soc. London B. Biol. Sci., 266(878) :307-340. Brandon, D. E. 1973. Waters of the Great Barrier Reef Province, in: Jones and Endean, pp. 187-232. Breder, C. M., and D. Rosen. 1966. Modes of reproduction in fishes. Am. Mus. Nat. Hist., The Natural History Press. 941 pp. Bricker, 0.P., ed. 1971. Carbonate cements. The Johns Hopkins Press, Baltimore, Md. 320 pp. Brock, V. E. 1954. A preliminary report on a method of estimating reef fish populations. Jour. Wildlife Management, 18:297-308. Calef, G. W. and G. D. Grice. 1966. Relationships between the blue-green alga Trichodesmium thiebautii and the copepod Marosetella gracilis in the plankton off northwest South America. Ecology, 47:855-856. Caperon, J., S. A. Cattell, and G. Krasnick. 1971. Phytoplankton kinetics in a subtropical estuary: eutrophication. Limnol. and Oceanogr., 16:599-607. Cary, L. R. 1931. Studies on the coral reefs of American Samoa, with special reference to the Alcyonaria. Carnegie Inst. Wash. Pub. 413, Dept. Mar. Biol. Pap. 27:53-98. Caswell, H., H. E. Keonig, J. A. Resh, and Q. E. Ross. 1972. An intro- duction to systems science for ecologists, in: Patten, 1972, pp.3-78. Chan, G. L. 1972. Use of potential lagoon pollutants to produce protein in the South Pacific, in: M. Ruivo, ed., Marine pollution and sea life, pp. 525-527, London, Fishing News (Books) Ltd. Chapman, F. 1901. Foraminifera from the lagoon at Funafuti. J. Linn. Socks Bet... 28: 161-201 . Chave, K. E. 1965. Calcium carbonate: association with organic matter in surface seawater. Science, 148:1723-1724. ---------- » ed. 1970. Fanning Island Expedition, January 1970. Hawaii Inst. Geophys. HIG-70-23:1-201. once n oe 1971. Fanning Island Expedition--1970. Pac, Sci., 25:188-190. Chave, K. E., S. V. Smith, and K. J. Roy. 1971. Carbonate production by coral reefs. Mar. Geol., 12:123-140. Chave, K. E., and E. Suess. 1967. Suspended minerals in seawater. Trans N. Y. Acad. Sci. Ser. 11, 29:991-1000. ee ee 144 Chave, K. E. and E. Suess. 1970. CaC0.,-saturation in seawater: Effects of dissolved organic matter. Limno?. and Oceanogr., 15:633-637. Chesher, R. H. 1969. Destruction of Pacific corals by the sea star Acanthaster planci. Science, 165:280-283. Clarke, W. D. and M. Neushul. 1967. Subtidal ecology of the southern California coast,: in T. A. Olson and F. J. Burgess, eds. Pollution and Marine Ecology, pp. 29-42. New York: Interscience. Clutter, R. I. 1972. Subtle effects of pollution on inshore tropical plankton, in: M. Ruivo, ed., Marine pollution and sea life, pp. 435-439, London, Fishing News (Books) Ltd. 7 , x. . Clymer, A. B. 1972. Next-generation models in ecology, in: Patten, 1972, pp. 533-577. Coles, S. L. 1969. Quantitative estimates of feeding and respiration for three scleractinian corals. Thesis, University of Georgia. Collette, B. B. and F. H. Talbot. 1972. Activity patterns of coral . | reef fishes with emphasis on nocturnal-diurnal changeover. Nat. q Hist. Mus. Los Angeles Cty. Sci. Bull., 14:98-124. Conard, R. A., L. M. Meyer et al. 1965. Medical survey of the people of Rongelap and Utirik islands nine and ten years after exposure to fallout radiation (March 1963 and March 1964). U.S. Brookhaven National Lab. BNL-780(T-296):1-67. Conard, R. A., J. E. Rall and W. W. Sutow. 1966. Thryoid nodules as a late sequela of radioactive fallout in a Marshall Island population exposed in 1954. New England Jour. Med., 274:1392-1399. ee a _- ¥ Conover, R. J. and E. D. S. Corner. 1968. Respiration and nitrogen excretion by some marine zooplankton in relation to their life cycles. J. Mar. Biol. Assoc. U.K., 48:49-75. P ODD € Cox, D. C. 1951. The hydrology of Arno Atoll, Marshall Islands. Atoll. Res. Bull., 8:1-29. Culberson, C., R. M. Pytkowicz, and J. E. Hawley. 1970. Seawater alkalinity determination by the pH method. J. Mar. Res., 28:15-21. Dahl, A. L. 1969. The effect of environment on growth and development of Zonaria farlowii. Proc. Int. Seaweed Symp., 6:123-132. worn o---- 1971. Development, form and environment in the brown alga Zonaria farlowii (Dictyotales). Botanica Marina, 14:76-112. allele! 1972. Ecology and community structure of some tropical reef algae in Samoa. Proc. Int. Seaweed Symp., 7:36-39. Daly, R. A. 1910. Pleistocene glaciation and the coral reef problem. Am. J. Sci., Ser. 4, 30:297-308. 145 Daly, R. A. 1915. The glacial control theory of coral reefs. Proc. Am. Acad. Arts Sci., 51:155-251. D'Angelo, H. 1970. Linear time-varying systems: Analysis and synthesis. Boston: Allyn and Bacon. Darwin, C. R. 1839. Journal and researches 1832-1836, vol. 3. ---------- 1842. The structure and distribution of coral reefs. London: Smith, Elder & Co., pp. 1-214. Yoeecrccoo-- 1962. Coral islands: with introduction, map, and remarks by D. R. Stoddart. Atoll. Res. Bull., 88:1-20. Davis, W. M. 1928. The coral reef problem. Spec. Publ. Amer. Geogr. Soc., 9:596 pp. : Denizot, M. 1969. Introduction de quelques algues en Polynésie et variations saisonniéres. Bull. Soc. Phyc. Fr., 13-14:33-35. =55555 5555 1972. Sur le role constructeur des algues en Polynésie frangaise, in: Mukundan and Pillai, pp. 497-505. Direction des Centres d'Expérimentations Nucléaires. 1969. Mururoa. [Papers on Mururoa investigations reprinted from Cahiers du Pacifique nos. 12-13.] Paris, pp. 1-333. DiSalvo, L. H. and K. Gundersen. 1971. Regenerative functions and microbial ecology of coral reefs. I. Assays for microbial population. Can. J. Microbiol., 17:1081-1089. Domm, S. B. and A. J. Domm. 1973. The sequence of appearance at dawn and disappearance at dusk of some coral reef fishes. Pac. Sci., 27(2):128-135. Doran, E., ed. 1961a. Land tenure in the Pacific. Atoll Res. Bull., 85:1-60. eocccn---- 1961b. Marshall Islands landscape. Atoll Res. Bull., 85:11-15. Doty, M. S. 1967. Pioneer intertidal population and the related general vertical distribution of marine algae in Hawaii. Blumea, 15:95-105. ---------- 1971. The productivity of benthic frondose algae at Waikiki Beach 1967-1968. Hawaii Bot. Sci. Pap., 22:1-118. Doty, M. S. and J. P. E. Morrison. 1954. Interrelationships of the Organisms on Raroia aside from man. Atoll Res. Bull., 35:1-61. 146 Duerden, J. E. 1902. West Indian Madreporarian Polyps. Mem. Nat. Acad. Sci., Washington, 8:410-597. Earle, S. A. 1972. The influence of herbivores on the marine plants of Great Lameshur Bay, with an annotated list of plants, in: B. C. Collette and S. A. Earle, eds., Results of the Tektite Program: Ecology of coral reef fishes. Nat. Hist. Mus. Los Angeles County, Sci. Bull. 14:17-44. Ehrenfeld, D. W. 1974. Conserving the edible sea turtle: Can mari- culture help? Amer. Sci., 62(1):23-31. Emery, A. R. 1968. Preliminary observations on coral reef plankton. Limnol. and Oceanogr., 13(2):293-303. Emery, K. 0., Tracey, J. I., Jr. and Ladd, H.S. 1954. Geology of Bikini and nearby atolls: Part 1, geology. U.S. Geol. Surv. Prof. Pap. 260-A:1-265. Fairbridge, R. W. 1950. Recent and Pleistocene coral reefs of Australia. J. Geol., 58:330-401. a--------- 1958. Dating the latest movements of the Quaternary sea level. Trans. New York Acad. Sci., II, 20:471-482. Forrester, J. W. 1961. Industrial Dynamics. Cambridge, Mass.: MIT Press, 464 pp. Forstner, H. 1970. Problem and methods of micro-climatic measurements in marine littoral habitats. Oceanogr. Mar. Biol. Ann. Rev. 1970, 8:225-248, 21 figs. ---------- 1971. Measurement of radiant energy in the littoral benthos. Thalassia Jugoslavica, 7(1):27-33. Fosberg, F. R. 1949. Atoll vegetation and salinity. Pac. Sci., 3:89-92. wocccecsos 1953. Vegetation of central Pacific atolls, a brief summary. Atoll Res. Bull., 23:1-26. ---------- 1954. Soils of the Northern Marshall Atolls, with special reference to the Jemo series. Soil Sci., 78:99-107. woo------- 1957a. Some geological processes at work on coral atolls. Trans. N.Y. Acad. Sci. II, 19:411-422. wooo nn--- 1957b. Description and occurrence of atoll phosphate rock in Micronesia. Am. J. Sci., 225:584-592. woccccon-- 1957c. Soils, vegetation and agriculture on coral atolls. Proc. 8th Pac. Sci. Cong., 3A:1037-1047. Fosberg, F. R. 1959. Darwin on atoll fresh-water lens. Atoll Res. ---------- 1961. Qualitative description of the coral atoll ecosystem. 147 Bull., 70:6-7. Atoll Res. Bull., 81:1-11- wont oi---- 1963a. Man's place in the island ecosystem: a symposium (Tenth Pac. Sci. Congr., Honolulu, 1961). Honolulu:Bish. Mus. Press, 1-264. woccrcoo-- 1963b. Qualitative description of the coral atoll ecosystem. Fosberg, F. R., T. Arnow, and F. S. MacNeil. 1956. Military geography Fosberg, F. R. and D. Carroll. 1965. Terrestrial sediments and soils of Gallagher, B. S., K. M. Shimada, F. I. Gonzalez, Jr., and E. D. Stroup. Proc. 9th Pac. Sci. Congr., 4:161-167. of the northern Marshalls. Tokyo: Off. Engineer, U.S. Army Forces Far East, pp. 1-320. the northern Marshall] Islands. Atoll Res. Bull., 113:1-156. 1971. Tides and currents in Fanning Atoll lagoon. Pac. Sci., 25:191-205. Gardiner, J. S. 1898. The coral reefs of Funafuti, Rotuma and Fiji together with some notes on the structure and formation of coral reefs in general. Proc. Camb. Phil. Soc. Math. Phys. Sci., 9:417-503. wooo n----- 1902. On the rate of growth of some corals from Fiji. Proc. Camb. Phil. Soc. Math. Phys. Sci., 11:214-219. soccco---- 1903. The Maldive and Laccadive groups, with notes on other coral formations in the Indian Ocean, in: J.S. Gardiner, ed., The Fauna and geography of the Maldive and the Laccadive Archipelagoes, 1:146-183, 313-346, 376-423. Cambridge: The University Press. tata 1907-1936. Reports of the Percy Sladen Trust Expedition to Gibson-Hill, C. A., ed. 1950. Papers on the fauna of the Cocos-Keeling Gilmartin, M. 1958. Some observations on the lagoon plankton of Eniwetok ---------- 1960. The ecological distribution of the deep water algae of Ginsburg, R. N. 1956. Environmental relationships of grain size and the Indian Ocean in 1905... Trans. Linn. Soc. Zool., II, vols. 12-19. Islands. Bull. Raffles Mus., 22:1-298. MEOW se hace SClas 2S S-sil6. Eniwetok Atoll. Ecology, 41:210-221. constituent particles in some south Florida carbonate sediments. Bull. Am. Assoc. Petroleum Geologists, 40:2384-2427. | 148 Ginsburg, R. N. and H. A. Lowenstam. 1958. The influence of marine bottom communities on the depositional environment of sediments. J. Geol., 66:310-318. Ginsburg, R. N., E. A. Shinn, and J. H. Schroeder. 1967. Submarine cementation and internal sedimentation within Bermuda reefs (Abstr.), Geol. Soc. America Spec. Pub. 115:78-79. Glynn, P. W. 1972. Observations on the ecology of the Caribbean and Pacific coasts of Panama. Bull. Biol. Soc. Washington [D.C. ], 2els=SOk alata! (in press). Aspects of the ecology of coral reefs in the western Atlantic region, in: R. Endean and 0. A. Jones, eds., Biology and geology of coral reefs. I. Biology. New York, Academic Press. Glynn, P. W., L. R. Almodévar, and J. G. Gonzalez. 1965. Effects of hurricane Edith on marine life in La Parguera, Puerto Rico. Carib. J. Sci., 4:335-345. Glynn, P. W. and R. H. Stewart. 1973. Distribution of coral reefs in the Pearl Islands (Gulf of Panama) in relation to thermal conditions. Limnol. and Oceanogr., 18:367-379. Glynn, P. W., R. H. Stewart, and J. E. McCosker. 1972. Pacific coral reefs of Panama: structure, distribution and predators. Geologische Rundschau, 61(2):483-519. Goering, J. J., R. C. Dugdal, and D. W. Menzel. 1966. Estimates of in situ rates of nitrogen uptake by Trichodesmium sp. in the tropical Atlantic Ocean. Limnol. and Oceanogr., 11(4):614-620. Goldberg, W. M. 1973. The ecology of the coral-octocoral communities off the southeast Florida coast: Geomorphology, species composition, and zonation. Bull. of Mar. Sci., 23(3):465-488. Goliman, G. V. 1969. Ecological aspects of some coral-boring gastropods and bivalves of the northwestern Red Sea. Am. Zool., 9:887-894. Gordon, M. S. and H. M. Kelly. 1962. Primary productivity of a Hawaiian coral reef: A critique of flow respirometry in turbulent waters. Ecology, 43:473-480. Goreau, T. F. 1959. The ecology of Jamaican coral reefs. I. Species composition and zonation. Ecology, 40:67-90. alltel! 1961a. Problems of growth and calcium deposition in corals. Endeavour, 20:32-39. woo------- 1961b. Geological aspects of the structure of Jamaican coral reef communities. Final Progress Report, Contract NONR(G)- 149 003060 (NR 104-556), Biol. Br., U.S. Navy Office of Naval Research. Goreau, T. F. and N. I. Goreau. 1959. The physiology of skeleton formation in corals. II. Calcium deposition by hermatypic corals under various conditions in the reef. Biol. Bull. Mar. Biol. Lab., Woods Hole, 117:239-250. U a--------- 1960. Distribution of labeled carbon in reef-building corals with and without Zooxanthellae. Science, 131(3401) :558-559. ,/Goreau, T. F. and W. D. Hartman. 1963. Boring sponges as controlling factors in the formation and maintenance of coral reefs, in:Mechanisms of Hard Tissue Destruction, Am. Assoc. Advmt. Sci., Pub. 75:25-54. ---------- 1966. Sponge: effect on the form of reef corals. Science, 151 :343-344. Goreau, T. F. and L. S. Land. 1974. Fore-reef morphology and depositional ‘processes, North Jamaica, in: Laporte, pp. 77-89. Greeve, W. 1968. The "Planktonxreisal", a new device for culturing zooplankton. Mar. Biol., 1(3):201-203. Gressitt, J. L., ed. 1954-date. Insects of Micronesia. Honolulu, Bernice P. Bishop Mus. Grieg-Smith, P. 1964. Quantitative Plant Ecology. 2nd ed. London: Butterworths, 256 pp. Gross, M. G., J. D. Milliman, J. I. Tracey, Jr., and H. S. Ladd. 1969. Marine geology of Kure and Midway Atolls, Hawaii. Pac. Sci. 23(1):17-25. Gulland, J. A. 1970. Food chain studies and some problems of world fisheries, in J. H. Steele, ed., Marine Food Chains, pp. 296-318. Edinburgh: Oliver and Boyd. Hartman, R. T. and D. Brown. 1967. Changes in the composition of the Internal atmosphere of submerged vascular plants in relation to photosynthesis. Ecology, 48(2):252-258. Hartman, W. D. and T. F. Goreau. 1970. Jamaican coralline sponges. their morphology, ecology and fossil relatives. Symp. Zool. Soc. Lond., No. 25: 205-243. Hartog, G. den. 1970. The sea-grasses of the world. Verhand. Kon. Ned. Akad. Wetens. Nat. II, 59(1):1-275. Hatheway, W. H. 1953. The land vegetation of Arno Atoll, Marshall Islands. Atoll Res. Bull., 16:1-68. ---------- 1955. The natural vegetation of Canton Island, an equatorial Pacific atoll. Atoll Res. Bull., 43:1-9. EE 150 Hautserre, M.A. d' 1960. Les perturbations atmosphériques et le climat de la Polynesie frangaise. Monog. Météor. Nat., 18:1-62. Hedgepeth, J. W., ed. 1957. Treatise on marine ecology and paleoecology, Vol. 1; Ecology. 1-1296, Washington, D. C. Geological Soc. Amer. Memoir 67. Hedley, C., ed. 1896-1900. The atoll of Funafuti, Ellice group; its zoology, botany, ethnology, and general structure. Austral. Mus. Mem. 3:1-609. Heinle, D. R. 1966. Production of a calanoid copepod, Acartia tonsa in the Patuxent River Estuary. Chesapeake Sci., 7(2):59-74. Held, E. E. 1960. Land crabs and fission products at Eniwetok Atoll. PEVeer Sen de WSIB=27/- Held, E. E., S. P. Gessel and R. B. Walker. 1965. Atoll soil types in relation to the distribution of fallout radionuclides. Univ. Washington Fish. Lab. UWFL-92:1-37. Hiatt, R. W. and D. W. Strasburg. 1960. Ecological relationships of the fish fauna on coral reefs of the Marshall Islands. Ecol. Monogr., 30:65-127. Hinckley, A. D. 1969. Ecology of terrestrial arthropods on the Tokelau atolls. Atoll Res. Bull., 124:1-18. Hirota, J. 1972. Laboratory culture and metabolism of the planktonic ctenophore, Pleurobrachia bachei A. Agassiz, in: A. Y. Takenouti, ed., Biological Oceanography of the Northern North Pacific Ocean, pp. 465-484, Tokyo: Idemitsu Shoten. Hobson, E. S. 1965. Diurnal-nocturnal activity of some inshore fishes in the Gulf of California. Copeia, 3:291-302. eocrcsccoH 1968. Predatory behavior of some shore fishes in the Gulf of California. Fish and Wildlife Serv., Bur. Sport Fish. Wildlife, Res. Rept. 73:92 pp. w--------- 1972. Activity of Hawaiian reef fishes during the evening and morning transitions between daylight and darkness. Fishery Bulletin 70(3):715-740. Hogman, W. 1968. Annulus formation on scales of 4 species of Coregonids reared under artificial conditions. J. Fish. Res. Bd. Canada, 25(10):2111-2122. Holling, C. S. 1963. An experimental component analysis of population processes. Mem. Ent. Soc. Canada, 32:22-32. wocccco-- 1964. The analysis of complex population processes. Can. Ent., 96:335-347. 151 Holling, C. S. 1965. The functional response of predators to prey density and its role in mimicry and population regulation. Mem. Ent. Soc. Canada, 45:1-60. ---------- 1966. The functional response of invertebrate predators to prey density. Mem. Ent. Soc. Canada, 48:1-86. Hoskin, C. M. 1963. Recent carbonate sedimentation on Alacran Reef, Yucatan, Mexico. Nat. Acad. Sci., NRC, Washington, D.C., Pub. 1089:160 pp. Hutchinson, G. E. 1950. The biogeochemistry of vertebrate excretion. Bull. Amer. Mus. Nat. Hist., 96:1-554. Iams, W. J. 1969. Boilers of Bermuda's South Shore. Seminar on organism-sediment interrelationships, R. N. Ginsburg and S. M. Stanley, eds. Bermuda Biol. Sta. Pub. Jackson, W. B. 1967. Productivity in high and low islands with special emphasis to rodent populations. Micronesica, 3:5-15. Jackson, W. B. and M. L. Carpenter. 1969. Radioisotope cycling in terrestrial communities at Eniwetok Atoll, in: Nelson and Evans, 1969, pp. 644-651. Jeffrey, S. W. 1968. Photosynthetic pigments of the phytoplankton of some coral reef waters. Limnol. and Oceanogr., 13(2):350-355. Jerlov, N. G. 1968. Optical Oceanography. Amsterdam: Elsevier, 194 pp. Johannes, R. E. 1964. Phosphorus excretion and body size in marine animals: Microzooplankton and nutrient regeneration. Science, 150:76-77. ao-------- 1967. Ecology of organic aggregates in the vicinity of a coral reef. Limnol. and Oceanogr., 12:189-195. wecnroe--- 1970. Coral reefs and pollution. FAO Tech. Conf. on Marine pollution and its effects on living resources and fishing, Rome, Italy, 9-18 Dec. 1970. FAO Pub. FIR:MP/70/R-14. ---------- 1970-71. How to kill a coral reef. Marine Pollution Bull., 1(12):186-187; 2(1):9-10. ata 1972. Coral reefs and pollution, in: M. Ruivo, 1972, pp. 364-375. Johannes, R. E., J. Alberts, and 21 others. 1972. The metabolism of some coral reef communities: A team study of nutrient and energy flux at Eniwetok. BioScience 22(9) :541-543. Johannes, R. E., S. L. Coles, and N. T. Kuenzel. 1970. The role of zooplankton in the nutrition of some scleractinian corals. Limnol. and Oceanogr., 15:579-586. 152 Johannes, R. E. and K. L. Webb. 1970. Release of dissolved organic compounds by marine and freshwater invertebrates. Proc. Symp. Organic Matter Natural Waters, D. W. Hood, ed., Univ. Alaska Press, pp. 257-274. Johnson, D. L., and M. E. Q. Pilson, 1972a. Spectrophotometric deter- mination of arsenite, arsenate and phosphate in natural waters. Analytica Chim. Acta, 58(2):289-299. ---------- 1972b. Arsenate in the western North Atlantic and adjacent regions. J. Mar. Res., 30(1):140-149. Johnson, J. H. 1961. Fossil algae from Eniwetok, Funafuti and Kita- Daito-Jima. Geol. Survey Prof. Pap. 260-Z:907-950. ---------- 1965. Coralline algae from the Cretaceous and early Tertiary of Greece. J. Pal., 39(5):802-814. Johnson, M. W. 1949. Zooplankton as an index of water exchange between Bikini Lagoon and the open sea. Trans. Am. Geophys. Union, 30(2): 238-244. w--------- 1954. Plankton of the northern Marshall Islands. Bikini and nearby atolls. U.S. Geol. Survey Prof. Pap. 260-F:301-314. Johnston, R. 1966. Determination of ammonia in seawater as rubazoic acid. ICES, C. M. Hydr. Com. Paper 10. Jones, J. A. 1968. Primary production by the tropical turtle grass, Thalassia testudinum Kénig, and its epiphytes. Ph.D. Diss., Univ. of Miami, Florida. Jones, 0. A., and R. Endean, eds. 1973. Biology and geology of coral reefs, Vol. I: Geology 1. New York and London, Academic Press. 410 pp. Kanda C. 1944. Ecological studies on marine algae from Kororu and adjacent islands in the South Sea Islands. Palao Trop. Biol. Sta. Studies, 2:733-800. Kanwisher, J. 1963. Effect of wind on CO, exchange across the sea surface. J. Geophys. Res., 68:3921-3927. Kawaguti, S. and D. Sakumoto. 1948. The effect of light on the calcium deposition of corals. Bull. Oceanogr. Inst. Taiwan, 4:65-70. King, W. B. 1967. Seabirds of the tropical Pacific Ocean. Washington, Smithsonian Institution, 126 pp. Kinne, 0. and H. P. Bulnheim, eds. 1973. Internation Helgoland Symposium: Man in the sea--in situ studies on life in oceans and coastal waters. Helgolander wissenschaftliche Meeresuntersuchungen, 24(1-4):1-535. 153 Kinsey, D. W. 1972. Preliminary observations on community metabolism and primary productivity of the pseudo-atoll reef at One Tree Island, Great Barrier Reef, in: Mukundan and Pillai, pp. 13-32. Kinsey, D. W. and R. A. Bottomley. 1963. Improved electrode system for the determination of oxygen tension in industrial applications. J. Inst. Brew., 69:164-176. Kohn, A. J. and P. Helfrich. 1957. Primary organic productivity of a Hawaiian coral reef. Limnol. and Oceanogr., 2:241-251. Koroleff, F. 1970. Direct determination of ammonia in natural waters as indophenol blue. Infor. Tech. Methods for Sea Water Analysis, No. 32. ICES Interlab. Rept., pp. 19-22. [Ladd, H.S., ed.] 1954-date. Bikini and nearby atolls, Marshall Islands. U.S. Geol. Survey Prof. Pap. 260, Parts 1-4. Ladd, H. S., ed. 1957. Treatise on marine ecology and paleoecology, Vol. 2: Paleoecology. 1-1077, Washington, D.C. Geological Soc. Amer. Memoir 67. ereccno-- 1961. Reef building. Science, 134:703-714. ---------- 1971. Existing reefs--geological aspects. Proc. No. Amer. Paleontol. Conv. Sept. 1969. J:1273-1300. ---------- 1973. Bikini and Eniwetok atolls, Marshall Islands, in: Jones and Endean, pp. 93-112. ---------- in press. Types of coral reefs and their distribution, in: 0.A. Jones and R. Endean, eds. Biology and geology of coral reefs, Vol. 2. Academic Press. Ladd, H. S., E. Ingerson, R. C. Townsend, M. Russell, and H. K. Stephenson. 1953. Drilling on Eniwetok Atoll. Bull. Am. Pal. Geol., 37:2257-2280. Ladd, H. S. and S. 0. Schlanger. 1960. Drilling operations on Eniwetok Atoll. U.S. Geol. Survey Prof. Pap. 260-Y:863-903. Ladd, H. S. and J. I. Tracey, Jr. 1949. The problem of coral reefs. Scient. Monthly, 69:297-305. Ladd, H. S., J. I. Tracey, Jr., and M. G. Gross. 1967. Drilling on Midway Atoll, Hawaii. Science, 156:1088-1094. wore n----- 1970. Deep drilling on Midway Atoll. Geol. Survey Prof. Pap. 680-A:1-22. Land, L. S. 1970. Carbonate mud: production by epibiont growth on Thalassia testudinum. J. Sed. Petrology, 40:1361-1363. 154 Land, L. S. and 7. F. Goreau. 1970. Submarine lithification of Jamaican reefs. J. Sed. Petrology, 38:457-462. Lang, J. C. 1970. Inter-specific aggression within the scleractinian reef corals. Ph.D. Diss., Yale University. wenn nn n-- 1971. Interspecific aggression by scleractinian corals. 1. The rediscovery of Scolymia cubensis (Milne Edwards & Haime). Bull. Mar. Sci., 21(4):952-959. Laporte, L. F., ed. 1974. Reefs in time and space; Selected examples from the recent and ancient. Soc. of Econ. Paleontol. & Mineral. Sp. Publ., 18:1-256. Lasker, R. 1966. Feeding, growth, respiration and carbon utilization of a euphauviid crustacean. J. Fish. Res. Bd. Canada, 23:1291-1314. Lavoie, R. L. 1963. Some aspects of the meteorology of the tropical Pacific viewed from an atoll. Atoll Res. Bull., 96:1-80. Lee, R. F., J. Hirota, and A. M. Barnett. 1971. Distribution and importance of wax esters in marine copepods and zooplankton. Deep-Sea Res., 18:1147-1165. Lemoine, P. 1939. Algues calcaires fossiles de 1'Algérie. Mat. Carte Géol. de 1'Algérie Sér. 1, No. 9:1-121. Likens, G. E., ed. 1972. Nutrients and eutrophication: The limiting- nutrient controversy. Amer. Soc. of Limnol. Oceanogr. Spec. Symposia 1:1-328. Lisco, H. and R. A. Conard. 1967. Chromosome studies on Marshall Islanders exposed to fallout radiation. Science, 157(3787) :445-447. Littler, M. M. 1971. Ph.D. Diss., Univ. of Hawaii. seceann--- 1973a. The population and community structure of Hawaiian fringing-reef crustose Corallinaceae (Cryptonemiales, Rhodophyta). Jour. Exp. Mar. Biol. Ecol., 11:103-120. tocrcccnn- 1973b. The productivity of Hawaiian fringing-reef crustose Corallinaceae and an experimental evaluation of production methodology. Limnol. and Oceanogr., 18(6):946-952. Livingston, R. J. 1968. A volumetric respirometer for long-term studies of small aquatic animals. J. Mar. Biol. Assoc. U. K., 48:485-497. Macintyre, I. G. 1970. Sediments off the west coast of Barbados: Diversity of origins. Mar. Geol., 9:5-23. waoo-- Ho - 1972. Submerged reefs of eastern Caribbean. Am. Assoc. Petroleum Geologists. Bull., 56(4):720-738. 155 Macintyre, I. G., S. V. Smith, and J. C. Zieman. 1974 (in press). Carbon flux through a coral-reef ecosystem: a conceptual model Jour. Geol., Vol. 82. MacNeil, F. S. 1972. Physical and biological aspects of atolls in the northern Marshalls, in: Mukundan and Pillai, pp. 507-567. Mahnken, C. V. W. 1966. Distribution of copepoda at Rongelap Atoll, Marshall Islands. M. S. Thesis, Univ. of Washington, Seattle. Maragos, J. E., G. B. K. Baines, and P. J. Beveridge. 1973. Tropical cyclone Bebe creates a new land formation on Funafuti Atoll. Science 181:1161-1164, 21 September. Marsh, J. A. 1970. Primary productivity of reef-building calcareous red algae. Ecology, 31(2):255-263. Marshall, J. T., Jr. 1951. Vertebrate ecology of Arno Atoll, Marshall Islands. Atoll Res. Bull., 3:1-38. Marshall, N. 1965. Detritus over the reef and its potential contribution to adjacent waters of Eniwetok Atoll. Ecology, 46:343-344. SSSSS555555 1972. Note on mucus and zooxanthellae discharged from reef corals, in: Mukundan and Pillai, pp, 59-65. Mathiesen, A. C., R. A. Fralick, R. Burns and W. Flahive. 1972. Comparative studies of subtidal vegetation in the Virgin Islands and the New England coastlines, in: J. W. Miller, J. G. VanDerWalker and R. A. Waller, eds., Scientists in the sea, pp. VI-106 to VI-176. Washington, D. C., U.S. Dept. of the Interior (antedated 1971). Mathisen, 0.A. 1964. Determination of plankton biomass in Rongelap Atoll, Marshall Islands, by the use of a multi-net plankton pump. Verhand]. Intern. Ver. flr theor. angew. Limnol. (Int. Assoc. Theor. Appl. Limnol. Proc.) 15(2):735-744. Mayer [later Mayor] , A. G. 1915. The lower temperature at which reef- corals lose their ability to capture food. Yearbook Carnegie Inst. Washington, 14:212 pp. ---------- 1918. Ecology of the Murray Island coral reef. Carnegie Inst. Washington Pub. 213, Pap. Dept. Mar. Biol., 9:48 pp. Mayor, A. G. 1924a. Structure and ecology of Samoan reefs. Carnegie Inst. Washington Pub. 340, Pap. Dept. Mar. Biol., 19:1-25. ---------- 1924b. Causes which produce stable conditions in the depth of the floors of Pacific fringing reef-flats. Carnegie Inst. Wash- ington Pub. 340, Pap. Dept. Mar. Biol., 19:27-36. 156 Mayor, A. G. 1924c. Growth-rate of Samoan corals. Carnegie Inst. Washington Publ. 340, Pap. Dept. Mar Biol., 19:51-72. McCreary, F. E., Jr. 1959. A study of the mean vertical wind structure over the Eniwetok Proving Ground area. U.S. Joint Task Force Seven JTFMC TP-15:1-65. McKee, E. D. 1958. Geology of Kapingamarangi Atoll, Caroline Islands. Bull. Geol. Soc. America, 69:241-278. won nc non-H 1959. Storm sediments on a Pacific atoll. J. Sed. Petrology, 29:354-364. McRoy, C. P., J. J. Goering and B. Chaney. 1973. Nitrogen fixation associated with seagrasses. Limnol. and Oceanogr. 18(6): 998-1002. Menzel, D. W., and N. Corwin. 1965. The measurement of total phosphorus in seawater based on the liberation of organically bound fractions by persulfate oxidation. Limnol. and Oceanogr., 10:280-282. Menzel, D. W., and R. F. Vaccaro. 1964. The measurement of dissolved organic and particulate carbon in sea water. Limnol. and Oceanogr., 9:138-142. Meyerhoff, H. A., and Macintyre, I. G. 1973. Submerged reefs of Eastern Caribbean: Discussion; [and] Reply. Am. Assoc. Petroleum Geologists Bull. 57(2):407-411. Milliman, J. D. 1965. An annotated bibliography of recent papers on corals and coral reefs. Atoll Res. Bull., 111:1-58. Milliman, J. D. and C. V. W. Mahnken. 1969. Reef productivity measurements. Atoll Res. Bull., 129:23-41. Moe, M. 1969. Biology of the Red Grouper Epinephelus mario (Valenciennes) from the eastern Gulf of Mexico. Florida Dept. Nat. Res., Mar. Res. Lab. Prof. Pap. 10, 95 pp. Mukundan, C., and C.S.G. Pillai, eds. 1972. Symposium on corals and coral reefs, Mandapam Camp, 1969. 1-591, Cochin, Marine Biological Association of India, 591 pp. Mullin, M., and E. R. Brooks. 1970a. The effect of concentration of food on body weight, cumulative ingestion, and rate of growth of the marine copepod Calanus helgolandicus. Limnol. and Oceanogr., 15(5):748-755. las atti 1970b. Growth and metabolism of two planktonic, marine copepods as influenced by temperature and type of food, in: J. H. Steele, ed., Marine Food Chains, Edinburgh: Oliver & Boyd. 157 Munk, W. H. and Sargent, M. C. 1954. Adjustment of Bikini Atoll to ocean waves. U.S. Geol. Surv. Prof. Pap. 260-C:275-280. Murphy, J. and J. P. Riley. 1962. A modified single solution method for the determination of phosphate in natural waters. Anal. Chim. Acta, 27:31-36. Muscatine, L. and C. Hand. 1958. Direct evidence for the transfer of materials from symbiotic algae to the tissues of a coelenterate. Proc. Nat. Acad. Sci., 44:1259-1263. Nehring, D. 1968. Some remarks on the determination of nitrate and ammonia in seawater. Infor. Tech. Methods for Sea Water Analysis, No. 2. ICES Interlab. Rept., pp. 16-17. Nelson, D. J. and F. C. Evans, eds. 1969. Proceedings of the Second National Symposium on Radioecology, Ann Arbor, Michigan, May 15-17, 1967. U.S. Atomic Energy Commission CONF-670503 Biology and Medicine (TID-4500). Oak Ridge, Tennessee, 774 pp. Nestéroff, W. D. 1956. Erosion et cimentation des calcaires dans les eaux tropicales. Res. de los Trabajos Presentados, 20 Congr. Geol. Intern. Mexico, :267. Neumann, A. C. 1965. Processes of recent carbonate sedimentation Harrington Sound, Bermuda. Bull. Mar. Sci. Gulf Carib., 15:987-1035. worn cnnn-- 1966. Observations on coastal erosion in Bermuda and measurements of the boring rate of the sponge Cliona lampa. Limnol. and Oceanogr., 11:92-108. Neumann, A. C. and L. S. Land. 1969. Algal production and lime mud deposition in the bight of Abaco: A budget (Abstract). Geol. Soc. America, Spec. Pap. 127:219. Neushul, M. 1967. Studies of subtidal marine vegetation in western Washington. Ecology, 48:83-94. Neushul, M., W. D. Clarke, and D. W. Brown. 1967. Subtidal plant and animal communities of the southern California Islands, in: R. N. Philbrick ed., Proc. Symp. Biol. California Islands. Santa Barbara Botanic Garden, Santa Barbara, Calif., pp. 37-55. Neushul, M. and A. L. Dahl. 1967. Composition and growth of subtidal parvosilvosa from Californian kelp forests. Helgoldnder Wiss. Meeresunters. 15:480-488. Newell, B. S. 1967. The determination of ammonia in sea water. J. Mar. Biol. Ass. U.K., 47:271-280. Newell, N. D. 1956. Geological reconnaissance of Raroia (Kon Tiki) Atoll, Tuamotu Archipelago. Bull. Am. Mus. Nat Hist., 109:311-372. 158 Newell, N. D. 1959. Questions of the coral reefs. Nat. Hist., 68:118-131. ---------- 1962. Questions of the coral reefs, in: J. F. White, ed., Study of the earth, pp. 121-136. Englewood Gifts Ned: Prentice Hall. (Reprinted from Newell, 1959) Newhouse, J. 1954. Ecological and floristic notes on the Myxophyta of Raroia. Atoll Res. Bull., 33:42-54. Newman, W. A. 1970. Acanthaster: a disaster? Science, 167:1274-1275. Niering, W. A. 1961. Observations on Puluwat and Gaferut, Caroline Islands, with historical and climatic information on Gaferut Island by M.-H. Sachet. Atoll Res. Bull., 76:1-15. ---------- 1963. Terrestrial ecology of Kapingamarangi Atoll, Caroline Islands. Ecol. Monogr., 33:131-160. Numata, M., convenor. 1967. Island ecosystems of the Pacific Basin, symposium at 11th Pac. Sci. Cong. Tokyo, 1966. Micronesica 3:1-54. Odum, E. P. 1971. Fundamentals of ecology. Philadelphia: Saunders & Co., 574 pp. Odum, H.T. 1957. Primary production measurements in 11 Florida springs and amarine turtle grass community. Limnol. and Oceanogr., 2:85-97. Odum, H. T., P. R. Burkholder, and J. Rivero. 1959. Measurements of productivity of turtle grass flats, reefs, and the Bahia Fosforescente of southern Puerto Rico. Bull. Inst. Mar. Sci., 6:159-170. Odum, H. T. and E. P. Odum. 1955. Trophic structure and productivity of a windward coral reef community. Ecol. Monogr., 25:291-320. Ogden, J. C., R. A. Brown, and N. Salesky. 1973. Grazing by the echinoid Diadema antillarum Philippi: Formation of halos around West Indian patch reefs. Science 182:715-717. Otter, G. W. 1937. Rock-destroying organisms in relation to coral reefs. Sci. Rep. Great Barrier Reef Exped., 1:323-352. Paine, R. T. and R. L. Vadas. 1970. The effects of grazing by sea urchins, Strongylocentrotus spp. on benthic algal populations. Limnol. and Oceanogr., .14:710-719. Paloheimo, J. E. and L. M. Dickie. 1966. Food and the growth of fishes. II. Effects of food and temperature on the relation between metabolism and body weight. J. Fish. Res. Bd. Canada, 23:869-908. Palumbo, R. F., A. H. Seymour, and A. D. Welander. 1966. Radionuclides in foods from the central Pacific, 1962. Nature 209(5029) :1190-1192. 159 Park, P. K. 1968. Seawater-hydrogen ion concentration: Vertical dis- tribution. Science, 162:358-359. ---------- 1969. Oceanic CO, system: An evaluation of ten methods of investigation. Limnol. and Oceanogr., 14:179-186. Patten, B. C., ed. 1971. Systems analysis and simulation in ecology, Vol. I. New York: Academic Press, 607 pp. ==-==>-=-- ed. 1972. Systems analysis and simulation in ecology, Vol. II. New York: Academic Press, 592 pp. Perkins, R. D. and P. Enos. 1968. Hurricane Betsy in the Florida-Bahama area--geologic effects and comparison with hurricane Donna. J. Geol., 76:710-717. Pichon, M. 1972. The coral reefs of Madagascar, in: R. Battistini and G. Richard-Vindard, eds. Biogeography and ecology in Madagascar, pp. 367-410, The Hague, W. Junk. Piggott, C. J. 1968. A soil survey of Seychelles. Dir. Overseas Surv. eehe= Bull ss2-—69- Pilson, M. E. Q. and S. B. Betzer. 1973. Phosphorus flux across a coral reef. Ecology, 54(3):581-588. Plessis, Y. 1972. Ichthyologie corallienne: écologie et exploitation, in: Mukundan and Pillai, pp 457-468. Pomeroy, L. R. 1960. Primary production of Boca Cieza Bay, Florida. Bull. Mar. Sci. Gulf and Carib., 10(1):1-10. Pomeroy, L. R., R. E. Johannes, E. P. Odum, and B. Roffman. 1969. The phosphorus and zinc cycles of a salt marsh, in: Nelson and Evans, 1969, mp. 412-419. Pomeroy, L. R. and E. J. Kuenzler. 1969. Phosphorus turnover by coral reef animals, in: Nelson and Evans, 1969,p. 474-482. Pomeroy, L. R., L. R. Shenton, R. D. H. Jones, and R. J. Reimold. 1972. Nutrient flux in estuaries, in: G. E. Likens, ed., Nutrients and eutrophication. Am. Soc. Limnol. Oceanogr. Spec. Symposia 1:274-293. Pomeroy, L. R., E. E. Smith, and C. M. Grant. 1965. The exchange of phosphate between estuarine water and sediments. Limnol. and Oceanogr., 10:167-172. Prabhu, M. S., S. Ramamurthy, M. D. K. Kuthalingham, and M. H. Dhulkhed. 1965. On an unusual swarming of the planktonic blue-green algae Trichodesmium spp., off Mangalore. Current Science (India) 34(3):95. Prosser, C. L. and F. A. Brown. 1961. Comparative animal physiology. 2nd ed. Philadelphia: W. B. Saunders, 688 pp. 160 Pugh, W. E. ed. 1950. Bibliography of organic reefs, bioherms and biostromes. Tulsa: Seismic Service Corp., 139 pp. Purdy, E. G. 1974. Reef configurations: cause and effect, in: Laporte, pp. 9-76. Qasim, S. Z. and V. N. Sankaranarayanan. 1970. Production of particulate organic matter by the reef on Kavaratti Atoll (Laccadives) . Limnol. and Oceanogr., 15:574-578. Quinn, W. H. and Burt, W. V. 1970. Prediction of abnormally heavy precipitation over the Equatorial Pacific Dry Zone. Jour. Applied Meteor. 9:20-28. Ramamurthy, V. D. and S. Krishnamurthy. 1967. Effects of N:P ratios on the uptake of nitrate and phosphate by laboratory cultures of Trichodesmium erythraeum (Ehr.). Proc. Ind. Acad. Sci. Sec. B, 65(2) :43-48. Ramamurthy, V. D. and R. Seshadri. 1966a. Effects of gibberellic acid (GA) on laboratory cultures of Trichodesmium erythraeum (Ehr.) and Melosira sulcata (Ehr.). Proc. Ind. Acad. Sci. Sec. B, 64(3):146-151. ---------- 1966b. Phosphorus concentration during red-water phenomenon in the near-shore waters of Porto-Novo (S. India). Current Science 35(4):100-101. Ramage, C. S. 1959a. Hurricane development. Jour. of Meteorol. 16:227-237. w--------~ » ed. 1959b. Notes on the meteorology of the tropical Pacific and Southeast Asia. 1-174, Honolulu, Hawaii Institute of Geophysics (AFCRC-TN-60-216). Ramage, C. S., F. R. Miller, and C. Jefferies. 1972. International Indian Ocean expedition meteorological atlas, Vol. 1, The surface climate of 1963 and 1964. 144 charts. Washington, National Science Foundation (NSF-IOE-2). Randall, J. E. 1961. Overgrazing of algae by herbivorous marine fishes. Ecology, 42(4):812. tated 1962. Tagging reef fishes in the Virgin Islands. Proc. Gulf and Carib. Fisher. Inst., Fourteenth Ann. Session, pp. 201-241. atta 1963a. Additional recoveries of tagged reef fishes from the Virgin Islands. Proc. Gulf and Carib. Fisher. Inst., Fifteenth Session, pp. 155-157. woenn----- 1963b. An analysis of the fish populations of artificial and natural reefs in the Virgin Islands. Carib. J. Sci., 3(1):31-47. ---------- 1965. Grazing effect on sea grasses by herbivorous reef fishes in the West Indies. Ecology, 46:255-260. 161 Randall, J. E. 1967. Food habits of reef fishes of the West Indies. Studies Trop. Oceanogr., 5:665-847. Randall, J. E. and W. D. Hartman. 1968. Sponge-feeding fishes of the West Indies. Mar. Biol. Berlin, 1:216-225. Randall, J. E. and H. A. Randall. 1963. The spawning and early devel- opment of the Atlantic parrot fish, Sparisoma rubripinne, with notes on other scarid and labrid fishes. Zoologica, 48(2):49-59. Ranson, G. 1958. Coraux et récifs coralliens (Bibliographie). Bull. Inst. Oceanogr. Monaco, 1121:80 pp. Reese, E. S. 1964. Ethology and marine zoology. Oceanogr. Mar. Biol. Ann. Rev., 2:455-488. Reeve, M. R. 1969. Growth, metamorphosis, and energy conversion in the larvae of the prawn, Palaemon serratus. J. Mar. Biol. Assoc. U.K., 49(1):77-96. Reinboth, R. 1973. Dualistic reproductive behavior in the protogynous wrasse Thalassoma bifasciatum and some observations on its day-night changeover. Helgoldander wiss. Meersunters. 24:174-191. Renaud-Mornant, J. C., B. Salvat, and C. Bossy. 1971. Macrobenthos and meiobenthos from the closed lagoon of a Polynesian atoll. Maturei Vavao (Tuamotu). Biotropica 3(1):36-55. Rice, M. E. 1969. Possible boring structures of sipunculids. Am. Zool., 9:803-812. Richards, F. A. and R. A. Kletsch. 1964. The spectrophotometric deter- mination of ammonia and labile amino compounds in fresh and seawater by oxidation to nitrite, in: Y. Miyake and T. Koyama, eds., Recent researches in the fields of hydrosphere, atmosphere and nuclear geochemistry, pp. 65-81, Nagoya, Editorial committee for Sugawara volume, Nagoya University. Risebrough, R. W., J. D. Davis, and D. W. Anderson. 1970. Effects of various chlorinated hydrocarbons, in: J. W. Gillett, ed., The biological impact of pesticides in the environment. Env. Health Series 1. Corvallis, Ore.: Oregon State Univ. Robbins, J., J. E. Rall, and R. A. Conard. 1967. Late effects of radioactive iodine in fallout. Ann. Intern. Medicine 66:1214-1242. Robertson, R. 1970. Review of the predators and parasites of stony corals, with special reference to symbiotic prosobranch gastropods. Pac. Sci. 24:43-54. Roy, K. J. 1970a. Change in bathymetric configuration, Kaneohe Bay, Oahu, 1882-1969. HIG Rep. No. 70-15. Univ. Hawaii, 26 pp. 162 Roy, K. J. 1970b. Note on phsophate rock at Fanning Island. Hawaii Inst. Geophys. Rep. 70-23:193-199. Roy, K. J. and S. V. Smith. 1971. Sedimentation and coral reef development in turbid water: Fanning lagoon. Pac. Sci., 25:234-248. Royal Society of London. 1904. The atoll of Funafuti: borings into a coral reef and the results. London, 428 pp. Ruetzler, K. 1971. Bredin-Archibold-Smithsonian Biological survey of Dominica: Burrowing sponges, genus Siphonodictyon Berggnist, from the Caribbean. Smithsonian Contr. Zool., 77:1-37. ---------- 1974. The burrowing sponges of Bermuda. Smithsonian Contr. Zool. 165:1-32. Ruivo, M., ed. 1972. Marine pollution and sea life. 1-624, London, Fishing News (Books) Ltd. Published by arrangement with FAO. Sachet, M.-H. 1957. Climate and meteorology of the Gilbert Islands. Atoll Res. Bull. 60:1-4. eorcrno--- 1962a. Flora and vegetation of Clipperton Island. Proc. Calif. Acad. Sci. IV, 31:249-307. worccnn--- 1962b. Monographie physique et biologique de 1'Tle Clipperton. Ann. Inst. Océanogr. n.s. 40(1):1-107. ---------- 1963. History of change in the biota of Clipperton Island, in: J.L. Gressitt, ed., Pacific basin biogeography, a symposium. Pp. 525-534, Honolulu, Bishop Museum Press. ----- ee 1967. Coral islands as ecological laboratories. Micronesica 745-49. Sachet, M.-H. and F. R. Fosberg. 1955. Island bibliographies: Micronesian botany, Land environment and ecology of coral atolls, Vegetation of tropical Pacific Islands. Nat. Acad. Sci., Washington, NAS-NCR Pub. 335:577 pp. eo-c------ 1971. Island bibliographies supplement. Pac. Sci. Bd., Nat. Acad. Sci., Washington, 427 pp. Sadler, J. C. and R. E. Gidley. 1973. Tropical cyclones of the North Indian Ocean. ENVPREDRSCHFAC Techn. Pap. 2-73:1-60. Monterey, California, U.S. Naval Postgraduate School. Salvat, B. 1970a. Etude quantitatives (comptages et biomasses) sur les mollusques récifaux de 1'atoll de Fangataufa (Tuamotu - Polynésie) . Cahiers du Pacifique 14:1-58. 163 Salvat, B. 1970b. Les mollusques des "récifs d'flots" du récif barriére des Tles Gambier (Polynésie); bionomie et densité de peuplement. Bull. Mus. Hist. Nat. Paris II, 42:525-542. Sargent, M. C. and T. S. Austin. 1949. Oceanic productivity of an atoll. Trans. Amer. Geophys. Union, 30(2):245-249. woo no-H---- 1954. Biologic economy of coral reefs, Bikini and nearby atolls, Marshall Islands. U.S. Geol. Surv. Prof. Pap. 260-E: 293-300. Schreiber, R. W. and N. P. Ashmole. 1970. Sea-bird breeding seasons on Christmas Island, Pacific Ocean. Ibis, 112:363-394. Setchell, W. A. 1926. Nullipore versus coral in reef-formation. Proc. Amer. Phil. Soc., 65(2):136-140. ---------- 1928. Coral reefs as zonational plant formations. Science, 68:119-121. Seymour, A. H., ed. 1971. Radioactivity in the marine environment. 272 pp. Washington: National Academy of Sciences. Panel on radioactivity in the marine environment. Shinn, E. A. 1963. Formation of spurs and grooves on the Florida reef tract. J. Sed. Petrology, 33:291-303. erccccooo- 1966. Coral growth rate, an environmental indicator. Jd. Palaeont., 40:233-240. oo-------- 1972. Coral reef recovery in Florida and the Persian Gulf. 1-9+22 figs., Houston, Shell Oil Co. Smith, C. L. and Tyler, J. C. 1972. Space resource sharing in a coral reef fish community. Sci. Bull. Nat. Hist. Mus. Los Angeles County 14:125-170. Smith, S. V. 197la. Budget of calcium carbonate, Southern California continental borderland. J. Sed. Petrology, 41:798-808. STSSSSSS5= 1971b. Factor analysis; a tool for environmental studies May. Tech... Soc..d., 5:15-19. ---------- 1972. Production of calcium carbonate on the mainland shelf of Southern California. Limnol. and Oceanogr. 17(1):28-41. atta 1973. Carbon dioxide dynamics: A record of organic carbon production, respiration, and calcification in the Eniwetok reef flat community. Limnol. and Oceanogr. 18:106-120. 164 Smith, S. V., K. E. Chave, and D. T. 0. Kam. 1973. Atlas of Kaneohe Bay: A reef ecosystem under stress. 1-128, Honolulu, University of Hawaii Seagrant Program (UNIHI-SEAGRANT-TR-72-01 ). Smith, S. V. and J. A. Marsh, Jr. 1973. Organic carbon production on the windward reef flat of Eniwetok Atoll. Limnol. and Oceanogr. 18(6) :953-961. Smith, S. V., K. J. Roy, K. E. Chave, J. E. Maragos, A. Soegiarto, G. Key, M. J. Gordon, and D. Kam. 1970. Calcium carbonate production and deposition in a modern barrier reef complex (Abs.). Pap. pres. Ann. Meeting Geol. Soc. America, Milwaukee. Smith, S. V., K. J. Roy, H. G. Schiesser, G. L. Shepherd, and K. E. Chave. 1971. Flux of suspended calcium carbonate (CaCO), Fanning Island Lagoon. Pac. Sci., 25:206-221. Soegiarto, A. 1972. The role of benthic algae in the carbonate budget of the modern reef complex, Kaneohe Bay. Ph.D. thesis, Univ. Hawaii. Solorzano, L. 1969. Determination of ammonia in natural waters by the phenolhypochlorite method. Limnol. and Oceanogr., 14:799-801. Sorokin, Y. I. 1973. On the feeding of some scleractinian corals with bacteria and dissolved organic matter. Limnol. and Oceanogr. 18(3) :380-385. South Pacific Commission. 1973. Regional Symposium on Conservation of Nature--Reefs and lagoons. Proceedings and Papers. i-xi, 1-156, 1-314, Noumea. Springer, V. G. and A. J. McErlean. 1962. A study of the behavior of some tagged south Florida coral reef fishes. Am. Mid]. Natur., 67(2) :286-297. Starck, W. A., II, and W. P. Davis. 1966. Night habits of fishes of Alligator Reef, Florida. Ichthyologica, The Aquarium Journal , 38(4) :313-356. Stockman, K. W., R. N. Ginsburg, and E. A. Shinn. 1967. The production of lime mud by algae in south Florida. J. Sed. Petrology, 37:633-638. Stoddart, D. R. 1962. Three Caribbean atolls: Turneffe Islands, Lighthouse Reef, and Glover's Reef, British Honduras. Atoll Res. Bull., 87:1-151. woo------- 1963. Effects of hurricane Hattie on the British Honduras reefs and cays, October 30-31, 1961. Atoll Res. Bull., 95:1-142. wooo ------ 1965. The shape of atolls. Marine Geol. 3:369-383. 165 Stoddart, D. R. 1968a. Catastrophic human interference with coral atoll ecosystems. Geography 53:25-40. ---------- 1968b. Isolated island communities. Sci. Jour. 4(4) :32-38. (peace enne- 1969a. Ecology and morphology of recent coral reefs. Biol. Rev., 44:433-498. a-o------- 1969b. Reconnaissance geomorphology of Rangiroa Atoll, Tuamotu Archipelago, with list of vascular flora of Rangiroa, by M.-H. Sachet. Atoll Res. Bull., 125:1-44. woot o---- 1969c. Post-hurricane changes on the British Honduras reefs and cays: Re-survey of 1965. Atoll Res. Bull. 131:1-25. SSSSSSS5=5 1969d. Islands as ecological laboratories. New Scientist 41(640-special sect.) :20-23. so-co----- 1971. Sea-level change and the origin of sand cays: Radiometric evidence. Jour. Mar. Biol. Assoc. India 11:44-58. worn onn-- 1972. Catastrophic damage to coral reef communities by earthquake. Nature 239(5366) :51-52. <==-====-~ 1973. Coral reefs: The last two million years. Geography 58(4) :313-323. Stone, E. L., Jr. 1951. The soils of Arno Atoll, Marshall Islands. Atoll] Res. Bull., 5:1-56. woo------- 1953. Summary of information on atoll soils. Atoll Res. Bull., 22:1-4. Strickland, J. D. H. 1960. Measuring the productivity of marine phytoplankton. Fish. Res. Bd. Canada Bull., No. 122:112 pp. Strickland, J. D. H. and T. R. Parsons. 1968. A practical handbook of seawater analysis. Fish. Res. Bd. Canada Bull. 167:143-147. Suess, E. 1968. Calcium carbonate interaction with organic compounds. Ph.D. Diss., Lehigh University. altel 1970. Interaction of organic compounds with calcium carbonate-- I. Association phenomena and geochemical implications. Geochim. Cosmochim., 34:157-168. Suyehiro, Y. 1942. A study on the digestive system and feeding habits of fish. Jap. J. Zool., 10(1):1-30. Talbot, F. H. and B. Goldman. 1972. A preliminary report on the diversity and feeding relationships of the reef fishes of One Tree Island, Great Barrier Reef system, in: Mukundan and Pillai, pp. 425-443. 166. Tayama, R. 1952. Coral reefs of the South Seas. Bull. Hydrogr. Dept., Tokyo, 11:1-292, and 2 vols. of plates and figures. Taylor, D. L. 1969a. On the regulation and maintenance of algal numbers in Zooxanthellae-coelenterate symbiosis, with a note on the nutritional relationships in Anemonia sulcata. J. Mar. Biol. Assoc. U.K. 49(4): 1057-1065. ---------- 1969b. The nutritional relationship of Anemonia sulcata (Pennant) and its dinoflagellate symbiont. Jour. Cell Sci. 4:751-762. wo-------- 1971. On the symbiosis between Amphidinium klebsii (Dinophyceae) and Amphiscolops langerhansi (Turbellaria: Acoela). J. Mar. Biol. Assoc. U. K., 51:301-313. ---------- 1973. The cellular interactions of algal-invertebrate symbiosis. Adv. Mar. Biol. 11:1-56. Taylor, R. C. 1973. An atlas of Pacific Islands rainfall. Hawaii Inst. Geophys. Data Rept. 25, HIG-73-9. Taylor, W. R. 1960. Marine algae of the eastern tropical and subtropical coasts of the Americas. Ann Arbor, Univ. of Michigan Press,870 pp. Templeton, W. L., R. E. Nakatani, and E. E. Held. 1971. Radiation effects, in: Seymour, 1971, pp. 223-239. Tercinier, G. 1956. Contribution a 1'étude des sols coralliens des atolls (les sols de Rangiroa: Touamotou). Vle Congrés International de la Science du Sol, Paris, 1956. Rapports présentés au Congrés V, 6:37-42. Deiat wig 1969. Note de synthése sur les sols du Motu Faucon (étude pédologique d'une portion représentative de 1'atoll de Mururoa). Cahiers du Pacifique, 13:17-46. Thomassin, B. 1972. Les biotopes de sables coralliens dérivant des appareils récifaux de la région de Tuléar (S.W. de Madagascar), in: Mukundan and Pillai, pp. 291-313. Tracey, J. I., Jr., D. P. Abbott, and T. Arnow. 1961. Natural history of Ifaluk Atoll: Physical environment. Bishop Mus. Bull., 222:1-75. Trichet, Jean. 1970. Etude des premiers stades d'évolution de la matiére organique dans des mares en milieu récifal (Polynésie frangaise), in: Hobson, G. D., ed., Advances in organic geochemistry 265-284. Tsuda, R. T. 1970. A floristic and distributional account of the Phaeophyta (brown algae) on Guam, Mariana Islands. Ph.D. Diss., Univ. Wisconsin. 167 Tsuda, R. T. 1972. Morphological, zonational, and seasonal studies of two species of Sargassum on the reefs of Guam. Proc. Intern. Seaweed Symp. 7:40-44. U.S. Joint Task Force Seven. 1956. Meteorological report on Operation Redwing, Part 1--Meteorological Data, Vol. 6--Wake. JTFMC TP-1:1-338, Pearl Harbor. (Period 15 April--31 July, 1956). ---------- 1958. Meteorological report on Operation Hardtack. JTFMC TP-8: 6 Vols., Pearl Harbor. ---------- 1959. A Christmas Island climatological study. JIFMC TP-11: 1-15+8, Pearl Harbor. U.S. Navy (Chief of Naval Operations). 1956-68. Marine climatic atlas of the world. Vol. II. North Pacific Ocean: NAVAER 50-1¢-529: i-xviii, 275 charts, 1956 [1957] ; III. Indian Ocean: NAVAER 50-1C-530: i-xvii, 267 charts, 1957 [1958] ; V. South Pacific Ocean: NAVAER 50-1C-532: i-xvi, 267 charts, 1959; VIII. The world: NAVAER 50-1C-54: i-xxi, 179 charts, 1968. Van Dorn, W. G. 1970. Tsunami response at Wake Island: A model study. Jour. Marine Res. 28:336-344. Van Oosten J. 1957. The skin and scales, in: M. Brown, ed., The physiology of fishes, pp. 207-244. New York: Academic Press. Van Overbeek, J. and R. E. Crist. 1947. The role of a tropical green alga in beach sand formation. Am. J. Bot., 34:299-300. Vary, W. E. 1970. Remote sensing by aerial color photography for water depth penetration and ocean bottom deta*]. Preprint, U. S. Naval Oceanographic Office, Washington, D.C. Vastano, A. C. and R. 0. Reid. 1970. Tsunami response at Wake Island. Comparison of the hydraulic and numerical approaches. Jour. Marine Res., 28:345-356. Vaughan, T. W. 1915. The geologic significance of the growth-rate of the Floridian and Bahaman shoal-water corals. J. Wash. Acad. Sci., 5:591-600. ---------- 1916a. Results of investigation of the ecology of the Floridian and Bahaman shoal-water corals. Proc. Nat. Acad. Sci., 2:95-100. nonce n---- 1916b. Some littoral and sublittoral physiographic features of the Virgin and Northern Leeward Islands and their bearing on the coral reef problem. J. Wash. Acad. Sci., 3:53-66. ---------- 1918. Some shoal-water corals from Murray Island (Australia), Cocos-Keeling Islands, and Fanning Island. Carnegie Inst. Wash. Pub. 213, Pap. Dep. Mar. Biol., 9:49-219. 168 Vaughan, T. W. and J. W. Wells. 1943. Revision of the suborders, families, and genera of the Scleractinia. Spec. Pap. Geol. Soc. America, 44:363 pp. Von Arx, W. S. 1954. Circulation systems of Bikini and Rongelap Lagoons, Bikini and nearby atolls, Marshall Islands. U.S. Geol. Surv. Prof. Pap. 260-B:215-373. Watson, G. E., Zusi, R. L., and Storer, R. E. 1963. Preliminary field guide to the birds of the Indian Ocean. Smithsonian Institution, 214 pp. Webb, K. L. and R. E. Johannes. 1969. Do marine crustaceans release dissolved amino acids? Comp. Bioch. Physiol. 29:875-878. Webb, K. L. and L. Wood. 1967. Improved techniques for analysis of free amino acids in seawater, in: Automation in analytical chemistry, Technicon Symposia, 1966, Vol. 1:440-444. White Plains, N.Y.: Mediad. Welander, A. D. 1969. Distribution of radionuclides in the environment of Eniwetok and Bikini atolls, August 1964, in: Nelson and Evans, 1969, pp. 346-354. Wells, J. W. 1954. Recent corals of the Marshall Islands. U.S. Geol. Survey Prof. Pap. 260-1:385-486. ---------- 1956. Scleractinia, in: R. C. Moore, ed., Treatise on invertebrate palaeontology, Part F (Coelenterata) :328-333. Geol. Soc. America, Univ. Kansas Press. ---------- 1957a. Coral reefs. Mem. Geol. Soc. America, 67(1):609-631. aaa 1957b. Annotated bibliography: corals. Mem. Geol. Soc. America, 67(1):1089-1104. Wentworth, C. K. 1931. Geology of the Pacific equatorial islands. Occ. Paps. Bernice P. Bishop Mus. 9(3):1-25. Westoll, T. S. and D. R. Stoddart, eds. 1971. A discussion on the results of the Royal Society Expedition to Aldabra 1967-68. Phil. Trans. Roy. Soc. London B. Biol. Sci. 260(836) :1-654. Wiens, H. J. 1962. Atoll environment and ecology. New Haven and London, Yale Univ. Press, 532 pp. Winn, H. E. and J. E. Bardach. 1960. Some aspects of the comparative biology of parrot fishes at Bermuda. Zoologica, 45(1):29-54. Wood, E. D., F. A. J. Armstrong, and F. A. Richards. 1967. Determination of nitrate in sea water by cadmium-copper reduction to nitrite. J. Mar. Biol. Assoc. U.K., 47:23-31. 169 Wood-Jones, F. 1910. Coral and atolls. London: L. Reeve, 392 pp. Woodwell, G. M., P. P. Craig, and H. A. Johnson. 1971. DDT in the biosphere: where does it go? Science, 174:1101-1107. Yonge, C. M. 1930a. Studies on the physiology of corals. I. Feeding mechanisms and food. Scient. Rep. Great Barrier Reef Exped., TP213=57. lata 1930b. Studies on the physiology of corals. II. Digestive enzymes. Scient. Rep. Great Barrier Reef Exped., 1:59-81. ---------- 1931. Studies on the physiology of corals. III. Assimilation and excretion. Scient. Rep. Great Barrier Reef Exped., 1:83-92. ---------- 1940. The biology of reef-building corals. Scient. Rep. Great Barrier Reef Exped., 1:353-391. ate! 1963. The biology of coral reefs. Adv. Mar. Biol., 1:209-260. ---------- 1968. Living corals. Proc. R. Soc. London, 169:329-344. Zadeh, L. A. and C. A. Desoer. 1963. Linear system theory, the state space approach. New York: McGraw Hill. Zeuthen, E. 1947. Body size and metabolic rate in the animal kingdom with special regard to the marine micro-fauna. Comp. Rend. Trav. Lab. Carlsberg (Copenhagen), Ser. Chim., 26:17-161. Zieman, J. C. 1968. A study of the growth and decomposition of the sea-grass, Thalassia testudinum, Kénig. M. S. Thesis, Inst. Mar. Sci., Univ. Miami, Florida. Zipser, E. J. and R. C. Taylor. 1968. A catalogue of meteorological data obtained during the Line Islands experiment February-April 1967. Nat. Center Atmospheric Res., Boulder, Colo., NCARTN-35, 362 pp. ATOLL RESEARCH BULLETIN No. 173 Preliminary checklist of the Marine Benthic Plants from Glover's Reef, British Honduras by Roy T. Tsuda and Clinton J. Dawes Issued by THE SMITHSONIAN INSTITUTION Washington, D. C., U.S.A. PRELIMINARY CHECKLIST OF THE MARINE BENTHIC PLANTS FROM GLOVER'S REEF, BRITISH HONDURASL/ by Roy T. TsudaZ/ and Clinton J. Terese INTRODUCTION The interest in Glover's Reef (16.5 N, 87.45 W), an atoll 25.6 km long and 10.4 km wide off the coast of British Honduras, has been in- tensified by its use as the planning workshop site for the Comparative Investigations of Tropical Reef Ecosystems (CITRE) program. One of the primary tasks of the individuals involved with the marine plants was the initiation of a checklist of the marine plants found on this atoll, since no published accounts were present in the literature. This paper is a summary of three collections made on the atoll by A. L. Dahl (June, 1971), C. J. Dawes (July, 1971), and R. T. Tsuda (November, 1971). Thirty-four stations were made where a total of 104 species of marine plants were collected. The checklist consists of 4 seagrasses, 8 blue-green algae, 39 green algae, 19 brown algae, and 34 red algae. The authors estimate that their present collections represent approximately 60 to 70 per cent of the number of marine plants which should occur on the atoll. Therefore, further collections are needed, especially of the epiphytic red algae. Persons interested in the des- criptive aspects of the algae listed here should consult Taylor (1960). 1/ Contribution No. 19, The Marine Laboratory, University of Guam. IMSWE Contribution No. 2. 2/ The Marine Laboratory, University of Guam, Agana, Guam 96910. 3/ Department of Biology, University of South Florida, Tampa, Florida 33620. (Manuscript received March 1972 --Eds.) Also, for further information on the description of Glover's Reef, see Stoddart (1962). All specimens collected by A. L. Dahl and R. T. Tsuda are deposited at the National Museum of Natural History, Smithsonian Institution, and those collected by C. J. Dawes at the University of South Florida Herbarium. , ACKNOWLEDGEMENTS Support for these collections was obtained for A. L. Dahl and R. T. Tsuda from the Smithsonian Environmental Sciences Program project of Investigations of Marine Shallow Water Ecosystems (IMSWE) and the NSF planning grant for the CITRE project to the Smithsonian Institution, and for C. J. Dawes from the Associated Universities for International Education (AUIE). The senior author is indebted to Dr. A. L. Dahl for the opportunity to examine his collections, to Dr. Michael S. Foster for serving as his diving companion as well as helping in certain of the collections, and to Dr. F. Raymond Fosberg and Dr. Charles Birkeland for some algal specimens included in this paper. The junior author wishes to thank Dr. Richard Keating and Dr. David Greenfield of AUIE for their support and diving companionship. STATION DESCRIPTIONS The station descriptions of the three separate collections are listed in consecutive order in terms of the dates the individual collections were made. Since all sorting, identification and processing of Dr. Dahl's collections were done by the senior author, his own number- ing system was used to identify each specimen. All of the collections were made by the respective collectors (A. L. Dahl, D. J. Dawes, or R. T. Tsuda), unless otherwise specified. Collections of A. L. Dahl, June 22-26, 1971 Station 1 - Windward reef, seaward reef slope, 5-10 m deep, 1.5 km north of Northeast Cay, June 22, 1971. (RT 4317-4331). Station 2 - Windward reef, seaward submarine terrace, 33 m deep, just north of West Entrance, June 23, 1971. (RT 4332 - 4342). Station 3 - Patch reef in lagoon, .5-1 m deep, near West Entrance, June’ 25, 197i- a(RL 4343 — Asa) Station 4 - Patch reef in lagoon, .2 m deep, near West Entrance, June: 25, £97. (i 4548) A555). Station 5 - Patch reef in lagoon, on silty slope with sand and coral rubble, 2-5 m deep, June 24, 1971. (RT 4354 - 4362). Station Station Station Station Station Station Station Station Station Station Station Station Station 10 jell WZ 15, 14 15 16 17 18 Patch reef in lagoon, on vertical surface of small algal- covered coral head, usually shaded, 1-2 m deep, June 24, 1971s ae (RE 4565 s= 2437/5), Patch reef in lagoon, reef rim, .3 m deep, June 23, 1971. (RES4S 7/605 2458): Patch reef in lagoon, top of dead coral branch, .2 m deep, June 24, 1971. (RT 4382 - 4383). Southwest outer reef, submarine cliff, 20-30 m deep, June 24, 1971. (RT 4384 - 4394). Patch reef in lagoon, center of tapered tubular sponge about 30 cm high, 3 m deep, June 24, 1971. (RT 4395). Seaward reef, submarine cliff, 25 m deep, just off Southwest Cay, June 25, 1971. (RT 4396 - 4398). Seaward reef, on coral rubble and in crevices, reef slope, 2-5 m deep, south of Southwest Cay, June 25, 1971. (RT 4399 - 4408). Seaward reef, on carbonate rock ridge in surf, intertidal zone, northeast of Long Cay, June 25, 1971. (RT 4409 - 4421). Seaward reef, on reef rock just below upper edge of submarine cliff, 25-33 m deep, outside Northeast Cay, June 26, 1971. (RT 4422 - 4429). Collections of C. J. Dawes, July 18-26, 1971 Windward reef, reef flat, 1-3 m deep, north seaward side of Long Cay, July 18, 1971. (CJD 6889 - 6900). Windward reef, 2-10 m deep, seaward side off Northeast Cay, July 19, 1971. (CJD 6901 - 6918). Windward reef, seaward side, 1.5 km south off Long Cay, JutyeZO O71. A. Reef flat, 0-1 m deep, (CJD 6919 - 6921, 6924, 6929). B. Outer reef slope, 1-40 m deep. (CJD 6922, 6923, 6926 - 6928, 6930). Windward reef, 0-2 m deep, south seaward side off Long Cay, a transect line, July 24, 1971. (CJD 6932 - 6961). Station Station Station Station Station Station Station Station Station Station Station Station Station 19 20 21 22 24 25 26 27 30 Si Patch reef in lagoon, 0-3 m deep, off northeast tip of Hong Cay, Uy 25). 0 homer (CN) TSG mm cen ACH ie Windward reef, seaward side, 26-35 m deep, 800 m north of Northeast Cay, July 24, 1971. (CJD 6977 - 6990). Patch reef in lagoon, base of coral patch, 2-10 m deep, about 3 km west of Long Cay, July 25, 1971. (CJD 6991 - 7009) . Windward reef, seaward side, 16-25 m deep, about 800 m south of Long Cay, July 26, 1971. (CJD 7010 - 7016). Collections of R. T. Tsuda, November 2-14, 1971 Lagoon, boat harbor at Long Cay, 3 m deep, November 2, 1971. (RE 4164" — 4166) ~ Patch reef in lagoon, 1-1.5 m deep, 200 m northeast of Long Cay, November 2, 1971. (RT 4167 - 4182). Patch reef in lagoon, 1-1.5 m deep, 500 m west of Long Cay, November 3, 1971. (RT 4183 - 4199). Patch reef in lagoon, 1-1.5 m deep, 1 km west of Long Cay, November 3, 1971. (RT 4200 - 4204). Northeast windward reef, November 3, 1971, collected by R. T. Tsuda and M. S. Foster. A. Lagoon slope, 2m deep. (RT 4205 - 4206). B. Reef flat, 1 m deep, (RT 4207 - 4209). C. Outer reef slope, 2-4 m deep. (RT 4210 - 4216). Patch reef in lagoon, 1-1.5 m deep, off Long Cay, November 3, 1971, collected by C. Birkeland. (RT 4217 - 4218): Beachdrift, lagoon side, Long Cay, November 3, 1971. (RT 4219). Patch reef in lagoon, 3-10 m deep, 2 km off Long Cay, November 4, 1971, collected by F. R. Fosberg. (RT 4220 - 4221). Windward reef, seaward side off Long Cay, November 5, 1971, collected by R. T. Tsuda and M. S. Foster. A. Reef flat, 1 m deep. (RT 4223 - 4224). | i | | | | B. Seaward slope, 3-5 m deep. (RT 4225 - 4226). C. Intertidal wave-washed bench at shoreline. (RT 4227 - 4239). Station 32 - Southeast leeward reef, November 6, 1971, collected by R. T. Tsuda and M. S. Foster. A. Lagoon slope, 2mdeep. (RT 4240 - 4241). B. Reef flat, 1m deep. (RT 4242 - 4243). C. Seaward reef terrace, 15-20 m deep. (RTI 4244 - 4250). D. Seaward reef slope, 25-30 m deep. (RT 4251 - 4254). Station 33 - Patch reef, lagoon slope, southeast sector of lagoon, November 9, 1971, collected by R. T. Tsuda and M. S. Foster. A. Coral substratum, 1 m deep. (RT 4263). B. Sand substratum, 3-15 m deep. (RT 4255 - 4262, 4264 - 4268). Station 34 - Southwest outer reef slope, 20 m deep, November 14, 1971, collected by M. S. Foster. (RT 4269). SPECIES LISTING DIVISION SPERMATOPHYTA (seagrasses) Hatlodule wrightit Ascherson Sta. 35B - RY 4256. Halophtla engelmannt Ascherson Sta. 33B - RT 4258. wt Syringodium filtforme Kutz. Stay 25.- Rt ANOS, Sta. 35B — RT 4257. Thalassia testudinum Konig. Sta. 23 - RT 4164, Sta. 33B - RT 4255. DIVISION CYANOPHYTA (blue-green algae) Order Oscillatoriales Calothrix econfervtcola (Roth) Ag. Sta. 25 - RT 4193 (epiphytic on Dictyosphaerta cavernosa) . Calothrtx crustacea Thuret Sta. 15 - CJD 6897. This alga forms black crusts on dead coral and limestone in the upper intertidal zone. Calothrtx partetina (Nageli) Thuret Sta. 22 - CJD 6898. Calothrtx ptlosa Harvey Stal. 1 - REAASAS Stak as) = Riera Stare 1 Shee OSI O) otc CID 69545, Stal 24 — RE any 4 Stal 26) = RA 204 sta. Gn shi ae tt Microcoleus lyngbyaceus (Kutz.) Crouan Sta. 10 - RT 4395. Rivularta nitida Bornet & Flahault Sta. 18 - CJD 6960. Schtzothrtx caletcola (Ag.) Gomont Stag 2 -REA40S, Sta. WS. CID IG895) Stan 18> (CIDEOISS = StapeolGa RT 4237 (intermixed with Sehtzothrix mextcana) . Sehtzothrix mexteana Gomont Sta. 1 - RT 4331, Sta. 25 - RT 4198 (numerous diatoms and sand grains interspersed among filaments), Sta. 31C - RI 4237 (inter- mixed with Sehizothrix calecicola) . DIVISION CHLOROPHYTA (green algae) Order Cladophorales af Chaetomorpha gracilis Kutz. Sta. 29 - RT 4219 (beachdrift). Cladophora frascatit Collins § Hervey Sta. 31€ - RY 4235b. ! Cladophora fultginosa Kutz. Sta. 18 - CJD 6958. Order Siphonales Avratnvillea asartfolia Boerg. Stak 26) -9Ri4200° Avrainvillea levts Howe Sta. 16 - CJD 6909, Sta. 17B - CJD 6930, Sta. 19 - CJD 6964, Sta. 21 - CJD 6997. Avrainvillea rawsont (Dickie) Howe Sitta.wG —) RIP4Z56. Bryopsts pennata Lamx. Sta. 31C - RT 4238 (epiphytic on Dietyota bartayresit) . Caulerpa cupressotdes (West) C. Ag. Sta. 5 - RT 4362 (war. serrata), Sta. 13 - RT 4419 (war. Lycopodium) , Stn ikh = CUD. CSUS5 Stale Ail = Ci) OSS Sey lO Se diz sehen lycopodtum), Sta. 33B - RT 4260 (war. serrata). Caulerpa mextcana (Sonder) J. Ag. Seal — CID) J005. Stal S5Br— REN Zo4 - Caulerpa mterophysa (Weber van Bosse) J. Feldmann Sta. 19) —1CIDAG975 sta 221 SCID, 7004. Caulerpa racemosa (Forssk.) J. Ag. Sta. 13)-)Rt 4414, Sta: 18 = CJD 6940; Sta. 21:- CJD 7002 and CJD 7003 (war. peltata), Sta. 22 - CJD 7016, Sta. 27C - RT 4215. Caulerpa sertulartoides (Gmelin) Howe Sta. 18 - CJD 6944. Caulerpa urvilltana Montagne Site Zee dete ite Codium intertextum Collins §& Hervey Sta. 6 - RT 4366. Codtum repens Crouan Siege IS) = (CJD) SA Codium taylort Silva sea. e 20) CID G98 1 Halimeda coptosa Goreau § Graham Sta. 14 - RT 4424, Sta. 26 - RT 4203, Sta. 27C - RT 4213. Halimeda dtscoidea Decaisne Siedee LOPE (527, Stao2) -—iRMMSser Sta. 59> RM4S565 sta. d4s- Re “44295 Sta. lop) Coy 691, Star WAS CID 6919,, RI4321% Sta..5'¢ REAS54b,-Sta-021 -\ GID 6994) Stare 2255 CJD: 7010.,8:Sta,0 50) -1RE A220. Stas SUS REA2Z46, Stas S2DAe Rt 4254, Sta. 33B - RT 4268. Udotea flabellum (Ellis §& Solander) Lamx. Sta. 5 - RT 4354a, Sta. 20 - CJD 6988, Sta-22i¥e! CID699S, StaleezieS RE 4208), Sta. SZC -e RT A244 eS tal 5ob) i Zone Udotea oectdentalis A §& E. S. Gepp. Sta. 25 - RT 4184. Order Siphonocladales Anadyomene stellata (Wulfen) C. Ag. Sta.0) SesvOeuotaer 9 eR) AS92 nota. AS — RT 44d, Stas 14. RE 4426, Sta. 25 - RT 4099, Stal 27G)- (RE, 42106. Cladophoropsts membranacea (C. Ag.) Boerg. Sta. 13 - RT 4421, Sta. 18 - CJD 6942, Sta. 31C - RT 4235a. Dietyosphaerta cavernosa (Forssk.) Boerg. Sta. 92 = RY AS55250 Sta. Si- oR 559 esta Oe PRI 4567 Sta nd 7Buc CID) 6927 > ssta. Ws) — CID G95iee Stacy 1S — CID: 6976.) Sta... 2 = CID 100i, sear, ZA Rig fee Stas 25. — Ri A1OS.4 Sta... SZB RT 4245b, Sta. 353A - RE 4263: Struvea anastomosans (Harvey) Piccone Sta. 14 - RT 4425. Valonta ocellata Howe Sta. 4 - RT 4351, Sta. 17B - CJD 6928, Sta. 19 - CJD 6974. Valonia ventricosa J. Siezis 1 = RIP Asal7/s Sta. 12 - RT 4408, Sta. 16 - CJD 6901, Sta. 19 - CJD 6975, Sta. 24 - RT 4168, Sta. 30 - RT 4221. DIVISION PHAEOPHYTA (brown algae) Order Sphacelariales Sphacelarta sp. Sta. 24 - RT 4177 (intermixed with Calothrix ptlosa). The few multiseriate filaments present lacked propagulae, thus making a specific identification impossible. Order Dictyotales Dietyopterts deltcatula Lamx. See Gr Ree Std.) SIBs = Rl 220. Dietyopterts jamatcensis Taylor Sta. 18 - CJD 6949. Dietyota bartayresit Lamx. Staal — RE AS2Zaeotde 7 - RI AS58i,) Sta. 9) = RY 4590a5, Sta. WZ = RE 4407, Sta. 13 - Ri 4416, Sta. 24 - RT,A172,. Sta... 25 -.RT 4191, Sta. 26 - RT 4202. W Dtetyota cervtcornts Kutz. Sta. 3ZA - RT 4241. 10 Dietyota dentata Lamx. Siezig. Is) = (CIID) OSIDS. Dtetyota dichotoma (Hudson) Lamx. Sita aie Ra aS 49F Stal VO CID O91) Mota alton Cio 4 8 motamen? OMe CJD 6985, Sta. 21 - CJD 7009. Ww Ditetyota divartcata Kutz. Stale Ri AS Oke stealen Ope RE AS OO, totale 5) ele AO Ocha ammenities Rip 42256 Lobophora vartegata (Lamx) Womersley Sta. 48 - Rie 458k" Stat 2 RASS stares ORT 4544) — Stan some ABS] Stas Oy RAS dee Sta. we, Riv ASS Stas, Oe ASs7 aS tae RE 4406, Stale, Sekt 4425 Stal al5 == (CJD) 68925 Sta, Lom Ciipoiler Stal.) A/7B SC JDN6925 3 Stays, SiCIDIG947.. tStaea20) — CID NG9SSreseas 25) RinAloone Stas 52D) — Rit 4255" Padina sanetae-cructs Boerg. Stale Rivas do. (Sea a/ Gaul PASii7 rota. Mle kr elo eo ect eee 68955) Sta lo) — CUD OSIAR Stale = Rie liiOr aS bden 25) leeoor Sta. 31C - RI 4231 (tetrasporic) . Padina vickerstae Hoyt Sede LES CIDEGIS ie Stypopoditum zonale (Lamx.) Papenfuss Sta. “9)-- RT 4385, ‘sta. 14 — sR) 4422... Stal. 15, CD WGsoik Staralione CID 6905 fista., 7A — CID O92 w Star LS" —"CIN6956,, Stake, Z0-eyy 6986, Sta. 21 - CJD 7012, Sta. 24 - RT 4167, Sta. 31A - RT 4224. Order Scytosiphonales Colpomenta sinuosa (Roth) Derbés § Solier Sta. 18 - CJD 6952. Hydroclathrus clathratus (Bory) Howe Stas 18) — GIDNG9SS ced SIG RA Sor Order Fucales Sargassum filtpendula C. Ag. Sta. 24 - RT 4181 (receptacles present). This species was never found attached but always floating. Sargassum polyceratium Montagne Sta. 1 - RI 4525, Sta. 2. - RY 4540, Stas 4 — RPS4546)) Sta. 0 aaa 4364, Sta. 9 - RT 4386, Sta. 13 - RT 4410, Sta. 14, RT 4427, Sta. IS, = CID 68905 iSta. 274 - CID 6929, Stal lk Ci GUS7 Rotana RT 4180, Sta. 27B - RT 4207, Sta. 32D - RT 4252. 11 '! Sargassum rigtdulum Kutz. Sta. 18 - GJD 6933. Turbinarta trtecostata Barton Sede Re Sod Ot mOM RI 4 505.1 ots Se CID O94 Turbinarta turbinata (L.) Kuntze Stdear® — Riv4355, ota. 15) - RE 44095 Sea. 15) =.GI)).6889, Sta. le = CJD 6918,.Sta- 19— CID -6962,,,Sta. 25..- RTA183. DIVISION RHODOPHYTA (red algae) Order Nemalionales Galaxaura comans Kjellman Sidon, Ries s/O, ota. 16)- CID69I5, Sta. 26.— RE AZO1. Galaxaura rugosa (Ellis § Solander) Lamx. Std.,9 — RE 4588, cca. 12 = 4404, Sta. 16 - CJD 0915, Sta. 19 = CD_.6965, Sta. 25.- RY 4192, Sta. SiR, > Ri,4225. Order Gelidiales Geltdtella acerosa (Forssk.) Feldmann § Hamel Spa. £5 — CINLo8d4, coca. 18 — CID 6945, Sta..5hG = RI 4226. Wurdemannta mintata (Draparnaud) Feldmann § Hamel Stan eles Rie 4 52 Sota Sun 4542 Sta 9> ehles4 595, ota. eG = RT 4230b. Order Cryptonemiales Amphtroa fragilissima (L.) Lamx. Sede Re aa2o sbi 2 RN AAS.) ota.) 18) CID 6957. Stas 25)— RT 4195, Sta. 28 — RY 4218. Amphtroa hancockti Taylor | Sta. 1 - RT 4326, Sta. 19 - CJD 6970, Sta. 20 - CJD 6977. Amphtroa rigida Lamx. Sta. 27C - RT 4214. Amphtroa trtbulus (Ellis §& Solander) Lamx. Stael2 Rie AGO2 ota. 20) - CID) 6978s Stas 276, — RP 4202. Fosliella fartnosa (Lamx.) Howe Sta. 23 - RT 4164b (epiphytic on Thalassta testudinum) . Halymenta agardhit De Toni Sta. 12 - RT 4400. 12 Janta captllacea Harvey Sta. 4 > RI 4352, Sta. 6 > RE 43575, Stas. US Ci nOo Soe totam ee RT 4177b (intermixed with Calothrix pilosa), Sta. 26 - RT 4204a (intermixed with Calothrix pilosa). Janta rubens (L.) Lamx. Sta on hid S65. Kallymenta limminghit Montagne Sta. 9 RI 43589, Sta. 1 - RE 4398, Sta. 54 — RE AZ69 (en gorgonian branch) . Metagontoltthon sp. Sta. 0) — RIV AS6cQeocan 7 hil Asal. These specimens represent the first record of this genus in the Caribbean region. Further studies are needed before a specific identification can be made. Peyssonnelta conehtcola Piccone §& Grunow Seals ZUG 2 IR 4lailile Order Gigartinales Eucheuma tstforme (C. Ag.) J. Ag. RT 4222 - This dehydrated specimen was found in the senior author's soup bowl. He was later told that the alga had been collected on a patch reef in the lagoon. Gelidiopsis intricata (Ag.) Vickers Stas 24 — RP 482), Sta. Zot RATO, Otdee obi te ae oe rt Hypnea sptnella (C. Ag.) Kutz. sta. 18 - CJD 6959. Order Rhodymeniales Botryocladia oeetdentalis (Boerg.) Kylin Seay eli RT T4596. Botryocladia pyriformts (Boerg.) Kylin Sta. 9 - RT 4394 (one pyriform thallus), Sta. 26 - RT 4204b (one spheroid thallus). Champta parvula (C. Ag.) Harvey Sta. 9 - RT 4390b. Order Ceramiales Acanthophora spictfera (Vahl) Boerg. Sta.’ 18° =" CJM 6950, Sta. 24.-" RT, Al75 Stan 27 Age alec. 13 Centroceras clavulatum (C. Ag.) Montagne Staeou— RE 4S47 5 ota. 27 =IRT 4206, Sta. SIG = RT 4253. Ceramiella huysmansti (Weber van Bosse) Boerg. Sta. 32B - RY 4243. Chondria floridana (Collins) Howe Sta. 12 - RT 4401. Chondrta leptaeremon (Melville) De Toni Sedo RD 44 Spauventile)), ota. 24.- RI 4176, ‘Sta. 25.-= RE MOG, (Sta SIG SER AZ27 Digenita simplex (Wulfen) C. Ag. Sta. 16) GIN 696, nota. 18) - CIDKG955,° Sta. SillC - Rh 4250: Laurencta intricata Lamx. Sta. 16 - CJD 6910, Sta. 18 - CJD 6946, Sta. 20 - CJD 6987. Laureneta paptllosa (Forssk.) Greville Stdee 4c RD 4550) sta. 6. RT AS7Zestay So RI 43582). Stalls 7- RT 4413a. Laureneta poitet (Lamx.) Howe Sta. Z1y- CID 7008. Martensta pavonia (J. Ag.) J. Ag. Sta. 1 - RT 4329 Polystphonta scopulorum Harvey Sta. 32C - RT 4245b (epiphytic on Udotea flabellum). Spyrtdta filamentosa (Wulfen) Herv. Stan Oo) — RASS. Wrangelta argus Montagne Sta. 13 - RT 4420. LITERATURE CITED Stoddart, D. R. 1962. Three Caribbean atolls: Turneffe Islands, Lighthouse Reef, and Glover's Reef, British Honduras. Atoll RES ee DUM. (Or: lols Dp. Taylor, W. R. 1960. Marine algae of the eastern tropical and sub- tropical coasts of the Americas. Univ. Michigan Press, Ann Arbor, ix + 870 pp., 80 pls. % U, S. GOVERNMENT PRINTING OFFICE : 1974 O - 561-274 0 December 31, 1974. No. 174. ATOLL RESEARCH ae BULLETIN 174. The Natural History of Pearl and Hermes Reef, Northwest- ern Hawaiian Islands by A. Binion Amerson, Jr., peeer B. Clapp, and William O. Wi irtz, I Issued by THE SMITHSONIAN INSTITUTION Washington, D.C., U.S.A. ~ ATOLL RESEARCH BULLETIN NO. 174 THE NATURAL HISTORY OF PEARL AND HERMES REEF, NORTHWESTERN HAWAIIAN ISLANDS by A. Binion Amerson, Jr., Roger C. Clapp, and William O. Wirtz, II 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, 1974 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 TABLE OF CONTENTS Page IESE C1 TIIGUINHS. 5.4 5 5a a4 Sd SO OO A OO CIO COO CICH RCHCROICRCY Cet CICLO CNC HERONCE CVcrer cur! at Ca E iO Bg PAIGE Spas oie steMe Reno ie elle evalieielelsin e's cle ieisra © 6 Sic eve sale.d eveyeyeteretas oe i foverave Vil LL IGIGSHE OUP APIO DEC NSIS 5 A aig 4 Old GG OO DIO COMO OMG OO OAC COC ce corey alnisn HAN OD) WG eON ET he eels te etame cole teteie tc a takcd a teitsieile)jeilelia\wei.s) os (e160), Sees ous ptetele miece eye levavateerare «Le DYES CRAMPS ON cs be to ts holis'selca'to’ clvsliee takes ie ous wols\ slce\ce eileviouo\eiisisice' ss ie) 6 e)ie!'s:oleleus te etadale-elereis ee erate so GHOIUCEN 4 6 5 Sic CAT 6 FRAG 46 OG 435: 0 OGG CCC TLOAA Gr COE RECTAN yea gt nd CILIUINIED 5 F8 cA Ha 06 OO 6000 diGH OOO. GOOG 6 COU CG Oe cic rete cre iD Eaeen rea LAE ig. Ante, at PS eels! a NUE elite le w whe tae lalagete rare me dohers oterace we ee CO Sepa pRB OM TiS TET i ali calza cay tee tous lente cases (oniete ie iaids telcos oo. .6ije @\0is0''s oeeeledeae wWibtanals ota eteee Ot Se eeeP EATON I OE OC RNIN MOR ie Rene eae ic alates olellSs aiowie wSaiseiare Cats oO Dieses clare ea TAT SUSE te ges anata cote vc io etlotn ye sous ie we oo ow oo wo Bi EsS RIERA ateve als iste GO Garis Salis yar veceveuaueusisustaledeustateletorekeusiorensieceleis:olalsis) sieeve: totetoiete coke iebeiereene OO. Resta ite NOab fa, sul rae eye (c ertede voy, ats ols ies eeetaue Wieiats oe SLO. 0 oie of aieele cere tO Nort esnSI aig ere yee tke ee eee AL oie oo oth hitiamelslavelepe te ataxepe Ube Beet Mise, (2. A vevete soyeleieynasteeeyotstote aslo aesiteus ceessco a0 eco wi oh DRMeioisre ole etsiem ereten kO OMENS as pal SMa oe. ee ee ieversicueseisieiieisie ie he Gene wo beaks oles Sie +O Bi islaessrepe eSNG S| 55 554515 pssojaqsyoceqesessuodsceieiesaiee sje ate valet bette tars breverrers DO SEMEN EAI MIMI pov sy spo gSiuct cy ahale Lee (ous peuecelsi,eapeyeys yacejeuetecsieloue. ous yore veue ee elelaietoleiee. afetaielatelatietoients DO SEMI PIP CIS UAH tops 5,5, «eho \Scogs anes o ayorebocet terol soles sie, ojatateceteh eke e/t w wyerereie fs OL epee sl tan, ge ae Ne IE os 050, ay oes 10\,610.0,0.0,0 0fe¥efatnfecere «cole lererent sO BBs TO AUG HA o> orn niayepeim ole cys veya e (le ropsrevete:essvess)o,eisvele.<.« attebtatofars apatousbeto lepers OO Residents Seabirds. vite. see yyetece cities sse see ent ols «lea Re pee bam kiokteiorecrwvars OF Mall ger ati, WS MORE DAT CIS. <<) svsyoressresesevelesereyeveye vevaiejeus. 6) 6, 6:0 ekitetepe tetalsieve\ cfs palaretete /'O Voerante Accidental, ands introduced Bird S\cpetepeyeyeisie efelielelcleie croketerovele((O ENeAY Abi tala Ory CCS patel fevalatotelel sf ciele| eis elerels\enaisicroferonsberote ey >) efenevoborcketeleits) cteketelctetereie (© Seaprrde Bree dain gee Cy. CS: <<) 0, oie; yee. si0) 5) ss eis a0 sel eieeitetekalartte atelate erevolonerete (Lk Wislanseig sacl Syousays IreSSoletes 5 coo coo OD DDG OOO DUO UU Ob edo Do OO aNdk Sprains. ang Summer BYCCCEIS cic, eleisiere\ aletolejereeRorekole oleteletetsveiioboveitelevercvete te (L- SuMMerwa nal Madi IPPSSCETS! 5 seve resieielsi's) oe) | s\.c:0) 6) «1 «\ elfe\oteleHetiotelatelale avefeveyele ( Haare WinniGer sre CGS). srcle ejere eisier sielel aie ess lelers ofepsvetete tele teiette: oketeropete |: ander bice edalmom CyiCWes hiejate «01 steleleyeletehe: se ctstals ia.eieic sbeteselolehe efetevensters (1 PO uENcLO Cypress - onoOD00 DON 00 DODD DODO GO DO OOD OOO OOUS Goon Soo ee os ROSTIGICING SiNSOLSS 4 SOHOOODO UD ODDO DOD OCOD OD OOO DODO OO on Obie ae pene Ms NOS Silden ms WC CCS lotslatetelea/ 66) eie)/aieleleialejeiais)«) shaletsiseteleliettohejaterejeiatovelevels (5 Miipateea Ae BA Cl as 86 ona Ya: 's a0) aie, Gay Sy 0, cleceyaieias ar eeisjack, oie: «i 4:s o, « ohare epee Ct Heplopical Distributcon Within the Atoll < .qety.lamnbemrests opelemcrententete (4: RESI dent aSeabardisn . tyes, «/heis nue, « sfalels alelelelsuateleratonetelPere e(eleicletcteietelelepere 15 Mirren Gate Oe WHC Sa, haves eyobepePopeuss =) acsie)/afelels, sy eieise) 86 6, ? as” a f E HLYON % Jon A i ee em ’ ) MARS q ae os i Seok “sh mages zy HLYON : he se ae ~ . es %, 5 Q i ee Se Eo rar) Ey, 8 a Ge So a et an ’ a “rus Wala ugrue rT.) vs 1 fn ar coe = fee \ Me (eee wig is pean Net are / I ¢ Say “fJooy soumzey pue Taeeg jo de -z Ws re" was bay > 29 FEMS q Reals hg, a! iia ot ee ae. Ssvuo. me ae daqd SANYGH GNV Tuvad PEARLanv HERMES GROUP with track of bark Gambia . Brooks Commander. Good Anchorage sutside or tv. Rise of tide 24 caches. Current NS 2kts. Seals tertile fish, fowls brakish water, old shipesod Ve vag Lys? yor { ( \\ 3. Early map of Pearl and Hermes Reef. Unpublished 1859 map by N.C. Brooks showing track of Gambia and wrecks of Pearl and Hermes (U.S. Nat. Archives, Cartogr. Div., R.G. 37). 1867 nae Pic ’ ee ue t) ir 2 g : \ . o. +, «&\ 4 a A i “AQ : a Z Pay) wie sl Reta aie wel a , eet it nc lena een Aas tee aad nautical miles ° 1 2 4 4, Map of Pearl and Hermes Reef, 1867. Redrawn from U.S. Naval Hydrographic Office chart 4, 1868. 5. Map of Pearl and Hermes Reef, 1924. Redrawn from U.S. Naval Hydrographic Office chart 4, reissued 192}. PEARL AND HERMES REEF wee NORTH |., =a 1924 Rae . a ae é ee j Pe >) x \ 2 SOUTHEAST ee SEAL |. GRAS oped Fm ae Ye RASS |. Pas eee Elm, oe TANT MD Nw mperpmeeO nautical miles ool 7 U.S. Naval Hydrographic Office chart 4 was reissued in August 1924 when additions and corrections made by the April 1923 Tanager Expedition were added. The western islands of Seal and Grass were named by the leader of that expedition, Alexander Wetmore (ms.), and Gregory (1924: 21) notes the naming of the two "hitherto uncharted" islands. This reissued version (Fig. 5) still indicated a land area southwest of North, but the four named islands had been reduced in size to their approximate present- day dimensions. No sandspits are indicated between Grass Island and Southeast Island although Wetmore (ms.) investigated this area by small boat. Tanager Expedition maps located in the Bishop Museum, Honolulu, show a small sandbar 500 yards northwest of Seal Island; this may have been the beginning of Kittery Island. The atoll was not charted again until 1930 (Galtsoff, 1933: 8). In addition to Grass, North, Seal, and Southeast Islands, Galtsoff's chart indicated two sandbars due south of North Island, three small sandbars due west of Southeast, and a small island about a half mile northwest of Seal in the position of what is now Kittery Island. U.S. Naval Hydrographic Office (USNHO) chart 5647,1 made from U.S. Navy hydrographic and topographic surveys in 1936, names Kittery Island and locates it in the position given by Galtsoff (1933: 8). Chart 5647 shows one island where Galtsoff indicated three sandspits and names it Bird. Another small island, Sand, is added west of Bird in an area where Galtsoff only indicated very shallow water. In the northeastern section, chart 5647 notes only one sandbar, awash at high tide, south of North Island. About three miles west of North, in roughly the position indicated as land by USNHO chart 4, is the notation, “sand bars awash at HW [High Water]." In 1963, when the POBSP first visited Pearl and Hermes, the sandbar south of North Island was at least 5 feet above sea level and 1,000 feet long. Four plant species were found, and a few albatrosses and Blue-faced Boobies were nesting. These data suggest that considerable changes have occurred in the topography of Pearl and Hermes Reef in the past one hundred years, and that minor changes are still occurring (see Geology section). Photographs taken by Walter F. Frear, former Governor of the Territory of Hawaii, and kindly lent to the POBSP by his daughter, and those lent by Alexander Wetmore and Paul S. Galtsoff support this hypothesis of change and provide further material for a discussion of vegetation changes in a following section. The photographs taken by Governor Frear in December 1912, though unlabeled, are probably all of Southeast Island. Wetmore has photographs of Grass, seal, and Southeast Islands taken in April 1923; Galtsoff's photographs are mainly of Southeast Island in the summer of 1930. lusNHO chart 5647 was published in February 1947; it is superseded by USCG chart 4175. Grass Island Grass Island (Fig. 6), at 27°46' N x 175°54'W is just inside the reef on the southern side of the lagoon 4.7 miles west of Southeast Island. It is 1,800 feet east to west, and only 400 feet wide at its broadest, near the western end. It has an area of 11 acres. A spur of reef rock extends westward from the eastern tip for 400 feet along the southern side of the island, but does not touch the beach. In 1965 the western end was being eroded by the sea, leaving a vertical drop of 5 feet to the water at that end of the island. Only a portion of the western end, less than 700 feet long east to west, is vegetated. The dominant plant species is the woody Solanum which grows in a dense mat to a height of about a foot in the central 400 feet of this area. Other herbs and grasses grow sparsely in the transition zone of coral rubble between the matted area and the sand beach. The rest of the island is composed of coarse to fine sand, with an area of coral rubble in the center of the eastern portion. In 1923 Wetmore (ms.), who named this island, noted that "it is about 450 yards long by 100 yards wide, 15 feet high,. surrounded by a beach of coral sand." The crest of the island was covered with bunch grass and a few of the shrubs recorded on Southeast. Cmdr. S.W. King sketched Grass Island in 1923 (Fig. 7). Kittery Island Kittery Island (Fig. 8), at 27°45' N x 175°56' W, is a low sand and coral rubble triangle less than half a mile northwest of Seal Island. It has no vegetation. Troughs eroded in the sand of the island's interior suggest that it is periodically inundated during severe weather. The island covers 11.9 acres, and its longest north-south and east-west axes are about 1,100 feet. The northwestern side is highest, about 5 feet above sea level, and the interior, southern, and eastern portions are just barely above normal high-water level. This island existed only as a small sandbar in 1923, was present in 1930 (Galtsoff, 1933: 8), and was finally named by the U.S. Navy survey in 1936. Little North Island Little North Island (Fig. 9), at 27°54' N x 175°44' W, is a half mile south of North Island. Its name was officially recognized 11 February 1969 by the U.S. Board of Geographic Names. Until then it had been nameless but was referred to by Clapp and Woodward (1968: 19-27) and Standen (1967: 29, 38) as Humphrey Island (after Philip S. Humphrey, Director of the POBSP). At low tide, it is presently less than 200 feet wide and is about 1,100 feet long in a north-south direction. The central portion of the main island, 400 feet long and 1.4 acres in area, is 6 to 10 feet above sea level, and has a meager flora of 4 species of grass and herbs. The long, low sandspit at its southern end appears to be extending itself southward and curving to the Aerial view of Grass Island, 5 June 1962. photograph. Official U.S. Navy Map of Grass Island, May 1923. Redrawn from B.P. Bishop Museum map by Cmdr. S.W. King. 0 100 200 ey yards Sand ere 7. CES, Low grass Y Long grass : : Rock 8. Map of Kittery Island, March 1965. Redrawn from POBSP map by WoO. Watricz, il. Aerial view of Little North Island, 5 June 1962. U.S. Navy photograph. Official 11 west; three small disconnected sandbars, presently just above high-tide level, extend the length of the island another 500 feet. Six to eight hundred feet west of the island is another series of sandbars which has not quite broken the surface of the water. The island may change from year to year or seasonally; however, recent observations indicate that the area is building up. North Island North Island (Fig. 10) lies in the northeastern corner of the lagoon at 27°56' N x 175°44! W, and is 10 miles north-northeast of Southeast. It is polliwog-shaped, with the tail extending southward. It has an area of 15.9 acres. The body is about 1,000 feet long north to south, and 800 feet wide; it is 10 feet above sea level. It has a steep narrow beach on its east, north, and west sides. The center of the body has a dense flora of herbs, and the remainder is a mixture of herbs and grasses, with some grasses growing among the coral rubble between the interior and the beach. There are some rotting timbers along the west side, and a few pilings in the lagoon to the southeast which were probably left by the mother-of-pearl oyster fisheries venture in the late 1920's. The tail extends southward from the body for 1,000 feet, and is about 100 feet wide throughout its length. Its proximal portion is 5 feet above sea level for 500 feet, and its distal portion is a low, shifting sandspit. One of the few previous descriptions of the island is that of Willett (in Bailey, 1956: 32), who mentions that in 1913 the island was "topped with a scanty growth of tough grasses." Elschner (1915: 62, 64) mapped it in 1914 and noted that it was "overgrown with high grass." Commander S.W. King of the Tanager Expedition also charted it (Fig. 11) in 1923. Seal Island Seal Island (Fig. 12) also lies just inside the reef, in the south- western section of the lagoon at 27°45' N x 175°56' W. It is 2 miles west-southwest of Grass Island. Seal is 1,400 feet from east to west, and 300 feet wide at its broadest point, with an area of 10.6 acres. The eastern half is composed of coral rubble and sand, with rocky ledges and reef sections on the southern side. Succulent plants grow sparsely throughout the area. This entire half is broken by numerous tidal pools and cuts in the rock. The intertidal zone on the northern side is of fine sand. A solid ledge of rock forms a northeast to southwest line across this half. The western half has a wide, sandy beach on the south, a narrow coarse rubble beach on the north, and a point of coarse coral rubble and sand curv- ing to the north at its western end. An area of the western half, measuring 600 feet east to west, and less than 150 feet wide, is about 6 feet above sea level. Almost all of the island's vegetation is confined to this area. 10. Aerial view of North Island, 5 June 1962. Official U.S. Navy L § photograph. 11. Map of North Island, May 1923. Redrawn from B.P. Bishop Museum map by Cmdr. S.W. King. : Sand me Dy 2 2 = sion ne DEPTS: 12. Aerial view of Seal Island, 5 June 1962. Official U.S. Navy photograph. 13. Map of Seal Island, May ‘1923. Redrawn from B.P. Bishop Museum map by Cmdr. S.W. King. th 0 100 200 \ Water eee : Sand yards \ 7 \ ed Coarse coral cobbles \ ; 7 4 ae rae a Y; Grass BE Solid coral rock 14 The dominant plant forms are vines and grasses. The central portion is fairly lush, giving way to a sparsely vegetated transition zone in coral rubble, and finally to beach sand. In 1923 this island, also named by Wetmore (ms.), was "elongate, 600 yards long by 300 yards wide, rising 15 feet above high tide, with a beach of. coral sand and a point of limestone rock in the east. The crest was covered with the bunch grass with much of the grayish leaved shrub." Figure 13 shows a sketch of Seal Island made at that time. . Southeast Island Southeast Island (Fig. V4), the largest of the group, lies in the eastern corner of the atoll at 27°47' N x 175°48! W. It is nearly cut into two unequal portions by a seaward extension of the lagoon. The smaller western section is connected to the eastern section by a narrow sandbar on its northern side. The entire island is 2,600 feet long east to west. Excluding a large ledge of reef rock along its southern side, the island has a maximum north to south width of 1,100 feet. It has a land area of 34 acres. The western section has a kidney-shaped central area 700 by 400 feet on which grows a sparse flora of grasses, decumbent herbs, and vines. A low flat shelf, or ledge, of reef rock, supporting a thick growth of a succulent herb, extends eastward from the southeastern corner of this sec- tion, and partially along the southern side of the large eastern portion of the island. A steep, narrow beach extends around the western and northern perimeters of the western section. A U.S. Coast and Geodetic Survey marker placed at the northwestern corner of the island in 1937 was toppled into the water in 1965 by erosion from the west. The larger eastern section is 1,800 feet from northeast to southwest, and is roughly triangular. There are up to three tidal pools, depending on tide and season, in the eastern half of this section; the depth of the brackish water rises and falls with the tide. The area in and around these pools is dom= inated by the succulent herb Sesuvium. a To the west of the pools are patches of grasses and decumbent herbs, almost completely surrounded by a large area of introduced mustard plant. From this central vegetated area to the beaches is relatively open coral sand and rock rubble with a sparse and patchy flora. The extent of these patches varies with location and season. A few small Scaevola bushes grow along the southwestern edge and a narrow sandy beach surrounds the entire section. A 15-foot-high steel frame tower near the lagoon side of the eastern section was built by U.S. Navy personnel in 1961, but by 1969 was badly = appease tc eae ennlitient 1ai though narrow and low, this sandbar has always been present. Bad storms from a northerly direction might, however, destroy it; north- westerly and northeasterly winds, on the other hand, would keep it intact. ht. oral rocks & Grass ig me} c a Sand & Grass Loy Coral rock-storm swept Tidal pools <7, CA, G; Grass 2 Tine 200 the r iew on Redrawn from B.P. Bishop 15 war (ope 4) jus Official U.S. Navy photograph. Aerial view of Southeast Island, 5 June 1962; a small-boat channel through the reef occurs Map of Southeast Island, May 1923. Museum map by Cmdr. S.W. King. 14, AUS ye 16 rusted and lacked the original wooden floor. Toward the middle of this section are two piles of rusting 55-gallon drums, and on the southwestern shore of the eastern portion is a large steel tank, the latter apparently washed up on the island by severe storms. There are little available data on early vegetative and physiographic changes on this, or any other, island of Pearl and Hermes Reef. There are apparently no notes from the ship wrecks of 1822. Brooks (1860: 502) re- ported that "the largest islands were covered with coarse grass and trees" in May 1859. Munro (1942: 12) mentions an island "about a mile long, with some low vegetation" seen during his visit in 1891. The Tanager Expedition mapped the island (Fig. 15) and on 26 April 1923 Wetmore (ms.) recorded that: the island is elongate, about 900-1000 yards long by 500 yards wide, rising 15 feet above high water. Along the western portion, the beach is of coral sand. Some- what west of the center there is a tiny, irregular lagoon. A band of blackened and eroded limestone forms the southern shore here and extends out in a broad hook to the eastward. The island rises between 10 and 18 feet at the highest point. Two ridges are covered with clumps of bunch grass one to two feet tall, and a sprawling shrub with thick rounded hirsute leaves of grayish-green eolor [Solanum] is abundant. There is a low spot with blackish soil on the eastern portion of the north side that evidently holds water after rains. It was now dry. Near this we found remains of an old camp. Upright sticks had apparently been used to support shelters of canvas, and some rubbish, iron cans, etc., lay about. Indications were that it was a camp of Japanese, perhaps from a wrecked sampan. The physiography of the island has not changed significantly in the hO years since that description, but the vegetation has been considerably altered. There is no sign of the camp mentioned by Wetmore, though there are still traces of the brief stay of the mother-of-pearl oyster fisheries --some rotting timbers, rusty corrugated roofing, and a pile of oyster shells. In calm weather, small boats can safely land on the island's southeast beach by way of a small-boat channel through the reef (see Fig. 14). The bottom near the beach in this area is coral-rock, not sand. Other Islands Bird Island (27°47' N x 175°50' W), Planetree Island© (27°k7' N x 175°49' W), and Sand Island (27°47' N x 175°52' W) are, though relatively lprobably refers to Scaevola. “Planetree Island was named by the crew of the U.S. Coast Guard buoy tender of that name in March 1964 ; it is not an official name recognized by the U.S. Board of Geographic Names. | | Li constant in position, continually changing sandspits along the inner margin of the southern reef between Southeast and Grass Islands. Rice (ms.,a) described them as "merely part of a three-mile chain of shifting sandspits just inside the south reef." On various aerial surveys in the fall and winter of 1956-57 and 1957-58 he found their number ranged from 4 to 12. They are presently nowhere wider than 100 feet, and vary in length, depending on winds and tide, from 500 to 1,500 feet in an east to west direction. POBSP personnel found no vegetation on them. A narrow, ship channel (Fig. 16) exists between Sand and Bird Islands (see also Fig. 2); its minimum depth is only 2 fathoms. A small-boat channel (Fig. 17) occurs between Bird and Planetree Islands; its minimum depth is 1.5 fathoms. GEOLOGY Today Pearl and Hermes Reef is but a group of tiny islands surrounded by a coral rim protruding above the water in the Pacific Ocean northwest of the Main Hawaiian Islands. The atoll is a remnant of a summit in a 2,000-mile range of volcanic mountains formed during the Tertiary Period. Erosion by wind, waves, and rain destroyed the volcanic domes, which in turn provided a suitable environment for the growth of coral. At equili- brium, a rim of coral remained, enclosing a lagoon on top of the peak (Stearns, 1966: 217-219; Wiens, 1962: 85-135). Gravity investigations to find the magnetic center of each of the Northwestern Hawaiian Islands by Kroenke and Woollard (1965: 363) show a +276 mgal Bouguer anomaly value on the southeast reef at Southeast Island. Gravity gradient values increase, at an average rate of 5 mgal per mile southwestward, to +285 mgal at Kittery Island. These gravity contours indicate that the highest reading, or the atoll's magnetic center, should be to the north and west of Kittery, with maximum Bouguer anomaly values possibly as high as +305 mgal. Reynolds (Anon., 1868: 272) first noted that "Pearl and Hermes Reef, like Ocean [Kure] and Brooks [Midway] Islands has a coral wall above water at its N.W. extreme." The reefs of all three are open only on the extreme west side. These features, as Standen (1967: 27-28) points out, are con- sistent with the theory of Wiens (1959: 39-40), who contends that reef growth is best where wave action is most active. Here on the northeast sector of the reef the trade winds produce the most persistent wave activity; the western side is in the lee of this activity. Standen (1967: 8, 87-89) points out that for the three northwestern- most Hawaiian Islands, which includes Pearl and Hermes, land distribution within the atoll is 99 percent in the southern half of the atoll. Further- more, he claims that land distribution is greatly affected by infrequent winter storms, with high winds from the northwest, and not by the normal Summer northeast trade winds. He concludes that this land is in the lee of such northwest winter winds, with their main axis perpendicular to the winds. 16. Aerial view of main ship channel between Sand and Bird Islands, 5 June 1962. Official U.S. Navy photograph. 17. Aerial view of small-boat channel between Bird and Planetree Islands, 5 June 1962. Official U.S. Navy photograph. 19 Standen (1967: 80-86) believes that "the shapes of the permanent islands are mere magnifications of smaller sand bars," and that "larger sections of land added...as sand spits during years with mild winters may become covered with vegetation and thus become part of the permanent island." Because of the atoll's large lagoon size, he suggests "that large [lagoon] waves breaking entirely over the small islets of Pearl and Hermes have hindered the growth of islands of substantial size." Standen further suggests that the small tidal lagoon in the depressed eastern section of Southeast Island may be an intermediate stage in the development of the island, and that eventually it will fill with debris and dry up. Similar dried areas are found at Green Island, Kure Atoll, and Kastern Island, Midway Atoll. Standen (1967: 71) analyzed the beach sand and found it to be com- posed of broken coral and sea shells; he noted that 86 percent of the particles had a diameter of from 0.5 to 4.0 mm. POBSP observers found that the soils ranged from pure coral sand and gravel on the beaches, through coarse coral rock rubble to mixtures of coral sand and some humus in the vegetated areas. The atoll does not appear to have any phosphatic guano deposits (Hutchinson, 1950: 207). Bryan (1942: 198) mentions that brackish water can be obtained by digging shallow wells. The efforts of Galtsoff (1933: 13) to find a fresh water-bearing stratum at Southeast Island were fruitless, although he found the slight depression in the eastern section briefly retained fresh water after a rain. It did not remain because of the porosity of the ground. Galtsoff (1933: 14) found that the depth of the lagoon varies from 1 to 104 feet, and at the western opening from 1 to 90 feet. Many small coral growths and reefs, all with a shallow central portion surrounded by a fringe of living coral, occur in the lagoon. Their center varies in depth from 3 to 8 feet and the bottom is covered with soft coral sand. The outer slope of the reefs supports living coral to a depth of 50 to 60 feet; past this depth coral growth is smothered by a very fine, sticky mud, with very few bottom organisms except foraminifera. Galtsoff (1933: 14) divided the lagoon into two sections on the basis of the prevailing character of the bottom; the central and northern parts are predominantly occupied by coral reefs, and the eastern and southern parts have predominantly sandy bottoms. Gross et al. (1969: 17-21) reported that the shallow (<5 meters deep) sediment-built lagoon terrace (Galtsoff's sandy bottom) occupies only 14 percent of the total lagoon. They suggest that this lagoon terrace has built leeward from the windward reef (northeastern side) and is encroach- ing upon the deeper parts of the lagoon. Thorp (1936: 109-118) studied the lagoon bottom samples collected in 1930 by Galtsoff. He found the four major constituents to be calcareous algae (48.5 percent), mollusks (17.8 percent), madreporarian corals (16.6 Pe ee 20 percent), and foraminifera (6.3 percent); the other organic groups, and minerals, silt, and clay contribute only 10.8 percent. A few volcanic rocks were mingled with the calcareous sediments. The larger of these fragments, Thorp believed, were derived from below the reef and show that the reef structure rests on a volcanic basement. Galtsoff (1933: 15) found the rocks in the northeastern section of the lagoon, several of which were 5 feet above water, to be of coral limestone, greatly resembling the fringing reef, but, as they were separated by several i hundred yards, the formation suggested an elevation of the reef or minor | changes in sea level during their formation. The outer reef is composed of large coral colonies and also solidified limestone rock.© Spaces between | and around the coral colonies are reinforced by lime deposited by the coraline _ algae, so that the entire structure is permeated and covered with a thick , layer of lime. The reef is slightly above sea level, and is covered by bar- nacles in places. : | Galtsoff (1933: 20-25) obtained temperature and salinity data from 22 July to 23 August 1930 (Fig. 18). His observations showed a gradual upward trend of surface water temperature from about 25° C. in July to about } ( 26.5° C. by the end of August; the extremes were 22.7° C. and 27.9° C. The temperature at the bottom differed only slightly from that at the surface; it was usually less than one degree Centigrade cooler (the difference at 90 feet was 1.6°C.). In comparison, Galtsoff pointed out that at-sea tempera- tures taken near Midway, 90 miles to the west-northwest, over a 10-year period varied from 18° C. in March to 25.1° C. in September, and that Honolulu fluctuations were from 22.5° C. to 25.2° C. He recorded no appar- ent trend in lagoon salinity; readings varied from 35.6 to 36.6 parts per thousand, but were usually closer to 36.0. His salinity distribution suggested that ocean water enters the lagoon primarily through the opening on the northwestern corner of the reef and the boat channel on the south side. Thorp (1936: 115) interpreted Galtsoff's data differently and wrote "apparently ocean water enters the lagoon principally CE Suge the southern boat channel and escapes through the open northwest side." Gross et al. (1969: 18, 24) examined a photograph of the area taken 9 December 1965 from the oe VII spacecraft and noted distinct plumes of discolored, pre- sumably turbid, water flowing westward out of Pearl and Hermes Reef and Midway Atoll. They suggest that the turbidity of the escaping water may be caused by suspended, silt-sized particles being carried out of both atolls. This recent finding thus supports Thorp's 1936 theory. Galtsoff (1933: 24-25) reported that the tidal range is about two feet and that the tide sets north and south with a velocity of two knots. The direction of tidal currents at specific localities within the reef is complex, and depends upon the configuration of the reef. lnadd, Tracey, and Gross (1957: 1088-1094) found a volcanic base on Midway using cone drillings. “Gross et al. (1969: 22) found the carbon-14 age of the emergent reef g | rocks at Midway to range from 1230) 4 250 hoe; 420 + 300 years. "(ne °d aqtsoddo ‘+ sans’tz :€€6T) FFOSITeN wWors uoTsstTu -Zoed UJTM peonporudsy °OC6T ysneny - Atne ‘ucoSeT 9uy utyaTM AjTUTTes pue sinzerzedme, jo uoTynqys4stp BSurmMoys yazeyO “eT SP-SLl (0S-SLI SS-SLI 00-941 . + a z 1 ott yt f <==. es S3TIN WOlLNYN Sele SS & ey o : o1 ms vse Y t USC oye recV H wor 952 ! wae’ pre i ose OS i = 52 : 02 = Eee ee ee = = Seal ee - i —— ose one y; son ! zee, ree zee" ase Vv oo co ut vee zs 7 wares 2 Ore ise eee, +92 a et" 992 Le yee opse Lae 99 | anne “Sit ove sw sae sniw suze reese 34 Ga wove Sh rs Gi "ts SE" ps7 ae 3 zp ene E8t+ 757 soc ‘Buipeas ge) 4ns 0} ssajos 1 ore *UdAIB St 24n3Y uo Aju ad94m suonEs Co] a yoedsa1 Suipeas Wo)j}oq pur Teese Oe onjans yuasaidas saanBy Jamo] pue haa o % pss aaddn ‘puesnoy) dad syed ui saniuipes Fonone ovis acre Wye << 7 4 Rati hertial tan dartorh cre ca due 9pis wuss ayy Ie saundy, Parue|s ‘ sauispeyos! yussesdos soAuN 90-82 00-p2 SP-SLi (8-821 ‘Ss-s21 00-911 vie 22 CLIMATE Climatic data for this area are available only from Midway Naval Station. There is no significant difference between the general weather conditions of the two areas. The following information is derived from a summary of the years 1953-1963 (Air Weather Service [MATS] Climatic Center USAF). The elimate is marine, influenced by marine tropical or marine Pacific air masses depending upon the season. During the summer the Pacific High becomes dominant, with the ridge line extending across the Pacific north of Kure and Midway. This places the region under the in- fluence of easterlies with marine tropical and trade winds prevailing. During the winter, especially from November through January, the Aleutian Low moves southward over the North Pacific, displacing the Pacific High before it. The Kure-Midway region is then affected by either marine Pacific or marine tropical air, depending upon the intensity of the Aleutian Low and/or the Pacific High. The monthly temperature means for the 10-year period and the range of the maximum and minimum temperatures for each month are shown in Figure 19. The temperature variation shown is indicative of a marine environment. The mean annual range is 16° F. From December through April the means are between 66° F. and 69° F., and during the remainder of the year between 70° F. and 81° F., the warmest months being July, August, and September, and the coolest January, February, and April. A 3/-degree difference exists between the absolute high of 89° F. and the absolute low Or 52° We Mean number of days with measurable precipitation and the mean monthly precipitation in inches are indicated in Figures 20 and 2l. Rain or drizzle most frequently occur from July through February, and least frequently in May and June. The mean annual precipitation for the period is 42.59 inches, with a maximum of 5.0/ inches occurring in January and August, and a mini- mum of 2.03 inches in November. A secondary high of 4.92 inches occurs in October. Combining amount of precipitation and days with measurable pre- cipitation shows May and June to be the driest months of the year. During the remaining months measurable rain falls on from 10 to 17 days. MThunder- storms have been recorded in all months except February, March, and April but peak activity seems to occur during August, September, and November. The annual average relative humidity is 76 percent, with a high monthly mean of 89 percent and a low of 62 percent. During the periods for which data are available, no tropical storm or typhoon passed through the area. Winter storms, however, are common, causing a noticeable increase in precipitation and winds, especially from September to December. Surface windspeeds and directions are shown in Figure 22. The pre- vailing wind direction 10 months of the year is easterly, and during December and January westerly. The annual mean windspeed is 10 knots, with 19. TEMPERATURE (°F) 5 g 3 a g aa Talay a ar LE se aoe a = ane Pe eas =| ar Tt gS ny a [eo Purser nane=djer ae, eer gn ES peau ao er ener BT presen le ee en Gr a) = eer Ta 9 oe Monthly temperature means and their maximum and minimum ranges for a 10-year period, 1953 to 1963, at Midway Atoll. c8 20 15 DAYS rs) J F M A M J J A S) (2) N D 20. Mean number of days with measurable precipitation for Mid- way Atoll, 1953 to 1963. 21. Mean monthly precipitation in inches for Midway Atoll, 1953 to 1963. 20 (7p) Ww a5 1) Zz 22. Wind direction and speed at Midway Atoll from 1953 to 1963. Length of directional line indicates percent of observa- tions from that direction; figure at end of directional line is mean wind speed in knots. 26 a range of 5 knots. Maximum winds occur generally from the east from i July through October, and from the west during the remainder of the year. | From May through August peaks range from 35 to 41 knots and in the re- | maining months from 42 to 55 knots. Peak gusts of 77 and 67 knots have 4 been recorded in December and January, respectively. The mean tenths of total sky cover is fairly uniform throughout the year, ranging from a low of 5.3 in August to a high of 7.3 in March. The yearly mean is 6.2. The occurrence of fog and haze is negligible, but highest in January and March. Minimum visibility caused by rain of less than 1 mile occurs rarely, most often from December through April. HISTORY 4 The atoll derives its name from those of two English whaling vessels, the Pearl and the Hermes, which ran aground at nearly the same time on the . § then unknown reef during the night of 25 April 1822. No lives were lost Z| and provisions and timber were salvaged and used to sustain the crews for | two months during which they built a schooner from the salvaged timbers. , 7 Shortly before the crews were ready to launch their new schooner, named the Deliverance, another ship--the Thames--was saved from disaster on the reef. Captain Phillips of the Hermes was able to warn her captain in time. While most of the two crews were safely taken off the reef by the Thames, » | 12 elected to sail the Deliverance into Honolulu (Hawaiian Mission Children's Society Library, Missionary Letters; Bryan, 1942: 197). The next recorded visit was that of Captain Benjamin Morrell, Jr., from 8 to 10 July 1825, who wrote of seeing "pearl-oysters and biuche de mer" as well as green turtles, seal elephants and sea leopards (Morrell, 1832: 217-218). Twenty-five years later Albert G. Osbun, aboard the brig Rodolph, visited the atoll on 11 August 1850 in search of sea turtles. Captain ~ 9 Perry and three crewmen landed on a "small island not 1 mile in circumference." They found the island covered with grass and a vine; nesting birds, seals, one small turtle, fish, and shell were noted. After killing 10 or 12 seal for food, they departed (Kemble, 1966: 154-156). Captain John Paty of the Hawaiian schooner Manokawai stopped at Pearl and Hermes in May 1857 to determine its position and map the islands (Paty, > 1857: 2-3; Bryan, 1942: 197). In 1859 Captain N.C. Brooks sailed the i Hawaiian bark Gambia there and on 5 July took possession, probably in the ® | name of Hawaii; he reported 12 islands (see Fig. 3), 6 more than Captain { Paty observed (Brooks, 1860: 502-503; U.S. Nat. Archives, Frear to Sec. a | Interior Dept. letter of 30 April 1909, R.G. 48). U.S. Naval Hydrographic ® | Office chart 4 (see Fig. 4) which resulted from the USS Lackawanna's hydro- | graphic observations in August 1867, Captain William Reynolds commanding, 7 eas shows the position of just five islands (U.S. Nat. Archives, Cartogr. Div., R.G. 37), not two as reported by Bryan (1937: 30; 1942: 197). Reynolds also took possession of the atoll for the United States.1 During the off season of sea otter hunting, the Japanese schooner Ada was charted by an American, George Mansfield, and his friends. They sailed from Yokahama, Japan, on 10 December 1881, bound first for the Bonin Islands and thence to ae Northwestern Heveihets hoping for a cargo of fish, shark, turtle and béche-de-mer. On 19 January 1882 the Ada, com- manded con, Hardy, anchored off Pearl and Hermes Reef and in the next two days her crew of 17 killed 28 turtles and collected 54 beche-de-mer and 43 pounds of albatross down. The down was obtained by killing - the chicks, dipping them in boiling water, and then stripping off the feathers; petrels, boobies, and frigates were treated in like fashion. The Ada visited the remaining islands down to French Frigate Shoals and stopped a second time at Midway in May 1882 to reprovision before returning to Japan (Hornell, 1934: 426-432). On 6-7 July 1891, Henry C. Palmer and George C. Munro, who were employed by the Honorable Walter Rothschild of England, the former as his bird collector, anchored off the Reef on the refitted schooner Kaalokai, Captain F.D. Walker in command. They did not land because of "submerged coral patches and sandbars" but described the largest island and the birds sighted (Munro, 1942: 12). Sometime during the late 1880's or early 1890's John Cameron reported pitching a tent on Pearl and Hermes, but did not remain long (Farrell, 1928: 415). During this entire period there was little political interest in Pearl and Hermes Reef. In fact, its name was omitted in various official listings of the Hawaiian Islands (Galtsoff, 1933: 11-12). On 15 February 1894, however, an indenture was entered into between J.A. King, Minister of the Interior for the Provisional Hawaiian Government, and a Hawaiian company then known as The North Pacific Phosphate and Fertilizer Company, © for the lthe question of who possessed Pearl and Hermes was resolved when Hawaii became a United States territory on 30 April 1900. The atoll became part of the State of Hawaii when the Territory of Hawaii was admitted as the 50th State in the Union on 21 August 1959 (Pearcy, 1959; U.S. Dept. of State, 1965). Presently, the City and County of Honolulu hold jurisdic- tion over Pearl and Hermes Reef by virture of Section 1717 of Chapter 118 in the Revised Laws of Hawaii dated 1925 (Morris, 1934). “The name changed to The Pacific Guano and Fertilizer Company in April 1894, but until its sale in 1904 the only island worked appears to have been Laysan (Anon., 1939: 2-22). 28 lease of Morrell, Ocean, Pearl and Hermes Reef, Midway, and French Frigate Shoals for 25 years at $1.00 per year. This company, which had been bring- ing guano into Hawaii from Laysan since 1891 (Anon., 1939: 2-22), was granted the exclusive right to remove guano, phosphate, fertilizers and other material, a royalty of 50 cents for each ton to be paid to the Hawaiian government semi-annually (U.S. Nat. Archives, King - Glade and Hackfeld indenture dated 15 Feb. 1894, R.G. 126). Beginning in 1902, Japanese feather poachers visited the Northwestern Hawaiian Islands and killed thousands of albatrosses but the extent of their poaching at Pearl and Hermes is not known. The USS Iroquois under Captain Niblack sailed past the atoll on 29 September 1904 and noted "wreckage...but no signs of human beings” (Wilder, 1905: 393). ‘Three Japanese were left there in July 1908 by a Japanese schooner; they were rescued in January 1909 by the schooner Florence Ward (Thrum, 1909: 176). By Executive Order No. 1019, Theodore Roosevelt made Pearl and Hermes a part of the Hawaiian Islands Reservation in February 1909, and placed it under the jurisdiction of the Bureau of Biological Survey, Department of Agriculture. " During the years 1910 to 1916 the U.S. Revenue Cutter Thetis visited Pearl and Hermes numerous times, sometimes landing, other times merely sailing around it. After arresting Japanese plumage hunters at both Laysan and Lisianski on 17 and 20 January 1910, Commander Jacobs proceeded to Pearl and Hermes but found "no evidence of it having been visited in recent times." On 24 December 1912 the Thetis returned and Commodore Salisbury visited "the sand islands inside the reef." In addition, Hawaii Governor W.F. Frear landed on one of the islands to take photographs (U.S. Nat. Archives, Thetis logs, R.G. 26). On 15 March 1913, the Thetis landed Alfred M. Bailey and George Willett at the northern island. In September of the following year Carl Elschner, a chemical engineer, although unable to land on the southwest island, visited the northeast island and found it "overgrown with high grass which offers attractive breeding places for the numerous Terns and Boobies.” He also observed other birds, seals, and turtles and made geological ob- servations (U.S. Nat. Archives, Thetis logs, R.G. 26; Elschner, 1915: 59- 60; Bailey, 1956: 30-31). In March 1915 and in February 1916 the Thetis checked the atoll for feather poachers but found none. During the latter visit, W.H. Munter reported birds, seals, turtles, and, on Southeast Island, rabbits. He also indicated that Southeast Island had "been recently visited by man; probably within a year and a half, judging by the conditions of the re- mains of a crude shelter that had been constructed on the north side of the island near its eastern end. They were Japanese fishermen most likely, ane 190, the preserve was transferred to the Department of the Interior. 29 for the reason that the shelter was constructed from bamboo and thatched straw or grasses....a number of upright poles was all that was left standing...[of] the shelter" (U.S. Nat. Archives, Thetis logs, R.G. 26; Munter, ms.). During April and May 1923, the Tanager Expedition visited the atoll to make scientific observations and collections for the Bureau of Biologi- eal Survey. The old Japanese camp was noted. They also reported rabbits on Southeast. In May all but one rabbit were killed and several kinds of plants and trees were planted. Two new islands, Grass and Seal, were charted in the lagoon (Gregory, 1924: 20-21; Wetmore, ms.) and the results of the expedition's observations were incorporated into U.S. Naval Hydro- graphic Office chart 4, revised August 1924 (see Fig. 5). Marine life at Pearl and Hermes was again studied in 1928 by Victor Pietschmann, a Bernice P. Bishop Museum fellow from Vienna (Bryan, 1942: 198). In May 1924 the USS Pelican, with Federal Game Warden Gerrit Wilder aboard, surveyed and photographed--with the aid of a seaplane--the atoll during its annual inspection. The north side of Southeast Island was determined to be the best place to land and beach seaplanes (U.S. Nat. Archives, Pelican log, R.G. 24; also USNHO corresp., R.G. 37). From 1926 to 1930 fishing operations became important in the history of the Reef. The Lanikai Fishing Company, Ltd., operated during 1926 and 1927 but by December 1928 had failed financially. Pearl oysters, which yield mother-of-pearl shell, had been discovered in May 1928 by Captain William B. Anderson who commanded the schooner Lanikai for the Lanikai Fishing Company. Because of its financial difficulties, the company trans- ferred its rights to the pearl beds to Hawaiian Tuna Packers, Ltd., with the two companies annually to divide the net profits. Another company, Hawaiian Sea Products Company, meanwhile, was quickly organized and on 3 December 1928 filed an application with the Department of Agriculture for a permit to establish a fishing station and to erect buildings on Pearl and Hermes. Anderson became vice president of Hawaiian Sea Products, as well as its general field manager and captain of the newly acquired schooner Lanikai. Hawaiian Tuna Packers subsequently made application in February 1929 to the Department of Agriculture for a license to develop the pearl beds (U.S. Nat. Archives, application of Hawaiian Sea Products Co. to USDA, 3 Dec. 1928; and letter from Hawaiian Tuna Packers to Sec. of USDA, 13 Feb. 1929, R.G. 22). The Department of Agriculture declined jurisdiction over the fisheries in waters adjacent to reefs and authority was given to the Governor of Hawaii to grant use and occupation permits for the fishing operations, pro- vided protection was accorded to wild animals and birds on the National Refuge (U.S. Nat. Archives, letter from R.W. Dunlap, Acting Sec. of USDA to E.C. Winston, Pres., Hawaiian Tuna Packers, 2 Mar. 1929; and copy of USDA Order signed by R.W. Dunlap, 15 May 1929; R.G. 22; memorandum on Administrative Control, 18 June 1929, R.G. 126). The newly organized Hawaiian Sea Products Company received permission to erect buildings (Figs. 23 and 2!) on Southeast Island and subsequently brought several tons of pearl shells to Honolulu. They later sold them to San Francisco and New York button manufac- turers (Bryan, 1942: 196). EE Ee 23). ok. Fishing camp of the Hawaiian Sea Products Company at South- east Island, summer 1930. Photograph by P.S. Galtsoff. One of three buildings at Southeast Island constructed by the Hawaiian Sea Products Company, summer 1930. Photograph by P.S. Galtsoff. 31 Because of the increased interest in the proposed establishment of a fishing station and cold storage plant and in the development of the pearl oyster beds, "the Territorial Government requested the U.S. Bureau of Fisheries to outline methods for [the] conservation and development" of the pearl oyster bottoms of the atoll. Acting on the Bureau's advice, the Territorial Government passed an act in May 1929 making it unlawful to take pearl oysters in Hawaiian waters and appropriated money to make a survey of the pearl oyster fisheries in Hawaiian territorial waters (Bryan, 1942: 196). This survey, conducted from 15 July to 1 September 1930 from the minesweeper USS Whippoorwill, Lt. M.M. Nelson Commanding Officer, under the direction of Paul S. Galtsoff of the Bureau of Fish- eries, spent 4 weeks at Pearl and Hermes. Galtsoff (1933: 47) recommended the prohibition of commercial fishing for at least 5 years and a resurvey of the atoll in 1935. In addition, he suggested the establishment of a pearl oyster reserve for transplantation and cultivation only, the con- tinuation of biological studies on the oysters, and the encouragement of cultivation of pearl oysters in suitable habitats by private citizens. Some fishing activity continued at Pearl and Hermes from the schooner Lanikai, but by October 1931 the fishing base operated by Hawaiian Sea Products was abandoned and the Lanikai was to be laid off (U.S. Nat. Archives, letter from F.L. Earnshaw to G.P. Wilder, 28 Oct. 1931, R.G. 22). During the mid-1930's regular inspection cruises, such as that of the USCGC Itasca in June 1934, were made to the Northwestern Hawaiians. In May 1935 the USS Lark, taking part in U.S. Naval war games, anchored at Pearl and Hermes on 15 May; she departed the next day. A hydrographic survey of the atoll was made in April 1935 by the USS Avocet, in company with the USS Quail, Tanager and Oglala. This resulted in the production of a modern chart of the atoll, but because of the political turmoil in the Pacific it was not officially published as USNHO chart 5647 until February 1947. In October 1937, another U.S. Navy exercise involved the USS Swan at Pearl and Hermes (Figs. 25, 26, and 27). Her log for the 26th records that 12 planes of Patrol Squadron 8 from French Frigate Shoals landed and tied up at temporary moorings in the lagoon; later in the day the planes returned to French Frigate. Ten planes of Patrol Squadron 4 also made a round trip visit from French Frigate Shoals that same day (U.S. Nat. Archives, Itasca log, R.G. 26; logs of the Avocet, Lark, and Swan, R.G. 2). Pearl andeHermes was involved in World War II in 1942. Acting under CINEPAC 190251 (192517), the USS Preble arrived on 19 April and at 1018 hours fired 4-inch batteries at the old fishing buildings on Southeast Island. With two U.S. Marine Corps VMF planes from Midway, they bombed and strafed the buildings; a subsequent landing party set them afire. All buildings were completely destroyed by 1408 hours when the Preble departed for Midway. No evidence was found on the island of recent habitation or landing (U.S. Nat. Archives, Preble log, R.G. 24+; U.S. Navy, Class. Oper. Archives, Preble war diary). In defense of Midway Island during May and June 1942, the oiler USS Kaloli, the civilian yacht Crystal, and the minesweeper USS Vireo were etn ae Bese Gade 6 2 One! ne Bee 3 Qrew's Mees é 4 Brady. og 5 Deck + Bos 5 i Oper OMe 6 Dock q Comp Detail EV Osficerd tent as agd Mbt 13,19, 15.5%, 17 Sad. personne} agd 1-4-6. 10 LAST < 1 Head 25. Drawing of U.S. Navy base camp at Southeast Island, drawn 6 August 1936. Official U.S. Navy photograph no. 80-CF- 79797-3 in the U.S. National Archives. 26. Chart of tender anchorages, plane moorings, and base camp for U.S. Navy exercise, 1 October 1937. Official U.S. Navy photograph no. 80-CF-79797-4 in the U.S. National Archives. Or SAE RET «onan 2 jodie ee Lege Avie BRUINS Toe *SOATUOTY TCUOTIEN °S°N 949 UT QLOTTI-D-0g °ou ydersoj0yd AAeN "a°g TetotssO ‘LE6T teqoq00 ATqeqoad ‘dweo aseq Jo sTTe dep SUTMOUS Je0F OOE WOT PUPTST JsSeayjinog fo META [etry *)2 34 deployed to Pearl and Hermes Reef. The Vireo, Lt. James C. Legg command- ing, was the first to reach the carrier USS Yorktown on 5 June 1942, after it was severely damaged during the Battle of Midway (Morison, 1949: 93, 154). Personnel of the Pacific Ocean Fisheries Investigations (hereafter re- ferred to as POFI) visited the atoll five times between June 1950 and May 1956. Aerial surveys were conducted in May 1949 by Alfred Bailey, during the 1950's by A.F. Carr, J.W. Aldrich, K.W. Kenyon, and D.W. Rice (see Scientific Visits section). On 12 March 1961, David H. Woodside and Raymond J. Kramer of the Division of Fish and Game of the State of Hawaii landed on Southeast Island and made general wildlife observations. They reported finding deep tracks made on the beach landing point by an amphibious tractor, as well as a 15- foot "steel observation tower...several 55 gallon drums, some apparently full of fuel...[and] hollow tile blocks." Subsequent investigation revealed these items had been left in 1960 by LORAC, an amphibious military operation from Midway which occupied the island without permit. The nature of this project is unknown. Also in 1961, personnel from another military project, HIRAN, which was engaged in plotting the exact location of Southeast, "landed by helicopter and camped for a few days" (HDFG, 1961). Beginning in February 1963 POBSP personnel made 12 trips to Pearl and Hermes Reef; BSFW personnel made 12 other trips. These are detailed in the Scientific Visits section. SCIENTIFIC VISITS Early information on the natural history of Pearl and Hermes Reef is available only from the reports made by various ships' captains who visited or were wrecked there. Even observations by scientists who visited the atoll prior to 1923 were limited to a few hours. Such early observations were made by Munro (1942: 12) and Palmer in July 1891, Bailey, Fullaway and Willett (Bailey 1956: 30-32) in March 1913, Elschner (1915: 59-67) in September 1914, and Munter (ms.) in February 1916. The Tanager Expedition visited from 26 through 28 April and again from 17 through 19 May in 1923. Hydrographic charts were made of the atoll and two previously uncharted islands were named. Collections were made of birds, mammals, insects, arachnids, plants, fish, and many varieties of marine invertebrates. In all, 21 species of birds including a mummified gull (Wetmore, ms.) were recorded, and the bird and mammal life was discussed by Wetmore (1925). Insects collected were reported in papers by Bryan et al. (1926), and Zimmerman (1948a, b, c, d). Eleven growing plant species and two represented in beach drift were discussed by Christophersen and Caum (1931). Myriapods were mentioned by Attems (1938). The Crustacea were treated by Edmondson (1925), the Echinodermata by Fisher (1925), and lela nes (1925) and A.H. Clark (1949), and the Foraminifera by Cushman (1925). 35 Collections of marine fauna were made between 1927 and 1930 by Captain Anderson and his associates, by Victor Pietschmann in 1928, and by Paul S. Galtsoff in 1930. Fish collections of the Tanager Expedition, as well as those of Anderson and Pietschmann, were treated by Pietschmann (1930 and 1938), Fowler and Ball (1925),1 and Fowler (1927, 1928, 1931, 1934, and 1949). Echinoderms collected from 1923 through 1930 were discussed by Holly (1932). Galtsoff (1933), who directed the studies of the pearl oyster fisheries in 1930, published identifications of several groups of marine invertebrates including Porifera, Coelenterata, Echino- dermata, Mollusca, and Crustacea, as well as fishes. His paper also discussed the oceanography of the lagoon and adjacent waters. An aerial seal survey was conducted by Alfred Bailey (1952: 16) in May 1949. Additional seal data were taken by Vernon Brock, who visited the atoll on a POFI survey in June 1950; other POFI surveys were made in May 1951,° January 1955, March 1956, and May 1956 (Ikehara, 1953; June and Reintjes, 1953; POFI, 1950, 1951, 1955, 1956a, b). Southeast Island was visited briefly in March 1961 by David H. Woodside and Raymond J. Kramer of the Division of Fish and Game of the State of Hawaii (HDFG, 1961). They made general observations on the wildlife, estab- lished photographic stations, and set up refuge signs. In addition to these landings, several aerial surveys have been made by the following personnel of the Bureau of Sport Fisheries and Wildlife: John W. Aldrich in December 1956; Karl W. Kenyon and Dale W. Rice in December 1956, January, April, and May 1957; and Dale W. Rice in October and January 1957, and January, April, May, and June 1958. Their observations were reported in three unpublished memoranda (Aldrich, Robbins, and Rice, ms.; Rice, ms. a and b) and papers on the monk seal (Kenyon and Rice, 1959; Rice, 1960a) and the albatrosses (Rice and Kenyon, 1962). An aerial turtle survey was conducted by Archie Poycare (ingMitt,) in January 1962, POBSP personnel spent 52 days at Pearl and Hermes Reef on 12 separate trips beginning in February 1963. BSFW and HDFG personnel were present on five of these. BSFW personnel spent at least an additional 43 days on 12 other trips. Dates and islands visited for each trip are listed in Table l. Personnel for these surveys, as well as all others, are presented in Appendix Table 1. POBSP and BSFW personnel banded thousands of birds and obtained many recaptures through 1969. Wildlife population estimates and notes on breed- ing status are available from each trip; all islands were not visited on each trip and on some islands no nocturnal activities of the avifauna were lranager collection only. “The George Vanderbilt Pacific Equatorial Expedition, concerned primarily with marine life, did not stop briefly at the Reef in 1951 as implied by Bailey (1952). x°4desg Tg “TEN oo “any QT “TeN +1 der LeueTd \O (99) “TEN oc “any QT "Ze 41 "IeK G p4ta *qdag TZ “TEN co “nV QT "TEN +L “Te G pues “deg gg “ZEW QT Cavols}s} Ait “any 61 eunLe Ge-€2 “IEW 9 49-10 eTV4TI *qdeg gg - rEW QL-LT *qdeg JT “Sny 0-61 sun Ge-€2g "IEW 9 W-20N “qdeg TZ “TEN dg "IeW QT *adag Tg “TEN gg * TeW 61-91 “any QT “TeW +1 aunP 9Z "TeW T8285 *qdeg TZ “TEN Ce “Ie 61 “ony QT "ZeW 41 aune 12-92 “IEW SG ssery 92-02 *Idy T “TEN co-ed - TOH 61-ST *qdeg OT “sny 61-91 Tey +1-€T aune og *€S-9L “IEW Q -"qad 92 qseoygnog MISH 996T dsd0d S961 MISd 496T dsd0d €96T Teak jeoay souzey pue [teed 1 spueTST go seqep Aeamms mMASG pue dSdOd °T eTGeL *qdeg TT *qdag ZT Ken 92 "rey TE eu Z1-OL *qdeg gZ *qdeg 9g *qdeg J2 “any 6¢ x ony 6c x ony 6g “ony 6¢ Ken TE Kew TE sor}, 0UeTd patg pueg U420N aTV44I *4dag 21 Ken 9g °TeN TE "qo 2T-OT “qdeg Je “ony O€-62 Y9-1ON “qdeg TT Key 92 gag ot-OL IEW +2 *4dag gz “sny 62 ATne 6 Aen TE 349TH *qdeg TT Ken 92 “qu eL-OT "ZEN He *qdeg gg “any 62 ATnL 9 Aken TE T8295 *f[TUO UuoTZeATESqO azZOUsSTIOx *q4dag *4d99 TT 61-OT ATnL gg aune ¢ Ley Key 92 TE-92 “ady g -"TeW TE *qad ZT-OT “any 9g * TEN “TEN td td-d¢ *qdas *q4desg gz 6c- 12 e Ssny *Sny 62 O€-92 ATNL Tue g 6-€ aun T Kew TE -Kew oF ‘ez ssezy 4seay nos (penutzu0o) MASa MASE MAS MASE MASH Masa Masa asdod “Masa MASE asdod MaSd dsdod qared aAINS TeaK 696T. g96T 1961 “T 9TdeL 38 observed. All of the islands were mapped and color and black and white photographs were taken showing the appearance of each. Papers resulting from POBSP activities at Pearl and Hermes Reef were published by Amerson | (1968); Amerson and Emerson (1971); Brennan (1965); Clapp and Woodward (1968); Kohls (1966); Kohls and Clifford (1967); Kohls, Sonnenshine, and Clifford (1965); Maa (1968); Sibley and McFarlane (1968); Standen (1967); Tsuda (1966); and Yocom (1965). In all, 106 scientific publications and manuscripts have resulted from the various surveys to Pearl and Hermes Reef. An annotated list of these is presented in Appendix Table 2. VEGETATION Osbun first recorded vegetation in August 1850; he noted an island "covered with coarse grass & a vine bearing prickly pod" (Kemble, 1966: 155). Brooks (1860: 500) next recorded vegetation in July 1859; he found the largest islands covered with coarse grass and trees. Bitter (1900) identified two species--Eragrostis hawaiiensis [=variabilis], and Solanum laysanense [=nelsoni]--from photographs taken by Walker in 1899. cin ee ee The vegetation was first detailed in a report by Christophersen and Caum (1931: 15-16, 20-41), who discussed the 11 species of vascular plants found growing by the Tanager Expedition in April 1923, and two species represented by seeds found in beach litter. More specimens were collected by Galtsoff in 1930. Photographs by Frear, Wetmore, and Galtsoff make it possible to reconstruct the vegetative history of the atoll in the present century. Vascular plant specimens were collected by Sibley in June 1963 and by Long in September 1964; other POBSP personnel have taken notes and photo- graphs on most visits. The islands and their major vegetation associa- tions were mapped in March 1965. Specimens have also been collected by Charles H. Lamoureux (date unknown) and Allen H. Young (August 1964) of the University of Hawaii, POBSP cooperators in botany. Lamoureux and Long prepared the basis for the annotated list of species presented in Appendix Table 3. Plant specimens may be found in the herbaria of the National Museum of Natural History (USNM), the Bernice P. Bishop Museum (BPBM), and the University of Hawaii (UH). Vascular Plants In all, 25 species of vascular plants, representing 17 families, have been recorded from 5 islands (Table 2 and Appendix Table 3). One species, Sicyos caumii, is endemic (St. John, 1970: 448-451). The following dis- cussion of the flora is based on all previous botanical accounts, as well as the data of the POBSP. Wherever plant associations are discussed, the species are listed in the order of decreasing abundance. Islands are listed in alphabetical order. 89 Table 2. Vascular plant distribution at Pearl and Hermes Reef Little Species Southeast North Grass Seal North Cynodon dactylon Eragrostis variabilis Lepturus repens Setaria verticellata Allium sp. Cocos sp. Pritchardia pacifica Casuarina equisetifolia Achyranthes splendens x x x Boerhavia repens Sesuvium portulacastrum Portulaca lutea Capparis sandwichiana x Brassica campestris Coronopus didymus Lepidium bidentatum Tribulus cistoides Malvastrum coromandelianum Hibiscus tiliaceus Tournefortia argentea x x x Solanum nelsoni Solanum nigrum Sicyos caumii Scaevola taccada Sonchus oleraceus MO Pt OM Od OM OM OS Mm OM Pas ba Pe teed ta Mm MM OM OM Pa ba Pa MM OM OM be te Total 22 12 aL ILL hh Grass Island Eleven species of vascular plants have been recorded from Grass Island (Table 2). Only a portion of the western end, 2.6 acres, is vegetated (Fig. 28). The interior of this vegetated portion is a dense patch of Solanum nelsoni about 400 feet long east to west, and 100 feet wide which grows to the height of a foot. Between the Solanum and the sandy beach is a sparsely vegetated area of Lepturus repens, Tribulus cistoides, Boerhavia repens, and §. nelsoni. About a dozen clumps of Eragrostis variabilis grow among the Solanum, An April 1923 photograph by Wetmore shows primarily dense Eragrostis (Fig. 29). Wetmore (ms.) noted that "the crest of the island was covered with bunch grass and a few of the shrubs recorded on Southeast Island." Eight species were recorded by Christophersen and Caum (1931: 15-16). The vegetation was dominated by Eragrostis, which was restricted to the central parts. lLepturus grew in a fringe around the Eragrostis. Two plants of SS Lepturus, Tribulus, Boerhavia NS (eae ee & Solanum nelsoni HH Solanum nelsoni iS Eragrostis Feet 28. Vegetation map of Grass Island, March 1965. Redrawn from POBSP map by W.O. Wirtz, II. 29. Grass Island covered with dense Eragrostis, 27 April 1923. Photograph by A. Wetmore. 30. Interior of Grass Island, 27 June 1963. lLaysan Albatross chicks (foreground ) in area of Setaria, Boerhavia and Tribulus; Great Frigatebirds nesting in Solanum. POBSP photograph by A.B. Amerson, Jr. he Achyranthes splendens were found. Boerhavia and Tribulus were present, but not abundant, and Lepidium o-waihiense [=bidentatum] was rare. A few small Scaevola taccada and Tournefortia argentea bushes were present. Solanum laysanense [=nelsoni] was present though not abundant. In 1963, the interior (Fig. 30) was covered by a dense mat of Solanum nelsoni. Lepturus formed a thick cover on the west end of the vegetated area, Lepidium was found along the southern side of the island, and Solanum nigrum was common. In all, 20 to 30 clumps of Eragrostis were found, and Boerhavia and Tribulus were widespread. Scattered clumps of Setaria verticellata were noted and there were no longer any plants of Scaevola or Achyranthes. In 1965 the Setaria was not found and the number of Eragrostis clumps had been reduced to 14. The vegetation change hag brought about a change in the bird nesting colony as well. For example, in April 1923 the Great Frigatebird (Fregata minor) was not breeding (Wetmore, ms.); in recent years 300 to 350 Great Frigatebirds roost in the Solanum, and 75 to 100 nest there. : } : The change from grasses to herbaceous creepers is, thus, well-documented. | Little North Island | Only four vascular plant species are known (Table 2); 0.5 acre is vegetated. The raised central portion, about 400 feet long north to south, presently supports a meager growth of one grass and two prostrate herbs GHgensi): Lepturus repens is represented by several clumps, small seed- lings of Boerhavia repens were found, as were a few sprigs of Tribulus cistoides. Small Tournefortia argentea plants were noted in June 1963, but were not found in March 1965. Three species were still present in August 1967 (Figs. 32 and 33). North Island Twelve species of vascular plants have been recorded (Table 2); 7.1 acres are vegetated. The center of the island is presently dominated by a lush growth of Sicyos caumii (Fig. 34). Some Boerhavia repens and Tribulus cistoides also grow in this area. At the northern perimeter is a nearly solid growth of Solanum nelsoni. East of this growth is an area of mixed Tribulus, Boerhavia, and S. nelsoni. At the northeast corner is a nearly pure stand of Eragrostis variabilis about 100 by 200 feet. The remainder of the vegetated area is a sparse mixture of Lepidium bidentatum, S. nelsoni, Sicyos, Tribulus, and Boerhavia, except for a nearly pure stand of Lepidium along the western side. Plant density decreases from the interior to the beaches, and Lepidium is the last species to be found in the coral rubble closest to the beaches and part way south on the southern tip. Several stunted and nearly dead Tournefortia argentea bushes were found along the southwestern edge of the island in March 1965. Several areas of Brassica campestris were found in August 1967. Other plants recorded in small numbers are Lepturus repens, Portulaca lutea, and Scaevola taccada. In March 1913 Willett (Bailey, 1956: 32) stated that "tussocks of } | bunch grass...covers [sic] the northern end of the island." Christophersen 1 NS Lepturus, Boerhavia & Tribulus | 31. Vegetation map of Little North Island, March 1965. Redrawn from POBSP map by W.O. Wirtz, II 32. Sparse vegetation at Little North Island, looking southeast (print reversed), 29 August 1967. POBSP photograph by C.A. Ely. ee aa gS,” NE A A 33. Scant vegetation at Little North Island, looking northwest (print reversed); North Island in background, 29 August 1967. POBSP photograph by C.A. Ely. Sand Coral rubble with Lepidum Lepidium, Tribulus, & coral rubble HE Solanum nelsoni NS Tribulus, Boerhavia, & N Sn aaa Solanum nelsoni Eragrostis Lepidium NY Sicyos, Boerhavia, & NN Tribulus 0 500 Feet 34. Vegetation map of North Island, March 1965. Redrawn from POBSP map by W.O. Wirtz, II. 46 and Caum (1931: 15-16) report that the Tanager Expedition collected Achyranthes splendens and Tribulus in 1923. As there are only a few patches of grass at present, with several species of creepers dominating the flora, apparently this island has under- gone the same type of successional changes as those on Grass and Southeast Islands. Seasonal changes also occur; the vegetation, in general, was more lush in August 1967 (Figs. 35 and 36) than in June 1963 (Figs. 37 and 38). Seal Island Eleven species of vascular plants have been recorded (Table 2); 2.5 acres are vegetated. The central portion of the western half, about 600 feet east to west and 150 feet wide, contains most of the vegetation (Fig. 39). The interior of this vegetated area is primarily Sicyos caumii, Solanum nelsoni, and Eragrostis variabilis (Fig. 40). Between the densely vegetated area and the beach is a transition zone of Tribulus cistoides, Boerhavia repens, Lepturus repens, and Eragrostis (Fig. Li). Growth of all but the Eragrostis extends out among the beach rubble. One clump of Achyranthes splendens grows at the eastern border of this vegetated area. Small Sesuvium portulacastrum plants are scattered on the rocky ledges of the eastern half. a In April 1923 Wetmore (ms.) noted that "the crest was covered with the bunch grass and much of the grayish leaved shrub" (Fig. 42). Christo- phersen and Caum (1931: 15-16) reported 11 species in 1923. Eragrostis and Lepturus were distributed indiscriminately (Fig. 43), Achyranthes was common, and Boerhavia. and Tribulus were present. Large flourishing plants of Sicyos grew on the eastern half of the western section (see also St. John, 1970: 450). Solanum nelsoni was present, though not abundant, and Lepidium o-waihiense [=bidentatum] was rare. Sesuvium grew on the raised reef and low wet flats of the eastern portion. Capparis sandwichiana was scattered over much of the island except for beach and reef. A few small Scaevola taccada bushes were also found. The same 11 species were present in 1963. Only one clump of Capparis and one Scaevola bush were found. Tribulus and Boerhavia were widespread, and Lepidium was common in the transition zone from bunchgrass to beach. There were 40 to 50 clumps of Eragrostis, Lepturus and Solanum nelsoni were common, and Sicyos was present. Sesuvium still occurred on the rocky and wet areas of the eastern portion. About five plants of Achyranthes were noted. Southeast Island Twenty-two vascular plants are known from Southeast Island (Table 2); 27.2 acres are vegetated. The vegetation is presently dominated by 10 species, two of which were introduced (Fig. 44). Eighteen species, including an onion (Allium sp.) growing on the refuse heap which was eradicated in March 1963, have been found by the POBSP. — 36. Low vegetation at North Island, from west-center looking north (print reversed), 29 August 1967. POBSP photograph by C.A. Ely. Creepers and a few patches of grass dominate the vegetation at North Island, from west-center looking east (print re- versed), 29 August 1967. POBSP photograph by C.A. Ely. 37. Extreme low vegetation at North Island, from west-center | looking north, 24 June 1963. POBSP photograph by A.B. Amerson, Jr. 38. Extreme low vegetation at North Island, from west-center looking south, 24 June 1963. POBSP photograph by A.B. Amerson, Jr. <- Tida flat Beach rubble & + Lepturus SS SS ° ° . ia, Tribulus, Boerhav 1s Lepturus,& Eragrost ‘, nelson Ane) yos, Solanum Tom Sic is ros Era 500 Redrawn from w o% ac c te) - ~ dl < ee UhNN Feet Vegetation map of Seal Island, March 1965. POBSP map by W.O. Wirtz, II. 39. ho. Sicyos, Solanum, and Eragrostis cover the interior of Seal Island, view looking south, 14 March 1964; Laysan Albatross adults and chicks. POBSP photograph by A.B. Amerson, Jr. 41, Transition zone of Tribulus, Boerhavia, Lepturus, and Eragrostis between dense interior and beach at Seal Island, view looking southeast, 14 March 1964. POBSP photograph by A.B. Amerson, Jr. 43. Grasses and low shrubs cover the crest of Seal Island on 2/7 April 1923; albatross chicks in foreground, Great Frigate- birds in flight. Photograph by A. Wetmore. Seals and albatrosses among scattered clumps of grass at seal Island, 2/7 April 1923. Photograph by A. Wetmore. ic ey a ca ay = 2 = > 2 ae o}| © > Sis > oS cu Pay £ €| > a 3] 9] 5 S} u 2 9 E) cs ft] SI c| c} -c — v > 6] 2 | s+ 2 Bee see el) clee see — =| a|.2] = ae! Ol Wal nial & ES don, & Cyno . Sicyos WN Brassica NS 4s} hy A 2254 z 4 Coronopu icyos /&S rostis Erag Hi n uit 500 Feet Redrawn Vegetation map of Southeast Island, March 1965. from POBSP map by W.O. Wirtz, II. hd, 53 The central portion of the western section (Figs. 45 and 46) has a sparse vegetation of Solanum nelsoni, Coronopus didymus, Boerhavia repens, Tribulus cistoides, and Lepidium bidentatum. Several clumps of Eragrostis variabilis grow to a height of about three feet near the center of this area. The ledge of reef rock extending southeastward from the western section has a large patch of Sesuvium portulacastrum in its interior. A patch of Sesuvium also grows on the reef rock extension at the southeastern corner of the eastern section. The area in and adjacent to the tidal pools in the eastern half of the eastern section is dominated by a lush growth of Sesuvium. West of these pools is a large area dominated by Coronopus. A large patch of Eragrostis and a smaller one of Cynodon dactylon, presumably introduced, grow adjacent to each other in the center of the Coronopus area. A few plants of Sonchus oleraceus and Solanum nigrum grow among the Cynodon and Coronopus. Surrounding the Coronopus area on the north and west is a steadily expanding area dominated by an introduced mustard, Brassica campestris. The area also has some Cynodon and some Sicyos caumii. From this central, heavily-vegetated area out to the beaches is a relatively open coral sand and rubble region with patches (Fig. 47) of Solanum nelsoni, Tribulus, Boerhavia, and Sicyos. The extent of these patches varies with location and season. Solanum nelsoni is especially dominant on the southern side and Tribulus on the north. Tribulus and Sicyos (Fig. 48) are mixed with Solanum nelsoni on the southern two-thirds, and Tribulus and Boerhavia are mixed with §. nelsoni on the northwestern side. A northern area dominated by Tribulus has one patch of pure Sicyos and a larger patch of Eragrostis. A second clump of this grass grows further east in the §S. nelsoni-Tribulus-Sicyos association. Several very stunted Scaevola taccada bushes grow along the margin between the vegetated area and the beach on the southwestern and southern sides of the eastern section. Other plant species recorded in small numbers are Setaria verticillata, evidently introduced in 1961 (HDFG, 1961); Lepturus repens; Portulaca lutea; Malvastrum coromandelianum; and Casuarina equisetifolia, introduced in 196 mostly dead in 1965, and absent in 1969. The photographs by Frear, taken in December 1912, indicate that the flora was chiefly tall bunchgrass, presumably Eragrostis (Fig. 50). Eleven years later, in April 1923, the photographs of Wetmore show the island to have a moderate, open cover of Eragrostis (Fig. 51). Sesuvium grew abundantly on the large reef rock ledge. The Tanager Expedition col- lected six species (Christophersen and Caum, 1931: ie-ie). Eragrostis and Lepturus were distributed indiscriminately on the island. Sesuvium grew on the reef rock areas and in the mud flats around the ponds at the eastern end of the island. Boerhavia was present, but not abundant, and Tribulus was present. Sicyos was represented by a few plants (see also St. John, 1970: 451). One seed each of Mucuna gigantea and M. urens was found on the beaches. K6, Laysan and Black-footed Albatrosses nesting among Solanum, Coronopus, Boerhavia, and Tribulus on western section at Southeast Island, view looking southwest, 13 March 1964; USCGC Planetree in left background. POBSP photograph by A.B. Amerson, Jr. Low vegetation on western section at Southeast Island, view looking southeast across interior toward reef-rock area, 13 March 1964. POBSP photograph by A.B. Amerson, Jr. 48, Eragrostis, Coronopus, Tribulus, and Boerhavia growing in the west-central portion of the eastern section at South- east Island, view looking west, June 1967; Laysan Albatross adults and chicks and Sooty Tern adults; U.S. Navy=built tower in center background. POBSP photograph by R.L. DeLong. Tribulus and Sicyos in the west-central portion of the eastern section of Southeast Island, 13 March 1964. POBSP photograph by A.B. Amerson, Jr. Black Noddies roosting in stunted Scaevola bushes on the south shore of the western portion of the eastern section at Southeast Island, view looking northwest, 28-30 August 1967. POBSP photograph by R.B. Clapp. 50s ibe Tall bunchgrass, presumably Eragrostis, growing on the crest of Southeast Island, December 1912; Hawaiian Monk Seal fore- ground, Black-footed Albatross nesting below crest. Photo- graph by W.F. Frear courtesy of Virginia Frear Wild. Open cover of Eragrostis at Southeast Island, 26 April 1923. Photograph by A. Wetmore. 58 In May 1923 the Tanager Expedition planted hau, Hibiscus tiliacea, and palm trees, Pritchardia pacifica, and scattered seeds (Gregory, 1924: 21; Wetmore, ms.). Though no record remains of what seeds were scattered, Wetmore has a list of the species given the expedition by the Territory of Hawaii and the Department of Agriculture for planting on the Reservation to replace vegetation destroyed by rabbits. With the exception of Scaevola none of the species on this list is presently growing on Southeast Island. Photographs taken by Galtsoff in 1930 show that the eastern section was primarily dense Eragrostis (Fig. 52) with some areas of Lepturus and Boer- havia. A photograph of the buildings of the pearl fisheries venture shows a Casuarinaabout five feet tall, many plants of Sonchus, and an unidentifi- able composite which is apparently no longer present. One view of the western section (Fig. 53) shows a flora of scattered Boerhavia and very sparse Lepturus. Galfsoff (1933: 16) mentions the planting of Casuarina and Cocos sp. in 1928, but further states that they were all dead or dying by 1930. Woodside and Kramer (HDFG, 1961) reported finding introduced Setaria verticillata in March 1961. Ironwood (Casuarina) trees were planted by the U.S. Navy sometime in 1963 to increase the islands' visibility from the ocean. As this was in violation of Refuge regulations, all trees which were not already dead were destroyed in 1964. Setaria was reintroduced with the ironwoods in 1963. A significant change has occurred in the flora since it was first described in 1923 and 1930 (Figs. 54 and 55). Eragrostis has been reduced from the status of a major plant cover to one of insignificance. Only a few isolated clumps remain on the eastern section, which in 1930 was nearly solid grass. Possibly the major vegetation change occurred in late 1930 when George Kaufman reported no live vegetation, only tall clumps of dead bunchgrass, after a severe storm (Fisher and Baldwin, 1945: 11). As information on plant succession in the Northwestern Hawaiian Islands is very limited, it is not known whether the floral change observed, from grasses to herbs and vines, is a result of natural succession or caused by some edaphic catastrophe. Other Islands POBSP and BSFW personnel found no plants on Bird, Kittery, Planetree, or Sand Islands on visits between 1963 and 1969, nor was vegetation listed for these islands in earlier reports (Wetmore, ms.; Galtsoff, 1933: 13-14; Christophersen and Caum, 1931: 15-16, 20-41). REPTILES Only one reptile--the Black Turtle of the Pacific (Chelonia agassizi)-- has been recorded. Perhaps the Pacific Hawksbill Turtle (Eretmochelys im- bricata), an uncommon species in the Hawaiian Islands, has visited the atoll, but no substantiated record exists of its occurrence. Common and scientific names follow Carr (1972). Dae Bureau of Fisheries personnel inspect Sooty Tern colony in dense Eragrostis of the eastern section at Southeast Island, summer 1930. Photograph by P.S. Galtsoff. Boerhavia and Lepturus growing on the western section at Southeast Island, view looking south, summer 1930. Photo- graph by P.S. Galtsoff. 54. Creepers and low shrubs cover most of the eastern section at Southeast Island, view from U.S. Navy-built tower looking east, 14 March 1964. POBSP photograph by A.B. Amerson, Jr. 55. Dense creepers, low shrubs, and some grasses cover most of the central portion of the eastern section at Southeast Island, view from tower looking east-northeast, 14 March 1964. POBSP photograph by A.B. Amerson, Jr. 61 BLACK TURTLE Chelonia agassizi Status Uncommon resident breeder; a few probably present year-round; frequents Little North, North, and Southeast Islands. Maximum recent POBSP and BSFW population estimate 55 in September 1966. Observations Morrell (1832: 218) first recorded sea turtles in 1825; he wrote that "great numbers of green turtles are found on the sand-beaches, where they come to deposit their eggs." He further noted that "the hawk's-bill turtle, also, sometimes visits this place, but in small numbers." Osbun (Kemble, 1966: 155) in August 1850 caught one small sea turtle on the reef, but saw "no Turtle nor the signs of any" on shore. Turtles were next recorded in 1857 by Paty (1857: 2-3) who wrote that the lagoon "seemed to abound with...turtle." Two years later Brooks (1860: 502) noted "plenty of...turtle," and in 1882 the crew of the Japanese schooner Ada killed 28 (Hornell, 1934: 432). Turtles were also reported to be plentiful in 1913 (Bailey, 1956: 30) and 1914 (Elschner, 1915: 60). Munter (ms.) noted eight on the beach in February 1916. Wetmore (ms.), however, did not observe them in April 1923. POBSP and BSFW personnel recorded turtles on most survey trips since early 1963. All turtle observations are summarized in Table 3. Table 3. Black Turtle observations at Pearl and Hermes Reef Population Date of Survey Tsland Estimate 3reeding Status, Remarks, References 1825 68-11 July ? great On beaches; notes eggs (Morrell, numbers 1832: 218). 1850 11 Aug. Southeast? O Saw "no Turtle nor the signs of any” on the island but caught "one small Green Turtle on the reef" (Kemble, TO 6ey 155). 1857 19-20 May ? 8 Lagoon abounded with turtle (Paty, 1857: 2-3). 1859 i duly ? % Plenty of turtle (Brooks, 1860: 502). 1882 19-21 Jan. 2 28 Killed by crew of schooner Ada (Hornell, 1934: 432). 1913 15 Mar. North ? Many large Green Turtles (Bailey, 1956: 30). 62 Table 3. (continued) Population . Date of Survey Island Estimate Breeding Status, Remarks, References 1914 Sept. North ? Good many turtles (Elschner, 1915: 60). 1916 4 Feb. Southeast 8 Hauled out on beach (Munter, ms.). 1923 26-28 Apr. 2 0 Did not record turtles (Wetmore, Msi) ie 1930 22 July- ? ? Notes presence of large amount of 23 Aug. algae (Codium) in turtle stomachs (Galtsoff, 1933: 16). 1950 27 June Southeast 12 Tagged (POFI, 1950). 1956 22 Mar. Southeast yy Counted (POFI, 1956a). 26 May Southeast 3 (POFI, 1956b). 26 May North il Digging nest (POFI, 1956b). MOS 4 Southeast 20-50 Basking on north beach (Parsons, 1962: 69-70). ? North 10-20 Basking on south beach (Parsons, 1962: 69-70). 1958 ? Southeast 20-50 Basking on north beach (Parsons, 1962: 69-70). 2 North 10-20 Basking on south beach (Parsons, 1962: 69-70). 1961 12 Mar Southeast } Adults; copulation observed in shallow water; no sign of egg laying (HDFG, 1961). * 1962 Jan Bird and 30 Seen from aerial survey; 8 9 bask- Sand raves ((Cehevae alla, Guibas, )) 2 1963 26 Feb.- Southeast is On beaches, mostly at night 8 Mar. (POBSP, 1964d). 6 Mar. North "le On east beach cove (POBSP, 1964d). *Between 14 and 28 January. 63 Table 3. (continued) Population Date of Survey Tsland Estimate Breeding Status, Remarks, References 1963 18-23, Southeast 20 On beaches, mostly nocturnal (POBSP, 25 June 1963). 23-25 June North 9 On beaches (POBSP, 1963). 23, 2 June Little North 2 On beaches, 1 15" yearling caught (POBSP, 1963). 1964 13-14 Mar. Southeast 8 Adults; 2c and 5 2 tagged (BSFW, 1964a; POBSP, 1964b). 16-19 Aug. Southeast 12 Noecturnally (POBSP, 1964a). 19-20 Aug. North 10 Nocturnally (POBSP, 1964a). 16 Sept. Southeast 31 Adults; 5 o and 14 9° tagged; nest pits, but no eggs (BSFW, 1964b; POBSP, 1964c). 17 Sept. North 2 Small 2' x 2' turtles; 8-10 pits, no eggs, 1 3" mummified young (BSFW, 1964b; POBSP, 1964c). 17 Sept. Little North 2 Adults; 1 @ on beach (BSFW, 1964b). 1965 15-19 Mar. Southeast 12 Counted on 15th (POBSP, 1965b). 17-18 Mar. North 5 Counted on 17th; 1 mummified hatch- ling (POBSP, 1965b). 21-22 Mar. Southeast 5 1o and 1? tagged; 1 recaptured (BSFW, 1965; POBSP, 1965a). 1966 1 Apr. Southeast 12 3 oc and 6 @ tagged (BSFW, 1966a). 20-26 Sept. Southeast 50 4h tagged, 6 recaptured (BSFW, 1966b). 22 Sept. North 2 Adults, 1c and 1 9 tagged (BSFW, 1966b) . 22 Sept. Little North 3 Adults, 2c and 1 2 tagged (BSFW, 1966b) . 1967 16-17 Mar. North 9 Counted on 16th (BSFW, 1967d). 64 Table 3. (continued) || ait Population || Date of Survey Island Estimate Breeding Status, Remarks, References | 1967 17 Mar. Southeast 8 Counted (BSFW, 1967d). | 21-23 Mar. Southeast 17 1 o and 10 2 tagged; 1c and 5 2 recaptured (BSFW, 1967a; POBSP, 19674). 28, 30 May- Southeast 5 h large, 1 small; copulation in 1 June water (POBSP, 1967b). 28-30 Aug. Southeast ? Present (POBSP, 1967a). 29-30 Aug. North 3-5 (POBSP, 1967a). 29 Aug. Little North 1 Counted (POBSP, 1967a). 27-29 Sept. Southeast 19 3 oc and 1 2 tagged at dusk (BSFW, 1967c). 1968 22-24 Mar. Southeast 22 10 tagged, 12 recaptured (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. Southeast 2 Unknown sex (BSFW, 1969a). 10-12 Feb. North aa 6 3, 19, 4 unknown (BSFW, 1969a). 10-12 Feb. Little North 2 Unknown sex (BSFW, 1969a). 31 Mar.- Southeast able 2? tagged, 9 recaptured (BSFW, 2 Apr. 1969b). 31 Mar. North 7 5 & and 2? tagged (BSFW, 1969b). 31 Mar. Little North 3 1c and 2Q tagged (BSFW, 1969b). 26-31 May Southeast 8 2 tagged, 6 recaptured (BSFW, 1969c). 26 May North 6 Counted (BSFW, 1969c). q 10-19 Sept. Southeast 27 Counted on the 10th; 3 o& and 9 ? tagged; 3 bo, 3 2 and 2 unknown recaptured; 10 nest pits on west portion, no eggs (BSFW, 19694). 65 Annual Cycle | Black Turtles have been observed in small numbers during most months of the year. Recent March population counts for the atoll range from 17 to 21. May and June counts range from 14 to 31, while September counts range from 35 to 55. Although turtles have been observed copulating in March, May and June, no egg laying has been observed. Small numbers of egg pits were recorded in September, but no eggs were found by digging. Mummified hatchlings were found in March and September. Turtles on other Northwestern Hawaiian Is- lands lay their eggs in the summer months with young hatching by late summer and fall. Ecological Distribution Turtles are attracted to the atoll for food and for breeding purposes. Adults and yearlings feed on algae growing in the lagoon. Tsuda (1966: 39) recorded 39 species of marine benthic algae from the lagoon. At least one of these, Codium arabicum, is frequently used as food by adult sea turtles. Turtles are known only from Bird, Little North, North, Sand, and South- east Islands; nesting occurs solely on North and Southeast Islands. None has been observed on the other islands. Little North Island: Turtles have been recorded on six oceasions (Table 3). The population, regardless of month, was always low and ranged from 1 to 3. North Island: Turtles have been observed on 16 surveys (Table 3). Population estimates have averaged 7 for all 11 recent surveys, and have ranged from 2 to ll. The southeast beach cove area is popular day and night with basking turtles (Fig. 56). In September 1964, 8 to 10 turtle nest pits were noted but digging produced no eggs; a mummified 3-inch hatchling was also found nearby. Another mummified hatchling was found in March 1965. Southeast Island: Of the three islands frequented by turtles, Southeast is the preferred. Turtles have been recorded there on 24 surveys (Table 3). Population estimates for 17 recent surveys average 15. March estimates average 13, and range from 8 to 22, while September estimates average 27, and range from 13 to 50. This range is similar to the 20 to 50 range found during 1957 and 1958 by Kenyon and Rice (Parsons, 1962: 69-70). Turtles are most frequently observed basking along the north lagoon beaches, especially on the east portion (Fig. 57). Turtles are usually present diurnally when a survey party arrives, but because of human activity (tagging of turtles and seals, and banding of birds) subsequently are present only nocturnally. Copulating pairs have been observed in offshore shallow waters in March, May and June; nest pits, but no eggs, have been found only in September. eit Bite S o P 56. The southeast beach-cove area at North Island is a popular place for sea turtles to bask in the sun, 29 August 1967 (print reversed). POBSP photograph by C.A. Ely. 57. Black Turtle sun basking on north beach at Southeast Island, June 1967. POBSP photograph by R.L. DeLong. 67 | Other Islands: Carr (in litt.) noted about 30 on Bird and Sand Islands in January 1962. None has been observed there since. Tagging and Movement Since 1964 BSFW personnel have tagged at least 137 Black Turtles (Table 3) with numbered Monel metal tags, usually placed in the trailing edge of the right front flipper. Of these, 122 were used at Southeast, 9 on North, and 6 on Little North Islands. Tagged individuals included 28 males and 55 females. Of the 137, 46 were recaptured on Southeast Island (Table 3). In addition, two females tagged at Whale-Skate Island, French Frigate Shoals, some 600 miles to the southeast, were recaptured at Southeast Island. Another turtle tagged at Southeast Island was captured at Whale-Skate Island, French Frigate Shoals. These data are being fruther analyzed by BSFW personnel. Amerson (1971: 80) noted that the French Frigate Shoals turtle popu- lation is the largest in the Hawaiian Islands. Hendrickson (1969: 94) theorized that "a double population nests" there. He suggested that the extreme Northwestern Hawaiian Islands may be the feeding ground for one group and that the other migrates eastward to feeding grounds around the inhabited islands. Early tag analysis seems to verify this theory (see also Amerson, 1971: 91-92). Carr (1964: 51-52) in January 1962! found the Pearl and Hermes Reef and French Frigate Shoals turtle populations to be predominantly dark and high- shelled despite single light-colored, flat yearlings at each atoll. He was unable to determine whether these individuals were variants of the local dark stock or visitors from some distant, genetically different, population. Carr (1972: 24-26) uses the name Chelonia agassizi for the mid-Pacific turtles, thus separating them taxonomically from the Atlantic green turtle, Chelonia mydas. C. agassizi also occurs in the eastern Pacific and in parts of the Indian Ocean. Carr suggests that "with its range extending through _ sO much territory,...the name...surely covers a number of hitherto unnamed races." He also notes that "some Pacific colonies that are obviously not C. agassizi...have to be grouped with the Atlantic green turtle as C. mydas." BIRDS Several sources were used in assumbling the common and scientific names of the birds occurring at Pearl and Hermes Reef. The names used in the American Ornithologists' Union's Check list of North American Birds, 1957, 5th edition, were followed for species occurring in North America. In the lyot August 1956 as noted by Amerson (1971: 61, 92, 340). 68 interest of consistency, seabird names agree with those which appear in Watson's Smithsonian Identification Manual: Seabirds of the Tropical Atlantic Ocean, and King's Smithsonian Identification Manual: Seabirds of the Tropical Pacific Ocean. Taxonomic order follows that of Peter's Check-list of Birds of the World, volumes I, II, and III, with the ex- ception of the Procellariiformes, which follow Alexander et al. (1965), the Anserformes, which follow Delacour (1954, 1959), and the Charadrii- formes, which follow Bock (1958). Introduction The 37 bird species recorded belong to 6 orders, 14 families, and 27 genera. In the following checklist, resident birds are unmarked, non- resident birds are marked with an *, and birds introduced by man are marked with a #. Order Procellariiformes Family Diomedeidae Diomedea nigripes Diomedea immutabilis Family Procellariidae Pterodroma hypoleuca Bulweria bulwerii Puffinus pacificus Puffinus nativitatus Family Hydrobatidae Oceanodroma tristrami Order Pelecaniformes Family Phaethontidae Phaethon rubricauda Family Sulidae Sula dactylatra Sula sula Sula leucogaster Family Fregatidae Fregata minor Order Anseriformes Family Anatidae Anas acuta* # Anas laysanensis Order Gruiformes Family Rallidae # Porzanula palmeri Order Charadriiformes Family Charadriidae Pluvialis dominica* Black-footed Albatross Laysan Albatross Bonin Petrel Bulwer's Petrel Wedge-tailed Shearwater Christmas Shearwater Sooty Storm Petrel Red-tailed Tropicbird Blue-faced Booby Red-focted Booby Brown Booby Great Frigatebird Pintail Laysan Teal Laysan Rail Golden Plover Family Scolopacidae Numenius tahitiensis* Heteroscelus incanum* Arenaria interpres* Capella sp.* Calidris canutus* Crocethia alba* Erolia acuminata* Erolia alpina* Philomachus pugnax* Family Phalaropodidae Phalaropus fulicarius* Family Laridae Larus delawarensis* Larus argentatus vegae* Larus glaucescens* Rissa tridactyla* Sterna lunata Sterna fuscata Anous stolidus Anous tenuirostris Gygis alba Family Alcidea Fratercula corniculata* 69 Bristle-thighed Curlew Wandering Tattler Ruddy Turnstone Snipe species Knot Sanderling Sharp-tailed Sandpiper Dunlin Ruff Red Phalarope Ring-billed Gull Herring Gull Glaucous-winged Gull Black-legged Kittiwake Gray-backed Tern Sooty Tern Brown Noddy Black Noddy White Tern Horned Puffin Order Passeriformes Family Drepaniidae Zeit irostrs cantans cantans Laysan Finch There are 17 species of resident seabirds, 5 regular migrant shore- birds, and 15 vagrant, accidental, and introduced birds. Resident Seabirds The 17 resident seabird species belong to 7 families--Diomedeidae, Procellariidae, Hydrobatidae, Phaethontidae, Sulidae, Fregatidae, and Laridae. These species all breed in the Hawaiian Islands. The Laysan Albatross breeds solely in the Hawaiian Islands. The Black-footed Albatross, Bonin Petrel, and Sooty Storm Petrel breed only in the Hawaiian and Bonin-Volcano Islands. The remaining species--Bulwer's Petrel, Wedge-tailed Shearwater, Christmas Shearwater, Red-tailed Tropicbird, Blue-faced Booby, Red-footed Booby, Brown Booby, Great Frigatebird, Gray-backed Tern, Sooty Tern, Brown Noddy, Black Noddy, and White Tern--breed in the Hawaiian Islands and other parts of the tropical Pacific. 70 Migrant Shorebirds Although over 30 shorebird species have been recorded from the Northwestern Hawaiian Islands (Bryan, 1958; Udvardy, 1961; Clapp, 1968; Clapp and Woodward, 1968; unpubl. POBSP data), only Golden Plover, Bristle-thighed Curlew, Wandering Tattler, Ruddy Turnstone, and Sanderling are considered regular migrants. They breed during the sum- mer in the Northern Hemisphere and migrate south for the winter; some use Pearl and Hermes for a "wintering ground.” Vagrant, Accidental, and Introduced Birds Fifteen species have been recorded, including four accidental gull species--Ring-billed, Herring, Glaucous-winged, and Black-legged Kittiwake. In general, Glaucous-winged Gulls breed in the area surrounding the extreme North Pacific, while Black-legged Kittiwakes breed circumpolarly in the Northern Hemisphere. The Herring Gull subspecies breeds in Siberia and the Ring-billed Gulls breed throughout most of northern North America. One vagrant alcid species, Horned Puffin, has been recorded. This species breeds in the area surrounding the extreme North Pacific. Six shorebird species--Snipe species, Knot, Sharp-tailed Sandpiper, Dunlin, Ruff, and Red Phalarope--are accidental visitors. The tidal pools on Southeast Island and the proximity to Midway Atoll, with its fresh-water puddles and large land area, may attract accidental shorebirds. The three introduced species all come from Laysan Island. The Laysan Rail, introduced in 1928, and the Laysan Teal, introduced in 1967, did not survive for long, but the Laysan Finch, also introduced in 1967, has sur- vived and is breeding very successfully. One accidental duck species, the Pintail, has been recorded. This species is a fairly regular migrant to the Main Hawaiian Islands which occasionally migrates to the Northwestern Hawaiian Islands. The shallow waters around Pearl and Hermes Reef offer an excellent feeding area for many seabirds, those which use, or breed on, the atoll as well as those which seldom occur there. The latter group includes both species which breed in the Hawaiian area (i.e., White-tailed Tropicbird) and pelagic birds which breed in other areas of the Pacific and migrate into or through the area during their non-breeding season. King (1967) records 50 seabird species classified as regular migrants, rare migrants, or vagrants to the Hawaiian area. Normally all of these birds stay at sea, but, because of sickness or bad weather, any could alight on the islands; none has been recorded at Pearl and Hermes to date. Annual Cycles Among the bird species annual population cycles and annual breeding cycles vary. These variations are discussed below. 71 Seabird Breeding Cycles Seabirds breed here during all seasons of the year (Fig. 58). Most have distinct breeding periods; some have extended breeding cycles. The 17 breeding seabird species are grouped, based on maximum breeding periods, as follows: 4 are winter and spring breeders, 11 are spring and summer breeders, and 2 are summar and fall breeders; none is a fall and winter breeder (Table 4). Of these, the cycles of only six agree with those suggested by Richardson (1957: 30). Winter and Spring Breeders: The four species are procellariiforms. The Black-footed and Laysan Albatrosses commence to nest in late fall, and the Sooty Storm Petrel and Bonin Petrel commence in early and mid- winter, respectively. The Albatrosses have usually fledged by late July. Sooty Storm Petrel and Bonin Petrel fledge in late spring and early sum- mer, respectively. Spring and Summer Breeders: Of the 11 species, 1 is a procellariiforn, 5 are pelecaniforms, and 5 are charadriiforms. Six species (55 percent), the Christmas Shearwater, Red-tailed Tropicbird, Gray-backed Tern, Sooty Tern, Black Noddy, and White Tern, commence laying during the spring months. Individual birds of the remaining five species (45 percent) start laying during the winter months: the Brown Noddy as early as December, the Red-footed and Brown Boobies in early January, and the Blue-faced Booby and Great Frigatebird in February. All young usually commence fledging in the summer and continue into the fall. Fledging of the Christ- mas Shearwater, Sooty Tern, and probably White Tern extends into early fall; of the Red-tailed Tropicbird, Blue-faced Booby, Red-footed Booby, Great Frigatebird, Gray-backed Tern, Brown Noddy, and Black Noddy into late fall; and of the Brown Booby even into early winter. Summer and Fall Breeders: Two procellariiforms, Bulwer's Petrel and Wedge-tailed Shearwater, start laying in early summer and usually finish fledging by mid- or late fall. Fall and Winter Breeders: Although no fall-winter breeders are known, virtually no late fall and early winter visits have been made. The Black Noddy, a fall-winter breeder on other Northwestern Hawaiian Islands, may breed during this period. Although all breeding seabird species have a breeding peak, four, the Blue-faced Booby, Red-footed Booby, Brown Booby, and Brown Noddy, have an extended breeding season with egg laying extending over three seasons. Land Bird Breeding Cycles The introduced Laysan Finch appears to have a winter and spring breed- ing cycle. Eggs and young are now known from February to May. Because of its recent introduction, however, the cycle may be erratic. The unsuccess- fully introduced Laysan Teal did not survive long enough to establish an annual breeding cycle. One pair, however, attempted to nest in mid-fall. Black-footed Albatross ee0 ec Laysan Albatross Bonin Petrel Bulwers Petrel Wedge-tailed Shearwater Christmas Shearwater Sooty Storm Petrel Red-tailed Tropicbird ee eee ee Blue-faced Booby Red-footed Booby Brown Booby as ste ceeeee Great Frigatebird Gray-backed Tern Sooty Tern Brown Noddy Black Noddy tet ag SE, li I PO ORB ALES it A EEE SII. White Tern 58. Breeding cycles of seabirds at Pearl and Hermes Reef; stip- pled areas represent eggs, barred areas young, and black dots non-breeding birds. a 73 Table 4. Maximum breeding periods of Pearl and Hermes Reef resident seabirds Winter-Spring Spring-Summer Summer-Fall Fall-Winter Black-footed Alba- Christmas Shear- Bulwer's Petrel None tross water* Wedge-tailed Laysan Albatross Red-tailed Tropic- Shearwater * Bonin Petrel bird* Sooty Storm Petrel Blue-faced Booby* Red-footed Booby* Brown Booby Great Frigatebird* Gray-backed Tern* Sooty Tern* Brown Noddy Black Noddy White Tern *Breeding period agrees with that suggested by Richardson (19572 4380)2 Population Cycles Perusal of the subsequent species accounts and inspection of Figure 58 and Table 5 reveal that many resident and non-resident species leave the atoll during part of each year. Even those that stay year-round have a population buildup sometime during the year. Resident Species: Of the resident seabird breeding species, 7 remain on the atoll year round: Blue-faced Booby, Red-footed Booby, Brown Booby, Great Frigatebird, Brown Noddy, Black Noddy, and probably White Tern. Their populations decrease during the non-breeding seasons in the late fall and early winter. The 10 resident seabird breeding species which do not stay on the atoll year-round leave just before or just after their young fledge. Some remain at sea in the general area of the Hawaiian Islands; others leave the area entirely and migrate to distant areas. The Black-footed Albatross, Laysan Albatross, Bonin Petrel, Red-tailed Tropicbird, and Gray-backed Tern are only away for two to three months. The Bulwer's Petrel, Wedge-tailed Shear- water, Christmas Shearwater, Sooty Storm Petrel, and Sooty Tern spend from 4 to 7 months away (Fig. 58). Non-resident Species: The monthly occurrence of the 17 non-residents-- regular migrant shorebirds, and vagrant and accidental species--is pre- sented in Table 5. Some species occur year-round, while others occur infrequently. Th The five regular migrant shorebird species are known from all seasons of the year. The six shorebirds classed as accidentals and vagrants are found irregularly during the year. The four gulls and the alcid are known from late winter or spring months. The accidental duck is known only from September. Introduced Birds: Of the three introduced bird species, only the Laysan Finch remains. It is found year-round on the atoll. Table 5. Monthly occurrence of the non-resident birds at Pearl and Hermes Reef * = 0 3 = 2 3 a q a oO Ss oO “d ) 3} p a en eee Species ee = = re «© 6 a (Wold) Te7OL 6 ec i -ec C Appon Yoeta GC Co) Appon umMorg 109 S € S ot 1 T PIT qazesTAq Peery a als qoog umozg Ge, at g fqoog pe o0os-pey S Coe iE Aqoog paoes-anTg prtgotdorz,, peTtey-pey IE TAVeMIesug Pellei-sspeM Tt Tatzeg utuog cea Z ssorjeqry ueskeT eal g SsorjzeqlTy pe ,oos- Yoel” $07 joey Souzloy pue [1eeg woly (OL) Te70L TL a Il 3 Appon Yoeta T AppoN uMorg udey, A1oos ¢ auo suing, Appny T ZTaACTd uepTtoy PIT QezVesTAA Jeary T qoog umorg Aqoog peyoos-pey Aqoog peoes-entg ZTaVemIeasys PeTtej-sspam Teryeg urtuog ssorpzeqrTy pe yoos- yoeld au I odio Ney Ht T HE oO Ne (oa) oa) = | Toa | a — To qu) ina) uw q — Way rc ro a ia} +tmnimMmotUnt ms 4 =I HM NONOM IS GUESS i N ioe) = font Fo | "IL WOT, joey Soultey pue [reed oO], Trud oyemM any Oo o Sy c LOIS 1E T-+OL JELPO FOTEAT Ad ueder 20TTTaA B98 4V uo1Suyo r Aemptn ueskeyT Tysue uoZSUTYUSeM -eTULO soutddt sTeoug 3a4e -3TIq youslr7 Jeo Soultoy] pue [Taeeg BUTATOAUT SpAitTq pepueq Fo JUSWeAOW pueTSTIeIUT “JT eTqeL Dee eee ———————————— ee ee 110 Table 18. Summary of POBSP records of birds from Pearl and Hermes Reef POBSP Records Clapp and Sibley and This Woodward, MacFarlane, Paper 1968 1968 Specimens Pre- POBSP POBSP Black-footed Albatross 2 Laysan Albatross 2 Bonin Petrel 4 sr Bulwer's Petrel r Wedge-tailed Shearwater 1 sr Christmas Shearwater 2 sr Sooty Storm Petrel 6 3 sr Red-tailed Tropicbird 3 sr ie Blue-faced Booby 3 sr Brown Booby 1 IL sr,br Great Frigatebird dL aL sr Pintail 12 Laysan Teal* rap og Golden Plover is Bristle-thighed Curlew iL sr Wandering Tattler 2 sr ig Ruddy Turnstone IL 2 sr Snipe sp. ig Knot dk SR Sanderling r Sharp-tailed Sandpiper 2 sr Dunlin ald sr Ruff 1 sr Red Phalarope aL sr Ring-billed Gull dk sr sr Herring Gull 2 sr sr Glaucous-winged Gull il 3 sr sr Black-legged Kittiwake 3 sr sr Gray-backed Tern 4 sr r Sooty Tern 2 al sr Brown Noddy 2 all sr Black Noddy 2 4 sr White Tern br Horned Puffin 2k* ie Laysan Finch* br Total Specimens Sif Sul Total POBSP Records 21 17 4 br = first breeding record; r = first sight record; sr = first specimen record; SR = first specimen confirmation of a species previously known only from sight records for both the main Hawaiian and Northwestern Hawaiian Islands. *Introduced. **Specimens not preserved. dLaLil BLACK-FOOTED ALBATROSS Diomedea nigripes Status Common breeding species; present from late October into July; occurs and nests on all islands except Planetree and Sand. Maximum recent popu- lation estimate 13,948 in March 1967. Observations Nesting Black-footed Albatross were first recorded in March 1913 (Bailey, 1956: 32; and Willett, ms.). Munter (ms.) found them again in 1916 and Wetmore (ms.) observed them in 1923. We have no data for the 1930's and 1940's. Richardson (1957: 16) noted that it "is abundant," but gave no data. Rice and Kenyon (1962: 369) reported this species con- centrated mainly at Grass, North and Southeast Islands during the 1956-67 and 1957-58 breeding seasons. POBSP and BSFW personnel observed this species on all major islands except Planetree and Sand. All observations are presented in Tables 19 to 25. Annual Cycle The local annual breeding cycle is presented in Figure 58. Adults arrive in early October and depart in late June or July; numbers increase in November and decline in late spring. The population fluctuation co- incides with breeding: most eggs are undoubtedly laid in mid- or late November and begin to hatch by late January or early February. Fledging occurs in late June and July. Rice and Kenyon (1962: 369) reported some 15,000 breeding Black-footed Albatross in December 1956 and 1957. No comparable data exist for recent Decembers. March surveys for 1963 to 1968 show an average of 7,900 breed- ing birds (range 6,600 to 8,900). Ecological Distribution The Black-footed Albatross nests on Bird, Grass, Kittery, Little North, North, Seal, and Southeast Islands. It has not been recorded at Planetree and Sand, both of which are small islands and awash at high tide. Bird Island: Rice and Kenyon (1962: 369) reported 126 breeding birds on "south sandspits," presumably Bird, during December 1956 and 1957. POBSP and BSFW personnel found this species nesting in small numbers (less than 20) only in 1964 and 1967 (Table 19). Nests are all in bare sand as there is no vegetation on the island. Grass Island: Munter (ms.) found nesting Black-footed Albatross very plentiful on Grass Island in February 1916. POBSP and BSFW personnel have recorded Black-footed Albatross each year since 1963 except 1966 when no spring visit was made (Table 20); from 300 to 1,466 young have been recorded in March. 2! This species nests mainly on the perimeter of the vegetated area; some nest in a sparsely vegetated area at the western end of the island. Kittery Island: Black-footed Albatross were first recorded by Rice and Kenyon Gece: 369), who counted 900 breeders from aerial photographic surveys in December 1956 and 1957. ‘The breeding population has been smaller in recent years. POBSP and BSFW figures based on the number of young present during March surveys ranged from 252 to 456 breeders between 1963 and 1968 (Table 21). Black-footed Albatross nest in bare sand as the island is unvegetated. Little North Island: Rice and Kenyon (1962: 369) in December 1956 and 1957 recorded the first Black-footed Albatross from Little North " Island, then referred to as "north sandspits." Their estimate of 240 ‘ breeders has not been equaled in recent years. POBSP and BSFW personnel recorded this species only in 1963 and 1965 r (Table 22). The nests were all in the raised central portion. North Island: Black-footed Albatross were first observed at North § Island in March 1913 (Bailey, 1956: 32; and Willett, ms.). Rice and Kenyon i (1962: 369) estimated 3,800 breeders in December 1956 and 1957. POBSP and BSFW personnel recorded this species nesting on all surveys (Table 23). March counts on young ranged from 500 to 1,200. Nests are concentrated on the east and west beaches, at the base of the southern tip, and on the raised middle portion of this elongated tip. Seal Island: Munter (ms.) first observed this species in February 1916. Wetmore (ms.) estimated 1,200 pairs in April 1923. With the exception of 1966 when no spring visits were made, POBSP and BSFW personnel recorded Black-footed Albatross each year since 1963 (Table 24). March counts of young range from 42 to 325. Nests are placed at the perimeter of the vegetated area with the ex- ception of a few placed in a depressed area on the eastern section. Southeast Island: Munter (ms.) recorded the first Black-footed Albatross in February 1916, and Wetmore (ms.) found it nesting in April 1923. Rice and Kenyon (1962: 369) estimated 4,600 breeding adults in December 1956 and 1957. POBSP and BSFW personnel recorded this species nesting each year from ai 1963 to 1969 (Table 25). March counts of young range from 1,560 to 2,500. ‘ Black-footed Albatross nest around the perimeter of the east and west sections, and on the coral rubble transition zone between the vegetated in- terior and the sand beach. They seldom nest in heavily vegetated areas. 113 Banding and Movements POBSP and BSFW personnel banded 9,008 Black-footed Albatross on six islands (Table 26). Of this total, 6,673 were banded by the POBSP and 2,335 were banded by the BSFW. POBSP and BSFW personnel have captured 390 previously banded Black- footed Albatross here. Three hundred and sixty-six were originally banded on the atoll; 24 originated elsewhere: Midway (14), Kure (6), at sea (3), and French Frigate Shoals (1). In addition, 55 birds originating at Pearl and Hermes have been captured elsewhere: at sea (34), Kure (17), French Frigate Shoals (2), and Midway and Japan (1 each). These movements are presented in Appendix Tables 4a and 4b. Specimens Non-POBSP: USNM 300815 and 300833, oo, collected 27 April 1923 by Wetmore. : Table 19. Observations of Black-footed Albatross at Bird Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1956 10 Dec. 168 126 breeders, 42 non-breeders (Rice and Kenyon, 1962: 369).* 1) Ut Oct 200 Aerial survey (Rice, ms. a). 18 Dec. 168 126 breeders, 42 non-breeders (Rice and Kenyon, 1962: 369).* 1963 5 Mar. 40-50 Roosting, none nesting (POBSP, 1964d). 1964 14 Mar. 43 30 adults, 13 live young counted; 1 dead (BSFW, 1964a; POBSP, 1964b). 1965 22 Mar. 5 Roosting, none nesting (POBSP, 1965a). 1967 31 May 19 Young counted (POBSP, 1967b). *Locality listed as "south sandspits." 114 Table 20. Observations of Black-footed Albatross at Grass Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. very Adults with eggs and young birds; ca. half plentiful as numerous as Laysan Albatross (Munter, iiaiSc, Ve 1923 27 Apr. 1,600 Ca. 800 pairs (Wetmore, ms.). 1956 10 Dec. Bip Uo) 3,800 breeders, 1,300 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1957 14 Oct. 4500 Aerial survey (Rice, ms. a). 18 Dec. 5,100 3,800 breeders, 1,300 non-breeders; based on aerial survey (Rice and Kenyon, 1962; 369). 1963 5 Mar. ? Nesting; ca. twice as numerous as Laysan Albatross (POBSP, 1964d). 26-27 June 263 Young counted (POBSP, 1963). 1964 14 Mar. 4275 3,000 adults, 1,275 young (BSFW, 1964a; : POBSP, 1964b). 1965 19 Mar. 1,100- 800-1,000 adults, 300-400 young (POBSP, 7 1,400 1965b). 7 22 Mar. 1,280 800 adults, 480 young (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 0 None seen; no dead young (BSFW, 1966b). 1967 22 Mar. 4, 398t 1,466 downy young counted; low mortality although 37 dead young counted (BSFW, 1967a; POBSP, 1967d). 31 May LL Young counted (POBSP, 1967b). 1968 24 Mar. BRO 2,380 breeders, 1,190 young counted (BSFW, 1968; POBSP, 1968). 115 Table 21. Observations of Black-footed Albatross at Kittery Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1956 10 Dec. 1,200 900 breeders, 300 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1957 14 Oct. 1,100 Aerial survey (Rice, ms. a). 18 Dec. 1,200 900 breeders, 300 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1963 5 Mar. several Nesting (POBSP, 1964d). hundred 26 June 48 124 young counted (POBSP, 1963). 1964 14 Mar. 426 300 adults, 126 young counted (BSFW, 1964a; POBSP, 1964b). 1965 18 Mar. 600-725 400-500 adults, 200-225 young (POBSP, 1965b). 22 Mar. 385 Ca. 205 adults, 180 young; 30 eggs ina windrow in east central area (BSFW, 1965; POBSP, 1965a). 1967 31 May 353 Young counted (POBSP, 1967b). 1968 2 Mar. 684 228 young counted (BSFW, 1968; POBSP, 1968). Table 22. Observations of Black-footed Albatross at Little North Island Population Date of Survey Estimate Breeding Status, Remarks, and References 156. 10 Dec. 320 240 breeders, 80 non-breeders (Rice and Kenyon, 1962: 369).* 1957 WF Oct. 0) Aerial survey (Rice, ms. a). 18 Dec. 320 240 breeders, 80 non-breeders (Rice and Kenyon, 1962: 369).* 1963 6 Mar. 100 Ca. 50 pairs nesting (POBSP, 1964d). 23, 25 June 64+ Young counted (POBSP, 1963). 1965 18 Mar. 56-66 40-50 adults, 16 young counted (POBSP, 1965b). 1969 26 May 0 None observed (BSFW, 1969c). *Locality listed as "north sandspits." 116 Table 23. Observations of Black-footed Albatross at North Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1913 15 Mar. 900-1, 050 Ca. 300-350 nestlings (Bailey, 1956: 32); 300 pairs (Willett, ms.). 1956 10 Dec. 5,060 3,800 breeders, 1,260 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1957 14 Oct. 4,500 Aerial survey (Rice, ms. a). 18 Dec. 5,060 3,800 breeders, 1,260 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1963 6 Mar. 100 Nestlings (POBSP, 1964d). 23-25 June Tou 1,500 adults, 751 young counted (POBSP, 1963). 1965 17 Mar. 1,500- 1,000-1,500 adults, 500-600 young 1/4 to 2,100 1/3 grown (POBSP, 1965b). 1967 16-17 Mar. BR Oe 1,150 t 50 chicks counted (BSFW, 19674). 1969 31 Mar. ? Nesting (BSFW, 1969c). Table 24. Observations of Black-footed Albatross at Seal Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. thickly Eggs and young birds (Munter, ms.). populated 1923 27 Apr. 2,400 About 1,200 pairs (Wetmore, ms.). 1956 10 Dec. 990 740 breeders, 250 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1957 14 Oct. 900 Aerial survey (Rice, ms. a). 18 Dec. 990 740 breeders, 250 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1963 5 Mar. ? Nesting in somewhat greater numbers than the Laysan Albatross (POBSP, 19644). 117 Table 24. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1963 26 June 97 Young counted (POBSP, 1963). 1964 14 Mar. 825 265 young counted; also 10 dead young (BSFW, 1964a; POBSP, 1964b). 1965 18-19 Mar. 150-250 100-200 adults, 42 young 1/4 to 1/3 grown counted (POBSP, 1965b). 22 Mar. 185 144 adults, 41 young counted (BSFW, 1965; POBSP, 1965a). 1967 22 Mar. 990 325 young counted, also 4 dead young (BSFW, 1967a; POBSP, 1967d). 31 May 310 Young counted (POBSP, 1967b). 1968 2h Mar. 660 439 breeders, 219 young counted (BSFW, 1968; POBSP, 1968). Table 25. Observations of Black-footed Albatross at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. thickly Eggs and very young birds present; less settled numerous than Laysan Albatross (Munter, ms.). 1923 26-28 Apr. 2, 000+ Adults; some young with developing breast feathers (Wetmore, ms.). 1956 10 Dec. 6, 100 Aerial survey estimated data: 4,600 breed- ing adults, 1,500 non-breeding adults (Rice and Kenyon, 1962: 369). 1957 14 Oct. 5,600 Aerial survey (Rice, ms. a). 18 Dec. 6,100 Aerial survey estimated data: 4,600 breed- ing adults, 1,500 non-breeding adults (Rice and Kenyon, 1962: 369). 1961 12 Mar. 2 Young present; much more abundant than Laysan Albatross (HDFG, 1961). 1963 26 Feb.- 5,000 Adults, all with young (POBSP, 19644). 8 Mar. 118 Table 25. (continued) Population ; Date of Survey Estimate Breeding Status, Remarks, and References 1963 18-23, 1, 200+ Few adults, 1,200 young counted (POBSP, : 25 June 1963). _ &§ 1964 13-14 Mar. 8,500 5,000 breeders, 1,000 non-breeders; 2,500 young, many dead young (BSFW, 1964a; POBSP, 1964b). 1965 15-19 Mar. 6, 000- 4, 000-5,000 adults, 2,000 young (POBSP, 7,000 1965b). 21-22 Mar. 3,250 2,000 adults, 1,250 young, low mortality | (BSFW, 1965; POBSP, 1965a). 1966 1 Apr. many Nesting (BSFW, 1966a). 1967 21-23 Mar. 4,710 1,560 young counted (BSFW, 1967a; POBSP, 1967d). 28, 30 May- 202i 50 adults, 1,971 fully grown young counted 1 June (POBSP, 1967b). 1968 22-24 Mar. 6, OOO+ 4, 000+ breeders, 2,002 young counted: 1,696 on eastern part and 306 on western part (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 11, 000 Most young less than 2 weeks old, few eggs (BSFW, 1969a). 31 Mar.- W435 1,478 young; little mortality (BSFW, 1969b). 2 Apr. 26-31 May 1, 964+ Young counted (1,598 on eastern half, 366 on western half); little mortality (BSFW, 1969c). LAYSAN ALBATROSS Diomedea immutabilis Status Abundant breeding species; present from early October to late July, a few stragglers into early August; absent during rest of year. Nests on all islands except Planetree and Sand. Maximum POBSP and BSFW population estimate 45,000 in February 1969. 119 Nie e 849 T 6€S ui 96€ (Se) On ~~ O|O oe) (ea) 6€ OOF *T 008 00S T2401 S96T 00S ‘T 000°T 00S MASE LTL gL9 6€ ut 96€ wae SS ee 6QT OST 6€ Oss 0 0 G96T bar 1=4/\ dsd0d Jooy Souzoy pue HOGI TOK MASE 9565S 954 ‘T 959°? 9ST oo€ * 0 +9 +9 OPE oS Al HoT 6 6 core ee © 0 €9¢ €9¢ €9¢ C92 ©) ) 659‘ 6ST 6Gean 6ST Oo *€ 0 Teq0L CO6T €96T aunt dSdod O O 00S‘ 00d *T oOo€ f€ €96T “Te -"qad dsdod :Te10L Sunox FINPY sunoz Suno_ sunox :Te10] Sunox TDPV > Te101 Bunox FTNPV T2201 Sumox TINY SSeTO -93V > Taepueg Teq101 UIION 9T99TT Y}.2ON £294.47M T8958 sseiry aseoygnos pues] [read Pe papueq ssouzeqly pe,oos-yoeTA *og aTqeL 120 Observations Nesting Laysan Albatross were recorded at Pearl and Hermes Reef as early as March 1913 (Bailey, 1956: 32; Willett, ms.). Anderson (1954: 84) saw this species in the late 1920's. Galtsoff (1933: 19) is credited, however, with the first published record from the atoll; he observed several hundred abandoned young at Southeast Island in summer 1930. : Richardson (1957: 16) noted that it "breeds abundantly," but gives no 4 data. POBSP and BSFW personnel found this species nesting each year from 1963 to 1969. All observations are shown in Tables 27 to 31. Annual Cycle The local annual breeding cycle is presented in Figure 58. Birds arrive in early October and egg-laying commences about early November. Hatching occurs in early and mid-January. Young begin to fledge in mid- June and have departed by late August. Adults commence to leave in late May and June and all have left by September. The Laysan Albatross population is the second highest in the atoll. Total young counts for five recent March visits average 7,872 and range from 5,802 to 12,840. Total POBSP and BSFW population estimates have been highest during February visits even though only Southeast Island was visited. The Rice and Kenyon (1962: 369) estimates for December 1956 and 1957 were higher for the entire atoll but lower for Southeast. Ecological Distribution Laysan Albatross nest on Bird, Grass, Kittery, Little North, North, Seal, and Southeast Islands; none is known from Planetree and Sand. Bird Island: Three or four adult roosting Laysan Albatross were observed by POBSP personnel in March 1963. None was found during two visits (March and August) in 1964 and one visit in 1965 (March). One chick was counted 31 May 1967 by POBSP personnel; this is the only nesting mHecordr. Grass Island: Munter (ms.) was the first to observe Laysan Albatross in early February 1916. All visitors since noted them. POBSP and BSFW personnel found Laysan Albatross nesting each year from 1963 to 1968 except in 1966 when the island was not visited during the albatross season (Table 27). Data for 1969 are unavailable. Recent March counts of young average 649, and range from 350 to 1,120. Laysan Albatross nest along both beaches but prefer the inland vege- tated area. Kitte Island: POBSP personnel first recorded nesting Laysan Alba- tross in March 1963, and, except for 1966 and 1969 when no data are ale available, the species has been recorded nesting ever since (Table 28). March population estimates for young average 28, and range from 4 to 52. At vegetationless Kittery, Laysan Albatross nest in the center portion of the island. Little North Island: The first few nesting Laysan Albatross were found on 6 March 1963 by POBSP personnel. Twenty young were counted when they next visited the island on 23 and 25 June 1963. On 18 March 1965, 8 to 10 adults and 4 young were counted on the crest of the island. The island was not visited during this species' breeding season in 1964, 1966- 68. BSFW personnel visited Little North on 31 March 1969; however, popu- lation data are not available. North Island: Bailey (1956: 32) and Willett (ms.) reported nesting Laysan Albatross in March 1913. It has been recorded breeding on all subsequent visits (Table 29). Available data from two recent March population counts show 750 young in 1965 and ca. 525 in 1967. Nests are predominately located in the vegetated northern portion. Seal Island: Munter (ms.) found Seal Island thickly populated with nesting Laysan Albatross in February 1916, and Wetmore (ms.) found 150 pairs in August 1923. Rice and Kenyon (1962: 369), however, first pub- lished their occurrence from aerial observations in 1956 and 1957. POBSP and BSFW personnel have recorded them nesting on all subsequent visits for which we have data (Table 30). Four recent March population counts show an average number of 151 young, and range from 40 to 399. Nests are located mainly in the vegetated western portion of the island. Southeast Island: Nesting Laysan Albatross were first observed by Munter (ms.) in February 1916. Galtsoff (1933: 19) first reported them after noting them in summer 1930. POBSP, BSFW, and HDFG personnel have collected breeding data for each year from 1961 to 1969, except in 1962 when the atoll was not visited (Table 31). Recent March population estimates of young average 6,950 and range from 4,600 to 11,269. lLaysan Albatross utilize the interior for nesting. Most prefer the eastern half of the island; in March 1968, 87 percent favored this half. Banding and Movements The POBSP and BSFW banded 10,375 Laysan Albatross on six islands since 1963, 9,625 by the POBSP and 750 by the BSFW (Table 32). POBSP and BSFW personnel have recaptured 196 Laysan Albatross, all originally banded on the atoll. In addition, 29 banded at Pearl and Hermes 122 have been captured elsewhere: 20 at Kure, 6 at sea, 2 at Laysan and 1 at Midway (see Appendix Table 5). Specimens Non-POBSP: USNM 300856, o, collected 26 April 1923 by Wetmore; USIM 289167, skull, collected 28 April 1923 by Wetmore. Table 27. Observations of Laysan Albatross at Grass Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. very Eggs and quite young birds; twice as plentiful numerous as Black-footed Albatross (Munter, ms.). 1923 27 Apr. 200 100 pairs (Wetmore, ms.). 1956 10 Dec. 3, 200 2,400 breeders, 800 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1957 4 Oct. 2,800 Observed from aerial survey (Rice, ms. a). 18 Dec. 3,200 2,400 breeders, 800 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1963 5 Mar. ? Two-thirds as numerous as Black-footed Albatross; breeding (POBSP, 1964d). 26-27 June 607 Young counted (POBSP, 1963). 1964 14 Mar. 2,750 2,000 adults, 750 young estimated (BSFW, 1964a; POBSP, 1964b). 18 Aug. 3 Young counted; no adults (POBSP, 1964a). 1965 19 Mar. 1,025 - 700-900 adults, 325-350 young (POBSP, il, 250 1965b). 22 Mar. 1, 100+ 850 adults, 250-260 young (BSFW, 1965; POBSP, 1965a). 1967 22 Mar. 660 220 young counted; 375 nests counted pre- viously destroyed (BSFW, 1967a; POBSP, 19674). 31 May 127 Young counted (POBSP, 1967b). 123 Table 27. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1967 8 July 96 Young counted (BSFW, 1967b). 1968 24 Mar. 3,360 2,240 breeders, 1,120 young counted (BSFW, 1968; POBSP, 1968). Table 28. Observations of Laysan Albatross at Kittery Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1956 10 Dec. none (Rice and Kenyon, 1962: 369). ne57 voMMeEc? none (Rice and Kenyon, 1962: 369). e637 5 Mar. few pairs Nesting (POBSP, 1964d). 26 June 27 Young counted (POBSP, 1963). 1964 14 Mar. 14-19 10-15 adults; 4 young counted (POBSP, 1964b). 1965 18 Mar. 17=27 10-20 adults; 7 young counted (POBSP, 1965b). 22 Mar.. yy 37 adults; 7 young counted (BSFW, 1965; POBSP, 1965a). 1967 22 Mar. 4h 48 young counted (BSFW, 1967a; POBSP, 1967d). 31 May 30 Young counted (POBSP, 1967b). 1968 2h Mar. 156 52 young counted (BSFW, 1968; POBSP, 1968). Table 29. Observations of Laysan Albatross. at North Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1913 15 Mar. 900-1,030 Ca. 300-350 nests with young (Bailey, 1956: 32); about 300 pairs (Willett, ms.). 1956 10 Dec. 5,600 4,200 breeders, 1,400 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 124 Table 29. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1957 14 Oct. 4, 800 Observed from aerial survey (Rice, ms. a). 18 Dec. 5,600 4,200 breeders, 1,400 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1963 6 Mar. few Nesting (POBSP, 1964d). 23-25 June 384 Young, all banded (POBSP, 1963). 1964 19-20 Aug. 9 8 large young, 1 flying immature (POBSP, 1964a). 17 Sept. O 10 long dead almost fully grown young (BSFW, 1964b; POBSP, 1964c). 1965 17 Mar. 2,250- 1,500-2,000 adults, 750 young (POBSP, 2,750 1965b). 1967 16-17 Mar. 1,575- 525 t 25 chicks counted (BSFW, 1967d). M625) 29-30 Aug. 1 Starving young banded in March, no adults (POBSP, 1967a). 1969 31 Mar. ? Nesting (BSFW, 1969b). Table 30. Observations of Laysan Albatross at Seal Island. Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. thickly Eggs and young birds (Munter, ms.). populated 1923 27 Apr. 300 150 pairs (Wetmore, ms.). 1956 10 Dec. 1, 200 900 breeders, 300 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1957 14 Oct. 1,000 Observed from aerial survey (Rice, ms. a). 18 Dec. 1, 200 900 breeders, 300 non-breeders; based on aerial survey (Rice and Kenyon, 1962: 369). 1963 5 Mar. ? Nesting (POBSP, 19644). 125) Table 30. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1963 26 June 186 Young counted (POBSP, 1963). 1964 14 Mar. 320-370 ~—« 250-300 adults, 70 live young, 5 dead young (BSFW, 1964a; POBSP, 1964b). 18 Aug. 1 Young (POBSP, 1964a). 1965 18-19 Mar. 191-241 100-150 adults; 91 1/4 to 1/3 grown young counted (POBSP, 1965b). 22 Mar. 300 138 adults; 100 young (BSFW, 1965; POBSP, 1965a). 1967 22 Mar. 102+ 34 live and 6 dead young counted; 175 ¢ additional nests previously destroyed (BSFW, 1967a; POBSP, 1967d). 31 May 27 Young counted (POBSP, 1967b). 6 July 23+ Few adults, 23 near-fledging young counted, also 1 dead chick (BSFW, 1967b). 1968 24 Mar. Toy, 798 breeders, 399 young counted (BSFW, 1968; POBSP, 1968). Table 31. Observations of Laysan Albatross at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. thickly Eggs and very young birds present; more settled abundant than Black-footed Albatross (Munter, ms.). 1923 26-28 Apr. 600 Adults; ca. 300 pairs on Southeast (Wetmore, ms.). 1930 23 July- several Abandoned young, all died (Galtsoff, Aug. hundred 1933): al9)).. 1956 10 Dec. 37,300 Aerial survey estimated data: 28,000 breeding adults; 9,300 non-breeding adults (Rice and Kenyon, 1962: 369). 1957 14 Oct. 33, 200 Observed from aerial survey (Rice, ms. a). 126 Table 31. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References Opi alto Dece 37,300 Aerial survey estimated data: 28,000 j | breeding adults, 9,300 non-breeding adults (Rice and Kenyon, 1962: 369). 1961 12 Mar. 2 With young; much more abundant than Laysan Albatross (HDFG, 1961). 1963 28 Feb.- 35, 000 All breeding birds with young (POBSP, 8 Mar. 1964d). 18-23, 8, 000- Young (POBSP, 1963). 25 June 10,000 1964 13-14 Mar. 24,500 15,000-15,500 breeders, 1,000-1,500 non- breeders, 7,500 young (BSFW, 1964a; POBSP, 1964b). 16-19 Aug. 39 Large young counted; 600-800 carcasses; no adults (POBSP, 1964a). 1965 15-19 Mar. 115}, OOOH 9,200 breeders, 800-1,800 non-breeders, 4,600* young: 1/4 to 1/3 grown (POBSP, 1965b). 21-22 Mar. 14,500 9,000-10,000 adults, 4,500 young (BSFW, 1965; POBSP, 1965a). 1966 1 Apr. many Nesting (BSFW, 1966a). 20-26 Sept. 0 450-500 large immatures had died (Kenyon and Kridler, 1969: 339). 1967 21-23 Mar. 15,900t 5,300 young counted (Kenyon and Kridler, 1969: 340); heavy mortality (BSFW, 1967a). 28, 30 May- 4, ooot 500f adults, 3,473 young counted (POBSP, 1 June 1967b). 3-9 July several Few adults, several hundred young; 426 hundred dead chicks (BSFW, 1967b). 1968 22-2 Mar. 33, 800 11,269 young counted (Kenyon and Kridler, 1969: 340); 9,573 on the eastern part of the island and 1,696 on the western part (BSFW, 1968; POBSP, 1968). 127 Table 31. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1969 10-12 Feb. 45,000 Most eggs hatched (BSFW, 1969a). 31 Mar.- 18, 200% 6,075 young (BSFW, 1969b). 2 Apr. 26-31 May 5,763 Young counted, 5,109 on the eastern part and 654 on the western part; little mor- tality noted (BSFW, 1969c). *Kenyon and Kridler (1969: 340) give a figure of 5,000 young for this visit. BONIN PETREL Pterodroma hypoleuca Status Common breeding species; present from late August to late June; absent during rest of year. Nests on the four major vegetated islands. Maximum POBSP and BSFW population estimate 1,000 in August 1967 and February 1969. Observations Bonin Petrels, with nests, were first observed in 1913 (Bailey, 1956: 32; Willett, ms.). POBSP and BSFW personnel have recorded them each year since 1963 (Tables 33 to 36). Annual Cycle Figure 58 presents the local annual breeding cycle. Adults are be- lieved to arrive in late August; egg-laying possibly commences as early as January. Eggs were known to occur as late as mid-March in 1964 and 1965. Hatching was recorded by mid-March in 1965. Large, fully-feathered juve- niles were recorded in early June 1967 and late June 1963; all young fledge by early July. Adults are probably absent during July and most of August. In recent years, approximately 1,000 Bonin Petrels are estimated to inhabit Pearl and Hermes. Ecological Distribution Bonin Petrels nest at Grass, North, Seal, and Southeast Islands. Grass Island: Munter (ms.) probably observed burrows of this species in February 1916. Wetmore (ms.) saw one individual in April 1923. POBSP GLE SOT 6r0T] 66S gle 6 0S6 00S 660 *T 66 66 ds 0 6€L *T OSh HE 0 0 922 © 0 S06 o) @) STeqOL Tei0L° L96T L96T Key dSdod 128 OS OSt OSt dsdod ete 8 Ore 8h @ 0) 0 0) L J ) 0) Tt TE € € Us [Je LEE LE 0) 0 Te70L° +967 496T ‘“sny dSdod jooy Ssouzey pue [ieeg 1e pepueq ssoreqty ueskeyT MASE 19 000 ST T2101 €96T E16 000‘T Qle*g O ) 000 ST 02g 0 Cis 10) 12 ) 98T O zo9 OO we 000 ‘T T9 0 0 000 *T €96T €96T eune * Te -"qaq dsd0d dsd0d T2207 Suno, 4Tapy Te {OL Uy4ON sunox STI4TT sunox Ud2ON sunojx £I94.9TY sunoz Tees sunox sse1y :Te10L Sunox FIOpy Os. FSBaYFNOSs SSeTO pueTst -o3V : Lapueg “ce eTaeL 129 and BSFW personnel have recorded small numbers nesting in most years since 1963 (Table 33). Burrows are located in the vegetated portion. North Island: Bonin Petrels nested in large numbers in March 1913 (Bailey, 1956: 32; Willett, ms.). In recent years (Table 34), POBSP and BSFW personnel observed this species in very small numbers only three times; none was nesting. Seal Island: Wetmore (ms.) found 1 adult and 2 young in April 1923. POBSP and BSFW personnel found it nesting in 1963 and 1965. It was present in 1969, and may have been overlooked in other years because of diurnal surveys (Table 35). Southeast Island: Munter (ms.) probably found a Bonin Petrel egg in February 1916 at Southeast. Wetmore noted the presence of Bonins in April 1923. POBSP and BSFW personnel recorded burrows of this species each year since 1963 (Table 36). The highest populations (1,000) occurred in August 1967 and February 1969. Nest burrows are found only on the east portion in, south, and west of the west-central Eragrostis area. Burrow length is approximately 3 feet and only one-half to one foot beneath the ground surface. Banding and Movements The POBSP banded 365 adult Bonin Petrels; 33 were banded on Seal Island in March 1965 and 332 were banded on Southeast Island (276 in Feb- ruary and March 1963, 5 in September 1964, 25 in March 1965, and 26 in September 1966). POBSP and BSFW personnel captured 15 Bonin Petrels at Pearl and Hermes Reef; 12 were originally banded on the atolls 3 were originally from Kure. In addition, 2 originating at Pearl and Hermes moved to Kure anc another moved to Laysan. These data are presented in Appendix Tables 6a and 6b. Specimens Non-POBSP: USNM 300667-69, 9, 9, oh, collected 27 April 1923 by Wetmore; USNM 289189, skull, collected 27 April 1923 by Wetmore. These are first specimen records. Table 33. Observations of Bonin Petrels at Grass Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. 2 "Island honeycombed with...petrel holes" (Munter, ms.). 1923 27 Apr. 1 (Wetmore, ms.). 130 Table 33. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1963 5 Mar. 4 At least 2 nests with eggs found (POBSP, 1964d). 26-27 June iL Near-fledging young (POBSP, 1963). 1964 14 Mar. 1-2 (BSFW, 1964a; POBSP, 1964b). 18 Aug. O (POBSP, 1964a). 1965 19 Mar. 100-200 Few nests contained eggs or small chicks (POBSP, 1965b). 22 Mar. 2 Some burrows with eggs (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 0 (BSFW, 1966a). 1967 22 Mar. B None seen during brief diurnal visit (BSFW, 1967a; POBSP, 1967d). 31 May 2 Young (POBSP, 1967b). 29 Aug. 0 (POBSP, 1967a). 1968 24 Mar. 2 None seen during brief diurnal visit (BSFW, 1968; POBSP, 1968). Table 34. Observations of Bonin Petrels at North Island Population Date of Survey Estimate Breeding Status, Remarks, and References Mele} aly Wevvae 3, 000 Nesting abundantly (Bailey, 1956: 32; Willett, ms.). 1963 6 Mar. 0) (POBSP, 1964d). 23-25 June ©) (POBSP, 1963). 1964 19-20 Aug. O (POBSP, 1964a). 17 Sept. 0) (BSFW, 1964b; POBSP, 1964c). 1965 17-18 Mar. few No burrows examined (POBSP, 1965b). 1966 22 Sept. 0) (BSFW, 1966b). 3\ab Table 34. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1967 16-17 Mar. 5 Minimum number present (BSFW, 1967d). 29-30 Aug. 0 (POBSP, 1967a). 1969 12 Sept. 10 (BSFW, 1969d). Table 35. Observations of Bonin Petrels at Seal Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1923 27 Apr. 3 1 adult, 2 young (Wetmore, ms.). 1963 5 Mar. z Nesting (POBSP, 1964d). 26 June O None seen during diurnal survey (POBSP, 1963). 1964 14 Mar. 2 Burrows (BSFW, 1964a; POBSP, 1964b). 18 Aug. 2 None seen during diurnal survey (POBSP, 1964a). 1965 18-19 Mar. 30-50 Heavily incubated to pipped eggs found; some small young may have been present (POBSP, 1965b). 22 Mar. 34 Adults handled; 1 hatching egg (BSFW, 1965; POBSP, 1965a). 1967 22 Mar. a None observed on diurnal survey; no fresh burrows (BSFW, 1967a; POBSP, 1967d). 31 May % None seen during diurnal survey (POBSP, 1967»). 28 Aug. 2 None seen during diurnal survey (POBSP, 1967a). 1968 2 Mar. 2 None seen during diurnal survey (BSFW, 1968: POBSP, 1968). 1969 11 Sept. 5 (BSFW, 1969d). SZ Table 36. Observations of Bonin Petrels at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. ? Fresh egg found in "Wedge-tailed Shear- water burrows" (Munter, ms.). [Probably Bonin egg and burrow]. 1923 26-28 Apr. 2 Presence noted (Wetmore, ms.). 1961 12 Mar. 2 None seen but "most likely they are present in small numbers" (HDFG, 1961). 1963 26 Feb- 800 All active burrows observed contained young 8 Mar. (POBSP, 19644). 1964 13-14 Mar. 150-200 Adults; 100 nests estimated, some birds in- cubating eggs, no young seen (BSFW, 1964a; POBSP, 1964b). 16 Sept. 25 6 adults seen (BSFW, 1964a; POBSP 1964b). 1965 15-19 Mar. 100-300 Nests with either well-incubated eggs or small downy young (POBSP, 1965b). 21-22 Mar. 200 No eggs seen (BSFW, 1965; POBSP, 1965a). 1966 20-26 Sept. 4OOt+ Nocturnal adults; no eggs or young (BSFW, 1966b). 25-27 Sept. 200 Adults digging burrows; no eggs (POBSP, 1966). 1967 21-23 Mar. 150 Nocturnal adults; many burrows inspected but none contained eggs or young (BSFW, 1967a; POBSP, 1967d). 28, 30 May- 5) Large, fully-feathered juveniles (POBSP, 1 June 1967b). 28-30 Aug. 1, 000% No eggs or young; most birds sitting quietly on ground surface (POBSP, 1967a). 1968 22-24 Mar. 25-50 Burrows not checked for contents (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 1, 000 Nocturnal adults; burrows present, but no eggs or young (BSFW, 1969a). 10-19 Sept. 300 Adults (BSFW, 1969d). 135 BULWER'S PETREL Bulweria bulwerii Status Uncommon breeding species; recorded only on Southeast Island during summer. Maximum POBSP population estimate 15 in June 1963. Observations Bulwer's Petrels were tentatively recorded 12 March 1961 when HDFG personnel found remains of several petrels on Southeast Island; these, however, were very likely Sooty Storm Petrels. A few nesting Bulwer's Petrels, with eggs, were found on Southeast by POBSP personnel in June 1963; several were observed in empty burrows in August 1964; an adult and one young bird were found in September 1966 (Clapp and Woodward, 1968: 7-8). None has been seen since. Annual Cycle The only positive evidence of breeding--eggs in June 1963 and a near-fledging young in September 1966--indicate that the Bulwer's Petrel may follow a spring and summer breeding season (see Fig. 58) as do other populations in the Northwestern Hawaiian Islands. The lack of observations from four fall-winter surveys and seven early spring surveys indicates these petrels are absent from October through March. The three positive observations suggest that the population is extremely. small. These low numbers and the absence of Bulwer's during the May to June 1967 and 1969 surveys when breeding should have occurred may indicate that the local population is gradually becoming extirpated. Ecological Distribution Bulwer's Petrels have been recorded only from Southeast Island. Because of its small population and the few nocturnal surveys of islands other than Southeast, the species may have been overlooked on Grass, North and Seal Islands. Southeast Island: POBSP personnel observed 15 Bulwer's Petrels and three nests with eggs at Southeast 18 to 23 and 25 June 1963. Five petrels were found in empty burrows 16 to 19 August 1964. An adult and a young bird, with only a trace of down on its nape, were observed by BSFW and POBSP personnel 20 to 27 September 1966. Most Bulwer's Petrels have been found near the south shore. The three nests containing eggs were observed under boards, and the fourth, containing a young bird, was found under a piece of corrugated tin. Other petrels have been observed in burrows beneath vegetation, and in one instance, beneath a Red-footed Booby nest in Solanum. 134 Banding and Movements Nine adult Bulwer's Petrels were banded on Southeast Island, 6 in June 1963 and 3 in August 1964. Three of the 6 banded in June 1963 were recaptured locally in August 1964. None has been recaptured elsewhere. WEDGE-TATLED SHEARWATER Puffinus pacificus Status Common breeding species; present usually from March to November. Nests and occurs only on the four major vegetated islands. Recent POBSP and BSFW maximum population estimate 26,500 in August 1964. Observations Munro (1942: 12) saw "numbers of Wedge-tailed Shearwaters" before sailing into the lagoon on 6 July 1891. No landing was made; he left early on the 7th. Munter (ms.) in February 1916 observed a fresh egg in a "Wedge- tailed Shearwater burrow." This observation is probably erroneous; this species consistently returns in March and lays eggs in June; the burrow was probably that of a Bonin Petrel. Wetmore (ms.) found adults, but no eggs in April 1923. Galtsoff (1933: 19) published the first observation from the atoll after finding them in large numbers in summer 1930. Richard- son (1957: 16) noted that it breeds here. POBSP, BSFW, and HDFG personnel olsoniee their presence on the atoll in 1961, and 1963 to 1969 (Tables 37 to 4O)r3 Annual Cycle Wedge-tailed Shearwaters at Pearl and Hermes are summer-fall breeders (see Fig. 58). Recent data show adult birds arrive in small numbers normally during early March. By mid- or late March the nocturnal population increases and some diurnal activity occurs. Egg laying begins in early June and most eggs are laid by late June. Hatching commences in late July and most eggs are hatched by mid-August. Young birds probably remain until mid-October and a few into early November. Adults more than likely start leaving in early October, prior to the departure of their young. The population is composed mainly of white-phased individuals. Ecological Distribution Wedge-tailed Shearwaters nest on Grass, North, Seal and Southeast Islands. Grass Island: Wetmore (ms.) first observed Wedge-tailed Shearwaters in April 1923. POBSP and BSFW personnel found them present in all years (Table 37). Population estimates for most recent visits are lacking because of the diurnal nature of the visits and shortness of the surveys. An estimated 135 1,000+ adults and 500 nest burrows with eggs, however, were recorded by POBSP personnel in June 1963. Nests were found only in the vegetated western part. North Island: POBSP personnel first observed nesting Wedge-tailed | Shearwaters in June 1963, and found them on most subsequent visits (Table 38). Summer estimates of nests with eggs or young ranged from 1,000 to 2,000. | Nest burrows were in the vegetated northern portion. Individual nest burrows were under dense Solanum, Sicyos, and Eragrostis. Seal Island: Wetmore (ms.) noted Wedge-tailed Shearwaters in April 1923. POBSP and BSFW personnel observed them on most recent visits (Table 39). The only breeding population estimate is of 500 nests with eggs. This species nests in burrows in the vegetated portion. Southeast Island: In April 1923 Wetmore (ms.) found Wedge-tailed Shearwaters to be fairly common. Galtsoff (1933: 19) wrote of seeing large numbers in summer 1930. All recent visitors have recorded them during their spring-summer breeding season (Table 40). Fall estimates of adults and young range from 3,000 and 800 respectively to 14,000 and 5,000. The population is distributed over the entire island, but is most numerous in the area of high grass in the northwest sector of the eastern portion. In areas of scant vegetation, the lack of extensive root systems causes loose sand; consequently, burrows frequently collapse. In June 1967 one percent of the population was estimated to be dark-phased indi- viduals. Banding and Movements POBSP personnel banded 3,2/1 adults on North, Seal, and Southeast Islands (Table 41). Of these, 148 have been recaptured on the atoll (Table 16). In addition, one adult (USFW band no. 605-12758) banded on Kure 12 March 1964 was captured on Southeast one day later. An adult (615-16431) banded 17 August 1964 on Southeast was found injured at Kure 23 September 1964. Another adult (615-17584) banded on Southeast 18 August 1964 was captured on Manana Island, Oahu 2 June 1966. Specimens POBSP: USNM 492958, co, collected 7 March 1963 on Southeast by Amerson. This is a first specimen record. 136 Table 37. Observations of Wedge-tailed Shearwaters at Grass Island Population S| Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. - Island honeycombed with shearwater* and &§ petrel holes (Munter, ms.). 1923 27 Apr. 100 (Wetmore, ms.). 1963 5 Mar. 0 (POBSP, 1964d). 26-27 June 1, 000+ An estimated 500 nest burrows with eggs (POBSP, 1963). | 1964 14 Mar. 2 No evidence of nesting (BSFW, 1964a; POBSP, e i 18 Aug. ? Adults with eggs and nestlings in burrows (POBSP, 1964a). 1965 19 Mar. 100-200 A pair seen; no active burrows (POBSP, 1965b). 22 Mar. 20 Many burrows observed; only 1 pair seen during diurnal survey (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 2 Active burrows (BSFW, 1966b). 1967 22 Mar. % None seen during brief diurnal visit; no fresh burrow digging (BSFW, 1967a; POBSP, 19674). 31 May 200 Diurnal survey (POBSP, 1967b). 29 Aug. ? 1 adult seen during diurnal survey; of 25 nests examined none was active (POBSP, 1967a). 1968 24 Mar. 2 1 adult seen on brief diurnal visit (BSFW, 1968; POBSP, 1968). *Probably Bonin Petrel burrows. tsi Table 38. Observations of Wedge-tailed Shearwaters at North Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1963 6 Mar. 0 (POBSP, 1964d). 23-25 June 5,000 Adults; 2,000f burrows with eggs (POBSP, 1963). 1964 19-20 Aug. epee) Adults; 1,000 burrows with eggs, and 1,500 with very small nestlings (POBSP, 1964a). a7. Sepitn 1, 000- Burrows contained downy young (BSFW, 1964b; 1,500 POBSP, 1964c). 1965 17 Mar. 100-200 No eggs or young (POBSP, 1965b). 1966 22 Sept. 0 (BSFW, 1966b). 1967 16-17 Mar. O (BSFW, 1967d). 29-30 Aug. 7, 000- 1,000-2,000 medium-sized downy young (POBSP, 8, 000 1967a). 1969 12 Sept. 10 (BSFW, 1969d). Table 39. Observations of Wedge-tailed Shearwaters at Seal Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. u "Shearwater holes were found thickly scat- tered about the island" (Munter, ms.).* 1923 27 Apr. 600 300 pairs (Wetmore, ms.). 1963 5 Mar. @) (POBSP, 1964d). 26 June 1, 200 Adults, 500 nest burrows with eggs (POBSP, 1963). 1964 14 Mar. @) (BSFW, 1964a; POBSP, 1964b). 18 Aug. 1 Adults with nestlings and eggs in burrows (POBSP, 1964a). 1965 18-19 Mar. 10-15 Adults; no eggs or young (POBSP, 1965b). 22 Mar. 5 Only 1 pair (BSFW, 1965; POBSP, 1965a). 138 Table 39. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1967 22 Mar. th None seen on diurnal survey; burrows showed no signs of fresh digging (BSFW, 1967a; POBSP, 1967d). 31 May 200 Diurnal survey (POBSP, 1967b). 29 Aug. it None seen during diurnal visit; 5 or 6 burrows investigated were inactive (POBSP, 1967a). 1968 24 Mar. 0 Diurnal survey (BSFW, 1968; POBSP, 1968). 1969 11 Sept. O Diurnal survey (BSFW, 1969d). *Probably Bonin Petrel burrows. Table 40. Observations of Wedge-tailed Shearwaters at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. rg Fresh egg found in a "Wedge-tailed Shear- water burrow" (Munter, ms.). [Observation probably erroneous; possibly refers to Bonin Petrel]. 1923 26-28 Apr. fairly Evidently no eggs laid (Wetmore, ms.). common J 1930 23 July-Aug. large (Galtsoff, 1933: 19). numbers 1961 12 Mar. small (HDFG, 1961). numbers 1963 26 Feb.- 5 First observed 6 March; no more than 3 8 Mar. nightly (POBSP, 1964d). 18-23, 5 5 000+ Almost all nesting, with fresh or nearly 25 June fresh eggs (POBSP, 1963). 1964 13-14 Mar. 200-300 Nocturnal, only hot diurnal; some birds paired, some excavating burrows (BSFW, 1964a; POBSP, 1964b). Table 40. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1964 16-19 Aug. 19, 000 14,000 adults, 5,000 young; almost all active burrows contained young chicks; only 2 nests with eggs found (POBSP, 1964a). 16 Sept. 3,500 Downy young (BSFW, 1964b; POBSP, 1964c). 1965 15-19 Mar. 100-300 No active burrows (POBSP, 1965b). 21-22 Mar. 700 No nests, most birds apparently paired (BSFW, 1965; POBSP, 1965a). 1966 20-26 Sept. 450-500 Young in late downy stages (BSFW, 1966b). 25-27 Sept. 3,800 3,000 adults, 800 mostly large downy young (POBSP, 1966). 1967 21-23 Mar. 40-50 Nocturnal; few paired, no active burrows (BSFW, 1967a; POBSP, 1967d). 28, 30 May- 1,400- Adults; some copulating and others digging 1 June 2,600 burrows; neither eggs nor young found (POBSP, 1967b). 28-30 Aug. 9, 000 8,000 adults, less than 1,000 young; all active burrows with medium-sized downy chicks (POBSP, 1967b). 1968 22-24 Mar. 50 Scattered individuals roosting quietly on surface of ground; no breeding activity (BSFW, 1968; POBSP, 1968). 1969 31 Mar.- 1, 000 (BSFW, 1969b). 2 Apr. 26-31 May it Adults calling and digging burrows; no evidence of egg-laying (BSFW, 1969c). 10-19 Sept. 4 Adults (BSFW, 1969d). CHRISTMAS SHEARWATER Status September and early October. 139 Puffinus nativitatus Uncommon breeding species; present from late February until late Nests at Southeast Island; occurs at North 140 *pepueq azeam Sunof ON, 9S Lie 0g T2701 0) 9€ 4320N ©) 1 Teas 9S Lté et qyseoyynos I961 9961 G96T T8701 +96T 4H96T T2401 96T 96T pueTst ‘ony °qdeg * Len +96T *qdag "ony €96T aunt “Le -" gai % dSd0d 2u} Aq gooy somzayH pue [1eveq 4e Ppapueq SZayemIeaYyg pPeTTey-sspeM JIMPY “Th eTAeL V41 Island. POBSP and BSFW maximum population estimate 50 in March 1965 and 1968. Observations Wetmore (ms.) had a Christmas Shearwater reported to him in April 1923. Clapp and Woodward (1968: 8-9) published the first atoll record from early POBSP data. POBSP, BSFW, and HDFG have noted its presenee in small numbers each year since 1963 except 1969 (Table 42). Annual Cycle The local annual breeding cycle is presented in Figure 58. Adults begin arriving by late February; egg laying probably begins in mid- or late April. Young usually hatch early in June with some eggs probably remaining until early July. Young probably fledge in late August and September. None occurs in late fall and early winter. March populations ranged from O to 50. Ecological Distribution Christmas Shearwaters nest only at Southeast Island; adults are known from North Island. North Island: Five adults were recorded by BSFW personnel 17 September 1964. This species may nest here. Southeast Island: Although Wetmore (ms.) recorded the first Christmas Shearwater in 1923, Clapp and Woodward (1968: 8-9) first published its occurrence from 1963 to 1966 POBSP data. POBSP and BSFW personnel have since recorded it in 1967 and 1968 (Table 42). The population is small with a maximum estimate of 50. Nests have been found under Scaevola and boards. Individuals have been sighted on open ground and in thick Solanum and Eragrostis. The species has been recorded from both the east and west portions of the island. Banding and Movement One adult was banded by POBSP personnel at Southeast in September 1966; no interisland movement exists for this species. Specimen Records POBSP: USNM 492965, do, collected on Southeast Island 7 March 1963, POBSP collector unknown; USNM 492966, o', collected on Southeast Island 26 February 1963, POBSP collector unknown. whe Table 42. Observations of Christmas Shearwaters at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. - Not mentioned in report (Munter, ms.). 1923 26-28 Apr. mn Reported to, but not seen by, Wetmore (ms.). 1930 23 July- - Not mentioned in report (Galtsoff, 1933). Aug. 1961 12 Mar. % Presence noted (HDFG, 1961). 1963 26 Feb.- . 5 No nests (Clapp and Woodward, 1968: 8-9; 8 Mar. POBSP, 1964d). 18-23, 5-10 Adults, 1 nest with egg (Clapp and Woodward, 25 June 1968: 8-9; POBSP, 1963). 1964 13-14 Mar. 12-15 2 diurnal, 15 nocturnal; no nests (BSFW, 1964a; POBSP, 1964b). 16-19 Aug. 10 Adults; only 2 found in a burrow which contained neither egg nor young (POBSP, 1964a). 16 Sept. 15-25 No evidence of nesting; 5 birds actually seen (BSFW, 1964b; POBSP, 1964c). 1965 15-19 Mar. 15-20 Adults sitting in pairs; no nests found (POBSP, 1965b). 21-22 Mar. 50 No active nests; 12 adults actually ob- served (BSFW, 1965; POBSP, 1965a). 1966, 1 Apr - Not mentioned in notes (BSFW, 1966a). 20-26 Sept. - Not mentioned in notes (BSFW, 1966b). 25-27 Sept. 20 3 large young (+21 days); only a few birds seen at sunset (Clapp and Woodward, 1968: 8-9; POBSP, 1966). ; 1967 21-23 Mar. 0 (BSFW, 1967a; POBSP, 19674d). 28, 30 May- <50 4 breeders: 2 nests with eggs (POBSP, 1 June 1967b). 28-30 Aug. 20 Adults found in pairs near empty burrows (POBSP, 1967a). 143 Table 42. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1968 22-24 Mar. 50 Ca. 5 pairs actually seen; no evidence of nesting (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. ) (BSFW, 1969a). 31 Mar.- ) (BSFW, 1969b). 2 Apr. 26-31 May O (BSFW, 1969c). . 10-19 Sept. O (BSFW, 1969d). SOOTY STORM PETREL Oceanodroma tristrami Status Common breeding species; present possibly late fall, winter, and spring. Nests at North, Seal, and Southeast Islands; known from Grass Island. Maximum POBSP and BSFW estimated population 7,500 in February 1969. Observations Willett (ms.) first recorded a nesting colony in March 1913 (see also Bailey, 1956: 60-61). Wetmore (ms.) observed young in April 1923. Richard- son (1957: 19) also records it as breeding. POBSP and BSFW personnel found moderate numbers on the four vegetated islands each year since 1963. Annual Cycle The Sooty Storm Petrel has a winter-spring breeding season locally (Fig. 58). Adults possibly arrive in late fall, with egg laying probably starting in late December. Young most likely begin hatching in late January; eggs are known as late as mid-March and large young are known as early as early March. Adults probably leave by mid-May, with young fledging by late May. Ecological Distribution Sooty Storm Petrels nest at North, Seal, and Southeast Islands, and occur at Grass Island. Grass Island: POBSP personnel found this species 5 March 1963; none has been observed since. ih North Island: Willett (ms.) found a small colony, with young, on 15 March 1913 (see also Bailey, 1956: 60-61). POBSP personnel recorded 50 to 75 nestling and 100-200 adult Sooty Storm Petrels 17-18 March 1965. Two adults were handled and another seen 16 to 17 March 1967 by BSFW personnel. Seal Island: Wetmore (ms.) observed a few young "Bulwer's Petrels" , 27 April 1923. These, however, were probably Sooty Storm Petrel young, ; for Bulwer's Petrel young do not usually start hatching until mid-summer. ; A few young and 15 to 25- adults were found 18 to 19 March 1965. ] Southeast Island: Young Sooty Storm Petrels were observed by Wetmore (ms.) in April 1923. POBSP and BSFW personnel recorded them nesting each spring since 1963 (Table 43). Although a high of 7,500 adults was esti- mated during the middle third of February 1969, the average March popula- tion (6 visits) is 441, and ranges from 200 to 1,000. Nest burrows are most abundant in the Bermuda grass (Cynodon) area on the west half of the eastern portion; they are also found in nearby open rubble and Boerhavia, and in an Eragrostis area on the south side of the eastern portion. On the western portion burrows are located in an area of open coral rubble and Coronopus in the west interior. Banding and Movement In all, 788 Sooty Storm Petrels have been banded by POBSP and BSFW personnel on three islands(Table 44). Of these, 20 have been recaptured on the atoll (Table 16). No inter-atoll movement is known. Specimens POBSP: USNM 496219, 21, 99, moderate light fat, respectively, col- lected at Southeast 14 March 1964 by B. Amerson; USNM 496220, o, light fat, collected at Southeast 14 March 1964 by B. Amerson. Non-POBSP: USNM 240027, ?, collected 15 May 1913 by Willett; USNM 28920}, skull, collected 27 April 1923 by Wetmore; USNM 300811-12, o,9, collected 27 April 1923 by Wetmore; USNM 300813, unsexed, collected 26 April 1923 by Wetmore; USNM 300814, 3s, collected 16 May 1923 by Reno. These are first specimen records. Table 43. Observations of Sooty Storm Petrels at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. - Not mentioned in report (Munter, ms.). : 1923 26-28 Apr. @ Young only (Wetmore, ms.). 145 Table 43. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1930 23 July- - Not mentioned in report (Galtsoff, 1933). Aug. 1961 12 Mar. O None seen, but 2 dead may have been this species (HDFG, 1961). 1963 26 Feb.- 1, 000 Eggs to chicks with well-developed pri- 8 Mar. maries, secondaries, and back feathers (POBSP, 1964d). 18-23, 0 (POBSP, 1963). 25 June 1964 13-14 Mar. 200+ 7 chicks (BSFW, 1964a; POBSP, 1964b). 16-19 Aug. 0 (POBSP, 1964a). 16 Sept. O (BSFW, 1964b; POBSP, 1964c). 1965 15-19 Mar. 600-700 400-500 adults, 200 chicks; 1 burrow with egg (POBSP, 1965b). 21-22 Mar. 700° Nocturnal adults; 5 young seen (BSFW, 1965; POBSP, 1965a). ag66. il Apr’, 2 2 chicks (BSFW, 1966a). 20-26 Sept. O (BSFW, 1966b). 25-27 Sept. 0 (POBSP, 1966). 1967 21-23 Mar. 200+ Nocturnal adults; ca. 10 chicks: all downy but with primaries just breaking out of sheaths (BSFW, 1967a; POBSP, 1967d). 28, 30 May- O (POBSP, 1967b). 1 June 28-30 Aug. 0) (POBSP, 1967a). 1969 10-12 Feb. 7,500 Adults, nocturnal only; eggs to large young (BSFW, 1969a). 31 Mar.= 250 (BSFW, 1969b). 2 Apr. 26-31 May 2 2 adults, 1 near fledging young seen (BSFW, 1969c). 10-19 Sept. ) (BSFW, 1969d). > TeL0OL Suno, 4IMpyY = TeqFOL :T e101 Sunox 4 Tpy YAIoN 410py Teas > T2101 Sunonz qInpy 4 4seeyynos STeqOL 696t 996t Lo6tT Te90L G96T = SOOT) 96 )©=— EQET = SSP TO pueTst "qed TN S961 5 i Sa -93y =" 99d ee BS 6 2 ee Oe ree el Misd MiSd Masa MiSd dSdod MASH dSd0d ‘aepueg Jooy Souey pue [ieeq 4e papueq sTazjeq wI04g AjooS “Ht ETAeL 146 147 RED-TAILED TROPICBIRD Phaethon rubricauda | Status Common breeding species; present from late winter probably through late fall; possible straggler rest of year. Nests at Grass, North, Sand, and Southeast Islands. Maximum POBSP and BSFW population estimate 159 in June 1963. Observations Munro (1942: 12) recorded a few tropicbirds, probably Red-tailed Tropicbirds, off Pearl and Hermes Reef on 6 July 1891. Wetmore (ms.) noted nesting in April 1923. Clapp and Woodward (1968: 11) published the first record of nesting Red-tailed Tropicbirds using POBSP data for 1963 to 1967. Tables 45 to 48 present all observations, including POBSP and BSFW for 1963 to 1969. Annual Cycle Figure 58 presents the local annual breeding cycle. Adults arrive in early February, with eggs appearing as early as mid-March. Most eggs are probably laid by late April; some are known into August. Young are known as early as early May with peak fledging probably in July and August; some have been recorded in late September. The young generally leave the atoll as soon as they fledge. Adults probably are not present during late fall and early winter. Ecological Distribution Red-tailed Tropicbirds are known to breed at Grass, North, Seal, and southeast Islands, and have been observed flying over Little North. Grass Island: Nesting Red-tailed Tropicbirds were observed by Wetmore (ms.) in April 1923. Clapp and Woodward (1968: 11) recorded their presence in 1963, 1964, and 1967 from POBSP data. POBSP and BSFW personnel noted the species in 1966, but failed to find it in 1965 and 1968 (Table 45). Red-tailed Tropicbirds nest under thick vegetation, especially high clumps of Eragrostis. The maximum number of nests in any year was two in 1963. North Island: Clapp and Woodward (1968: 11) published 1963 to 1965 data on nesting Red-tailed Tropicbirds; their March 1967 data, however, should have referred to Southeast Island. All POBSP and BSFW data are shown in Table 46. The maximum number of nests was 19 in June 1963, which coincided with the peak population of 64. All nests were placed under clumps of Eragrostis or Solanum. Seal Island: In April 1923 Wetmore (ms.) observed nesting. Clapp and Woodward (1968: 11) published part of the 1963 and 1964 POBSP data. All data are presented in Table 47. 148 Wetmore's (ms.) 1923 Seal population estimate of 30 Red-tailed Tropic- birds has not been equaled; 20 adults and 6 nests was the largest number present in recent years. Eragrostis clumps are favored vegetation under which this species nests. Southeast Island: Wetmore (ms.) observed the species in April 1923 and POBSP and BSFW noted its presence from 1963 to 1966 (Clapp and Woodward, 1968: 11). These and other data are presented in Table 48. March population estimates average 17 and range from 6 to 25. The two June visits have produced the highest populations, 70 and 59. Most nest under large Eragrostis clumps located west of the tidal pools, under low Scaevola, and in and under old oil drums scattered over the island. Other Islands: Two adults flew over Little North Island on 27 August 1967; no nests or previously used nest sites were found. This species may occasionally fly over the remaining four sandy islands. Banding and Movements The POBSP banded 171 Red-tailed Tropicbirds on four islands (Table 49). Of these, 56 have been recaptured on the atoll (Table 16). An adult, sex unknown, banded at Southeast 28 August 1967 (USFW band no. 705-12788) was captured by POBSP personnel 18 May 1969 on Johnston Atoll where it was on an egg. Specimens Non-POBSP: USNM 301000-02, 99, collected 27, 27, and 26 April 1923 by Wetmore. These are first specimen records. Table 45. Observations of Red-tailed Tropicbirds at Grass Island Population Date of Survey Estimate Breeding Status, Remarks, and References TO 23 eimai. 4 Nesting among high grass clumps (Wetmore, ms.). F 1963 5 Mar. 2 Present (POBSP, 196d). y ® é 26-27 June 5) Adults, 2 nests with eggs (Clapp and Woodward, 1968: 11; POBSP, 1963). 4 1964 14 Mar. iD Flying over; no nests (BSFW, 1964a; POBSP, 5 1964b). , @ 18 Aug. 9 8 adults, 1 nest with large young (Clapp and Woodward, 1968: 11; POBSP, 1964a). 1965 19 Mar. 0 (POBSP, 1965b). 149 Table 45. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1965 22 Mar. 0 (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 2 Adults; no nests (BSFW, 1966b). 1967 22 Mar. 1 Adult on ground and flying; no nests (Clapp and Woodward, 1968: 11; BSFW, 1967a; POBSP, 19674). 31 May O (POBSP, 1967b). 29 Aug. 2 Adults in courtship display over island; no nests found (POBSP, 1967a). 1968 24 Mar. 0 (BSFW, 1968; POBSP, 1968). Table 46. Observations of Red-tailed Tropicbirds at North Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1963 6 Mar. 1 Flying over; no nests (POBSP, 1964d). 23-25 June 64 60 adults; 4 nestlings, 38 breeders; 19 nests: 15 (79%) with eggs, 4 (21%) with young (Clapp and Woodward, 1968: 11; POBSP, 1963). 1964 19-20 Aug. 55 4O adults, 32 breeders, 7 non-breeders, 15 young; 16 nests: 1 (6%) with an egg, 5 (31%) with small downy young, 10 (63%) with larger young (Clapp and Woodward, 1968: 11; POBSP, 1964a). 17 Sept. Tet 10 adults, 2 breeders, 1 large dependent young (Clapp and Woodward, 1968: 11; BSFW, 1964b; POBSP, 1964c). 1965 17-18 Mar. 5-10 Adults, 4 breeders, 2 nests with eggs (Clapp and Woodward, 1968: 11; POBSP, 1965b). 1966 22 Sept. 3 Adults flying overhead; no nests (BSFW, 1966b). 1967 16-17 Mar. fo) (BSFW, 1967d). 29-30 Aug. Wo+ 20 breeders, 15 non-breeders; from large downy young to dependent immatures; sample nest count of 10 nests: 1 (10%) with a large downy young, 9 (90%) with dependent immatures (POBSP, 1967a). 150 Table 46. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1969 12 Sept. i 3 adults, 1 young (BSFW, 19694). Table 47. Observations of Red-tailed Tropicbirds at Seal Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1923 2/7 Apr. 30 15 pairs nesting among high grass clumps (Wetmore, ms.). 1963. 5 Mar. O (POBSP, 1964d). 26 June 20 12 breeders, 8 non-breeders; 6 nests with eggs (Clapp and Woodward, 1968: 11; POBSP, 1963). 1964 14 Mar. 6 Flying over, courtship behavior; no nests (BSFW, 1964a; POBSP, 1964b). 18 Aug. 17-18 15 adults; 8 breeders; 4 nests: 1 leg, 1 small downy young, 1 medium-sized young, and 1 dependent immature (Clapp and Woodward, 1968: 11; POBSP, 1964a). 1965 18-19 Mar. 0 (POBSP, 1965b). { 22 Mar. 0 (BSFW, 1965; POBSP, 1965a). 1966121 Sept. {HP 6 4 breeders (only 3 adults seen), 2 nearly full-grown young (BSFW, 1966b). 1967 22 Mar. ee O (BSFW, 1967a; POBSP, 19674). 31 May 6 Breeders, 3 nests with eggs (POBSP, 1967b). 29 Aug. 2 Flying over; no nests (POBSP, 1967a). 1968 24 Mar. 0) (BSFW, 1968; POBSP, 1968). Table 48. Observations of Red-tailed Tropicbirds at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1923 26-28 Apr. 20 [Presumably nesting.] (Wetmore, ms.). iljyil Table 48. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1961 12 Mar. - Not mentioned in report (HDFG, 1961). 1963 26 Feb.- 15 Adults; up to 11 in air at once, some on 8 Mar. ground; no eggs (Clapp and Woodward, 1968: 11; POBSP, 1964d). 18-23, 70 65 adults, 58 breeders; 29 nests counted: 25 June 24 (83%) with eggs; 3 (10%) with small downy young; and 2 (7%) with larger young (Clapp and Woodward, 1968: 11; POBSP, 1963). 1964 13-14 Mar. 6 h breeders; 2 nests with eggs, 1 of which was abandoned; 6 seen flying at once (Clapp and Woodward, 1968: 11; BSFW, 1964a; POBSP, 1964b). 16-19 Aug. 18 15 adults, 12 breeders; 6 nests counted: 3 (50%) with eggs, 1 (17%) with a small downy young, 1 (17%) with dependent imma- ture; 1 (17%) with young of unrecorded size (Clapp and Woodward, 1968: 11; POBSP, 1964a). 16 Sept. 15 6 breeders; 3 large young (Clapp and Woodward, 1968: 11; BSFW, 1964b; POBSP, 1964c). 1965 15-19 Mar. 15-20 8 breeders; 4 nests with eggs (Clapp and Woodward, 1968: 11; POBSP, 1965b). 21-22 Mar. 2 Breeders with an egg (BSFW, 1965; POBSP, 1965a). 1966 1 Apr. 12 As many as 12 flying at one time (BSFW, 1966a). 20-26 Sept. 2 Adults; no nests (BSFW, 1966b). 25-27 Sept. several Flying over; no nests (POBSP, 1966). 1967 21-23 Mar. 15 Adults; 4 breeders; 2 nests with eggs (BSFW, 1967a; POBSP, 1967d).* 28, 30 May- 59 56 breeders; 28 nests counted: 25 (89%) with 1 June eggs, 3 (10%) with small downy young; 1 nest also empty (POBSP, 1967b). 28-30 Aug. 39 20 breeders; 10 nests counted: 1 (10%) with eggs 3 (30%) with large downy young; 6 (60%) with dependent immatures (POBSP, 1967a). 152 Table 48. (continued) Population , Date of Survey Estimate Breeding Status, Remarks, and References : 1968 22-24 Mar. 20-25 Adults; 4 breeders: 2 nests with eggs; at least 11 birds on ground and 9 in air at one time (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 7 Adults; 1 on ground (BSFW, 1969a). 31 Mar.- 22 11 nests counted (BSFW, 1969b). 2 Apr. 26-31 May 40 36 breeding adults; 18 nests counted: 16 (89%) with eggs and 2 (11%) with half-grown young; 2 empty nests (BSFW, 1969c). 10-19 Sept. 6 4 breeders, 2 young (BSFW, 1969d). - *Erroneously listed by Clapp and Woodward (1968: 11) under North Island. BLUE-FACED BOOBY Sula dactylatra Status i Common breeding species; present year-round with possible population i decrease in late fall and early winter; occurs and nests on all islands except Sand and Planetree where it roosts only. Maximum POBSP and BSFW population estimate 585 in June 1963. Observations Blue-faced Boobies, with nests, were first recorded in March 1913 (Bailey, 1956: 32; Willett, ms.). Richardson (1957: 20) also notes that it breeds here, but gives no data. They have been recorded by all observers, including POBSP and BSFW personnel, since (Tables 50 to 56). Annual Cycle The local annual breeding cycle is presented in Figure 58. Some remain on the atoll year-round; numbers increase in winter and decrease in late sum- mer and fall. This fluctuation coincides with the breeding cycle. On the basis of recent data, eggs are laid as early as February; most eggs are laid by the end of March. Although hatching occurs as early as 1 April, most occurs in May and very few eggs remain into late June or July. Fledging takes place in late summer and fall. The total number of active nests in March has ranged from 66 to 10%. 153 :Te10L SUTTISON FINDY TeIOL >Te101, SUTTISON JIMDY Y9-ZONl o) :Te101, 0 BUTTISON 0 41Npy Teas ) :Te201, 0 BUTT ISON 0 qINpy sseiy N OIN co Olco dr >Te10L 0 BUTTISON L 410PY qseayynos gt 61 0 6 61 T8401 L961 L961 GO6L sseTo pueTst ),96T “Sny Key “TEN “Sny €OQ6T aune * Tey -o3V aAING FO potszag said oa) BS MTs MN ON mM Im fe oa) Mm Olm a mr AHI + Aho 2S 5 \O o =! oO p [@) i=l (oe) \O od (99) \O od asdod eu} Aq gooy soulzey pue Taveq 4e pepueq spxtqoTdoazy peTtei-pay “6h eTaeL 154 Ecological Distribution Blue-faced Boobies nest at Bird, Grass, Kittery, Little North, North, Seal, and Southeast Islands; in addition, they roost on Planetree and Sand. Grass Island: Munter (ms.) first recorded Blue-faced Boobies in February 1916. Galtsoff's 1930 photographs (in litt.) show nesting Blue- faced Boobies at either Grass or Seal. In each year since 1963 (Table 50) except 1966 and 1969, POBSP and BSFW personnel found them nesting in small numbers. The most birds seen were in March 1965; the most nests were found in March 1963. This species nests on the upper sand beaches and eggs are laid in a slight depression in the bare sand. Kittery Island: POBSP personnel recorded the first Blue-faced Boobies, with 24 nests, in March 1963. Since, they have been recorded each year until 1969 (Table 51). The March population has fluctuated from 22 to 80; March nests have varied from 11 to 32. Nests are primarily near the perimeters and especially along the north- west edge. A few, however, nest as much as 150 yards inland from the water- line of this large sand island. Little North Island: Rice (ms. a) first recorded Blue-faced Boobies on 14 October 1957 while flying over the atoll. POBSP and BSFW personnel have subsequently found them nesting each year since March 1963, except in 1968 when the island was not visited (Table 52). A high nest count of 29 was reached in June 1963. In August 1967 a large nocturnal roosting club of 150 formed at dusk. Nests are placed on the raised, central portion. North Island: Nesting Blue-faced Boobies first were recorded in March 1913 (Bailey, 1956: 32; Willett, ms.). Rice (ms. a) next noted them in October 1957. None was observed again until March 1963 when POBSP personnel found them nesting. POBSP and BSFW personnel have since found them nesting each year except 1966 (Table 53); no visits were made in 1968. The highest nest count was 29 in June 1963; the highest population estimate, 204, was also made at that time. This species nests on the upper beach crest on all sides of the island. Nocturnal roosting birds are usually present in a club on the south tip. Seal Island: Munter (ms.) observed several non-nesting Blue-faced Boobies in February 1916. Galtsoff's photographs (in litt.) show nests in summer 1930. POBSP personnel recorded it nesting each year since 1963 except 1969 (Table 54). March nest counts average 25 and range from 14 to 33. The highest March population estimate of 100-125 in 1965 equals that of May 1967 when 53 nests were counted. Blue-faced Boobies nest on the upper beach crest of the vegetated por- tion, but prefer the rock ledge and open, coral rubble of the east portion. 1p Southeast Island: Blue-faced Boobies with eggs were first noted in February 1916 by Munter (ms.). They have been recorded on all visits since (Table 55). March nest counts made by POBSP and BSFW personnel average 26 and range from 6 to 40. March population estimates average 61 and range from 20 to 100; a peak population of 105 occurred in June 1963. This species restricts nesting to the coral sand area just above the shoreline. The majority of the population nests on the west shore of the east section, on the seaward rock ledge of the western section, and on the south and southwest beaches of both sections. Other Islands: A small population nests at Bird Island. Rice (ms. a) noted the species there in October 1957; POBSP and BSFW personnel found them nesting in 1964, 1965, and 1967. The maximum population estimate at Bird is 10 birds (Table 56). POBSP and BSFW personnel also noted small numbers of roosting Blue- faced Boobies at Sand and Planetree Islands (Table 56). Banding and Movements POBSP personnel have banded 587 and BSFW personnel 6 Blue-faced Boobies on 8 islands (Table 57); of these, 249 have been recaptured on the atoll (Table 16). In addition, 3 Blue-faced Boobies--2 banded at Kure and 1 at French Frigate Shoals--were captured at Pearl and Hermes. Furthermore, 12 banded on the atoll moved to other atolls: 5 to Kure, 3 to Laysan, 2 to Lisianski, and 1 each to French Frigate Shoals and Johnston; these data are presented in Appendix Tables 7a and 7b. Specimens Non-POBSP: USNM 300951, 2, collected April 1923 by Wetmore; USNM 300952-53, &, 2, collected 28 and 26 April 1923 by Wetmore. These are first specimen records. Table 50. Observations of Blue-faced Boobies at Grass Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. few Nesting had not begun (Munter, ms.). 1923, 27 Apr. - Not mentioned in notes (Wetmore, ms.). 1957 4 Oct. iL Observed from aerial survey (Rice, ms. a). 1963 5 Mar. 12 Breeding adults; 6 nests (POBSP, 1964d). 26-27 June 15) 10 breeders; 5 nests: 1 with a small downy young, 4 with medium-sized or larger young (POBSP, 1963). 156 Table 50. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1964 14 Mar. 4 2 breeders; 1 nest with eggs (BSFW, 1964a; POBSP, 1964b). 18 Aug. 2 Immatures (POBSP, 1964a). 1965 19 Mar. 15-20 Adults; 4 breeders: 2 nests with eggs (POBSP, 1965b). 22 Mar. 4 Adults on 2 nests with eggs (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. - None mentioned in notes (BSFW, 1966b). 1967 22 Mar. al Adult; no nests (BSFW, 1967a; POBSP, 1967d). 31 May 6 4 breeding adults; 2 nests: 1 with a small . 9 downy young, 1 with a medium-sized downy j young (POBSP, 1967b). 29 Aug. 3 No adults observed but 1 dependent immature seen (POBSP, 1967a). 4 1968 2 Mar. 4 2 breeders: one nest with egg; 2 other a adults flying over (BSFW, 1968; POBSP, 1968). q 1969 11 Sept. O (BSFW, 1969d). Table 51. Observations of Blue-faced Boobies at Kittery Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1963 5 Mar. 4gt Ca. 24 nests with eggs (POBSP, 19644). 26 June 84 60 adults; 50 breeders: 25 nests counted: 1 (4%) with eggs, 24 (96%) with young (POBSP, 1963). 1964 14 Mar. 70-80 Adults, 64 breeders: 32 nests with eggs (BSFW, 1964a; POBSP, 1964b). 18 Aug. 5) 2 adults and 3 immatures seen from Seal I. (POBSP, 1964a). 1965 18 Mar. 25-35 22 breeders; 11 nests with eggs (POBSP, 1965b). ii Table 51. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1965 22 Mar. ho 34 breeders; 17 nests with eggs (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 11 9 adults and 2 flying immatures (BSFW, | 1966b). 1967 22 Mar. 22 Adult breeders; 11 nests (BSFW, 1967a; POBSP, 1967d). 31 May (ale 50 adult breeders; 25 nests: 4 (16%) with eggs, 4 (16%) with small downy young, 17 (68%) with medium-sized or large downy young (POBSP, 1967b). 29 Aug. 30 14 adults, 10 immatures counted, some of which were still being fed by their parents (POBSP, 1967a). 1968 24 Mar. Ne) 32 breeders; 16 nests with eggs counted (BSFW, 1968; POBSP, 1968). Table 52. Observations of Blue-faced Boobies at Little North Island Population Date of Survey Estimate Breeding Status, Remarks, and References 177 ot KOck. ah Observed from aerial survey (Rice, ms. a). 1963. +6 Mar. ? Nesting (POBSP, 196d). 23, 25 June 73 58 breeders; 29 nests counted: 1 (44%) with eggs, 5 (17%) with small young, 23 (79%) with medium- or large-sized downy young (POBSP, 1963). 1964 19 Aug. ial 10 adults; 2 breeders, 1 downy young (POBSP, 1964a). 17 Sept. te 51 adults counted; 2 immatures, 1 large de- pendent young (BSFW, 1964b; POBSP, 1964c). 1965 18 Mar. 20-30 18-20 breeders; 9-10 nests with eggs (POBSP, 1965b). 1966 22 Sept. 19 16 adults, 3 flying immatures (BSFW, 1966b). 158 Table 52. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1967 29 Aug. 200 20 breeders, 10 nests counted: 2 (20%) 7 with eggs; 1 (10%) with a small downy young, 5 (50%) with medium-sized or large downy young, 2 (20%) with large dependent young; 150 in a nocturnal roosting club (POBSP, 1967a). 1969 31 Mar. 25 Breeders, most nests contained eggs (BSFW, 1969b). 12 Sept. 10 Adults (BSFW, 1969d). Table 53. Observations of Blue-faced Boobies at North Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1913 15 Mar. 100 Breeders; most nests with eggs, 2-3 with newly hatched young (Bailey, 1956: 32; Willett, ms.). 15a We Oct 12 Observed from aerial survey (Rice, ms. a). 1963 6 Mar. 2 Nests with eggs (POBSP, 196d). 23-25 June 204 175 adults; 58 breeders; 29 nests: 6 (21%) with small downy young, 23 (79%) with medium- sized or larger young (POBSP, 1963). 1964 19-20 Aug. os Adults, some dependent immatures; nocturnal roosting clubs (POBSP, 1964a). 17 Sept. 30 20 adults and 1 flying immature counted (BSFW, 1964b; POBSP, 1964c). 1965 17-18 Mar. 100-150 Adults; 32-34 breeders: 16 or 17 nests, all with eggs (POBSP, 1965b). 1966 22 Sept. 6 Adults; no young (BSFW, 1966b). 1967 16-17 Mar. 36+ 34 breeders: 16 nests with eggs, 1 with a 4 small young; 20 nocturnal roosting adults (BSFW, 1967d). 29-30 Aug. 60= 4O adults; 20 immatures, many of them still dependent (POBSP, 1967a). 159 Table 53. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1969 31 Mar. 14 6 nests: 4 with eggs and 2 with young (BSFW, 1969b). 12 Sept. 4 Adults (BSFW, 1969d). Table 54. Observations of Blue-faced Boobies at Seal Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. several No nests (Munter, ms.). seen 1923 27 Apr. - Not mentioned in notes (Wetmore, ms.). 1930 July-Aug. few Nesting (Galtsoff, photograph). 1963 5 Mar. ? Nesting on the beach; some with eggs (POBSP, 1964d). 26 June 98 56 breeders; 12 non-breeders; 28 nests counted: 3 (11%) small young, 25 (89%) medium-sized or large downy young (POBSP, 1963). 1964 14 Mar. 70 Adults; 66 breeders: 33 nests, all with eggs (BSFW, 1964a; POBSP, 1964b). 18 Aug. 18 9 adults counted; 12 breeders: 6 dependent immatures counted (POBSP, 1964a). 1965 18-19 Mar. 100-125 Adults only; 28 breeders: 14 nests counted, all with eggs (POBSP, 1965b). 22 Mar. 28 Breeders; 13 adults observed; 14 nests counted, all with eggs (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. a4 16 adults, 8 immatures capable of flight (BSFW, 1966b). 1967 22 Mar. 46 Breeders; 23 nests counted, all with eggs (BSFW, 1967a; POBSP, 1967d). 31 May 125 106 breeders; 53 nests counted: 34 (64%) with eges; 4 (8%) with small downy young; 15 (28%) with medium-sized or large downy young (POBSP, 1967 b). 160 Table 54. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1967 29 Aug. 62 90 flying birds; 24 breeders: 10 large downy young, 2 dependent immatures (POBSP, 1967a). 1969 24 Mar. 60 50 breeders: 25 nests counted, all with eggs (BSFW, 1968; POBSP, 1968). Table 55. Observations of Blue-faced Boobies at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks , and References 1916 4 Feb. 70 Eggs (Munter, ms.). 1923 26-28 Apr. 100 50 nesting pairs; 1 nest with an egg (Wet- more, ms.) 1930 23 July- large (Galtsoff, 1933: 19). Aug. numbers 1957 14 Oct. 8 Observed from aerial survey (Rice, ms. a). 1961 12 Mar. not With eggs or small young (HDFG, 1961). abundant 1963 26 Feb.- 100 50 non-breeders; 50 breeders: 2 nests with 8 Mar. 1 or 2 eggs (POBSP, 1964d). 18-23, 105 Te breeders, 33 young; 36 nests counted: 3 25 June (8%) with eggs; 1 (3%) with a recently hatched young; 6 (17%) with small downy young; 26 (72%) with larger young (POBSP, 1963). 1964 13-14 Mar. 70-80 Adults; 35-40 nests contained eggs (BSEW, 1964a; POBSP, 1964b). 16-19 Aug. 50 ho adults, 2 dependent immatures, 8 other im- matures (POBSP, 1964a). 16 Sept. 20-30 Adults, 1 flying immature; no nests (BSFW, 1964b; POBSP, 1964¢), 1965 15-19 Mar. 60-70 Adults; 34 breeders: 17 nests counted, all with eggs (POBSP, 1965p). 21-22 Mar. 35 Adults; 24 breeders: 12 nests counted, all with eggs (BSFW, 1965; POBSP, 1965a). 161 Table 55. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1966) 1 Apr. 36 18 nests counted, all with eggs; 1 egg hatching (BSFW, 1966a). 20-26 Sept. 46 40 adults; 12 breeders with dependent young, | each capable of flight (BSFW, 1966b). | 25-27 Sept. 54 50 adults; 8 breeding adults with 4 depen- dent immatures (POBSP, 1966). 1967 21-23 Mar. 50 Adults; 32 breeders with 16 nests, all with eggs (BSFW, 1967a; POBSP, 19674d). 28, 30 May- 95 78 breeders, 17 young; 39 nests counted; 1 June sample of 22 nests: 5 (23%) with eggs; 1 (5%) with small downy young, 15 (68%) with medium or large downy young, 1 (5%) depen- dent immature (POBSP, 1967b). 28-30 Aug. 19 15 flying birds; 8 breeders: 2 nests with large downy young, 2 dependent immatures (POBSP, 1967a). 1968 22-24 Mar. 50 4h breeders with 22 nests; 48% of nests with fresh or very slightly incubated eggs, the rest more heavily incubated; 2 pre-nesting pairs (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 30 Adults; 6 breeders: 3 nests with eggs (BSFW, 1969a). 31 Mar.- 20 Adults; 12 breeders: 6 nests, all with eggs; 2 Apr. 4 other pre-laying pairs (BSFW, 1969b). 26-31 May 52 38 breeding adults; 19 nests counted: 5 (26%) with eggs and 14 (74%) with small to large downy young; no nocturnal roosting clubs (BSFW, 1969c). 10-19 Sept. 26 22 adults, 4 young (BSFW, 1969d). 162 Table 56. Observations of Blue-faced Boobies on other islands at Pearl and Hermes Reef Population Date of Survey Island Estimate Breeding Status, Remarks, and References 1957 Ws Oct. | Bard 4 Observed from aerial survey (Rice, ms. a). 1963 5 Mar. Bird 6 Roosting (POBSP, 1964d). 1964 14 Mar. Bird 6 Adults; 3 nests (BSFW, 1967a; POBSP, 1964b). 18 Aug. Bird 3 2 adults, 1 dependent immature (POBSP, 1964a). 18 Aug. Sand 9 Adults roosting (POBSP, 1964a). ICES) 22 Were, — Balizsl 6 Adults; 3 nests wigh eggs (POBSP, 1965a). 1966 21 Sept. Planetree iL Adult roosting (BSFW, 1966b). 4 1967 31 May Bird 10 8 adults; 4 nests counted: 2 with eggs, q 1 with a small downy young, 1 with a medium-sized downy young (POBSP, 1967b). 29 Aug. Bird fe) 4 adults, 4 subadults, 1 dependent imma - ture roosting (POBSP, 1967a). 29 Aug. Sand 3 Adults roosting (POBSP, 1967a). RED-FOOTED BOOBY Sula sula Status Common breeding species; present year-round with possible population decrease in winter; occurs and nests on the four vegetated islands: Grass, North, Seal, and Southeast. Maximum recent population estimate 200 in February and March 1963. Observations Wetmore (ms.) noted Red-footed Boobies in April 1923. Galtsoff (1933: 19) recorded their presence in the summer of 1930. No data exist for the 1940's. Rice (ms. a) noted the species from aerial surveys in the fall of 1957. Woodside and Kramer (HDFG, 1961) found it nesting in spring 1961. POBSP and BSFW personnel have recorded Red-footed Boobies each year on every visit since February 1963. These observations are presented in Tables 58 to 61. 163 OT +E 9g 9 + 89 LE On HE 98T T2701 0 6 6 0 I 0 T 9eT 9gT 0 T?59N (0) GT GT 0) € 0) +2 (0) (0) (0) wu 0) 3] Z 0 () 0 © ©) 0 © peqns OT 8 g 9 0) 99 cL +E Get 98T qIupy Te70L T 0 0 ww PITa Tt 0) ) 71INpy pues Ot Ot aul ie) Shs BS ie) TET od I; Z 0 ) is) €2 0 T?S9N T T (0) (0) (©) ©) (0) wu U4ION g Z + 0 Ze AS 0 4T0py Seay at gt Ge 2 L cee ect i a Sie Ome “esa 0 0 0 0 TL €g €2 0 TdS9N eL A (©) 0 cI 0) 0 0 wu, 9 0 9 d L reff gel fe) FINpYy UZtON 9 9 9 Q i) oD Ze a, Higebteth 0 0 0 0 0 €g €2 0 TUSEN ©) 0 0 ©) ©) et 6 +E 71Npy £2944-70 Q io) ie) o) 2S 8) OS atu ) PJ Nek 0 0 O 0 0 0 l2 12 0 THS9N 0 0 0 (0) 0) 9 0 0 (0) wu, (0) ©) 0 0 AS (0) €2 Ni 9 J TNpy Tees Q Q o Q @ Q tt g ) T2400 0 0 0 0 0 0 + + 0 TIS9N 0) ) (0) 0) @ 0 OT + 9 4T0py sseiy Qt 2 2) i) nue & 9 OTT of 0g T2701, © g Zé 0) i 0) 0) 92 92 0 TISEN 0 re g 0 € (©) 2 © (0) 0 WUT @) 3g g (0) 0 (0) @) (o) @) 0) peqns OT 0 0 (0) 0) S + +9 + 0g qInpy ysSeeynos Coot = TEC LOGIN ECON OS60 Te10L TIS59N wal peqns FTO :Te10] TF59N wu peqns FTINPY TeyOL ssery 49-ION qseayynosg SSeTO -33V jooy Souzey pue [Azveg 1% pepueq setqoog pe,oos-pey pUeTST "29 eTqeL 171 Southeast Island: Wetmore (ms.) recorded breeding Brown Boobies in April 1923. Galtsorrl (in litt.) photographed them in summer 1930; HDFG personnel noted them in March 1961. POBSP and BSFW personnel have recorded this species breeding each year since 1963 (Table 63). March nest counts from 1963 through 1969 (no count in 1966) average 25 and range from 10 to 36. Most nests are placed inland on grassy, Sesuvium, or Chenopodium areas; a few are placed on bare areas of coral rubble. Most twig-lined nests are on the eastern section, with a few on the western. The primary area for nocturnal roosting was on the exposed rocks seaward from the western section. Other roosts are on the oil drums, and, on one occasion, the tower. Other Islands: Brown Boobies have been recorded roosting on the three other vegetated islands, and on sand-covered Kittery (Table 64). North and Seal Islands are the most popular roosting islands; the species has been seen on Grass and Kittery only twice each. Banding and Movements POBSP and BSFW personnel banded 313 Brown Boobies (Table 65). Of these, 145 have been recaptured on the atoll (Table 16). In addition, 2 birds originating from Kure and one each from Wake and Laysan have been captured at Pearl and Hermes. Seven banded at Pearl and Hermes have been captured elsewhere: Kure (4), and Laysan, Wake, and the Ellice Islands (1 each). Details on these movements are found in Appendix Tables 9a and Qb. Specimens POBSP: USNM 494128, 9, collected 15 March 1965 on Southeast by Amerman. Non-POBSP: USNM 300879, 9, collected 26 April 1923 by Wetmore. These are first specimen records. Table 63. Observations of Brown Boobies at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1923 26-28 Apr. eal 20 breeding adults; 9 nests with eggs, 1 with small young (Wetmore, ms.). 1930 23 July- few Adults with downy young (Galtsoff, photo- Aug. graph). Igaltsoff's 1930 photographs show about a dozen nesting Brown Boobies with chicks on the rocky western section of Southeast Island. Another picture, showing an adult with 2 chicks on rocky terrain, is labeled Seal or Grass Island but probably was Southeast (see Fig. 67). 172 Table 63. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References i | 1961 12 Mar. not Eggs or small young (HDFG, 1961). ; abundant H 1963 26 Feb.- 100 60 breeders; some subadults; 30 nests : 8 Mar. counted in February, 26 of these deserted q by March; sample count of 28 nests: 23 : (82%) with eggs, 5 (18%) with young (POBSP, 1964d). 18-23, 195+ 144 breeding adults; 72 nests: 19 (26%) 25 June with eggs, 11 (15%) with recently hatched young, 12 (17%) with small downy young, 28 (39%) with larger young; 2 (3%) nests with no contents but with attendant adults; many subadults (POBSP, 1963). 1964 13-14 Mar. 30 27 adults, 2 subadults; 10 nests counted: 9 (90%) with eggs, 1 (10%) with young (BSFW, 1964a; POBSP, 1964b). 16-19 Aug. 125 74 adult breeders, 37 nests: 8 (22%) with eggs, 29 (78%) with varying sizes of young; about 20 flying immatures (POBSP, 1964a). 16 Sept. 100-125 50 breeders; 6 of 25 nests (24%) with eggs, 19 (76%) with young; 10 flying immatures (BSFW, 1964b; POBSP, 1964c). 1965 15-19 Mar. 40-50 eh breeders; 12 nests with eggs (POBSP, 1965b). 21-22 Mar. yy 26 breeders; 13 nests with eggs (BSFW, 1965; POBSP, 1965a). 1966 1 Apr. - Not mentioned in notes (BSFW, 1966a). 20-26 Sept. hO+ 20 nests with eggs to flying young counted (BSFW, 1966b). J 25-27 Sept. 120 hot breeders; count of 19 nests: 3 (16%) Hi with eggs, 5 (26%) with small young, 11 (58%) { with larger young (POBSP, 1966). ; 1967 21-23 Mar. 50+ hO breeding adults; count of 21 nests: 2 (10%) empty but active, 18 (86%) with eggs, and 1 (5%) with a recently hatched young; several fledged immatures (BSFW, 1967a; POBSP, 1967d). LS Table 63. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1967 28, 30 May- 139 114 breeders; 57 nests: 32 (56%) with 1 June eges, 16 (29%) with downy young, 9 (15%) with medium-sized or large downy young (POBSP, 1967b). 28-30 Aug. 17 68 breeders; 34 nests: 7 (21%) with eggs; 2 (6%) with recently hatched young; 3 (9%) with small downy young, 10 (29%) with large downy young, 12 (35%) dependent immatures (POBSP, 1967a). 1968 22-2) Mar. 100+ 90 breeders; 41 nests: 1 (2%) empty but active, 36 (88%) with eggs, 3 (7%) with recently hatched young, and 1 (2%) with a large downy young (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. Ne) 18 breeders; 9 nests with eggs (BSFW, 1969a). 31 Mar.- 8h 72 breeders; 36 nests: 24 (66%) with eggs, 2 Apr. 4 (11%) with an egg and young, and 8 (22%) : with 1 young (BSFW, 1969b). 26-31 May 107 82 breeders; 28 nests: 3 (10%) contained eggs, and 25 (90%) contained recently hatched to near-fledging young (BSFW, 1969c). 10-19 Sept. 99 90 adults, 9 young (BSFW, 1969d). Table 64. Observations of Brown Boobies on other islands at Pearl and Hermes Reef Population Date of Survey Island Estimate Breeding Status, Remarks, and References 1913 15 Mar. North 2-3 (Bailey, 1956: 32). Hos Te Oct. Seal i (Rice, ms. a). 14 Oct. Northeast 2 In flight (Rice, ms. a). Reef 1963 5 Mar. Seal ig Roosting on rock ledges; no nests (POBSP, 1964d). 5 Mar. Grass ni Roosting on beach; no nesting (POBSP, 1964d). 174 Table 64. (continued) Population Date of Survey Island Estimate Breeding Status, Remarks, and References 1963 6 Mar. North few None nesting (POBSP, 196d). 23-25 June North few Flying by island each day (POBSP, 1963). 26 June Kittery ab (POBSP, 1963). 26-27 June Grass a Roosting adult; banded (POBSP, 1963). 1964 14 Mar. Seal i Flying over (BSFW, 1964a; POBSP, 1964b). 18 Aug. Kittery 1 Seen from Seal Island (POBSP, 1964a). 19-20 Aug. North 1 Adult; none nesting (POBSP, 196a). 1965 17-18 Mar. North an Flock of 22 adults and 2 subadults; none nesting (POBSP, 1965b). 18-19 Mar. Seal 30-35 Birds roosting; none nesting (POBSP, 1965b). 22 Mar. Seal 1 Roosting immature (BSFW, 1965; POBSP, 1965a). 1967 16-17 Mar. North a In flight (BSFW, 1967d). 22 Mar. Seal 3 Flying from island; no nests (BSFW, 1967a; POBSP, 1967d). 29-30 Aug. North i Adult roosting on rocks at dusk (POBSP, 1967a). 1968 24 Mar. Seal 14 13 adults and 1 immature or subadult flying from island; paint on 2 birds indicated they had been on Southeast Island 1-2 days previously (BSFW, 1968; POBSP, 1968). GREAT FRIGATEBIRD Fregata minor Status Common breeding species; present year-round; nests at Grass, North, and Southeast Islands; in the past bred at Seal Island; occurs at Kittery. Maximum POBSP and BSFW population estimate 890 in March 1965. 175 /h, > TeIOL TIS9N wu peqng yInpy Te,0L 0) F1NpV Y4tON T @) 0) 0) i 0) 710pY sseip Ore o€ at ite zZ u Ld 99S LB40L 88 T Ze HL 0 0 62 0 TIS0N ge € TI 8 I ©) 0) 0) wu €T i @) 0) 0) 0) ©) OT peqns ea G2 Wl 6 T 1 St 9S 41npyY 4Seau}Nog STeq0L 8961 L961 9961 Teq0L 96T G96T HOOT Teq0L €96T €961 SS®TO pueTsT * TaN “ony *qdag G96T * TEN * LEW “Sny €96T oun * TEN -33V =" 9a. KeaAing fO potted dSd0d dSd0d dSdod MaSd dSdod dSdod dSd0d dSdOd :zepueg jaoy Souley pue [Taveg 4e papueq setTqoog uMozg "GQ aTqeL 176 Observations Munro (1942: 12) and Palmer observed a few frigates offshore on 7 July 1891; no landing was made on the atoll. Elschner (1915: 60) in 1912 found some 20 live and 100 dead frigatebirds. A small nesting colony was noted in March 1913 (Bailey, 1956: 32; Willett, ms.). Wetmore (ms.) found Great Frigatebirds, some nesting, on most islands in April 1923. The species was present in large numbers in late summer 1930 (Galtsoff, 1933: 19). Rice (ms. a) counted 770 on an aerial survey in October 1957. Nesting pairs were found by Woodside and Kramer (HDFG, 1961) in March 1961. POBSP and BSFW personnel recorded Great Frigatebirds on every visit since 1963. Tables 66 to 69 present all observations. Annual Cycle The local annual breeding cycle is presented in Figure 58. Great Frigatebirds remain on the atoll throughout the year; numbers are probably highest from late winter until late fall. This coincides with the breeding period. Eggs are laid as early as late February, but most are laid during March. Eggs may be present as late as late June. Hatching probably com= mences in early May, with most young hatched by late June. Fledging probably starts in October, but most young birds, even though able to fly, probably sit on or near their nest sites into late fall or early December; some may remain into the following breeding season. In March 1965 the three breeding islands yielded 120 to 175 active nests; the number of active March nests probably varies from year to year. The number of young during August and September averaged 118, and ranged from 65 to 187. Ecological Distribution Great Frigatebirds are known to nest on Grass, North, Seal, and Southeast Islands, and to roost on Kittery Island. Grass Island: Wetmore (ms.) noted non-breeding Great Frigatebirds in April 1923. Rice (ms. a) estimated 200 on an aerial survey in October 1957. POBSP and BSFW personnel found them nesting each year (Table 66). Active March nest counts averaged 71, and ranged from 37 to 123. August-September young counts averaged 34, and ranged from 20 to 48. Nests are placed on the dense vegetation, especially Solanum. Nor Sland: Elschner (1915: 60) noted non-breeding frigatebirds in 1912; frigatebirds were observed nesting in March 1913 (Bailey, 1956: 32; Willett, ms.). Rice (ms. a) found them in October 1957 and POBSP and BSFW personnel have recorded them breeding each year since 1963 when visits were made (Table 67). Only a few nests were found on March visits. August- September young counts averaged 41 and ranged from 14 to 80. All nests were in Solanum in the central and northwestern parts of the island; all nests were within a foot of the ground. 177 Seal Island: Munter (ms.) noted a few female Great Frigatebirds in February 1916. In April 1923, Wetmore (ms.) recorded this species breeding. POBSP and BSFW personnel found roosting frigatebirds on most visits (Table 68). Wo evidence of recent nesting was found. Southeast Island: A few Great Frigatebirds were observed by Munter (ms.) in February 1916. Wetmore (ms.) found them nesting in April 1923. They have been recorded by all subsequent visitors, and have been seen nesting by POBSP and BSFW personnel each year (Table 69). March nest counts averaged 52, and ranged from 13 to 83. August- September young counts averaged 43, and ranged from 20 to 67. Nests were all built in low, shrubby vegetation, especially Solanum, on the south side of the eastern portion. All nests were within two feet of the ground and consisted of simple platforms of Boerhavia, Solanum, and Tribulus. Roosting birds utilized the low vegetation, the tower, and oil drums scattered about the island. Other Islands: POBSP personnel recorded one Great Frigatebird at Kittery Island on 18 August 1964. A single female roosted on the Kittery Fish and Wildlife sign 29 August 1967. This species may also fly over or roost on the other four sandy islands. Rice (ms. a) noted 210 Great Frigate- birds in flight over various parts of the atoll on 14 October 1957. Banding and Movements POBSP and BSFW personnel banded 539 Great Frigatebirds (Table 70). Of these, 53 were recaptured on the atoll (Table 16). Fifty others were recaptured elsewhere, on Kure (42 birds), Johnston and the Philippines (3 each), and French Frigate Shoals and Lisianski (1 each); data are presented in Appendix Table 10. In addition, two originating elsewhere were captured at Pearl and Hermes. A subadult (USFW band no. 767-45468), sex unknown, banded on Kure 6 May 1966 was captured on Southeast as an adult female 31 May 1967 on a nest containing a naked chick. The other, an adult female (737- 37124)* captured 4 October 1966 at Sand Island, Johnston Atoll, was re- captured at Southeast 30 May 1967 on a nest with an egg, and recaptured again at Southeast 3 July 1967 on a nest with a week-old chick. Specimens POBSP: USNM 494129, o, collected 15 March 1965 on Southeast by Amerman, Non-POBSP: USNM 300968, o, collected 26 April 1923 by Wetmore. These are first specimen records. *Great Frigatebird number 737-37124 was originally banded on Southeast 21 June 1963 by POBSP personnel. 178 Table 66. Observations of Great Frigatebirds at Grass Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. - Not mentioned in report (Munter, ms.). 1923 27 Apr. ? Present but not breeding (Wetmore, MSiep)ic 1957 14 Oct. 200 Aerial survey (Rice, ms. a). 1963 5 Mar. 100+ Most breeding birds on nests with eggs (POBSP, 19644). 26-27 June 350 250 adults, 226 breeders; 113 nests: 17 (15%) with eggs, 43 (38%) with recently hatched and small downy young, 53 (47%) with larger young (POBSP, 1963). | 1964 14 Mar. 300 246 breeders; 123 nests, most with eggs, no young (BSFW, 1964a; POBSP, 1964b). 18 Aug. 298 250 adults; 96 breeders, 48 nests with young (POBSP, 1964a). 1965 19 Mar. 250-350 100-150 breeders: 50-75 nests, most with eggs; immatures from the previous breeding season still present (POBSP, 1965b). 22 Mar. eu5+ 2) 24 breeders, 62 nests counted with eggs (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 87 58 breeders; 29 nests with young (BSFW, 1966b). 1967 22 Mar. - Not mentioned in notes (BSFW, 1967a; POBSP, 1967d). 31 May 130+ 104 breeders; 52 nests: 27 (52%) with eggs, 14 (27%) with recently hatched young, 10 (19%) with small downy young, and 1 (2%) with medium- sized downy young (POBSP, 1967b). 29 Aug. 60 No adults present; 40 breeders, 20 nests with large downy young (POBSP, 1967a). 1968 24 Mar. 150 86 breeders; 37 nests: 24+ (65%) empty but active, 13 (35%) with eggs; 6 partially built nests also seen; many courting males present (BSFW, 1968; POBSP, 1968). 1969 11 Sept. 120 80 breeders; 40 nests with young; no adults seen (BSFW, 1969d). 179 Table 67. Observations of Great Frigatebirds at North Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1912 ? 20+ Non-breeding; 100¢ dead along the beach (Elschner, 1915: 60). 1913 15 Mar. few Nesting (Bailey, 1956: 32); small colony (Willett, ms.) 1957 14 Oct. 60 Aerial survey (Rice, ms. a). 1963 6 Mar. ? Probably more nesting on North than on Southeast (POBSP, 196d). 23-25 June 280 225 adults; 124 breeders, 62 nests: 7 (114%) with eggs, 34 (55%) with recently hatched and small downy young, 21 (34%) with larger young (POBSP, 1963). 1964 19-20 Aug. 200-350 96 breeders; 48 nests with young; none of the young appeared to be as old as the oldest on Southeast (POBSP, 1964a). 17 Sept. 200 80 counted in air; 72 breeders; 36 large de- pendent young and 7 flying immatures counted (BSFW, 1964b; POBSP, 1964c). 1965 17-18 Mar. 100-200 60-80 breeders: 30-40 nests, many with eggs; some immatures from the previous breeding season still present (POBSP, 1965b). 1966 22 Sept. 63 1 adult seen; 42 breeders: 21 nests counted, each containing 1/4- to 1/2-grown young (BSFW, 1966b). 1967 16-17 Mar. 80 Adults and immatures; 30 breeders: 15 nests with eggs (BSFW, 19674). 29-30 Aug. 100 28 breeders; 14 nests with large downy young (POBSP, 1967a). 1969 12 Sept. 240 160 breeders, 80 young; only 60 adults seen (BSFW, 1969d). Table 68. Observations of Great Frigatebirds at Seal Island 180 : - Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. few Females (Munter, ms.). O23 27 kpre 160 Breeding (Wetmore, ms.). 1Q6S 5 Were. 2 Roosting (POBSP, 1964d). 26 June 50-75 Roosting; none nesting (POBSP, 1963). 1964 14 Mar. ©) (BSFW, 1964a; POBSP, 1964b). 18 Aug. © Roosting; none nesting (POBSP, 1964a). 1965 18-19 Mar. 30-40 Roosting adults; none nesting (POBSP, 1965b). } 22 Mar. 0 (BSFW, 1965; POBSP, 1965a). : 1966 21 Sept. 6 Adults seen flying over; no nests (BSFW, : 1966b). : 1967 22 Mar. ©) (BSFW, 1967a; POBSP, 1967d). 31 May 1 Adult male roosting on beach (POBSP, 1967b). | | 29 Aug. 10 Mostly roosting subadults; none nesting (POBSP, 1967a). : 1968 24 Mar. ©) (BSFW, 1968; POBSP, 1968). | | Table 69. Observations of Great Frigatebirds at Southeast Island . Population Date of Survey Estimate Breeding Status, Remarks, and References | 1916 4 Feb. few (Munter, ms.). 1923 26-28 Apr. 300 Breeders; nests either empty or containing | eggs (Wetmore, ms.). . 1930 23 July- large (Galtsoff, 1933: 19). Aug. numbers 1957 14 Oct. 300 Aerial survey (Rice, ms. a). 1961 12 Mar. not Nesting on low vegetation (HDFG, 1961). abundant 181 Table 69. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1963 26 Feb.- 100 Nesting; sample count of 13 nests: 2 (15%) 8 Mar. empty but active, 11 (85%) with eggs; 10 deserted by end of survey period (POBSP, 19644). 18-23, 81 58 breeders; 29 nests: 6 (21%) with eggs, 25 June 8 (28%) with recently hatched young, 6 (21%) with small downy young, and 9 (31%) with large downy young (POBSP, 1963). 1964 13-14 Mar. 125-150 112 breeders; 56 nests counted; sample count of 27 nests: 7 (26%) empty but active, 20 (74%) with eggs (BSFW, 1964a; POBSP, 1964b). 16-19 Aug. 150 130 adults; ho breeders; 20 nests with half- grown to nearly fledged young (POBSP, 1964a). 16 Sept. 100 15 adults and 20 large young unable to fly counted; 40 breeders (BSFW, 1964b; POBSP, 1964c). 1965 15-19 Mar. 200-300 80-120 breeders; 40-60 nests, most contain- ing eggs (POBSP, 1965b). 21-22 Mar. 149 82 breeders; 41 nests, most containing eggs; 8 flying immatures (BSFW, 1965; POBSP, 1965a). 1966 1 Apr. ? Incubating eggs (BSFW, 1966a). 20-26 Sept. 159+ 106 breeders; 53 nests with young from 2/3 to almost full grown; an undetermined number of adults roosting at night (BSFW, 1966b). 25-27 Sept. 250 98 breeders; 49 nests with large downy young to well-feathered birds (POBSP, 1966). 1967 21-23 Mar. 96 Breeders; 48 nests: 22 (46%) empty but active; 26 (54%) with eggs (BSFW, 1967a; POBSP, 1967d). 28, 30 May- 270 24k2 breeders; 121 nests counted; sample count 1 June of 4O nests: 30 (75%) with eggs; 3 (8%) with recently hatched young; and 7 (17%) with small downy young (POBSP, 1967b). 28-30 Aug. 100 Flying; 62 breeders; 31 nests counted, all containing large downy young (POBSP, 1967a). 182 Table 69. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1968 22-24 Mar. 125 102 breeders; 51 nests; sample count of 38 nests: 17 (45%) empty but active, 21 (55%) with eggs; about 85% of eggs fresh or very slightly incubated, 15% slightly incubated (POBSP, 1968). 1969 10-11 Feb. 25 Mostly subadults; 1 displaying adult male (BSFW, 1969a). 31 Mar.- 176 Breeders; 83 nests counted: 19 (23%) empty 2 Apr. but active, 64 (77%) with eggs (BSFW, 1969b). 26-31 May 146 Breeders; 73 nests counted: 55 (75%) with eggs and 18 (25%) with small downy young (BSFW, 1969c). 10-19 Sept. 201 134 breeders; 67 young, only 33 adults seen (BSFW, 1969d). PINTAIL Anas acuta Status Very uncommon migrant; one sight record. Observations An immature male Pintail was observed at Southeast Island 13 September 1969 by BSFW personnel. Sex and age were verified when the bird was cap- tured on the 14th in an emaciated condition. This sighting is a new species record for Pearl and Hermes Reef. Pintails are common migrants to the Main Hawaiian Islands, and are known to have occurred at most of the Northwestern Hawaiian Islands as well (Amerson, 1971: 228). LAYSAN TEAL Anas laysanensis Status Recent unsuccessful introduction. 183 =F Qi Oe Nind oe A In| S OO O10 oOmMoO Cle — LN [e701 L961 OT € 8 6S oi Z ST 9eT 892 €6T GL ?T870L i 0 0) ©) 0) 0) 0) 0) 0 0 QO uUMoUyUn 0 ©) @) Ot 0 ) 0) HIT 19 18 @) T359N T @) ©) + T ©) T T 0) 0) ) wu, G 0) ©) 0) g 1 T ©) 9€ OT 9g peqns TI et 8 1, al IL €T TI sal 96 64 qInpy = Teq0L OL 8 Ot QL £6 £9) a) Q * [83 OL T 0 0 0 0) 0 0) QO uUMmMouyUn ) 0) ©) ©) Lt the TZ 0) TIS9N T ©) 0) @) 0 ©) 0) 0) wy og O T T 0 S S 0 peqns 9 g 6 6 8 6€ 6€ ) ZINPY YON io) Q io) ie) io) so) Q gt uaa Z6 ecg Skee 0 0 0 0 0 0 0 On SG €S 0 T75eN ©) 0) @) 0) ©) 0) @) 0 8 ) 8 peqns 0 ©) 0 ©) 0 0 O ©) €S 6€ 4 TINY ssp SaSsbte Ong Bale Fly eee 6 feo (een | ge Es gs urenor 0 0 0 om 0 0 0 61 CT CT 0 TUS9N 0) 0 ©) + It 0) "t i 0) 0) @) ww € @) (e) ) T iT 0 ) €2 S QT peqns S et 0 ), S T + € €S QT GE qInpy yseeyynOS Z96t L96t 2961 9961 TeIoL 961 So6t 4OOT TeI0OL ©9061 ©961 SS¥TO pueTsT “ony Kew “sey *ydeg G96T “Teh “se “any €96T eur “ae -o3y -° Qad RoAang JO potted dsdod dSd0d MaSd dSdOd MASH dSd0d dSdod dsd0d dSd0d :tepueg joeoy Somzey pue [azeeg 1e pepueq spitqeyeStaq yeery ‘*o/, eTqeL 184 Observations Fourteen Laysan Teal--9 female and 5 male--were captured on Laysan 18 March 1967 by BSFW personnel and banded and released on Southeast Island, Pearl and Hermes Reef, 21 March. The first two ducks released, a female and a male, soon took flight, left the atoll, and headed out to sea. The remaining birds were wing-clipped before release. On 22 March a wing-clipped drake was seen on a sandspit near Seal Island about five miles from the release point. Four others were observed roosting on Southeast Island that night. The following morning a dead drake was found on the north shore of Southeast, reducing the surviving population to 5 drakes and 4 hens. When the island was visited by POBSP personnel in late May and early June 1967 only one live bird--a drake seen near the lagoon at Southeast-- was observed. A dead teal was found on Southeast, reducing the possible number of surviving birds to eight. No teal were seen on any of the other islets visited. BSFW personnel visited Southeast 3 to 9 July 1967 but no biological report was written on this trip. POBSP personnel observed a pair of teal at their campsite on the east point of Southeast Island 28 to 30 August 1967. On their 27 to 29 September 1967 visit to Southeast, BSFW (1967c) personnel "searched for [a Teal]...nest...[and] found evidence it did not hatch." This refers to a teal nest with an egg, presumably seen at Southeast in July, that did not hatch. They did, however, observe the pair of teal, both of which were banded. No trace of these adults could be found when Southeast was visited in 1968 and 1969 by POBSP and BSFW personnel. Banding and Movements The 14 Laysan Teal introduced in 1967 were banded by BSFW personnel. Two were recaptured later in 1967. LAYSAN RATL Porzanula palmeri Status Unsuccessful introduced species; now extinct. Observations yin June 1929 Captain William G. Anderson released seven pairs of Laysan Rail from Sand Island, Midway Atoll, on an island, most likely Southeast, at *Laysan Rails were native to Laysan Island. In 1891 rails were first trans- ported to Midway, where they multiplied. They probably became extinct on Laysan soon after 1923. The last record of the Laysan Rail occurred at Midway in June 1944, or possibly June 1945 (Baldwin, 1945; 343-348, and 1947: 14-21; Bailey, 1956: 84-90; Ely and Clapp, in press). 185 Pearl and Hermes Reef (Munro, 1945: 13-14). George Kaufmann, who lived at Midway from 1929 to 1931, visited Pearl and Hermes in 1930 and found no rails; in addition, he saw no live vegetation, and found that storms had left only tall clumps of dead bunchgrass. He saw only 3 or 4 large sea- birds during his visit. Since the rail depended upon insects, birds' eggs, and meat scraps for food, Fisher and Baldwin (1945: 11-12) believed it im- possible for the transplanted colony to have survived, even if Kaufmann had overlooked living specimens in 1930. Fisher and Baldwin (1946: 8) further- more, pointed out that Galtsoff (1933: 19) did not observe the Laysan Rail during his late summer 1930 visit to Pearl and Hermes. GOLDEN PLOVER Pluvialis dominica Status Common migrant; usually found in small to large flocks on beaches and rock outcrops. Most abundant during early spring, late summer, and fall; a few individuals present throughout the year. POBSP and BSFW maximum population estimate 230 in March 1968. Observations Willett (ms.) first recorded the Golden Plover in March 1913. Munter (ms.) next observed a few hundred in February 1916, and Wetmore (ms.) re- corded it in April 1923. POBSP and BSFW personnel have recorded Golden Plovers on most of the islands since 1963. All observations are presented in Tables 71 to 74. Annual Cycle Gol@en Plovers may be present throughout the year. Although observations are lacking for five months, available data indicate a peak population in early spring, late summer, and fall. This fits with the known migration pattern on other Northwestern Hawaiian Islands. The arrival and departure of new birds during migration periods may cause the daily population to vary. Ecological Distribution Golden Plovers have been recorded from all four major islands and the two larger sand islands; it has not been recorded at Bird, Planetree, and Sand Islands. Grass Island: Wetmore (ms.) observed Golden Plovers in 1923. POBSP and BSFW personnel recorded it infrequently and in low numbers (Table 71). This species prefers the beach areas of the island. North Island: Following Willett's (ms.) 1913 observations, none was recorded until POBSP personnel noted 70 in March 1963. BSFW and POBSP per- sonnel have found it only six times since and in much smaller numbers (Table T2)- 186 Seal Island: Wetmore (ms.) observed a few in 1923. None was recorded again until 1965. Since then POBSP and BSFW personnel have only seen this species on five visits. Usually the population has been low (2-10) but in March 1969 the population was 200 (Table 73). At that time birds were most numerous around the central inlet and around the tidal pools. Southeast Island: Munter (ms.) first recorded Golden Plovers in 1916. POBSP, BSFW, and HDFG personnel recorded them each year during the 1960's when visits were made (Table 74). March populations average 14, with a range of from 0 to 25; September populations average 81, and range from 18 to 175. Golden Plovers are usually very abundant around the cove that separates the two sections of the island, and on the seaward rocky ledge of the west- ern section. Other Islands: POBSP personnel recorded 2 Golden Plovers from Kittery Island 31 May 1967 and 1 from Little North 29 August 1967. Banding and Movements Five Golden Plovers have been banded on Southeast Island by POBSP and BSFW personnel: 2 by the POBSP in February 1963; 1 each in March 1964, March 1965, and February 1969 by the BSFW. POBSP and BSFW personnel have captured one Golden Plover, originally banded on Lisianski, at Pearl and Hermes. The unsexed adult (USFW band no. 662-06097 ) was banded 11 March 1964 by BSFW personnel, captured at Kure on 1 September 1964, and found dead on Southeast Island 15 March 1965. Table 71. Observations of Golden Plovers at Grass Island Population - Date of Survey Estimate Remarks and References 1923 27 Apr. few (Wetmore, ms.). 1963 5 Mar. few (POBSP, 1964d). 26-27 June O (POBSP, 1963). 1964 14 Mar. 0 (BSFW, 1964a; POBSP, 1964b). 18 Aug. 0 (POBSP, 1964a). 1965 19 Mar. few (Clapp and Woodward, 1968: 17; POBSP, 1965b). 22 Mar. 0 (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 0 (BSFW, 1966b). 187 Table 71. (continued) Population Date of Survey Estimate Remarks and References 1967 22 Mar. 2 (Clapp and Woodward, 1968: 17; BSFW, 1967a; POBSP, 19674). 31 May il (POBSP, 1967b). 29 Aug. O (POBSP, 1967a). 1968 24 Mar. 5) 2 seen in a small flock of turnstones; 3 others seen flying along beach (POBSP, 1968). Table 72. Observations of Golden Plovers at North Island Population Date of Survey Estimate Remarks and References 1913 15 Mar. plentiful (Willett, ms.). 1963 6 Mar. 70 (POBSP, 1964d). 23-25 June 2-3 Daily (Clapp and Woodward, 1968: 17; POBSP, 1963). 1964 19-20 Aug. 0 (POBSP, 1964a). 17 Sept. 1 (Clapp and Woodward, 1968: 17; BSFW, 1964b; POBSP, 1964c). 1965 17-18 Mar. few (POBSP, 1965b). 1966 22 Sept. 1 (BSFW, 1966b). 1967 16-17 Mar. 5 (BSFW, 1967d). 29-30 Aug. 10 (POBSP, 1967a). Table 73. Observations of Golden Plovers at Seal Island Population Date of Survey Estimate Remarks and References 1923 27 Apr. few (Wetmore, ms.). 1963 5 Mar. 0 (POBSP, 1964d). 26 June @) (POBSP, 1963). 188 Table 73. (continued) Population Date of Survey Estimate Remarks and References 1964 14 Mar. 0 (BSFW, 1964a; POBSP, 1964b). 18 Aug. 0 (POBSP, 1964a). 1965 18-19 Mar. few (Clapp and Woodward, 1968: 17; POBSP, 1965b). 22 Mar. 0 (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 10 (BSFW, 1966b). 1967 22 Mar. 2 (Clapp and Woodward, 1968: 17; BSFW, 1967a; POBSP, 1967d). 31 May 2 (POBSP, 1967b). 29 Aug. 10 Flock of 7 and a single bird flying with turnstones (POBSP, 1967a). 1968 2k Mar. 200 Most numerous around tidal pools (BSFW, 1968; POBSP, 1968). Table 74. Observations of Golden Plovers at Southeast Island Population Date of Survey Estimate Remarks and References 1916 4 Feb. few (Munter, ms.). hundred 1923 26-28 Apr. few (Wetmore, ms.). 1930 23 July- O Not mentioned (Galtsoff, 1933). Aug. 1961 12 Mar. small (HDFG, 1961). numbers 1963 25 Feb.- 4O Widespread (Clapp and Woodward, 1968: 16; 8 Mar. POBSP, 1964d). 18-23, 5 Less than 5 daily (Clapp and Woodward, 1968: 25 June 16; POBSP, 1963). 1964 13-14 Mar. 3 (Clapp and Woodward, 1968: 16; BSFW, 1964a; POBSP, 1964b). 189 Table 74. (continued) Population Date of Survey Estimate Remarks and References 1964 16-19 Aug. al (Clapp and Woodward, 1968: 16; POBSP, 1964a). 16 Sept. 150 (Clapp and Woodward, 1968: 16; BSFW, 1964b; POBSP, 1964c). 1965 15-19 Mar. few (Clapp and Woodward, 1968: 16; POBSP, 1965b). 21-22 Mar. 9 (Clapp and Woodward, 1968: 16; BSFW, 1965; POBSP, 1965a). 1966 1 Apr. 0 Not mentioned in notes (BSFW, 1966a). 20-26 Sept. 125 (BSFW, 1966b). 25-27 Sept. 175 Most common in area between interior lagoon and south rock ledge (Clapp and Woodward, 1968: 16; POBSP, 1966). 1967 21-23 Mar. 15-20 Along reef and around lagoon (Clapp and Wood- ward, 1968: 16; BSFW, 1967a; POBSP, 1967d). 28, 30 May- 2 Actual count; about half molting into or in 1 June breeding plumage; all seen in 1 flock near lagoon (POBSP, 1967b). 28-30 Aug. 100 Most foraging in central Sesuvium depressions and in areas of low density grass and Tribulus (POBSP, 1967a). 1968 22-24 Mar. 25 Most in 1 flock on rock ledge west of entrance to interior lagoon (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 10 (BSFW, 1969a). 31 Mar.- 0 (BSFW, 1969b). 2 Apr. 26-31 May 6 Maximum count during survey period (BSFW, 1969e). 10-19 Sept. 18 (BSFW, 1969d). 190 BRISTLE-THIGHED CURLEW Numenius tahitiensis Status Common migrant; usually recorded in small to medium-sized flocks in rocky areas. Most abundant during late summer and fall; some may be present throughout the year. POBSP and BSFW maximum population estimate 30 in August 1967 and September 1966. Observations Bristle-thighed Curlews were first noted in March 1913 (Willett, ms.). Munter (ms.) observed large numbers in February 1916; Wetmore (ms.) col- lected one in April 1923. POBSP, BSFW, and HDFG personnel found this species present each year during the 1960's when visits were made. All observations are presented in Tables 75 and 76. Annual Cycle Bristle-thighed Curlews are known from all seasons. Recent March pop- ulation counts average 19, and range from 3 to 28; September counts average 2/, and range from 20 to 30. Ecological Distribution The Bristle-thighed Curlew is known only from Grass, North, Seal, and Southeast Islands. These four islands are the only ones in the atoll that are vegetated and have rocky portions. southeast Island: Bristle-thighed Curlews are more frequently found at Southeast than on all the other islands combined (Tables 75 and 76). South- east is much larger than the other islands. Its size and its expansive seaward rock ledges and Sesuvium-lined, brackish tidal pools apparently attract this species, which normally forages in such areas. Other Islands: Curlews utilize the rocky eastern portions of Seal and Grass. At North they have been seen in the vegetated portion eating cater- pillars. On most visits to these islands curlews were not noted. Banding and Movements Four Bristle-thighed Curlews were banded on Southeast Island by POBSP and BSFW personnel: 1 in March 1963 (POBSP), 2 in March 1965 (BSFW), and 1 in May 1967 (POBSP). One adult Bristle-thighed Curlew, originally banded (USFW no. 6445-12620) at Southeast 31 May 1967, was captured on Laysan 27 September 1967 by BSFW personnel. 191 Specimens Non-POBSP: USNM 301042, 9, collected at Southeast 19 May 1923 by Wetmore. This is a first specimen record. Table 75. Observations of Bristle-thighed Curlews at Southeast Island Population Date of Survey Estimate Remarks and References 1916 4 Feb. 50 (Munter, ms.). 1923 26-28 Apr. 2 1 collected 26 April; 1 or more seen 28 April (Wetmore, ms.). 1930 23 July- - Not mentioned in report (Galtsoff, 1933). Aug. 1961 12 Mar. iy (HDFG, 1961). 1963 26 Feb.- 5 (POBSP, 1964d). 8 Mar. 18-23, 4 (POBSP, 1963). 25> June 1964 13-14 Mar. 6 (BSFW, 1964a; POBSP, 1964b). 16-19 Aug. 15 Counted about the inlet (POBSP, 1964a). 16 Sept. 29 Count (BSFW, 1964b; POBSP, 1964c). 1965 15-19 Mar. 15 Flock seen 16 March (POBSP, 1965b). 21-22 Mar. 13 (BSFW, 1965; POBSP, 1965a). 1966 1 Apr. - Not mentioned in notes (BSFW, 1966a). 20-26 Sept. 28-30 Maximum of 28 seen at one time; found par- ticularly in Sesuvium areas, especially around interior lagoon (BSFW, 1966b). 25-27 Sept. 27 Most often on tank ledge (POBSP, 1966). 1967 21-23 Mar. 20-25 Flock of 14 on 23 March (BSFW, 1967a; POBSP, 1967d). 28, 30 May- 2-3 (POBSP, 1967b). 1 June 192 Table 75. (continued) Population Date of Survey Estimate Remarks and References 1967 28-30 Aug. 30 Most in 1 flock of 22 birds; appeared to have a distinct preference for the open flat in vicinity of large bank on southeast cor- ner of lagoon and rocky ledge opposite it; isolated individuals frequented the interior; 4-5 in largest interior tidal pool (POBSP, 1967a). 1968 22-2) Mar. 5) Foraged on beaches and interior, but most common on tank ledge where 10 were seen in 1 flock on 23 March (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 20 Feeding along tidal pools (BSFW, 1969a). 31 Mar.- 3 Count (BSFW, 1969b). 2 Apr. 26-31 May T Count (BSFW, 1969c). 10-19 Sept. 20 (BSFW, 1969d). Table 76. Observations of Bristle-thighed Curlews on other islands at Pearl and Hermes Reef Population Date of Survey Island Estimate Remarks and References 1913 15 Mar. North several (Willett, ms.). 1923 27 Apr. Seal 2 Presence noted (Wetmore, ms.). 27 Apr. Grass 2 Presence noted (Wetmore, ms.). 1963 5 Mar. Seal 9 (POBSP, 1964d). 5 Mar. Grass 2 (POBSP, 1964d). 23-25 June North 4 1 eating large caterpillar (POBSP, 1963). 26 June Seal 2 (POBSP, 1963). 26-27 June Grass 3 On the reef (POBSP, 1963). 1964 14 Mar. Seal 16 (BSFW, 1964a; POBSP, 1964b). 193 Table 76. (continued) Population Date of Survey Island Estimate Remarks and References 1964 19-20 Aug. North 2 Count (POBSP, 1964a). 17 Sept. North al Count (BSFW, 1964b; POBSP, 1964c). 1965 18-19 Mar. Seal 13 1 flock (POBSP, 1965b). 1967 31 May Grass 1 (POBSP, 1967b). 31 May Seal 1 (POBSP, 1967b). 29 Aug. Seal 8 On rocky eastern half (POBSP, 1967a). 1968 24 Mar. Grass 2 On unvegetated areas (BSFW, 1968; POBSP, 1968). WANDERING TATTLER Heteroscelus incanum Status Common migrant; usually recorded as singles or in small flocks on beaches. Most abundant during late summer and fall; a few birds probably present throughout the year. POBSP and BSFW maximum population count 29 in May and June 1967. Observations Wandering Tattlers were first recorded in April 1923 by Wetmore (ms.). HDFG personnel noted them next in small numbers in March 1961. POBSP and BSFW personnel on 15 of 18 visits observed the species on all the islands except Planetree and Sand (Tables 77 and 78). Annual Cycle Wandering Tattlers can probably be found throughout the year. Observa- tions are lacking for five months, but available data indicate a peak popu- lation in summer. Ecological Distribution The Wandering Tattler has been sighted on Bird (1 time), Grass (4), Kittery (1), Little North (1), North (5), Seal (9), and Southeast (14) (Tables 77 and 78). This species is commonly seen as singles or in small flocks on the beaches. At Southeast it prefers the western rocky outcropp- ings, especially along the water's edge. It also frequents the rocky ledges of Grass and Seal Islands. 194 1916 1923 1930 1961 1963 1964, 1965 1966 1967 Table 77. specimens Non-POBSP: Feb. 26-28 Apr. 23 July- Aug. 12 Mar. 26 Feb.- 8 Mar. USS23, 25 June 13-14 Mar. 16-19 Aug. 16 Sept. 15-19 Mar. 21-22 Mar. 1 Apr. 20-26 Sept. 25-27 Sept. 21-23 Mar. 28, 30 May- 1 June Population Date of Survey Estimate Remarks and References 12-15 small numbers 3) 1-2 - ©) 25 USNM 301024-25, 2, o', collected 26 April 1923 by Wetmore. These are first specimen records. Observations of Wandering Tattlers at Southeast Island Not mentioned in report (Munter, ms.). In a flock on bare rock at the eastern end (Wetmore, ms.). Not mentioned in report (Galtsoff, 1933). (HDFG, 1961). Not seen some days (Clapp and Woodward, 1968: 19; POBSP, 1964d). (Clapp and Woodward, 1968: 19; POBSP, 1963). (Clapp and Woodward, 1968: 19; BSFW, 196a; POBSP, 1964b). ; (Clapp and Woodward, 1968: 19; POBSP, 1964a). (Clapp and Woodward, 1968: 19; BSFW, 1964b; POBSP, 196lc). (POBSP, 1965b). (Clapp and Woodward, 1968: 19; BSFW, 1965; POBSP, 1965a). Not mentioned in notes (BSFW, 1966a). (BSFW, 1966b). Together on northwest beach 25 September (Clapp and Woodward, 1968: 19; POBSP, 1966). (Clapp and Woodward, 1968: 19; BSFW, 1967a; POBSP, 1967d). 15 counted; most singly (POBSP, 1967b). ee — 195 Table 77. (continued) Population Date of Survey Estimate Remarks and References 1967 28-30 Aug. 10-15 Maximum number seen at once: 4; none inland, most foraging on rocky outcroppings (POBSP, 1967a). 1968 22-24 Mar. 0 (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 2 (BSFW, 1969a). 31 Mar.- @) (BSFW, 1969b). 2 Apr. 26-31 May 0 (BSFW, 1969c). 10-19 Sept. i (BSFW, 1969d). Table 78. Observations of Wandering Tattlers on other islands at Pearl and Hermes Reef Population Date of Survey Island Estimate Remarks and References 192327 Apr’. Grass few (Wetmore, ms.). 27 Apr. Seal few (Wetmore, ms.). 1963 23-25 June North iL Daily (Clapp and Woodward, 1968: 19; POBSP, 1963). 26-27 June Grass i (Clapp and Woodward, 1968: 19; POBSP, 1963). 1964 14 Mar. Seal i (Clapp and Woodward, 1968: 19; BSFW, 1964a; POBSP, 1964b). 18 Aug. Bird i (POBSP, 1964a). 19-20 Aug. North 1-2 (Clapp and Woodward, 1968: 19; POBSP, 1964a). 17 Sept. Little North 1 (BSFW, 1964b; POBSP, 1964c). 1965 18 Mar. Little North 1 (Clapp and Woodward, 1968: 19; POBSP, 1965b). 18-19 Mar. Seal 1 (Clapp and Woodward, 1968: 19; POBSP, 1965b). 196 Table 78. (continued) Population Date of Survey Island Estimate Remarks and References 1965 22 Mar. Seal i (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. Seal 2 (BSFW, 1966b). 22 Sept. North 1 (BSFW, 1966b). 1967 16-17 Mar. North de (BSFW, 1967d). 22 Mar. Kittery 1 (Clapp and Woodward, 1968: 19; BSFW, 1967a; POBSP, 1967d). 22 Mar. Seal at (Clapp and Woodward, 1968: 19; BSFW, 1967a; POBSP, 1967d). 31 May Seal 3 (POBSP, 1967b). 31 May Grass iL (POBSP, 1967b). 29 Aug. Seal 6 (POBSP, 1967a). 29-30 Aug. North 3 (POBSP, 1967a). 1968 24 Mar. Grass iL On the rocky ledge at southwest corner (BSFW, 1968; POBSP, 1968). 24 Mar. Seal 5) Foraging on rocky south perimeter (BSFW, 1968; POBSP, 1968). RUDDY TURNSTONE Arenaria interpres Status Common migrant; usually recorded in flocks on beaches or inland tidal pools. Most abundant during late summer and fall; some birds present through- out the year. POBSP and BSFW maximum population estimate 670 in August 1967. Observations Ruddy Turnstones were first observed in March 1913 (Willett, ms.). On 14 October 1957 Rice (ms. a) observed about 30 during an aerial survey but gave no specific locations. The first published record appeared after POBSP personnel began visiting the atoll in 1963 (Clapp and Woodward, 1968: 20). It has been observed each year since (Tables 79 to 83). j ; 197 Annual Cycle Ruddy Turnstones, the predominant shorebird on the atoll, can be found throughout the year. Available data indicate the highest population is in late summer and fall. During the migratory period, the population fluctuates from day to day as birds arrive and depart. Eeological Distribution The Ruddy Turnstone is known from all islands. Southeast, the largest island, is by far the most popular; it is commonly seen there in small to large flocks on the beaches, rock flats, and around the central tidal pools. Banding and Movements POBSP and BSFW personnel banded 46 Ruddy Turnstones on Pearl and Hermes Reef. The POBSP banded 2 on Seal Island in March 1965, 26 on Southeast Island in February 1963, and 1 on Southeast in August 1964. All banded by the BSFW were on Southeast: 3 in March 1965, 9 in March 1968, and 5 in February 1969. Three Ruddy Turnstones originally banded on Pearl and Hermes have been recaptured on the atoll (Table 16). In addition, three originally banded by POBSP personnel on the Pribilof Islands have been captured on the atoll; these data are presented in Appendix Table ll. Specimens POBSP: USNM 494155, 2, collected 17 August 1964 on Southeast by Woodward; USNM 497545, 9, (USFW band # 722-14020), collected 28 May 1967 on Southeast by DeLong. Non-POBSP: USNM 301061, o, collected 26 April 1923 by Wetmore. Table 79. Observations of Ruddy Turnstones at Grass Island Population Date of Survey Estimate Remarks and References 1923 27 Apr. 2 Present (Wetmore, ms.). 1963 5 Mar. several (POBSP, 1964d). 26-27 June al - On reef (Clapp and Woodward, 1968: 20; POBSP, 1963). 1964 14 Mar. 20 (Clapp and Woodward, 1968: 20; BSFW, 1964a; POBSP, 1964b). 18 Aug. 3 (POBSP, 1964a). 1965 19 Mar. 50-60 Flock of 39, as well as scattered individuals (POBSP, 1965b). 198 Table 79. (continued) Population Date of Survey Estimate Remarks and References 1965 22 Mar. 80 (Clapp and Woodward, 1968: 20; BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 30 (BSFW, 1966b). 1967 22 Mar. 80 1 flock (BSFW, 1967a; POBSP, 1967d). 31 May 2 (POBSP, 1967b). 29 Aug. 35 Flocks of 13 and 18 (POBSP, 1967a). 1968 24 Mar. 100 Seen along beach perimeter and in sand and rubble at east end; flocks of 30, 32, 40, and 18; some probably seen in more than 1 flock (BSFW, 1968; POBSP, 1968). 1969 11 Sept. ©) (BSFW, 1969d). Table 80. Observations of Ruddy Turnstones at North Island Population Date of Survey Estimate Remarks and References 1ST Mere common (Willett, ms.). 1963 23-25 June 12-15 Daily (Clapp and Woodward, 1968: 20; POBSP, 1963). 1964 19-20 Aug. O (POBSP, 1964a). 17 Sept. 15 Count (Glapp and Woodward, 1968: 20; BSFW, 1964b; POBSP, 1964c). 1965 17-18 Mar. 75-100 (Clapp and Woodward, 1968: 20; POBSP, 1965b). 1966 22 Sept. ho (BSFW, 1966b). 1967 16-17 Mar. 75 (BSFW, 1967d). 29-30 Aug. 60 51 counted (POBSP, 1967a). 1969 12 Sept. 35 (BSFW, 1969d). 199 Table 81. Observations of Ruddy Turnstones at Seal Island Population Date of Survey Estimate Remarks and References 1923 27 Apr. ? Presence noted (Wetmore, ms.). 1963 5 Mar. ? Presence noted (POBSP, 196d). 26 June O (POBSP, 1963). 1964 14 Mar. O (BSFW, 1964a; POBSP, 1964b). 18 Aug. 9 Count (POBSP, 1964a). 1965 18-19 Mar. ? Presence noted (POBSP, 1965b). 22 Mar. 50 In 1 flock (Clapp and Woodward, 1928: 20; BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 30 (BSFW, 1966b). 1967 22 Mar. 45 In 1 flock (BSFW, 1967a; POBSP, 1967d). 31 May 5 (POBSP, 1967b). 29 Aug. 65 Flock of 50 around tidal pools (POBSP, 1967a). 1968 24 Mar. 35-40 Most numerous around tidal pools (BSFW, 1968; POBSP, 1968). 1969 11 Sept. 0 (BSFW, 1969d). Table 82. Observations of Ruddy Turnstones at Southeast Island Population Date of Survey Estimate Remarks and References 1916 4 Feb. large (Munter, ms.). numbers 1923 26-28 Apr. common (Wetmore, ms.). 1961 12 Mar. small (HDFG, 1961). numbers 1963 26 Feb.- 150-200 Throughout the island; roosting on rocky 8 Mar. reefs nocturnally (Clapp and Woodward, 1968: 20; POBSP, 1964d). 200 Table 82. Date of Survey Estimate Remarks and References 1963 1964 1965 1966 1967 1968 1969 (continued) Population 18-23, 15-20 25 June 13-14 Mar. 200 16-19 Aug. 1) 16 Sept. 500 15-19 Mar. few 21-22 Mar. 86 20-26 Sept. 200 25-27 Sept. 350 21-23 Mar. 100+ 28, 30 May- 116 1 June 28-30 Aug. 500 22-24 Mar. 100 10-12 Feb. 50 31 Mar.- 26 2 Apr. 26-31 May 16 10-19 Sept. 225 Daily (Clapp and Woodward, 1968: 20; POBSP, 1963). Count of 178 (Clapp and Woodward, 1968: 20; BSFW, 1964a; POBSP, 1964b). In several flocks along south shore and around interior lagoon (Clapp and Woodward, 1968: 20; POBSP, 1964a). (Clapp and Woodward, 1968: 20; BSFW, 1964b; POBSP, 1964c). (POBSP, 1965b). Count (Clapp and Woodward, 1968: 20; BSFW, 1965; POBSP, 1965a). (BSFW, 1966b). 75-100 new birds arrived on night of 26 Sept. (Clapp and Woodward, 1968: 20; POBSP, 1966). Scattered around (Clapp and Woodward, 1968: 20; BSFW, 1967a; POBSP, 1967d).. Count (POBSP, 1967b). Flocks of 50-70 frequently flushed from rock ledges on south side; ca. 30 birds feeding at once in central tidal pools; flocks of 10-30 frequented beaches (POBSP, 1967a). Feeding in small numbers in interior tidal pools but most numerous on rocky ledges along south side (BSFW, 1968; POBSP, 1968). Common along beaches (BSFW, 1969a). Count (BSFW, 1969b). Maximum number during survey (BSFW, 1969c). (BSFW, 1969d). L j 201 Table 83. Observations of Ruddy Turnstones on other islands at Pearl and Hermes Reef Population Date of Survey Tsland Estimate Remarks and References 1964 18 Aug. 18 Aug. 18 Aug. 17 Sept. 1965 18 Mar. 18 Mar. 22 Mar. ee Mar. 22 Mar. 22 Mar. 1966 21 Sept. 1967 22 Mar. 29 Aug. 1968 24 Mar. SNIPE species Status Accidental; one sight record. Bird Sand Kittery Little North Little North Kittery Planetree Sand Bird Kittery Kittery Kittery Little North Kittery 3 2 dL 5 30 iy (POBSP, 1964a). (POBSP, 1964a). Seen from Seal Island (POBSP, 1964a). Count (Clapp and Woodward, 1968: 20; BSFW, 1964b; POBSP, 1964c). (Clapp and Woodward, 1968: 20). Flock flying over island (Clapp and Woodward, 1968: 20; POBSP, 1965b). (Clapp and Woodward, 1968: 20; BSFW, 1965; POBSP, 1965a). (Clapp and Woodward, 1968: 20; BSFW, 1965; POBSP, 1965a). 1 long-dead banded turnstone found (BSFW, 1965; POBSP, 1965a). (Clapp and Woodward, 1968: 20; BSFW, 1965; POBSP, 1965a). (BSFW, 1966b). (Clapp and Woodward, 1968: 20; BSFW, 1967d). Count; 1 flock (POBSP, 1967a). 1 flock (BSFW, 1968; POBSP, 1968). Capella sp. 202 Observations An unusual shorebird, identified by Kridler (BSFW, 1964a) as a Common (Wilson's) Snipe, was seen on Southeast Island 13 March 1964. Both Common Snipe, Capella gallinago delicata, and Pintail Snipe, ~te stenura, have been collected on Kure Atoll (Clapp and Woodward, 1960: 21). This is a first sight record for Pearl and Hermes. KNOT Calidris canutus Status Accidental; one specimen record. Observations A very fat female Knot, collected 15 March 1965, constitutes the only record; it was the first specimen taken in the entire Hawaiian area. Knots have been seen at Midway and Oahu (Clapp and Woodward, 1968: 21-22); one was recently collected on Oahu (Clapp and Pyle, 1968: 38). It was first seen in the afternoon as it fed along the Sesuvium- bordered edge of a small brackish pond in the center of Southeast Island. The bird was very wary, flushing when approached; it was later observed on rocky rubble with a flock of Golden Plovers. After flushing once more, the Knot returned to the margin of the central pond where it was collected (POBSP, 1965b). Specimens POBSP: USNM 494130, 9, collected on Southeast Island 15 March 1965 by Clapp. SANDERLING Crocethia alba Status Common migrant; usually recorded in singles or in small flocks on beaches. A few birds present throughout the year. POBSP and BSFW maximum population count 7 in March 1963 and August 1967. Observations HDFG personnel first recorded Sanderlings in March 1961. POBSP and BSFW personnel recorded them in 1963, and have found them each year except 1966. Clapp and Woodward (1968: 22) reported some of these observations; all Sanderling observations are presented in Tables 84 to 86. 203 Annual Cycle Sanderlings can be found on the atoll in small numbers throughout the year. March population counts average 5 and range from 3 to 7. Ecological Distribution Sanderlings have been recorded on Grass (7 times), Kittery (2), Little North (1), North (1), Seal (3), and Southeast (9). This species is commonly seen as singles, or in small flocks on beaches, where it frequently associates with Ruddy Turnstone. Table 84. Observations of Sanderlings at Grass Island Population Date of Survey Estimate Remarks and References 1963 5 Mar. 1-2 (Clapp and Woodward, 1968: 22; POBSP, 19644). 26-27 June 0 (POBSP, 1963). 1964 14 Mar. 3 (Clapp and Woodward, 1968: 22; BSFW, 1964a; POBSP, 1964b). 18 Aug. 2 (Clapp and Woodward, 1968: 22; POBSP, 1964a). 1965 19 Mar. 3 In flock of turnstones at southwest corner — (Clapp and Woodward, 1968: 22; POBSP, 1965b). 22 Mar. 3 (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 0 (BSFW, 1966b). 1967 22 Mar. O (BSFW, 1967a; POBSP, 1967d). 31 May ) (POBSP, 1967b). 29 Aug. 2 (POBSP, 1967a). 1968 2h Mar. 3 Feeding with turnstones along sandy strip connecting 2 halves of island (BSFW, 1968; POBSP, 1968). 0 (BSFW, 1969d). 204 Table 85. Observations of Sanderlings at Southeast Island Population Date of Survey Estimate Remarks and References 1923 26-28 Apr. O (Wetmore, ms.). 1961 12 Mar. small (HDFG, 1961). numbers 1963 26 Feb.- 3 (Clapp and Woodward, 1968: 22; POBSP, 1964d). 8 Mar. 18-23, @) (POBSP, 1963). 25 June 1964 13-14 Mar. 2 (Clapp and Woodward, 1968: 22; BSFW, 1964a; POBSP, 1964b). 16-19 Aug. O (POBSP, 1964a). 16 Sept. @) (BSFW, 1964b; POBSP, 1964c). 1965 15-19 Mar. 1 On several occasions (Clapp and Woodward, 1968: 22; POBSP, 1965b). 21-22 Mar. iL (Clapp and Woodward, 1968: 22; BSFW, 1965; POBSP, 1965a). 1966 25-27 Sept. O (POBSP, 1966). 1967 21-23 Mar. 2 (BSFW, 1967a; POBSP, 1967d). 28, 30 May- 0 (POBSP, 1967b). 1 June 28-30 Aug. 2-3 Most frequently along sandy north beach, also foraging along east shore of lagoon; 1 feeding inland in a tidal pool (POBSP, 1967a). 1968 22-2 Mar. 1-2 Most frequently on sandy north beach and spit connecting two halves of island; 1 on southwest beach (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 0 (BSFW, 1969a). 31 Mar.- 0 (BSFW, 1969b). 2 Apr. 26-31 May @) (BSFW, 1969c). 10-19 Sept. 2 (BSFW, 1969d). nN ee 205 Table 86. Observations of Sanderlings on other islands at Pearl and Hermes Reef Population Date of Survey Island Estimate Remarks and References 1963 5 Mar. Seal few (Clapp and Woodward, 1968: 22; POBSP, 1964d). 1964 19-20 Aug. North iL (Clapp and Woodward, 1968: 22; POBSP, 1964a). 17 Sept. Little 1 (BSFW, 1964b; POBSP, 1964c). North 1965 22 Mar. Kittery i (BSFW, 1965; POBSP, 1965a). 1967 22 Mar. Seal il In flock of Ruddy Turnstones : (Clapp and Woodward, 1968: 22; BSFW, 1967a; POBSP, 1967d). 29 Aug. Seal 2 Foraging around tidal pools (POBSP, 1967a). 1968 24 Mar. Kittery 1 In flock of turnstones (BSFW, 1968; POBSP, 1968). SHARP-TATLED SANDPIPER Erolia acuminata Status Accidental; two specimen records. Observations On 27 September 1966 POBSP personnel collected two female Sharp-tailed Sandpipers at the pond on the east side of Southeast Island. Both had ex- tremely heavy fat deposits and were not molting. These birds, and one seen the preceding day, comprise the only records. The species is also known from Laysan, Midway, and Kure (Clapp and Woodward, 1968: 23-24; POBSP, 1966). Specimens POBSP: USNM 497216-17, 99, collected on Southeast Island 27 September 1966 by Lewis. DUNLIN Erolia alpina sakhalina Status Accidental; one specimen record. 206 Observations A very fat male Dunlin, with medium to heavy body molt, was collected by POBSP personnel on 15 March 1965 as it fed in a small Sesuvium-bordered brackish pond on Southeast Island. This specimen, the only record for Pearl and Hermes Reef, was subsequently identified as Erolia alpina sak- halina by Mrs. Roxie C. Laybourne of the U.S. Fish and Wildlife Service. Dunlins have also been recorded from Kure, Laysan, and Midway (Clapp and Woodward, 1968: 24; POBSP, 1965b). Spe imens POBSP: USNM 494127, o, collected on Southeast Island 15 March 1965 by Clapp. RUFF Philomachus pugnax Status Accidental; one specimen record. Observations A Ruff was seen on the morning of 28 August 1967 as it fed with several Ruddy Turnstones in the largest tidal pool in the Sesuvium-filled depression on Southeast Island. It was subsequently collected and found to be an immature bird, not molting, with light fat deposits (POBSP, 1967a). This is the third record from the Central Pacific. Ruffs have been reported from Kure Atoll (Clapp and Woodward, 1968: 24-25) and from Johnston Atoll (Shelton, in prep.). Specimens POBSP: USNM 543042, immature 9, collected 28 August 1967 on Southeast by Clapp. This is a first specimen record. RED PHALAROPE Phalaropus fulicarius Status Accidental; one specimen record. Observations On 22 March 1967 POBSP personnel found the dried carcass of a Red Phalarope in a stand of wild mustard in the interior of Southeast Island (POBSP, 1967d). This is the only record from Pearl and Hermes Reef; this species has also been taken from Kure and the main Hawaiian area (Clapp and Woodward, 1968: 25). 207 Specimens POBSP: USNM 497293, collected 22 March 1967 on Southeast Island by Hackman. RING-BILLED GULL Larus delawarensis Status Straggler; one specimen and one sight record. Observations A Ring-billed Gull in second winter plumage was collected 5 March 1963 on Little North Island by POBSP (1964d) personnel. This specimen constituted the first record from Pearl and Hermes Reef; this species is also known from Kure Atoll (Clapp and Woodward, 1968: 25; Sibley and McFarlane, 1968: 315). On the afternoon of 23 March 1967 BSFW personnel reported a Ring-billed Gull flying off Southeast Island. This bird was not seen by the POBSP ob- server present on the survey and no corroboratory details of the sighting are available (BSFW, 1967a; POBSP, 1967d). Specimens POBSP: USNM 493342, unsexed, collected 5 March 1963 on Little North Island by Sibley. HERRING GULL Larus argentatus vegae Status Rare visitor; two specimen records. Observations The only specimens of Herring Gulls were collected by POBSP personnel in 1963: a young male in first winter plumage was shot on Southeast on 2/ February; a young female in first nuptial plumage was shot on Kittery on 5 March (POBSP, 1964d). Both specimens were subsequently referred to Dr. Lester L. Short, Jr., of the American Museum of Natural History, and Mrs. Roxie C. Laybourne, of the U.S. Fish and Wildlife Service, for subspecific determination. Both were identified as Larus argentatus vegae, a race that breeds in Siberia. Herring Gulls have also been recorded from Kure, Laysan, Lisianski, and Midway (Clapp and Woodward, 1968: 26; Sibley and McFarlane, 1968: 315). 208 Specimens POBSP: USNM 493346, co, collected on Southeast Island 27 February 1963 by Sibley; USNM 493347, 9, collected on Kittery Island 5 March 1963 by Sibley. GLAUCOUS-WINGED GULL Larus glaucescens Status Irregular visitor; four specimens and three sight records from February, March, and April. Observations The first record was the carcass of an adult bird collected on Seal Island 27 April 1923 by Alexander Wetmore (Clapp and Woodward, 1968: 27). Another Glaucous-winged Gull, one of four gulls present on the atoll, was shot on Southeast Island by POBSP personnel on 26 February 1963 (Sibley and McFarlane, 1968: 315; POBSP, 1964d). This bird was a male in first nuptial plumage. POBSP personnel saw two more Glaucous-winged Gulls on the southern tip of North Island on 17 March 1965. An unsuccessful attempt was made to col- lect these gulls which flushed and flew to nearby Little North Island where they were collected the following morning (Sibley and McFarlane, 1968: 315; POBSP, 1965b). Both specimens were young birds in first nuptial plumage. Their stomachs contained fish remains; one contained a gastropod and several foraminifera as well. ae | el Specimens POBSP: USNM 493345, o, collected 26 February 1963 on Southeast by Sibley. USNM 494131-32, 99, collected 18 March 1965 on Little North by Hoeman. Non-POBSP: USNM 489330, collected 27 April 1923 on Seal by Wetmore. BLACK-LEGGED KITTIWAKE Rissa tridactyla Status ee ee ee a eT ey Rare visitor; three specimen records. _ Observations On 15 March 1965 POBSP personnel found the remains of two Black-legged Kittiwakes on Southeast Island: one nearly whole carcass of an adult in winter plumage, and a wing from an immature bird on the beach near the western 209 end of the island. Partial remains of another immature bird were found 19 March 1965 on the north beach of Grass Island near the vegetation line (POBSP, 1965b). Markings on the wings of the two immature birds indicate that both were less than two years old. Black-legged Kittiwakes have been reported from Kure and Laysan (Clapp and Woodward, 1968: 28-29). Specimens POBSP: USNM 496205 and 497375, adult and immature respectively, col- lected 15 March 1965 on Southeast Island by Stadel; USNM 496206, immature, collected 19 March 1965 on Grass Island by Wirtz. GRAY-BACKED TERN Sterna lunata Status Common breeding species; present spring, summer, and early fall; absent rest of year. Nests during spring and summer on Grass, Seal, and Southeast Islands; occasionally seen on other islands. POBSP and BSFW maximum popula- tion estimate 1,860t in May and June 1967. Observations Wetmore (ms.) recorded nesting Gray-backed Terns in April 1923. Anderson (1954: 82) saw a colony during his late 1920's visits. On 14 October 1957, while surveying the atoll from the air, Rice (ms. a) noted several uniformly gray terns in bright sunlight flying near White Terns over the large protrud- ing rocks at the northeastern end of the reef. He tentatively identified these as Blue-gray Noddies (Procelsterna cerulea), but they were more likely to have been Gray-backed Terns. Woodside and Kramer (HDFG, 1961) found Gray- backed Terns in March 1961. POBSP and BSFW personnel recorded nesting gray- backs each year since 1963: All Gray-backed Tern observations are presented in Tables 87 to 89. Annual Cycle The local annual breeding cycle is shown in Figure 58. Adults return as early as early February after being absent during late fall and winter. Eggs are known as early as mid-March; peak egg laying must be during April, and eggs are known as late as mid-August. Hatching began as early as late April in 1923, and flying immatures were seen in late May 1969. Most young fledge by late August and September. Ecological Distribution Gray-backed Terns nest on Grass, Seal, and Southeast Islands, and have been seen at Bird, Kittery, and North Islands. 210 Grass Island: POBSP personnel first recorded a few adults but no nests in May 1967. Several unfledged young were found on the north beach and in low vegetation in August 1967; no nests were seen then. BSFW and POBSP personnel again noted a few adults in March 1968 (Table 89). Seal Island: Wetmore (ms.) found nesting Gray-backed Terns in April 1923. Clapp and Woodward (1968: 29) recorded their presence in 1963 to 1965 and 1967 from POBSP data. Table 87 presents all observations. March maximum population estimates average 32, and range from O to 300. The peak population estimate was 350, which coincided with the peak nest count of 1754. This species nests on the rocky east portion of the island; a few birds may nest on the ground in the vegetated west portion. Southeast Island: Although Wetmore (ms.) noted breeding Gray-backed Terns in April 1923 and Woodside and Kramer (HDFG, 1961) found some in March 1961, Clapp and Woodward (1968: 29) first published records of their occurrence using 1963 to 1967 POBSP data. All observations are presented in Table 88. March maximum population estimates average 209, and range from 30 to 600. The peak population of 1,500 occurred in May and June 1967; coinciding with this was a peak nest estimate of 450. Nests are found on both east and west portions. On the east portion small groups of nests are scattered about the vegetated portions, around the central tidal pools, and along the perimeter of the northeast, South, and southeast beaches. Some nests are found around the perimeter of vegetation on the west portion. Usually, the nests are on the edge of the Sooty Tern sub-colonies. Other Islands: POBSP personnel have recorded Gray-backed Terns roost= ing on Bird and Kittery Islands and flying over North Island (Clapp and Woodward, 1968: 29). These data are presented in Table 89. Banding and Movements POBSP and BSFW personnel banded 338 Gray-backed Terns (Table 90). Kleven have been recaptured on the atoll. There have been no interisland movements involving Pearl and Hermes Reef. Specimens Non-POBSP: USNM 300633-34, o, 2, collected 27 April 1923 by Wetmore; USNM 300635 -36, 3, oh, collected 26 April 1923 by Wetmore. These are first specimen records. 211 Table 87. Observations of Gray-backed Terns at Seal Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. - Not mentioned in report (Munter, ms.). 1923 27 Apr. 300 Breeders with eggs and newly hatched young (Wetmore, ms.). 1963 5 Mar. ©) (POBSP, 1964d). 26 June 120-140 100 adults, 20-40 young (Clapp and Woodward, 1968: 29; POBSP, 1963). 1964 14 Mar. a5 No nests (Clapp and Woodward, 1968: 29; BSFW, 1964a; POBSP, 1964b). 18 Aug. iS) All adults; no evidence of breeding (Clapp and Woodward, 1968: 29; POBSP, 1964a). 1965 18-19 Mar. 200-300 Roosting at night; 40-100 breeders; 20-50 nests with eggs; just beginning to lay (Clapp and Woodward, 1968: 29; POBSP, 1965b). 22 Mar. 150 Mostly adults present nocturnally; 4 nests with eggs (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 7+ 5 flying immatures and 2 large dependent young (BSFW, 1966b). 1967 22 Mar. 25 Roosting on rocky area at north end; no nests found (Clapp and Woodward, 1968: 29; BSFW, 1967a; POBSP, 1967d). 31 May 350+ Breeders; 175¢ nests: most with eggs, a few with chicks up to 10 days old (POBSP, 1967b). 29 Aug. 10 1 adult; 9 fledged immatures (POBSP, 1967a). 1968 24 Mar. 100 Adults; 30-40 breeders; 15-20 nests with eggs (BSFW, 1968; POBSP, 1968). 1969 11 Sept. 12 (BSFW, 1969d). Table 88. Observations of Gray-backed Terns at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. Not mentioned in report (Munter, ms.). ale Table 88. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1923 26-28 Apr. 600 Breeders, with eggs or newly hatched young (Wetmore, ms.) 1961 12 Mar. ig Apparently just arriving; on the ground but no eggs yet laid (HDFG, 1961). 1963 26 Feb.- 200 Adults; lst arrived 3 March; gradually in- 8 Mar. creased in numbers throughout survey; no eggs laid (Clapp and Woodward, 1968: 29; POBSP, 1964d). 18-23, 1, o40 1,000 adults, most breeding birds on eggs, 25 June but 40 pairs with small chicks (Clapp and Woodward, 1968: 29; POBSP, 1963). 1964 13-14 Mar. 20-30 Adults on the ground; no nests (Clapp and Woodward, 1968: 29; BSFW, 1964a; POBSP, 1964b). 16-19 Aug. 850-950 600-700 adults; 500 breeders with 250% near- fledged young, a few 1/4-grown chicks, 1 egg (Clapp and Woodward, 1968: 29; POBSP, 1964a). 16 Sept. 50 40 adults, ca. 10 flying immatures (Clapp and Woodward, 1968: 29; BSFW, 1964b; POBSP, 1964c). 1965 15-19 Mar. 500-600 Adults on ground but no eggs (Clapp and Woodward, 1968: 29; POBSP, 1965b). 21-22 Mar. 300 Adults, no nests (BSFW, 1965; POBSP, 1965a). 1966. 1 Apr. - Not mentioned in notes (BSFW, 1966a). 20-26 Sept. 10 8 adults, 2 flying immatures (BSFW, 1966b). 25-27 Sept. 47 40 adults, 144 breeders, ca. 7 flying im- matures still being fed (Clapp and Woodward, 1968: 29; POBSP, 1966). 1967 21-23 Mar. TS) Adults on ground by day, but no nests with eggs (Clapp and Woodward, 1968: 29; BSFW, 1967a; POBSP, 19674). 28-30 May- 1,500 900-1,000 breeders; 450 nests: mostly with 1 June fresh eggs, 10-day-old chicks (POBSP, 1967b). a RN 213 Table 88. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1967 28-30 Aug. 15 50 breeders, ca. 25 dependent young, some flying (POBSP, 1967a). 1968 22-24 Mar. 100-150 Adults on ground by day, but no eggs (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 50 Adults, nocturnal only (BSFW, 1969a). 31 Mar.- 190 Breeding adults, 95 nests (BSFW, 1969b). 2 Apr. 26-31 May 150 100 on east portion, 50 on west; eggs to flying young (BSFW, 1969c). 10-19 Sept. 4 (BSFW, 1969d). Table 89. Observations of Gray-backed Terns on other islands at Pearl and Hermes Reef Population Breeding Date of Survey Island Estimate Status, Remarks, and References 1963 23-25 June North 1-2 Flying over daily (Clapp and Wood- ward, 1968: 29; POBSP, 1963). 1964 18 Aug. Bird 1 (Clapp and Woodward, 1968: 29; POBSP, 1964a). 18 Aug. Kittery i Seen from Seal Island (Clapp and Woodward, 1968: 29; POBSP, 1964a). 19-20 Aug. North 6 Adults flying over (Clapp and Wood- ward, 1968: 29; POBSP, 1964a). 1967 31 May Grass 6-8 Adults flying over; their behavior indicated nests but none was found (POBSP, 1967b). 29 Aug. Grass 7-10 1 unfledged young; remainder were recently fledged young (POBSP, 1967a). 1968 2h Mar. Grass 4-6 On the ground but no eggs (BSFW, 1968; POBSP, 1968). 214 AG Ln G oL it iw) +1d Tog €T TeI0L G 0 S ) 0 0 0 0 INV Tees Li dig? aL in Wh ie ee Tea 0 0) © - Gh i +t iT Tr ©) Sunox Lt Ly 0 l2 O 12 eT? 00g €T JIMpy 4S8eeyynog Te40L G96 G96 T2701 HO6L HO6L T2401 CO6T C961 sseTo pueTST G96T “Ze “Tey 496T ‘* deg “ony €96T aunt “Ten -93y =°geaq Kaaing {oO potaog T7177 <= eee MASE dSd0d dSd0d dsd0d dSd0d dSd0d ‘topueg Joy Souza pue [1teeq 42 paepueq suze], peyoeq-Aery °O6 ataeL 215 SOOTY TERN Sterna fuscata Status Abundant breeding species; present usually from mid-February through early October; absent during rest of year. Nests on the ground at Seal and Southeast Islands; known from Grass and North Islands. Maximum POBSP and BSFW population estimate 80,000+ in May and June 1967. Observations Munro (1942: 12), on 6 July 1891, was the first to see a few Sooty Terns off Pearl and Hermes Reef; he did not go ashore. One was seen by Willett in mid-March 1913 (Bailey, 1956: 32). Wetmore (ms.) observed Sooty Terns with eggs in April 1923, and Galtsoff (1933: 19) noted them in large numbers in late summer 1930. Richardson (1967: 24) noted that they bred here, but gave no data. POBSP, HDFG, and BSFW personnel observed this species in 1961, and 1963 to 1969. All observations are summarized in Tables 91 to 93. Annual Cycle The local breeding cycle is illustrated in Figure 58. Adults begin to arrive in mid-February; after swirling overhead diurnally and alighting on the ground nocturnally during late February and March, they usually settle on the ground both day and night by late March. Egg laying probably commences in mid-April and reaches a high from late April to mid-May. Hatching may be as early as mid-May, but reaches a high in late May and early June. Fledging occurs by mid-August; some young remain into late September, with a few stragglers remaining into October. The majority of adults and immatures usually departs by mid-September. The Sooty Tern is the most abundant species at Pearl and Hermes Reef. The peak population occurs in late May and June: 80,000 in 1967 and 40,000 in 1963 and 1969. The peak nest counts also occur during May-June: 38,700+ in 1967, ca. 20,000 in 1963 and 1969. Ecological Distribution The Sooty Tern nests at Seal and Southeast Islands, and is known from Grass and North Islands. Seal Island: Wetmore (ms.) first observed nesting Sooty Terns in April 1923. POBSP and BSFW personnel observed nesting in most years since 1963 (Table 91). The highest population was 1,000 to 1,500 roosting birds in August 1964. Nine nests were found in late May 1967, and 50 in late June 1963. Nest sites in 1967 were in an area of Tribulus, Boerhavia, Lepidium, and Rragrostis. Southeast Island: Southeast is the major breeding island at Pearl and Hermes Reef. Wetmore (ms.) first recorded nesting in April 1923. Galtsoff 216 (1933: 19) published the first record of their occurrence from observa- tions taken in late summer 1930. Since 1963 BSFW and POBSP personnel observed nesting Sooty Terns each year (Table 92). A peak population of 80,000 occurred in May and June 1967. Nests are usually scattered over the entire island. The May and June 1967 population contained definite subpopulations, groups of birds that had begun to nest at different times; some of these subpopulations were isolated, but others were contiguous. Density of nesting birds and age of eggs (using flotation method) were determined for all the subpopulations. From this, the entire island's Sooty Tern population size was calculated (Table 94). Other Islands: Willett observed a Sooty Tern at North Island in March 1913 (Bailey, 1956: 32). In recent years, POBSP and BSFW personnel noted sooties flying over in very small numbers. From 2 to 20 individuals were also recorded over Grass Island by POBSP and BSFW personnel (Table 93). Sooty Terns probably fly over all the other islands in the atoll during the spring and summer. Banding and Movements POBSP personnel banded 4,327 Sooty Terns (Table 95). Of these, 343 have been recaptured on the atoll (see Table 16); another 8 have been cap- tured (2 each) at Lisianski, Midway, Kure, and Johnston. Furthermore, 14 Sooty Terns, 7 from Johnston, 5 from Kure and 1 each from Lisianski and Midway, have been captured on Pearl and Hermes. Data are presented in Appendix Tables 12a and 12b. Specimens POBSP: USNM 494934, 9, collected on Southeast Island 17 August 1964 by Marshall (USFW 753-24662). Non-POBSP: USNM 300543-44, o, o, collected 26 April 1923 by Wetmore. These are first specimen records. Table 91. Observations of Sooty Terns at Seal Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1923 27 Apr. 300 Adults, few with eggs (Wetmore, ms.). 1963 5 Mar. 2 Flying over (POBSP, 1964d). 26 June 100 Adult breeders, 50 nests with eggs (POBSP, 1963). 1964 14 Mar. 0 (BSFW, 1964a; POBSP, 1964b). ZANT Table 91. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1964 18 Aug. 1, 000- Adults roosting on ground (POBSP, 1964a). 1,500 1965 18-19 Mar. 200-300 Swirling overhead at night; none on ground (POBSP, 1965b). 22 Mar. 100 Flying over by day (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 0 (BSFW, 1966b). 1967 22 Mar. ) (BSFW, 1967a; POBSP, 19674). 31 May 18 Adult breeders; 9 on eggs among nesting Gray-backed Terns (POBSP, 1967b). 29 Aug. ) (POBSP, 1967a). 1968 24 Mar. 2 Adults flying offshore (BSFW, 1968; POBSP, 1968). 1969 11 Sept. 20 (BSFW, 1969d). Table 92. Observations of Sooty Terns at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. - Not mentioned in report (Munter, ms.). 1923 26-28 Apr. 200 Adults, few with eggs (Wetmore, ms.). 1930 23 July- large (Galtsoff, 1933: 19). Aug. numbers 1961 12 Mar. a Apparently just arriving; none on ground (HDFG, 1961). 1963 26 Feb.- 5, 000- Adults, on ground by day and night; no 8 Mar. 7, 000 eggs (POBSP, 1964d). 18-23, 40,000 Most with eggs but a few recently hatched 25 June young present; very few with large chicks (POBSP, 1963). 218 Table 92. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1964 13-14 Mar. 2, 000 Adults: about 300 in air at noon, 700 at dark; about 1,000 on ground at night with more overhead; no eggs (BSFW, 1964a; POBSP, 1964b). 16-19 Aug. 25 , 000 Most with fledged young but some unfledged birds; 6,000 young (POBSP, 1964a). 16 Sept. 5, 000- Adults and immatures, some still incapable 6, 000 of flight (BSFW, 1964b; POBSP, 1964c). 1965 15-19 Mar. 1,500- Adults flocking overhead, some on ground 2,000 by day; no eggs (POBSP, 1965b). 21-22 Mar. 3, 000- Adults, some on ground by day, many more 4,000 at night; no eggs (BSFW, 1965; POBSP, 1965a). 1666 Ball Apr thousands Circling; few on ground (BSFW, 1966a). 20-26 Sept. 2 Few heard overhead at night; 1 emaciated immature incapable of flight (BSFW, 1966b). 25-27 Sept. 5 Few flying over (POBSP, 1966). 1967 21-23 Mar. 2, 000- About 2,000 in swirling flocks by day, 5,000 5,000 at night; 1,200 on ground at night; no eggs (BSFW, 1967a; POBSP, 1967d). 28, 30 May- 80, 000 38,707 nests; fresh eggs to recently hatched 1 June young; most with moderately or heavily in- cubated eggs; estimate based on density-area calculations (POBSP, 1967b). 28-30 Aug. 650 Adults and flying young; some 200 young; most fledged but some still being fed by parents (POBSP, 1967a). 1968 22-24 Mar. 1, 000 Swirling by day, numbers increasing towards dusk; about 150 on the ground by day; no eggs (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 5 Heard calling at night, none on ground (BSFW, 1969a). 31 Mar.- 3, 000 Adults; numbers increased towards evening; 2 Apr. no nests (BSFW, 1969b). ER 219 Table 92. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1969 26-31 May 40, 000 A number of sub-colonies: the great majority (20,000) with eggs or recently hatched young; 1 small colony of about 400 contained near-fledged chicks; sample nest counts in 2 other colonies suggest 87% eggs and 13% young (BSFW, 1969c). 10-19 Sept. IDO (BSFW, 1969d). Table 93. Observations of Sooty Terns on other islands at Pearl and Hermes Reef Population Breeding Date of Survey Island Estimate Status, Remarks, and References 1891 6 July Offshore few (Munro, 1942: 12). 1913 15 Mar. North il (Bailey, 1956: 32). 1963 23-25 June North few Flying over (POBSP, 1963). 26-27 June Grass several Flying over (POBSP, 1963). 1964 18 Aug. Grass 20 Flying around (POBSP, 1964a). 19-20 Aug. North few Flying over (POBSP, 1964a). 1967 16-17 Mar. North 2 Flew over by day (BSFW, 19674). 29-30 Aug. North several Flew over late afternoon and evening (POBSP, 1967a). 1968 24 Mar. Grass 2 Adults flying offshore (BSFW, 1968; POBSP, 1968). Table 94. Density and status of subpopulations of Sooty Terns on Southeast Island, 28, 30 May-1 June 1967 Subpopulation Area* Density** Nest Sites Status 1 4815 400 15,600 18-day eggs and hatchlings 2 1,780 Why By, 2a l-day eggs to hatchlings 220 Table 94. (continued) Subpopulation Area* Density** Nest Sites Status 3 2,676 576 6,621 8-14-day eggs 4 500 900 978 fresh eggs 5 675 BIT) 1,124 14-day eggs to hatchlings 6 6,075 576 aya 14-day eggs to hatchlings i 2,774 SD) DRoao 14-day eggs to hatchlings 8 800 576 1,800 7-day eggs to hatchlings Total 20, 095 CM hxaxas 38,703 l-day eggs to hatchlings *Subpopulation area in square yards. *XMean individual territory sizes in square inches. *x* Average. Table 95. Sooty Terns banded at Pearl and Hermes Reef by the POBSP Period of Survey Age- June Aug. Mar. Aug. Island Class 1963 1964 1965 1967 Totals Southeast Adult 1,300 2, 000 O 25 Be) Young O 1, 000 O LO) 1, 000 Total: 1,300 3, 000 O 25 F325 Seal Adult O O 2 O 2 Total 1,300 3, 000 2 25 1327 BROWN NODDY Anous stolidus Status Common breeding species; present year-round, lower numbers in late fall and winter; breeding season may extent throughout the year. Nests at 2el1 Grass, North, Seal, and Southeast Islands; roosts on most other islands. Maximum POBSP and BSFW population estimate 8,395 in August 1964. Observations Brown Noddies, with eggs and young, were first recorded in March 1913 (Bailey, 1956: 32). Munter (ms.) found non-nesting flocks of various sizes in February 1916, and Wetmore (ms.) noted nesting birds in April 1923. In recent years, Rice (ms. a) saw low numbers in October 1957 and POBSP and BSFW personnel observed nesting Brown Noddies each year since 1963. All observations are presented in Tables 96 to 99. Annual Cycle The local annual breeding cycle is shown in Figure 58. Observations and interpolations of data reveal that eggs and young are known or are possible throughout the year. Eggs and young are, however, probably found infrequently during the late fall and winter. In most years the first eggs are laid in March; eggs have occurred as late as September. Although a few young occur in March, most usually hatch in April and continue into August. Fledging is known from spring months, but is more common in August and September. The peak population in recent years has been in August: 8,395 in 1964, and 4,010 in 1967. The peak nest count was in May and August 1967, 663% and 990, respectively. Ecological Distribution Brown Noddies nest at Grass, North, Seal, and Southeast Islands, and are known to roost at Kittery, Little North, and Sand Islands. Grass Island: Munter (ms.) first recorded small numbers in February 1916. Wetmore (ms.) found them nesting in April 1923. POBSP and BSFW personnel observed this species each year (Table 96). March population estimates average 10, and range from O to 30, whereas August and September estimates average 872, and range from O to 3,000. A peak nest,count of 263 was recorded in May 1967. Nests are located in the vegetated west part of the island. They are found on the ground in associa- tion with Tribulus, Boerhavia, and Solanum. Roosting birds utilize the northwest beach and the rocky east portion of the island. North Island: Eggs and young were first recorded in March 1913 (Bailey, 1956: 32). POBSP and BSFW personnel observed them on each visit made since 1963 (Table 97). March population estimates are few, but range from O to 200. Peak populations of adults and flying immatures as well as a peak nest count occurred in August 1964; September estimates ranged from 45 to 1,300. Nesting occurs over the entire vegetated part of the island; nests are 222 concentrated, however, around the periphery, particularly along the north- east and northwest beaches. Nests are located on the ground, primarily in association with Tribulus, Boerhavia, and Lepturus; smaller numbers are in areas of Eragrostis, Sicyos, and Solanum. Seal Island: Brown Noddies were first observed by Munter (ms.) in February 1916. Wetmore (ms.) found the species nesting in April 1923. POBSP and BSFW personnel recorded Brown Noddies each year except 1969 (Table 98). March maximum population estimates averaged 58, and ranged from O to 150; August and September estimates ranged from 0 to 1,500. A peak nest count of 200 was made in May 1967. Nests are mostly in the areas of low Tribulus and Lepturus bordering higher growth on the west half of the island. Roosting birds primarily utilize the beaches and the east rocky area. Southeast Island: Munter (ms.) found large flocks in February 1916. Some 30 nesting pairs, with almost fledged young, were recorded by Wetmore (ms.) in April 1923. POBSP and BSFW personnel observed nesting Brown Noddies each year (Table 99). March maximum population estimates averaged 37, and ranged from 10 to 100; June maximum estimates averaged 500; August and September maximum esti- mates averaged 2,040, and ranged from 50 to 2,600. Nests are located on both halves of the island. On the east portion, nests are around the Scaevola on the perimeter of the island, in Sesuvium around the central tidal pools, in the Tribulus and Lepturus areas south and east of those pools, and a few are in association with Solanum. On the west portion, this species nests in association with Tribulus, Lepturus, Brasica, and a few with Solanum and Boerhavia. In May and June 1967 30 nests were examined; all contained Tribulus, 96 percent contained grass, 70 percent feathers, 67 percent bones, and 27 percent seaweed (Table 100). Other Islands: POBSP personnel observed 20 roosting at Sand Island 18 August 196% Three were noted flying over Little North Island 22 September 1966 by BSFW personnel. At Kittery Island 29 August 1967, 10, including 4 to 5, mostly immatures, roosting on the beach and 2 flying offshore, were esti- mated by POBSP personnel. Brown Noddies probably also roost on the two other sandy islands. Banding and Movements The POBSP and BSFW banded 695 Brown Noddies (Table 101). Of these, 17 have been recaptured on the atoll (Table 16); 2 others travelled to other atolls. One (USFW 723-65116) unsexed adult banded on Southeast 20 June 1963 was found dead at Sand Island, Johnston Atoll 24 February 1964. The other (723-63485), also an unsexed adult, was banded on North Island 24 June 1963 and was captured at Sand Island, Johnston 11 February 1966. Still another Sooty Tern (753-24902) banded as an adult on Sand Island, Johnston Atoll 20 December 1963 was collected at North Island 19 August 1964. 223 Specimens POBSP: USNM 494154, 9, collected 19 August 1964 on North Island by Woodward (USFW 753-24902). Non-POBSP: USNM 300502-03, o, oh, collected 26 April 1923 by Wetmore. These are first specimen records. Table 96. Observations of Brown Noddies at Grass Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. small No eggs or young (Munter, ms.). number 1923 27 Apr. ho Breeders, young well grown (Wetmore, ms.). 1963 5 Mar. O (POBSP, 1964d). 26-27 June 500% 200f nests with eggs; few young (POBSP, 1963). 1964 14 Mar. 0 (BSFW, 1964a; POBSP, 1964b). 18 Aug. 3, 000 Adults, not breeding (POBSP, 1964a). 1965 19 Mar. 15-20 12 breeders; 6 nests with eggs (POBSP, 1965b). 22 Mar. 10 1 nest with egg and 1 with a very small chick (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 90 Adults and flying immatures; no nests (BSFW, 1966b). 1967 22 Mar. 2 Breeding adults; 1 nest with egg (BSFW, 1967a; POBSP, 19674). 31 May 650 526 breeders; 263 nests counted: 95% with eggs, 5% with small downy young; ca. 100 birds roosting on beach (POBSP, 1967b). 29 Aug. 400 4h breeders; 22 nests; sample count of 20 nests: 10 (50%) with eggs, 4 (20%) with small downy young, and 6 (30%) with near- fledging young (POBSP, 1967a). 1968 2 Mar. 30 Mostly roosting birds; 1 medium-sized downy young (BSFW, 1968; POBSP, 1968). 1969 11 Sept. ©) (BSFW, 1969d). EES S9$ T2401 Jie BuUNOA 9EE ZIMpy y940N S 41I0py Tees OOT INV ssery LET > T2101, iS : sunoz HET 410py qseayynos STeVOL L961 GO6L T2101 961 HOG Teq0L CO6T €96T SSeTO pueTsl “any Tey 4H96T “ony “Tey €96T eune “ley -a3y -" 99d dS$d0d dSd0d d$q0d MASE dSd0d_ dSd0d :tapueg aAING JO potied 228 joey Ssowzey pue [1leeg 1e pepueq SeTppoN UMOZG “TOT eTqdeL 229 late as late August. Young may hatch as early as mid-April, but most hatch in late May and June, and probably into July. Fledging normally occurs in summer and fall, but may take place in spring. March maximum population estimates for the atoll averaged 430, and ranged from 25 to 900+; May-June population counts averaged 2,023, and ranged from 1,400 to 3,070; August and September maximum counts averaged 1,761, and ranged from 1,036 to 4,300. The peak nest count of 55 to 75 occurred in June. Ecological Distribution Black Noddies nest at North and Southeast Islands; roosting birds are known from Sand Island. Grass Island: Wetmore (ms.) found nesting Black Noddies with young in April 1923. No evidence of nesting was found by POBSP and BSFW personnel since 1963 (Table 102). Recent population estimates ranged from O on several visits to 400 in September 1969. North Island: Small colonies of Black Noddies with eggs and young were first recorded in March 1913 (Bailey, 1956: 32). POBSP and BSFW per- sonnel observed evidence of breeding in 1963, 1964, and probably 1967. Most counts were low. All observations are given in Table 103. A peak population of 1,100 roosting Black Noddies was present in August 1967. Wests are placed in Eragrostis clumps. Most roosting birds utilize the beaches. Seal Island: Wetmore (ms.) first recorded breeding Black Noddies with young in April 1923. POBSP and BSFW observers noted this species roosting but not nesting on six visits since 1963. The roosting population has varied from 1 in March to 700 in August; roosting occurs on the beaches. southeast Island: Breeding Black Noddies were first noted by Wetmore (ms.) in April 1923. POBSP and BSFW personnel observed this species breeding each year. All observations are presented in Table 105. March maximum population estimates averaged 174, and ranged from 40 to 400; May and Juné estimates ranged from 1,000 to 3,000; August and September estimates averaged 1,354, and ranged from 550 to 3,000. A high nest count of 50 occurred in June 1963. Most nests have been located in Eragrostis clumps; a few have been in solanum. Most are on the east half. Nests are constructed with Tribulus, grasses, and leaves and may be within 6 inches of the ground. Roosting birds utilize the tall Eragrostis, Brassica, and Scaevola. Other Islands: Rice (ms. a) observed 25 Black Noddies on the "south sandspits"--Bird, Planetree and Sand Islands--on 14 October 1957; five additional birds were sighted flying along the northeast reef. POBSP 230 personnel recorded 100 roosting at Sand Island on 18 August 1964; another 100 roosting birds were noted on 31 May 1967. Black Noddies probably also occasionally visit Kittery and Little North Islands. Banding and Movements POBSP and BSFW personnel banded 377 Black Noddies (Table 106); of these, 20 have been recaptured on the atoll (Table 16); 9 others moved elsewhere, 3 each to Kure and French Frigate Shoals, 2 to Lisianski, and 1 to Midway. In addition, 1 each from French Frigate Shoals, Laysan, and Midway travelled to Pearl and Hermes Reef. All movements are presented in Appendix Tables 13a and 13b. Specimens POBSP: USNM 497922-25, 9, o, ob, 2, collected 18 June 1963 on Southeast by Sibley and Amerson. Non-POBSP: USNM 300464-65, ot, o, collected 27 April 1923 by Wetmore. These are first specimen records. Table 102. Observations of Black Noddies at Grass Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. - Not mentioned in report (Munter, ms.). 1923 27 Apr. ho 20 pairs with well-grown young (Wetmore, ms.). 1963 5 Mar. 0 (POBSP, 19644). 26-27 June 0 (POBSP, 1963). 1964 14 Mar. 0) (BSFW, 1964a; POBSP, 1964b). 18 Aug. 3 Not nesting (POBSP, 1964a). 1965 19 Mar. a Roosting (POBSP, 1965b). 22 Mar. @) (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. ) (BSFW, 1966b). 1967 22 Mar. 0 (BSFW, 1967a; POBSP, 1967d). 31 May 200 Diurnal roosting; no evidence of nesting (POBSP, 1967b). 29 Aug. 50 1 flock roosting on the vegetated portion; few roosting with Brown Noddies; none nesting (POBSP, 1967a). 231 Table 102. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1968 24 Mar. 0 (BSFW, 1968; POBSP, 1968). 1969 11 Sept. 400 (BSFW, 19694). Table 103. Observations of Black Noddies at North Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1913 15 Mar. small Eggs and young (Bailey, 1956: 32). colonies 1963 6 Mar. O (POBSP, 1964d). 23-25 June 70 20-60 breeders; 10-30 nests (POBSP, 1963). 1964 19-20 Aug. 303+ 300 adults; 6 breeders: 3 young counted (POBSP, 1964a). 17 Sept. 100 No nests (BSFW, 1964b; POBSP, 1964c). 1965 17-18 Mar. 200-300 Mostly roosting at night; no nests (POBSP, 1965b). 1966 22 Sept. - Not mentioned in notes (BSFW, 1966b). 1967 16-17 Mar. goo+ Maximum of 60 on ground by day, an additional 850 arriving at dusk from the east and settl- ing in one large flock (BSFW, 1967d). 29-30 Aug. 1,100 Adults, mostly nocturnal, but flocks of 20- 350 along beaches during the day; no active nests, but 1 found that may have been used earlier in the year (POBSP, 1967a). 1969 12 Sept. O (BSFW, 1969d). Table 104. Observations of Black Noddies at Seal Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. - Not mentioned in report (Munter, ms.). 1923 27 Apr. ho Breeders; young well-grown (Wetmore, ms.). 232 Table 104. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1963 5 Mar. 0) (POBSP, 19644). 26 June 0 (POBSP, 1963). 1964 14 Mar. ) (BSFW, 1964a; POBSP, 1964b). 18 Aug. 700 Roosting adults; no nests (POBSP, 1964a). 1965 18-19 Mar. 50-100 Roosting adults at night; no nests found (POBSP, 1965b). 22 Mar. 1 Diurnal; no nests (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 36 Flying about; no nests (BSFW, 1966b). 1967 22 Mar. O (BSFW, 1967a; POBSP, 1967d). 31 May 200 Roosting; no nests (POBSP, 1967b). 29 Aug. 150 Roosting on the open beach towards the end of the reef; no evidence of nesting (POBSP, 1967a). 1968 24 Mar. ©) (BSFW, 1968; POBSP, 1968). 1969 11 Sept. @) (BSFW, 1969d). Table 105. Observations of Black Noddies at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1916 4 Feb. - Not mentioned in report (Munter, ms.). 1923 26-28 Apr. 100 Breeders: 1 fresh egg collected, but most nests apparently contained young, some near fledging (Wetmore, ms.). 1930 23 July- - Not mentioned in report (Galtsoff, 1933). Aug. 1961 12 Mar. - Not mentioned in report (HDFG, 1961). 1963 26 Feb.- ho Mostly present at night; no nests (POBSP, 8 Mar. 1964d). 233 Table 105. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1963 18-23, 2, 000- 100 breeders: 45 nests with eggs, 5 with 25 June 3, 000 young; remainder roosting non-breeding population (POBSP, 1963). 1964 13-14 Mar. 300-400 Mostly nocturnal; no nests (BSFW, 1964a; POBSP, 1964b). 16-19 Aug. 550 Adults; 16 breeders: 6 nests with eggs and 2 with medium-sized young (POBSP, 1964a). 16 Sept. 800-900 Roosting population that became larger at night; no nests (BSFW, 1964b; POBSP, 1964c). 15-19 Mar. 300-400 Mostly roosting birds; 12-14 breeders: 6-7 nests with eggs (POBSP, 1965b). 21-22 Mar. 0 (BSFW, 1965; POBSP, 1965a). iP Aor. - Not mentioned in notes (BSFW, 1966a). 20-26 Sept. 1, 000 Ca. 400 by day increasing to 1,000 at night; no eggs or young (BSFW, 1966b). 25-27 Sept. 1, 000 All non-breeding, except 2 breeders with 1 large chick (POBSP, 1966). 21-23 Mar. 5 Roosting at night; occasional during day; no nests (BSFW, 1967a; POBSP, 19674). 28, 30 May- 1, 000+ Nocturnally; only 200 diurnally; 22 breed- 1 June ers: 11 nests, 4 with eggs and 7 with small downy young (POBSP, 1967b). 28-30 Aug. 3,000 Mostly roosting non-breeders; 8-20 breeders: 2 nests with eggs and 2 with small downy young (POBSP, 1967a). 22-2 Mar. 25 16-20 breeders: 8-10 nests, 1 with fresh egg, rest recently built but empty (BSFW, 1968; POBSP, 1968). 10-12 Feb. 50 Nocturnal, no evidence of nesting (BSFW, 1969a). 31 Mar.- - Not listed in report (BSFW, 1969b). 2 Apr. 234 Table 105. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1969 26-31 May 1,600 Nocturnally; 50 breeders; 25 nests counted; of 22 nests sampled: 7 (32%) contained eggs and 15 (68%) contained small downy chicks (BSFW, 1969c). 10-19 Sept. IS 320 (BSFW, 19694). WHITE TERN Gygis alba Status Irregular breeding species; present irregularly year-round; likely to occur on most islands, known to nest only at Southeast. Maximum POBSP and BSFW population estimate 10 in September 1969. Observations A single White Tern at North Island in March 1913 was the first record at Pearl and Hermes Reef (Bailey, 1956: 32). Rice (ms. a) observed 25 on, and flying near, the northeast reef on an aerial survey 14 October 1957. He suggested they probably nest there but no other observers have suggested this nor have such numbers been seen subsequently. POBSP and BSFW personnel recorded small numbers each year from 1963 to 1969 (Table 107); nesting was first recorded at Southeast in 1969. Annual Cycle White Terns were observed during all survey periods except February (see Fig. 58). Few birds were seen in March; most birds were seen in late spring and summer. BSFW personnel found a White Tern incubating an egg at Southeast on 26 May 1969; although this is the first breeding record on the atoll, per- haps it has bred in other years and has been overlooked because of its extremely small population. Ecological Distribution White Terns have been recorded as flying over or roosting on Grass, Little North, North, Seal, and Southeast Islands. Wetmore (ms.) was the first to observe White Terns on Southeast; all other observations were by POBSP and BSFW personnel. Recorded populations have ranged from 1 to 7 birds. A pair nested in May 1969; its egg, pre- ecariously placed on a small piece of raised coral near the southwest shore, was subsequently destroyed by a wave. 259) Idd 12 *TeI0L sunox FTOPYV T2301 ~m co >Te10, Sunoz 10py Y320N dz >Te10L 0 DUNO 12 qINpY qsBayynos T®4OL €96T €96T sseTo pueTst IL ON €96T aune * LEW -o5y KeaAIng jo potazed dsd0d dsdod MISE dsd0d dSsdod + topued ZOT fs) 3 + Ol+ HOA]! MO Ne) qd re O g ‘ 4 = oat \O Or oS ap 5 have been recaptured. The finches frequent the tall Eragrostis west of the central depression, as well as the low-growing Solanum and the Brassica. Most nests are in lperger (1970: 38) thought the population numbered between 75 and 100 birds. 239 Eragrostis clumps and a few are in the moderately dense Solanum. Food con- sists of seeds of Setaria, Tribulus, Eragrostis, Boerhavia, and Sonchus. Banding and Movements BSFW personnel banded 220 Laysan Finches at Southeast Island: 109 adults in March 1967 (original introduction), 17 young in March 1968, 34 adults in February 1969, 19 adults and 6 young in March and April 1969, and 12 adults and 23 young in May 1969. Of these, 25 have been recaptured on the atoll (Table 16). These data are being analyzed by BSFW personnel. Table 108. Observations of Laysan Finches at Southeast Island Population Date of Survey Estimate Remarks and References 1967 21-23 Mar. 110 59 males and 51 females, all but one banded, introduced from Laysan (BSFW, 1967a; POBSP, 1967d). 28, 30 May- 30-50 Successfully adapted to local conditions; 1 June 16 nests counted in Eragrostis clumps: 3 old, 9 recently completed, 1 with 4 eggs, 1 with 3 eggs, and 2 with 2 eggs (POBSP, 1967b). 1968 22-24 Mar. 27 h* Q nests: 4 with 2-3 eggs, 3 recently com- pleted, and 2 old (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 100 5 nests: 1 with 2 downy young and an egg, 1 with 2 eggs, 1 new but empty, and 2 old (BSFW, 1969a). 31 Mar.- 165 13 nests: 2 with 1 egg, 1 with 2 eggs, 1 2 Apr. with 3 eggs, 3 with 4 eggs, 2 with 1 young, 2 with 2 young, and 2 with 3 young (BSFW, 1969b). 26-31 May 2 No estimate taken; 17 active nests: 1 with 1 egg, 1 with 2 eggs, 1 with 1 egg and 1 young, 3 with 1 young, 6 with 2 young, 4 with 3 young, and 1 empty; 7 old nests (BSFW, 1969c). * This estimate is possibly high. 20 MAMMALS Besides man, four mammalian species have been recorded from Pearl and Hermes Reef. The Hawaiian Monk Seal is a resident breeding species found on most islands. The Hawaiian Spinner Dolphin, a new record for the atoll, and the Bottle-nosed Dolphin are visitors to the lagoon waters. The Euro- pean Rabbit, introduced between 1913 and 1916, was exterminated in 1928. No mice, cats, or dogs are known to have occurred. For descriptions of these species, the reader is referred to Tomich (1969). HAWAIIAN SPINNER DOLPHIN Stenella roseiventris Status Visitor; one definite record. Observations POBSP and BSFW personnel sighted 8 dolphins offshore Southeast Island on 24 March 1968. Photographs by Kosaka were later examined by Brownell who identified them as Stenella roseiventris. This dolphin is common in Hawaiian waters (Tomich, 1969: LLY, but has not been previously recorded from Pearl and Hermes Reef. BOTTLE-NOSED DOLPHIN Tursiops truncatus Status Visitor; two definite records. Observations On 14 March 1958 Rice (1960b: 407) observed from the air about 30 Bottle-nosed Dolphins in shallow water just outside the southwest reef. His data led Tomich (1969: 46) to suggest that this species is regularly associated with shallow waters from French Frigate Shoals to Kure Atoll. Hershkovitz (1966: 48-53) thought the dolphins Rice sighted belonged to the race Tursiops truncatus aduncus. Bailey (1918: 398) first recorded porpoises on 15 March 1913; he wrote that "a school of porpoises played off our bow and came in so close as almost to splash in the boat as they cut water.” On 22 March 1956 POFI personnel sighted "3 large bottle nose porpoise in front of [the] bow” of the USFW John R. Manning on the western side of the reef (POFI, 1956a). POBSP and BSFW personnel noted 15 to 20 unidentified porpoises inside the lagoon between Southeast and Grass Islands 14 March 1964. Lewis sighted eran 40 small unidentified porpoises inside the lagoon near Southeast as he approached the island in a helicopter on 25 September 1966. HAWAIIAN MONK SEAL Monachus schauinslandi Status Common resident breeder; present year-round; occurs and breeds on most islands. Maximum recent POBSP and BSFW population estimate 195 in February and March 1963. Observations Hawaiian Monk Seals were first recorded by Morrell (1832: 217-218) 8-11 July 1825, who wrote "the sea-elephant and sea-leopard resort to the islands in the summer season, in large rookeries, and the former are perfectly tame." Osbun, who visited in August 1850, wrote of killing 10 or 12 seal for their livers and hearts; he also noted their tameness and estimated their weight at 500 pounds (Kemble, 1966: 155). Seals were next recorded in 1857 by Paty (1857: 2-3) who noted that the "islets...seemed to abound with...seals." Brooks, who visited Pearl and Hermes and other Northwestern Hawaiian Islands in July 1859, may have found seals here; he returned to Honolulu with a cargo of 240 barrels of seal oil and 1,500 seal skins (Anon., 1859a, 1859b). Brooks (1960: 502) failed, however, to mention seals in his subsequent description of the atoll, though he noted birds and turtles (see also Kenyon and Rice, 1959: 215). Frear's notation of 35 seals and a newborn pup in December 1912 prompted Atkinson and Bryan (1913: 1050-1051) to write that "Pearl and Hermes Reef seem [sic] to have been the place where they were most abundant.” In March 1913 Willett and Bailey found seals with pups on North Island and a nearby sandbar (Bailey, 1918: 399; 1956: 30), and Elschner (1915: 60) reported many there in September 1914. Munter (ms.) found 30 or 35, including several pups, in February 1916. Wetmore (ms.) estimated 125 on Grass, Seal and Southeast Islands in April 1923, while Galtsoff (1933: 19) counted 68 in the lagoon in summer 1930. Bailey (1952: 16) estimated over 100 seals when he flew over the atoll in May 1949. A POFI survey in June 1950 found 100 seals during the day and noted that females had pups that were a few days or weeks old (POFI, 1950); Brock esti- mated 180 seals (Bailey, 1952: 25). Dumont and Neff (Svihla, 1959: 227) reported 96 in November 1954. Kenyon and Rice (1959: 221), on aerial surveys, counted 257 adults and subadults on 17 December 1956, and counted 33 pups on 15 April and 14 May 1957. Using the same type surveys, Rice (1960a: 377) counted 286 adults and subadults and 52 pups in spring 1958. Hawaiian Monk Seals have been counted, sexed, and tagged on most BSFW, POBSP, and HDFG survey trips during the 1960's. Tables 109 to 115 present all seal observation data for Pearl and Hermes. eke Annual Cycle Hawaiian Monk Seals are found at Pearl and Hermes year-round. Kenyon and Rice (1959: 236) found that pups are born throughout the entire Hawaiian chain from late December until the end of June, with most born from mid- March through May. Bailey (1952: 5) recorded a newborn pup at Pearl and Hermes in late December 1912. Only one pup was found in two recent Tee surveys. The maximum March countl was 5 pups, while the highest June count was 28. The highest August countl was 17, and the highest September count was 13. Recent pupulation counts! produce a high of 195 in late February and early March, while five March counts] average 140. Two August counts! num- bered 116 and 115; and two September countsl gave 105 and 119. The average population count! from 11 recent surveys was 128. Kenyon (in prep.) recently pointed out, using data from Midway Atoll, Kure Atoll, French Frigate Shoals, and Pearl and Hermes Reef (1968 data only), that Hawaiian Monk Seal populations will not tolerate repeated disturbance by man. He found that the Midway population has almost disappeared and the Kure population has decreased. The Pearl and Hermes Reef data obtained over the last seven years seem to indicate a declining population, both in total size and in number of young produced. The 1963 February population contained 195 and produced at least 28 pups by June; the 1969 February population contained 152 adults and produced at least 17 pups by late May. Ecological Distribution Hawaiian Monk Seals have been recorded from all islands and sandbars. Breeding occurs on all islands except Planetree and Sand. Movement from islet to islet within the atoll is common (see Tagging and Movement section). Females with newborn pups, however, do not usually move from islet to islet. Seals are found predominately on the beaches, but may be found in the vegetated areas. Grass Island: Hawaiian Monk Seals have been recorded on at least 16 POBSP and BSFW surveys (Table 109). The average population was 18 seals, and ranged from 9 to 39. Six pups are known to have been born since 1963. Kitte Island: BSFW and POBSP personnel recorded seals on 14 surveys (Table Tio), The population count averaged 24, and ranged from 2 to 43. Eleven pups are known to have been born since 1963. Little North Island: Seals were noted by POBSP and BSFW personnel on 10: surveys (Table 111). The population count averaged 19 and ranged from 4+ to 29. In all, 25 pups are known to have been born since 1963. lusing only surveys where counts were taken on six or more islands. O43 North Island: POBSP and BSFW personnel observed seals on 13 surveys (Table 112). The average population count of 28 was higher than on any of the other islands in the atoll; the population has ranged from 15 to 53. The number of pups produced since 1963 is 24, which is also a high for the atoll. Pup counts for both Little North and North Islands are high since they are isolated and not greatly affected by human disturbance. Seal Island: Seals were recorded on 15 surveys by POBSP and BSFW per- sonnel (Table 113). The average population count was 14, and ranged from 9 to 25. Seventeen pups are knwon to have been born since 1963. Southeast Island: Seals were recorded on 19 POBSP and BSFW surveys (Table way). Southeast, the atoll's largest island, had the highest island population, 58+. However, because of human disturbance (most campsites were here), the average population was only 26. Average winter and spring counts are higher (29 seals) than summer and fall counts (21 seals). Only 10 pups were observed from 1963 through 1969. Seals are most common along the lagoon beach (Fig. 79). Some occur on the seaward beach and a few have been found in the vegetated areas (Fig. 80) and the tidal pools. Other Islands: Small numbers (average 5) were noted on Bird, Planetree, and Sand Islands during visits by POBSP and BSFW personnel (Table 115). A single pup is known from Bird. Tagging and Movement Since 1963 POBSP and BSFW tagged at least 197 seals, including 94 in 1963, 33 in 1966, 33 in 1967, 8 in 1968, and 29 in 1969. Tagging showed that movement within the atoll was common. Seven seals tagged on Pearl and Hermes have been recaptured on Laysan, 4 on Lisianski, 2 on French Frigate Shoals, and 1 on Kure. These data are being analyzed by BSFW personnel. Specimens POBSP: USNM 334576, dead newborn pup, alcoholic, collected on Grass Island 5 March 1963 by Wirtz; USNM 334577, adult skull and skeleton, col- lected on beach at Southeast Island 6 March 1963 by Wirtz. Non-POBSP: USNM 181251, ? pup, collected on North Island 15 March 1913 by Willett for the U.S. Bureau of Biological Survey; USNM 243843-4, skin and skull, collected 28 April 1923 by Wetmore; USNM 243845, skull, collected 28 April 1923 by Wetmore. Table 109. Observations of Hawaiian Monk Seals at Grass Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1950 27 June present (POFI, 1950). 2k Table 109. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1951 14 May present Adults with pups (POFI, 1951; Svihla, 1959: 227). 1956 22 Mar. 18 Adults (POFI, 1956a). 26 May iG Adults (POFI, 1956b). 1963 5 Mar. 39 16 tagged--13 adults: 5 o, 8 9; 3 sub- adults: 20, 19; 8 repeats; 13 adults and 1 subadult untagged; 1 dead pup (POBSP, 19644). 26-27 June 8-10 Daily (POBSP, 1963). 1964 14 Mar. 27 15 adults: 30, 3 2, 9 unk*; 12 subadults: 9, 3 2 (BSFW, 1964a; POBSP, 1964b). 18 Aug. 20+ Adults (POBSP, 1964a). 1965 19 Mar. 29 Adults (POBSP, 1965b). 22 Mar. 27 19 adults: 70, 89, 4 unk; 8 subadults: 20, 29, 2 unk (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 12 7 adults, 4 subadults, 1 pup (BSFW, 1966b). 1967 17 Mar. 19 Includes counts for Bird, Planetree, and Seal Islands (BSFW, 1967d). 22 Mar. 9 Adults (BSFW, 1967a; POBSP, 1967d). 31 May 13 12 adults: 40, 4 9, 4 unk; 1 unk year- ling (POBSP, 1967c). 29 Aug. 19 16 adults: 80, 7, 1 unk; 3 pups (POBSP, 1967a). 28 Sept. 10 9 adults, 1 yearling (BSFW, 1967c). 1968 24 Mar. 5) 12 adults® 6.6, 6 9; subadults: lcs aug. 1 o yearling (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 18 17 unk adults, 1 unk subadult (BSFW, 1969a). 26 May 8 7 adults: 29, 5 unk; 1 ? pup (BSFW, 1969c). 11 Sept. 12 li adults: 3 6) °2°9/ 6 unk; 19subaduls x unk = unknown. (BSFW, 1969d). | ts ahs Table 110. Observations of Hawaiian Monk Seals at Kittery Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1956 22 Mar. 30 Adults (POFI, 1956a). 26 May U5) Adults, no pups (POFI, 1956b). 1963 5 Mar. 32 10 tagged--7 adults: 5 oh, 293 3 subadults: 1c, 29 (POBSP, 1964d). 1964 14 Mar. 38 25 adults: 11 oc, 9 9, 5 unk; 13 subadults: ho, 5 9, 4 unk (BSFW, 1964a; POBSP, 1964b). 18 Aug. 2 Adults (POBSP, 1964a). 1965 18 Mar. 22 Adults (POBSP, 1965b). 22 Mar. 35 26 adults: 15 oh, 10 9, 1 unk; 9 subadults: 26, 5 2, 2 unk (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. el 16 adults, 5 subadults (BSFW, 1966b). 1967 22 Mar. 43 4] adults, 1 subadult, 1 yearling (BSFW, 1967a; POBSP, 19674). 31 May 20 17 adults: 140, 19, 2 unk; 3 yearlings: 2d, 1 (POBSP, 1967c). 29 Aug. 26 16 adults, 10 pups (POBSP, 1967a). 28 Sept. 18 16 adults, 1 subadult, 1 yearling (BSFW, 1967c). 1968 2 Mar. 28 21 adults: 160, 4 9, 1 unk; 7 subadults: 5 oh, 22 (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 10 9 adults: 4 oh, 3 9, 2 unk; 1c pup (BSFW, 1969a). 26 May 23 22 adults: 19, 21 unk; 1 9 pup (BSFW, 1969c). 11 Sept. A183 T2vadults:;"3 co, i 9, 8 unk; 1 o subadult (BSFW, 1969d). Table 111. Observations of Hawaiian Monk Seals at Little North Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1963 23, 25 June 13 Pups (POBSP, 1963). 46 Table 111. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1964 19 Aug. 25 -28 Adults (POBSP, 1964a). 17 Sept. 19 9 adults, 7 yearlings, 3 unk (BSFW, 1964b; POBSP, 1964c). 1965 18 Mar. 5 23 adults, 2 pups (POBSP, 1965b). 1966 22 Sept. Tate (BSFW, 1966b). 1967 17 Mar. 4 Aerial count (BSFW, 19674). 29 Aug. 20 12 adults, 4 yearlings, 4 pups (POBSP, 1967a). 1969 10-12 Feb. 29 Unclassified (BSFW, 1969a). 26 May 19 13 adults -fal2) Oo. unk sAGe pupsi:norCemmlane (BSFW, 1969c). 12 Sept. 21 17 adults: 4 oh, 29, 11 unk; 4) pups-eleee 3 unk (BSFW, 1969d). Table 112. Observations of Hawaiian Monk Seals at North Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1913 15 Mar. 50-60 About 15-20 2 with young, few co, several half-grown (Bailey, 1956: 30). 1914 Sept. many "Sea lions" (Elschner, 1915: 60). 1950 27 June present (POFI, 1950). 1956 25 May 31 18 adults, 13 pups; several born previous night (POFI, 1956b). 1963 6 Mar. 27 4 tagged--2 adults: 1o, 19; 2@ sub- adults; 1 dead pup (POBSP, 1964d). 23-25 June 13-15 Pups (POBSP, 1963). 1964 19-20 Aug. 22 20 adults, 2 pups (POBSP, 1964a). Ui SS eGc 36 14 adults: 16, 3 9, 10 unk; 19 yearlings: 6 o, 4 9, 9 unk; 3 unclassified (BSFW, 1964b; ' POBSP, 1964c). 247 Table 112. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1965 17-18 Mar. 53 52 adults, 1 dead pup (POBSP, 1965b). 1966 22 Sept. 26 (BSFW, 1966b). 1967 16-17 Mar. 30 11 adults, 19 subadults; 2 recaptured (BSFW, 1967d). 29-30 Aug. al 15 adults, 6 subadults (POBSP, 1967a). 27 Sept. 29 21 adults, 2 subadults, 6 yearlings (BSFW, 1967c). 1969 10-12 Feb. yy 33 unclassified on North; 11 unclassified on nearby sandbars (BSFW, 1969a). 31 Mar. 27 ipaamlts: 30, 72. © unk, Iedead 955 subadults: 10, 192, 3 unk; 1 yearling; 4 pups (BSFW, 1969b). 26 May ay 9 adults: 76, 2 unk; 2 unk subadults; 6 pups: 20, 3 9, 1 unk (BSFW, 1969c). 12 Sept. 21 13 adults: 606, 4 9, 3 unk; 2 stbadults: IG, L283) ey joujosis 3) ecluittsg io, 2 walk on nearby sandbars (BSFW, 1969d). Table 113. Observations of Hawaiian Monk Seals at Seal Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1956 22-2) Mar. 16 12 adults, 3 pups, 1 dead pup (POFI, 1956a). 26 May (2\n. Bei Ca. 20 adults, 15 pups (POFI, 1956b). 1963 5 Mar. 19 2 subadult 2 tagged; 5 repeats (POBSP, 19644). 26 June ast (POBSP, 1963). 1964 14 Mar. aly 8 adults: 4 of, 19, 3 unk; 9 subadults: 4 ot, 5 2 (BSFW, 1964a; POBSP, 1964b). 18 Aug. 15-20 Adults (POBSP, 1964a). 1965 18-19 Mar. 20 18 adults, 2 dead pups (POBSP, 1965b). 248 Table 113. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1965 22 Mar. 10 2 adults, 5 subadults, 2 pups, 1 unk (BSFW, 1965; POBSP, 1965a). 1966 21 Sept. 8 3 adults, 1 subadult, 4 pups (BSFW, 1966b). 1967 22 Mar. h 3 adults, 1 pup (BSFW, 1967a; POBSP, 19674). 31 May 18 liyadulits: 26, 7<2, 2 unk; 1.° subaduiltoe 6 pups: 3 9, 3 unk; 5 pups tagged on the 28th (POBSP, 1967c). 29 Aug. 7 4 adults: 30, 19, 3 pups (POBSP, 1967a). 28 Sept. 105) 9 adults, 6 subadults (BSFW, 1967c). 1968 24 Mar. 16 Quadults: 6 6, 3495.3 subadulitss, aac, ae ee 2 yearlings: 1c, 193; 29 pups (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 9 8 adults: 10, 3 9, 4 unk; 1 ? subadult (BSFW, 1969a). 26 May 10 Ssaduilltsize. 7, Ps: lyunks 2) pups syliac eae (BSFW, 1969c). 11 Sept. 12 9 adults: 1o, 4 9, 4 unk; 3 pups: 1d, 2? (BSFW, 1969da). Table 114. Observations of Hawaiian Monk Seals at Southeast Island Population Date of Survey Estimate Breeding Status, Remarks, and References 1950 27 June present (POFI, 1950). 1956 22 Mar. O No adults seen (POFI, 1956a). 26 May 20 Adults, no pups (POFI, 1956b). 1963 26 Feb.- 58+ 58 tagged--32 adults: 120, 18 9, 2 unk; 8 Mar. 26 yearlings: 14 oc, 129 (POBSP, 1964a). 18-23, 12 Counted on 18th; 6-8 other days (POBSP, 25 June 1963). 1964 13-14 Mar. 22 Mvadults:), .5 6, 509; 11 subaduitss are cs 2? (BSFW, 1964a; POBSP, 1964b). 24g- Table 114. (continued) Population Date of Survey Estimate Breeding Status, Remarks, and References 1964 16-19 Aug. 15 Adults (POBSP, 1964a). 16 Sept. 33 16 adults, 16 subadults (BSFW, 1964b; POBSP, 1964). 1965 15-19 Mar. ail Adults (POBSP, 1965b). 21-22 Mar. 4s 24 adults: 100, 14 9; 18 subadults: 54, 13 9; 1 unk pup; 2 unclassified (BSFW, 1965; POBSP, 1965a). 1966 1) Apr. 18 4 adults, 14 subadults (BSFW, 1966a). 20-26 Sept. 27 8 adults, 14 subadults, 5 pups counted on 20th; 33 tagged (Kenyon, in prep.; BSFW, 1966b). 1967 17 Mar. 4 8 adults, 6 subadults, 1 dead (BSFW, 1967d). 21-23 Mar. oh 15 adults, 1 subadult, 8 yearlings (BSFW, 1967a; POBSP, 1967d). 28-30 May- 21 IB Vadulte- 23 Co, 7 9. SB unk; 77 yearlings: 1 June 1o, 29, 4 unk; 1 @ pup; counted on 28 May (POBSP, 1967c). 28-30 Aug. iMG) 12 adults, 3 yearlings (POBSP, 1967a). 27-29 Sept. Tinh 10 adults, 1 yearling (BSFW, 1967c). 1968 22-2) Mar. 37 19 adults: llc, 7 9, 1 unk; 11 subadults: 6c, 59; 7 yearlings: 4c, 3 2 (BSFW, 1968; POBSP, 1968). 1969 10-12 Feb. 23 15 adults: 506, 8 9, 2 unk; 3 unk subadults; 5 unclassified (BSFW, 1969a). 31 Mar.- Hy 17 adults: 66, 4 9, 5 unk, 2 dead; 20 sub- 2 Apr. adults: 306, 99, 8 unk (BSFW, 1969b). 26-31 May 16 ieedulie io. 5n9.) 5 unks 2 yank! subadults ; 1? pup (BSFW, 1969c). 10-19 Sept. Bub 18 adults: 4 o, 5 9, 9 unk; 10 subadults: 29, 8 unk; 3 unk pups (BSFW, 1969d). 19. 80. Yearling Hawaiian Monk Seal on lagoon beach at Southeast Island, March 1963. POBSP photograph by W.O. Wirtz, II. Adult Hawaiian Monk Seals basking in the sun on the vegetated portion at Southeast Island, 28 August 1967. POBSP photograph lf7 Della lnlostat Table 115. Pearl and Hermes Reef Population 251 Observations of Hawaiian Monk Seals on other islands at Date of Survey Island Estimate Breeding Status, Remarks, and References 1825 1850 1857 1950 1956 1963 1964 1965 8-11 July ? 2 Large rookeries (Morrell, 1832: 217-218). Lda Greg ? few Killed 10 or 12 for food (Kemble, 1966: 155). 19-20 May 2 ? Islets abound with seal (Paty, 1857: 2-3). 27 June Bird present (POFI, 1950). 27 June Sand present (POFI, 1950). 22 Mar. Bird 11 Adults (POFI, 1956a). 22 Mar. Sand 6 Adults (POFI, 1956a). 26 May Bird 5 Adults (POFI, 1956b). 5 Mar. Bird 13 3 tagged: 2c adults, 1c subadult (POBSP, 196d). 5 Mar. Sand if 1 unk adult tagged (POBSP, 1964d). 14 Mar. Bird 3 Subadults: 19, 2 unk (BSFW, 1964a; POBSP, 1964b). 14 Mar. Planetree T 6 adults: 4 do, 19, 1 unk; 1c subadult (BSFW, 1964a; POBSP, 1964b). 14 Mar. Sand 7 Peagmihess Jeo. Ose subadultss 3.01 2 unk (BSFW, 1964a; POBSP, 1964b). 18 Aug. Bird 5 Adults (POBSP, 1964a). 18 Aug. Planetree 3 Adults (POBSP, 1964a). 18 Aug. Sand a) Adult (POBSP, 1964a). 22 Mar. Bird 8 5 adults, 3 subadults (BSFW, 1965; POBSP, 1965a). 22 Mar. Planetree a4 6 adults: 3 do, 1 9, 2 unk; 1 subadult (BSFW, 1965; POBSP, 1965a). 22 Mar. Sand 6 4 adults: 36, 19; 29? subadults (BSFW, 1965; POBSP, 1965a). 252 Table 115. (continued) Population Date of Survey Island Estimate Breeding Status, Remarks, and References 1967 31 May Bird 7 5 adults: 2c, 3 unk; 2 subadults: 1c, 1 2 (POBSP, 1967c). 29 Aug. Bird 6 5 adults: 4 9, 1 unk; 1 pup (POBSP, 1967a). 29 Aug. Planetree al Adult (POBSP, 1967a). 1969 10-12 Bird, 20 Unclassified (BSFW, 1969a). Feb. Planetree and Sand 26 May Bird, 7 Adults (BSFW, 1969c). Planetree and Sand 11 Sept. Bird 4 3 adults: 20, 1 unk; 1 9 subadult (BSW, 1969d). EUROPEAN RABBIT Oryctolagus cuniculus Status Introduced prior to 1916; exterminated in 1928. Observations Munter (ms.) lst lieutenant aboard the USCGC Thetis, first recordedl rabbits on Southeast Island on 4 February 1916. Since they were of the do- mestic variety, Munter believed they had "been brought to the island from Laysan or Lisiansky [sic]...by fishermen."2 This was suggested when the remains of a crude shelter which Munter believed to be less than a year and a half old and which probably was built by Japanese 3 fishermen, were found. Munter caught four of the rabbits. Jouring Jacobs' 21 January 1910 visit to Pearl and Hermes, no rabbits were noted. Salisbury also found none there 15 March 1913 (U.S. Nat. Archives, R.G. 26, letter from Jacobs to Secretary of Treasury dated 2 February 1910; R.G. 22, letter from Salisbury to Palmer dated 20 March 1913). 2rabbits were introduced to Laysan about 1903 by Max Schlemmer to start a rabbit canning business (Dill and Bryan, 1912: 9-10). 3 Three Japanese were rescued from Pearl and Hermes in January 1909 by the schooner Florence Ward. They had been left there 7 July 1908 with a 253 Tanager Expedition personnel next recorded "a number of rabbits and the remains of a camp" on Southeast in April 1923 (Bryan et al., 1926: 6). On 26 April, Wetmore (ms.) wrote: "When we camped we were astonished to find rabbit dung scattered about and later found the animals fairly common in the bunch grass. Apparently they had been introduced three or four years ago by the crew that had camped there. Indications were that the animals had come from Laysan as they were of the same size and mixed colors. I shot a considerable number and Dr. Wilson more so that we killed 25 before I was out of shells." On 27 April he noted "no sign of rabbits on either Grass or Seal Islands." Rabbits were hunted on Southeast on the 28th; afterward Wetmore wrote "the number killed today and on the 26th made a total of 90 rabbits." He estimated "that perhaps 30 remain," and observed they "were wilder than on Laysan and usually ran as soon as they saw us.” The Tanager personnel left Southeast on the 28th (Wetmore, ms.), leav- ing some 30 live rabbits. Additional Tanager personnel returned on 17 May; prior to leaving on the 19th they had "killed all but one of the rabbits seen" (Gregory, 1924: 21). Two of three rabbits on Southeast were shot by G.P. Wilder on 2 May 1924. Rabbits (20?) were subsequently collected in May 1927 and returned in several crates to Honolulu by Captain Peterson of the Lanikai. During April 1928 the Lanikai crew shot three rabbits on Southeast. In June or July 1928 a diligent search was made by Captain Anderson of the Lanikat, but none was found. None has been found since (U.S. Nat. Archives, R.G. 22, letters from Wilder to Nelson dated 19 April 1926, and Wilder to Goldman dated 20 July 1928; Anon., 1927). SUMMARY Pearl and Hermes Reef is an uninhabited atoll containing nine islands in the Northwestern Hawaiian Islands. Wildlife is protected as the atoll is part of the Hawaiian Islands National Wildlife Refuge. Personnel of the Pacific Ocean Biological Survey Program (POBSP), Smithsonian Institution, Washington, D.C., made 12 biological survey trips to the atoll from February 1963 to March 1968. Data from these visits, as well as from visits through 1969 by Bureau of Sport Fisheries and Wildlife (BSFW) personnel, Hawaiian Division of Fish and Game personnel, Pacific Ocean Fisheries Investigation personnel, and from all previously published litera- ture are summarized and discussed. Emphasis is placed on the vascular flora and the vertebrate terrestrial fauna. month's supply of food by a Japanese schooner; the vessel was supposed to have been lost in a storm (Thrum, 1909: 176). Japanese feather poachers were on Laysan and Lisianski at this same time (Ely and Clapp, in press); the three on Pearl and Hermes were probably part of that group. It is most likely that rabbits were introduced during this period. 254 Twenty-five species of vascular plants, representing 17 families, have been observed or collected from five islands. Only one breeding reptile species is known. Since 1964 BSFW personnel have tagged at least 137 Black Sea Turtles; 46 of these were recaptured on the atoll, and one was captured elsewhere in the Hawaiian Islands. In addition, two tagged elsewhere were captured on the atoll. Of the 37 species of birds recorded, 17 are resident seabirds, 5 are regular migrant shorebirds, and 15 are vagrant, accidental, or introduced. In all, 68 specimens of 26 species have been collected. Before the POBSP, 37 specimens of 16 species were collected. POBSP personnel collected 31 specimens of 18 species, 14 of which represent new specimen records. An additional three species represent new sight records, and four represent new breeding records. Of the 31,661 birds of 22 species banded on the atoll, 1,790 of 18 species were recaptured on the atoll, and 211 of 13 species were captured at other Pacific localities. In addition, 70 birds of 12 species banded at other Pacific localities were captured on the atoll. Beside man, four mammalian species have been re- corded. Since 1963, POBSP and BSFW personnel have tagged at least 197 Hawaiian Monk Seals; many recaptures are known, and 14 inter-atoll move- ments exist between other Hawaiian Islands. ACKNOWLEDGMENTS Acknowledgment is first and foremost made to Eugene Kridler, Refuge Manager, Hawaiian Islands National Wildlife Refuge, Kailua, Oahu, and Michio Takata, Director, Hawaiian Division of Fish and Game, for granting permission for POBSP personnel to visit Pearl and Hermes Reef and for pro- viding unpublished field notes and preliminary reports of U.S. Fish and Wildlife Service and State of Hawaii survey trips to the atoll. We would also like to thank the U.S. Navy and U.S. Coast Guard for full cooperation and assistance in providing transportation and in gathering scientific and historical data. We are very grateful for the full cooperation and assistance in obtain- ing POBSP band recapture information from the Bird Banding Center, Yamashina Institute for Ornithology, Tokyo, Japan, and The Centre of Ringing and Marking of Birds and Terrestrial Mammals, USSR Academy of Sciences, Zoological Insti- tute, Moscow, USSR, as well as the Bird Banding Laboratory, Migratory Bird Populations Station, Bureau of Sport Fisheries and Wildlife, Laurel, Maryland. It is difficult to give adequate acknowledgment to all individuals who helped gather historical and scientific data. Those who deserve special q acknowledgment include: Ronald R. Amerson, U.S. Navy, Honolulu, Hawaii; Edwin H. Bryan, Jr., Pacific Scientific Information Center, Bernice P. Bishop Museum, Honolulu, Hawaii; Archie F. Carr, University of Florida, Gainesville, Florida; Jean Dabagh, Hawaii State Archives, Honolulu, Hawaii; Paul S. Galtsoff, Woods Hole, Massachusetts; B.C, Jones, Bureau of Commercial Fisheries, Hono- lulu, Hawaii; Chandler §. Robbins, Bureau of Sport Fisheries and Wildlife, Patuxent, Maryland; Harry Schwartz, Modern Military History Division, U.S. National Archives, Washington, D.C.; Alexander Wetmore, Smithsonian Institu- tion, Washington, D.C.; and Virginia Frear Wild, Honolulu, Hawaii. We are 255 especially grateful to Kay B. Forck, Redwood City, California, for as- sistance in assemblage of the history section. We are deeply indebted to the following POBSP personnel who have contributed to this manuscript: Kenneth E. Amerman, Alan H. Anderson, Winston E. Banko, F. Allen Blagden, David L. Burckhalter, Robert L. DeLong, Charles A. Ely, Robert R. Fleet, C. Douglas Hackman, the late J. Vincent Hoeman, T. James Lewis, Charles R. Long, Peter Marshall, Robert W. McFarlane, Richard W. Merrill, Fred C. Sibley, Dennis L. Stadel, F. Christian Thompson, Robert Tuxson, J. Douglas Whitman, George S. Wislocki, and Paul W. Woodward who spent long hours collecting field data; Anne Keenan Poulson who care- fully drafted the maps and charts; Mae H. Esterline who gave editorial advice on early drafts of the manuscript; Jane P. Church who edited the final draft and attended to all details of producing the final copy; and Philip S. Humphrey who encouraged and supervised the entire POBSP field and writing effort. We also wish to thank Robert L. Brownell, POBSP; F. Raymond Fosberg, Smithsonian Institution; Eugene Kridler, BSFW; Robert M. Mengel, University of Kansas; Philip C. Shelton, POBSP; and George E. Watson, Smithsonian Insti- tution, for critically reading various sections of the manuscript. The camera copy was typed by Barbara B. Anderson with funding through a contract with the Bureau of Sport Fisheries and Wildlife, Department of Interior (con- tract number 14-16-008-596, February 3, 1971). 256 LITERATURE CITED Air Weather Service [MATS] Climatic Center, USAF. Pearl and Hermes Reef, Hawaii. USN. Summaries 1953-1963. Asheville, North Carolina. Aldrich, J.W., C.S. Robbins, and D.W. Rice. (ms.). Investigations of bird hazards to aircraft of Midway Island - 1956. 4 February 1957 Progress Report. Smithsonian Institution, Washington. 11 pp. Alexander, W.B. et al. 1965. The families and genera of the petrels and their names. Ibis 107: 401-405. Allen, G.M. 1942. Extinct and vanishing mammals of the Western Hemisphere. Amer. Comm, Int. Wildlife Prot., Spec. Publ. 11: 1-620. American Ornithologists’ Union. 1957. Checklist of North American birds. 5th ed. Lord Baltimore Press, Baltimore. ' ) i" Amerson, A.B., Jr. 1968. Tick distribution in the Central Pacific as in- fluenced by sea bird movement. J. Med. Ent. 5: 332-339. a le ----1971. The natural history of French Frigate Shoals, Northwestern Hawaiian Islands. Atoll Res. Bull. 150: 1-383. Amerson, A.B., Jr., and K.C. Emerson. 1971. Records of Mallophaga from Pacific birds. Atoll Res. Bull. 146: 1-30. ; Anderson, W.G. 1954. Notes on food habits of sea birds of the Pacific. Elepaio 14: 80-84. Anonymous. 1859a. The Friend, 13 August 1859. ----1859b. Polynesian, August 1859. ----1868. North Pacific Ocean. Report of Captain Wm. Reynolds, U.S. Ship "Lackawanna," on Midway or Brooks Islands, Ocean Islands, and Pearl and Hermes Reefs--30th September 1867. Naut. Mag. 37: 269-275. ----1927. Fishing and shell hunting expedition returns from Pearl and Hermes Reef. Honolulu Advertiser, 13 May 1927, 1: 1-6, 2; 3-4. ----1939. Manure. Sales Builder, January 1939, 12(1): 2-22. Atkinson, A.L.C., and W.A. Bryan. 1913. A rare seal. N.Y. Zool. Soc. i Bull. 16: 1050-1051. - Attems, C.G. 1938. Myriopods of Hawaii [English translation]. Zool. Soc. London, Proc., B, 108: 365-387. Bailey, A.M. 1918. The monk seal of the Southern Pacific. Nat. Hist. 18: 396-399. =i ----1952. The Hawaiian monk seal. Denver Mus. Nat. Hist., Mus. Pict. 7: 1-30. ----1956. Birds of Midway and Laysan Islands. Denver Mus. Nat. Hist., Mus. Pict. 12: 1-130. Baldwin, P.H. 1945. Fate of the Laysan rail. Audubon Mag. 47: 343-348. ----1947. The life history of the Laysan rail. Condor 49: 14-21. Bartsch, P. 1931. A new pearl oyster from the Hawaiian Islands. U.S. Nat. Mus., Proc. 79(12): 1-2. Beardsley, J.W. 1966. Insects and other terrestrial arthropods from the Leeward Hawaiian Islands. Pac. Sci. 19: 157-185. Berger, A.J. 1970. The present status of the birds of Hawaii. Pac. Sci. a4: 29-h2, Bitter, G. 1900. Die phanerogamische Pflanzenwelt der Insel Laysan. Abh. Nat. Ver. Brem. 16: 430-439. Bock, W.J. 1958. A generic review of the plovers (Charadriinae, Aves). Mus. Comp. Zool. Bull. 118: 27-97. Brennan, J.M. 1965. A small collection of chiggers (Acarina: Trombiculidae) from the North Central Pacific. J. Parasitol. 51: 888-892. Brennan, J.M., and A.B. Amerson, Jr. 1971. Six new species and additional records of chiggers from the Central Pacific (Acarina: Trombiculidae). J, PEWQe Sr OG 77s Isllosly Brooks, N.C. 1860. Islands and reefs west-northwest of the Sandwich Islands, Pacific. Naut. Mag. 29: 499-504. Bryan, E.H., Jr. 1937. Pearl and Hermes Reef. Parad. Pac. 49(10): 19, 30. ----1942. American Polynesia and the Hawaiian chain. Tongg Publ. Cony Honolulu. 253 pp. ----1954. The Hawaiian chain. B.P. Bishop Mus. Press, Honolulu. 71 pp. ----1958. Check list and summary of Hawaiian birds. Books about Hawaii. Honolulu, Hawaii. 28 pp. Bryan, E.H., Jr. et al. 1926. Insects of Hawaii, Johnston Island and Wake Island. B.P. Bishop Mus. Bull. 31: 1-94. BSFW--Bureau of Sport Fisheries and Wildlife, U.S. Fish and Wildlife Service, Dept. of Interior, Kailua, Hawaii [unpublished reports]: 1964a. Kridler, E. Hawaiian Islands National Wildlife Refuge, March 1964. 258 1964b. ---- Hawaiian Islands National Wildlife Refuge, September 16 through 27, 1964. 30 pp. 1965. ---- Hawaiian Islands National Wildlife Refuge, March 11-23, 1965. “2O0epp: 1966a. ---- Hawaiian Islands National Wildlife Refuge trip report, March 17 - April 16, 1966. 24 pp. 1966b. ---- Hawaiian Islands National Wildlife Refuge trip--September 8-28, 1966. 34 pp. 1967a. ---- Refuge log book for 6 March - 1 April 1967. 39 pp. 1967b. ---- Pers. comm., July 1967. 1967c. ---- Refuge log book for 19-29 September 1967. 12 pp. 1967d. Robbins, C.S. Pers. corresp., March 1967. 1968. Kridler, E. Pers. conm., March 1968. 1969a. ---- Field trip report, Pearl and Hermes Reef, Hawaiian Islands National Wildlife Refuge, 9-13 February 1969. 7 pp. 1969b. ---- Hawaiian Islands National Wildlife Refuge spring trip, March 19 - April 6, 1969. 32 pp. 1969c. ---- Hawaiian Islands National Wildlife Refuge, Kailua, Hawaii, field trip report, May 25 - June 1, 1969. 8 pp. 1969d. ---- Hawaiian Islands National Wildlife Refuge, field trip report. August 19 - September 23, 1969. 32 pp. 1969e. Olsen, D.L. Hawaiian Islands National Wildlife Refuge tenia trip May 28- June 11, 1969. 18 pp. Canaan MoM a5 dies 1964. Transoceanic migrations of the green turtle. Bioscience 14(8): 49-52. ----1972. Great reptiles, great enigmas. Audubon Mag. 74(2): 24-35. Christophersen, E., and E.L. Caum. 1931. Vascular plants of the Leeward Islands, Hawaii. B.P. Bishop Mus. Bull. 81: 1-41. Clapp, R.B. 1968. Three unusual shorebirds from Midway Atoll, Pacific Ocean. Elepaio 28: 76-77. Clapp, Rebs, andeReie. Pyle 1968. Noteworthy records of waterbirds from Oahu. Elepaio 29: 37-39. Clapp, R.B., and P.W. Woodward. 1968. New records of birds from the Hawaiian Leeward Islands. U.S. Nat. Mus., Proc. 124(3640): 1-39. Bo) Clark, A.H. 1949. Ophiuroidea of the Hawaiian Islands. B.P. Bishop Mus. Bull. 195: 1-133. Clark, H.L. 1925. Echinoderms other than sea stars. In Edmondson et al., 1925. B.P. Bishop Mus. Bull. 27: 89-111. Cushman, J.L. 1925. Foraminifera. In Edmondson et al., 1925. B.P. Bishop Mus. Bull. 27: 121-144. Dall, W.H., P. Bartsch, and H.A. Rehder. 1938. A manual of the recent and fossil pelecypod mollusks of the Hawaiian Islands. B.P. Bishop Mus. Bul 153) 233 \s Delacour, J. 1954. The waterfowl of the world. Vol. 1. Country Life Ltd., London. ----1959. The waterfowl of the world. Vol. 3. Country Life Ltd., London. Dill, H.R., and W.A. Bryan. 1912. Report of an expedition to Laysan Island in 1911 under the joint auspices of the United States Department of Agriculture and the University of Iowa. U.S. Dept. Agr., Biol. Surv. Bull. 42: 1-30. Edmondson, C.H. 1925. Crustacea. In Edmondson et al., 1925. B.P. Bishop Mus. Bull. 27: 3-62. ----1930. New Hawaiian Crustacea. B.P, Bishop Mus., Occ. Pap. 9(10): 1-18. ----1935. New and rare Polynesian Crustacea. B.P. Bishop Mus., Occ. Pap. 10(24): 3-40. Elschner, C. 1915. ‘The Leeward Islands of the Hawaiian Group. Honolulu Advertiser, Honolulu. 68 pp. Ely, C.A., and R.B. Clapp. In press. The natural history of Laysan Island, Northwestern Hawaiian Islands. Atoll Res. Bull. Farrell, A. [ed.]. 1928. John Cameron's odyssey. The Macmillan Co., New York. 461 pp. Fisher, H.I., and P.H, Baldwin. 1945. A recent trip to Midway Islands, Pacific Ocean. Elepaio 6: 11-13. ----1946. War and the birds of Midway Atoll. Condor 48: 3-15. Fisher, W.K. 1925. Sea stars. In Edmondson etralts<1925. “BLP. Bishop Mus. Bull. 27: 63-88. Fowler, H.W. 1927. Fishes of the tropical Central Pacific. B.P. Bishop Mus. Bull. 38: 1-32. ----1928. The fishes of Oceania. B.P. Bishop Mus., Mem. 10: 1-540. 260 ----1931. The fishes of Oceania. Supplement 1. B.P. Bishop Mus., Mem. 11: 313-381. ----1934. The fishes of Oceania. Supplement 2. B.P. Bishop Mus., Mem. 11: 385-466. ----1949. The fishes of Oceania. Supplement 3. B.P. Bishop Mus., Mem. 12: 37-186. Fowler, H.W., and S.C. Ball. 1925. Fishes of Hawaii, Johnston Island, and Wake Island. B.P. Bishop Mus. Bull. 26: 1-31. Galtsoff, P.S. 1933. Pearl and Hermes Reef, Hawaii, hydrographical and biological observations. B.P. Bishop Mus. Bull. 107: 1-49. Garrett, L.E., and F.H. Haramoto. 1967. A catalog of Hawaiian Acarina. Haw. Ent. Soc., Proc. 19: 381-414. Goff, M.L. 1971. New records of chiggers (Acarina, Trombiculidae) from the Northwestern Hawaiian Islands. J. Med. Ent. 8: 456. Gosline, W.A., and V.E. Brock. 1965. Handbook of Hawaiian fishes. Univ. of Hawaii Press, Honolulu. 372 pp. Gregory, H.E. 192+. Report of the director for 1923. B.P. Bishop Mus. Bull. 10: 1-38. Gross, M.G., J.D. Milliman, J.I. Tracy, Jr., and H.S. Ladd. 1969. Marine geology of Kure and Midway Atolls, Hawaii; a preliminary report. Pac. Sei. 23: 17-25. Hartman, O. 1966. Polychaetous annelids of the Hawaiian Islands. B.P. Bishop Mus., Occ. Pap. 23: 163-252. Hawaiian Mission Children's Society Library, Honolulu [unpublished letters]: Missionary Letters. 1: 20-21: Letter from Bingham, Thurston, and Loomis dated 12 October 1822; letter from E. Loomis to his brother dated October 1822. HDFG--Hawaiian Division of Fish and Game, State of Hawaii, Honolulu [unpub- lished reports]: 1961. Woodside, D.H., and R.J. Kramer. A report on a survey trip to the Hawaiian Islands National Wildlife Refuge, March 1961. 32 pp. 1964. Walker, R.L. Notes on a survey of the Hawaiian Leeward Islands, March 1964. 35 pp. Hershkovitz, P. 1966. Catalog of the living whales. U.S. Nat. Mus. Bull. 246: 1-259. Holly, M. 1932. Echinodermata from Pearl and Hermes Reef. B.P. Bishop Mus., Occ. Pap. 10(1): 1-9. 261 ----1935. Polychaeta from Hawaii. B.P. Bishop Mus. Bull. 129: 1-33. Hornell, J. 1934. Log of the schooner Ada on a fishing cruise in the North Pacific, 1882. Mariner's Mirror 20: 426-437. Howe, M.A. 1934. Hawaiian algae collected by Dr. Paul C. Galtsoff. J. Wash. Acad. Sci. 4: 32-2, Hutchinson, G.E. 1950. Survey of existing knowledge of biogeochemistry. 3. The biogeochemistry of vertebrate excretion. Amer. Mus. Nat. Hist. Bull. 96: 1-554. Tkehara, I.I. 1953. Live-bait fishing for tuna in the Central Pacific. U.S. Fish and Wildl. Serv., Spec. Sci. Rep.: Fisheries 107: 1-20. June, F.C., and J.W. Reintjes. 1953. Common tuna-bait fishes of the Central Pacific. U.S. Fish and Wildl. Serv., Res. Rep. 34: 1-54. Kemble, J.H. [ed.]. 1966. To California and the south seas: the diary of Albert G. Osbun, 1849-1851. The Huntington Library, San Marino, Cart. eo S PDs Kenyon, K.W. In prep. Man versus the monk seal. Kenyon, K.W., and E. Kridler. 1969. lLaysan Albatrosses swallow indigestible matter. Auk 86: 339-343. Kenyon, K.W., and D.W. Rice. 1959. Life history of the Hawaiian monk seal. Pac. Sei. 13 5¢2h5=-252¢ King, J.E. 1956. The monk seals (genus Monachus). Brit. Mus. (Nat. Hist.) Zool. Bull. 3: 201-256. King, W.B. 1967. Preliminary Smithsonian identification manual. Seabirds of the tropical Pacific Ocean. Smithsonian Institution, Washington. 126 pp. Kohls, G.M. 1966. A new sea bird tick, Ixodes amersoni, from Phoenix Island. J. Med. Ent. 3: 38-40. Kohls, G.M., and C.M. Clifford. 1967. The male and larva of Ixodes laysan- ensis Wilson with notes on rearing (Acarina, Ixodidae). J. Med. Ent. 4: 83-86. Kohls, G.M., D.E. Sonenshine, and C.M. Clifford. 1965. The systematics of the subfamily Ornithodorinae (Acarina: Argasidae). II. Identification of the larvae of the Western Hemisphere and description of three new species. Ent. Soc. Amer., Ann. 58: 331-364. Kroenke, L.W., and G.P. Woollard. 1965. Gravity investigations on the Leeward Islands of the Hawaiian Ridge and Johnston Island. Pac. Sci. 19: 361-366. 262 Ladd, H.S., J.1. Tracey, Jr., and M:Gy Gross. 1957. Drilling Yon) Midway Atoll, Hawaii. Science 156: 1088-1094. Lamoureux, C.H. 1964. The Leeward Hawaiian Islands. Haw. Bot. Soc., Newsl. 3(2): 7-11. Laycock, G. 1970a. The Hawaiian islands of birds. Audubon Mag. 72(1): ; Wh -61, j , ----1970b. Haunted sands of Laysan. Audubon Mag. 72(2): 42-9. Lopes, H. de Souza. 1938. On the genus Goniophyto Townsend, 1927, with description of a new species from Hawaii. B.P. Bishop Mus., Occ. Pap. 14: 193-197. Maa, T.C. 1968. Records of Hippoboscidae (Diptera) from the Central Pacific. J. Med. Ent. 5: 325-328. Morison, S.E. 1949. History of United States Naval Operations in World War II. Vol. IV. Coral Sea, Midway, and submarine actions, May 1942 - August 1942, Little, Brown and Co., Boston. 307 pp. Morrell, B., Jr. 1832. A narrative of four voyages, to the South Sea, North and South Pacific Ocean, Chinese Sea, Ethiopic and Southern Atlantic Ocean, Indian and Antarctic Ocean. Harper and Brothers, New York. 492 pp. Morris, P.C. 1934. How the Territory of Hawaii grew and what domain it covers. Haw. Hist. Soc. 42nd Ann. Rep. for 1933: 13-39. Munro, G.C. 1942. Birds of Hawaii and adventures in bird study; an ocean CmUtse, MO WO, wilejato© 32 23}. ----1945. The small birds of Midway. Elepaio 6: 13-14. ----1917. Effect of a tidal wave on some sea bird nesting islands. Elepaio 7: 43-44, 51-52. Munter, W.H. (ms.). Report to the captain of the Thetis of bird observations made during a cruise to the Hawaiian Bird Reservation in January and Feb- ruary 1916. U.S. Nat. Archives. Rec. Group 26. 19 pp. Office of Geography. U.S. Dept. of the Interior. 1956. NIS [National In- telligence Survey] Gazetteer, Hawaiian Islands. Central Intelligence Agency, Washington. 89 pp. Parsons, J.J. 1962. The green turtle and man. Univ. of Florida Press, Gainesville. 126 pp. Paty, J. 1857. Account of the Manuokawai--interesting account of her explorations. The Polynesian. 6 June 1857, 40: 2-3. Pearcy, G.E. 1959. Hawaii's territorial sea. Prof. Geog. 11(6): 2-6. 263 Peters, J.L. 1931. Check-list of birds of the world. Vol. I. Harv. Univ. Press, Cambridge, Mass. 345 pp. ----1934. Check-list of birds of the world. Vol. II. Harv. Univ. Press, Cambridge, Mass. 401 pp. ----1937. Check-list of birds of the world. Vol. III. Harv. Univ. Press, Cambridge, Mass. 311 pp. Pettingill, O.A., Jr. 1964. Midway Islands: man and birds in conflict. Audubon Mag. 66: 154-159. Pietschmann, V. 1930. Remarks on Pacific fishes. B.P. Bishop Mus. Bull. 73: 1-24. ----1938. Hawaiian shore fishes. B.P. Bishop Mus. Bull. 156: 1-55. Pilsbry, H.A. 1917. Marine molluses of Hawaii, IV-VII. Acad. Nat. Sci. Phila., Proc. 69: 309-333. ----1920. Marine molluscs of Hawaii, VII-XIII. Acad. Nat. Sci. Phila., Proc. 72: 296-328. ----1927. Littoral barnacles of the Hawaiian Islands and Japan. Acad. Nat. Sci. Phila., Proc. 79: 305-317. POBSP=--Pacific Ocean Biological Survey Program, Smithsonian Institution, Washington [unpublished reports]: 1963. Sibley, F.C. Pacific Project field trip--June 1963. [Leeward Survey No. 2]. 9 pp. 1964a. Amerman, K.E., Leeward Islands survey: Pearl and Hermes Reef, Lisianski Island, August 1964. [Leeward Survey No. 4]. 11 pp. 1964b. Amerson, A.B., Jr. Northwest Hawaiian Islands trip report, March 1964. [Leeward Survey No. 3]. 20 pp. 1964c. Fleet, R.R. Leeward Islands survey no. 5, September 1964. 15 pp. 1964d. Sibley, F.C. Preliminary report on ATF trip no. 1, February- March 1963. [Leeward Survey No. 1]. 13 pp. 1965a. Banko, W.E. Trip report, Hawaiian Leeward Islands survey, March 11-23, 1965. [Leeward Survey No. 8]. 11 pp. 1965b. Wirtz, W.O., Il. Leeward Islands survey: Pearl and Hermes Reef, Laysan Island, Lisianski Island, March 1965. [Leeward Survey Noe ike6 Apps 1966. Lewis, T.J. Trip report, Pearl and Hermes Reef, September 25- 27, 1966. [Leeward Survey No. 16]. 10 pp. 264 1967a. Clapp, R.B., and C.A. Ely. Leeward survey no. 20, pre- liminary report, Pearl and Hermes Reef, 28-30 August 1967. 16 pp. 1967b. DeLong, R.L. Preliminary report: Pearl and Hermes Reef survey, May 28 - June 1, 1967. [Leeward Survey No. 19]. 12 pp. 1967c. ---- Census and observations of Hawaiian monk seal on Pearl and Hermes Reef, Lisianski, and Laysan Islands, May 31 to June ll, 1967. [Leeward Survey No. 19]. 8 pp. 1967d. Hackman, C.D. Preliminary report of Leeward Island survey no. 18, March 6 to March 27, 1967. 12 pp. 1968. Clapp, R.B. Leeward survey no. 22, preliminary report, Pearl and Hermes Reef, 22-24 March 1968. 20 pp. POFI--Pacific Ocean Fisheries Investigations, Bureau of Commercial Fisheries, Honolulu [unpublished notes and reports]: 1950. Narrative report; Hugh M. Smith - cruise V; 16 June -6 August. 1951. Narrative report; Hugh M. Smith, cruise IX; 5 May - 2 July. 1955. Cruise report; FSW M/V John R. Manning, cruise 23, Part II; 5 January - 6 February. 1956a. Cruise report; FWS M/V John R. Manning, cruise 30; 15 March - 20 April. 1956b. Cruise report; FWS M/V John R. Manning, cruise 31; 19 May - 22 June. Rice, D.W. 1960a. Population dynamics of the Hawaiian monk seal. J. Mammal. 41: 376-385. ----1960b. Distribution of the bottle-nosed dolphin in the Leeward Hawaiian Islands. J. Mammal. 41: 407-408. ----1964. The Hawaiian monk seal. Nat. Hist. 72: 48-55. ----(ms. a). Sea bird populations on Pearl and Hermes Reef - 1957. 15 October 1957 memorandum to John W. Aldrich. Smithsonian Institu- tion, Washington. 5 pp. ----(ms. b). Findings of the 1957-58 investigations of the bird problem at Midway Island. Undated summary report. Smithsonian Institution, Washington. 3 pp. Rice, D.W., and K.W. Kenyon. 1962. Breeding distribution, history and populations of North Pacific albatrosses. Auk 79: 365-386. Richardson, F. 1957. The breeding cycles of Hawaiian seabirds. B.P. Bishop Mus. Bull. 218: 1-41. 265 St. John, H. 1970. The genus Sicyos (Cucurbitaceae ) on the Hawaiian Leeward Islands, Hawaiian plant studies 35. Pac. Sci. 24: 439-456. Schilder, F.A. 1933. Cypraeacea from Hawaii. B.P. Bishop Mus., Occ. Pap. 10(3): 1-22. Shelton, P.C. In prep. The natural history of Johnston Atoll, Central Pacific Ocean. Atoll Res. Bull. Sibley, F.C., and R.W. McFarlane. 1968. Gulls in the Central Pacific. Pac. Sci. 22: 314-321. Standen, R.S. 1967. An explanatory description of the sand islands of Kure, Midway, and Pearl and Hermes Atolls: Hawaiian Islands. MA thesis, Calif. State Coll., Los Angeles. 93 pp. Stearns, H.T. 1966. Geology of the state of Hawaii. Pacific Books, Palo Mito. Callie.) 266 pps Svihla, A. 1959. Notes on the Hawaiian monk seal. J. Mammal. 40: 226-229. Taylor, W.R. 1964. The genus Turbinaria in eastern seas. J. Linn. Soc. (Bot.) 58: 471-487. Thompson, G.B. 1948. Mallophaga collected by the Tanager expedition. B.P. Bishop Mus., Occ. Pap. 19: 195-200. Thorp, E.M. 1936. The sediments of the Pearl and Hermes Reef. J. Sed. Petrol. 6: 109-118. Thrum, T.G. [comp.]. 1909. Hawaiian almanac and annual for 1910. T.G. Thrum, Honolulu. 216 pp. Tomich, P.Q. 1969. Mammals of Hawaii. A synopsis and notational biblio- graphy. B.P. Bishop Mus., Spec. Publ. 57: 1-238. Treadwell, A.L. 1925. Polychaetous annelids. In Edmondson et al., 1925. B.P. Bishop Mus. Bull. 27: 113-119. Tsuda, R.T. 1966. Marine benthic algae from the leeward Hawaiian group. Atoll Res. Bull. 115: 1-13. Udvardy, M.D.F. 1961. Additions to the check list of Hawaiian birds. Elepaio 21: 83-90. USCGS--U.S. Coast and Geodetic Survey, Washington. Chart 4175. U.S. Department of State. 1965. United States and outlying areas. Geog. Bull. 5: 1-15 ° a U.S. National Archives, Washington [unpublished letters, documents, and reports |: Record Group 22. U.S. Fish and Wildlife Service. Record Group 2+. U.S. Navy personnel records and ships' logs. Record Group 26. U.S. Coast Guard. Record Group 37. U.S. Hydrographic Office. Record Group 48. U.S. Department of Interior. Record Group 126. Records of the Office of Territories. USNHO--U.S. Naval Hydrographic Office, Washington. Chart 4, Chart 5647. U.S. Navy, Classified Operational Archives. Navy Yard, Washington [un- published]: War Diary, USS Preble, 1942. Usinger, R.L. 1942. The genus Nysius and its allies in the Hawaiian Islands. B.P. Bishop Mus. Bull. 173: 1-167. Watson, G.E., III. 1966. Smithsonian identification manual: seabirds of the tropical Atlantic Ocean. Smithsonian Institution, Washington. ----Wetmore, A. 1925. Bird life among lava rock and coral sand. Nat. Geogr. Mag. 48: 77-108. ----(ms.). Field notes, 1923. Smithsonian Institution, Washington. Wheeler, W.M. 1934. Revised list of Hawaiian ants. B.P. Bishop Mus., Occ. Pap. 10(21): 3-21. Wiens, H.J. 1959. Atoll development and morphology. Assoc, Amer. Geogr., Ann. 49: 31-54. ----1962. Atoll environment and ecology. Yale Univ. Press, New Haven. 532 pp. Wilder, G.P. 1905. A short trip to Midway Islands with Capt. A.P. Nidblack in the U.S.S. "Iroquois." Haw. For. Agr. 2: 390-396. Willett, G. 1919. Notes on the nesting of two little-known species of petrel. Condor 21: 60-61. 267 ----(ms.). Report on the Bureau of the Biological Survey Expedition to the Northwestern Hawaiian Islands in 1912-1913. Bureau of Sport Fisheries and Wildlife, Kailua, Hawaii. Yocom, C.F. 1965. Longevity record of a Black-footed Albatross. Condor 67: 187-188. Zimmerman, E.C. 1940. Studies of Hawaiian Neuroptera. Haw. Ent. Soc., Proc. 10: 487-510. ----1948a. Insects of Hawaii. Vol. 1. Introduction. Univ. of Hawaii Press, Honolulu. 191 pp. ----1948b. Insects of Hawaii. Vol. 2. Apterygota to Thysanoptera. Univ. of Hawaii Press, Honolulu. viii and 475 pp. ----1948c. Insects of Hawaii. Vol. 3. Heteroptera. Univ. of Hawaii Press, Honolulu. v and 255 pp. ----1948d. Insects of Hawaii. Vol. 4. Homoptera: Anchenorhyncha. Univ. of Hawaii Press, Honolulu. vii and 268 pp. ----1957. Insects of Hawaii. Vol. 6. Ephemeroptera, Neuroptera, Tri- droptera, and supplements to Vols. 1 to 5. Univ. of Hawaii Press, Honolulu. ix and 209 pp. ----1958. Insects of Hawaii. Vol. 7. Macrolepidoptera. Univ. of Hawaii Press, Honolulu. ix and 542 pp. 268 Appendix Table 1. Date of Visit 1857 19-20 May 1859 5 July 1891 6-7 July 1912 2h Dec. 1913 15 Mar. 1OMe ) 2iSsept. 1916 & Feb. 1923 26-28 Apr. 17-19 May 1927- 1931 1O2BIn 2 1930 22 July- *XSymbols and abbreviations: Scientific visits to Pearl and Hermes Reef, 1857-1969 ** Vessel Manoukawai Gambia Kaalakai USRC Thetis USRC Thetis USRC Thetis USRC Thetis USS Tanager USS Tanager Hermes (renamed Lanikai) Lanikai USS Whippoor- will Personnel John Paty N.C. Brooks Rothschild Expedition: George C. Munro, Henry C. Palmer USN: Cmdr. G.R. Salisbury BBS: Alfred M. Bailey, George Willett, David T. Fullaway Carl Elschner W.H. Munter Tanager Expedition: Edwin L. Caum, Chapman Grant, David T. Fullaway, Eric L. Schlemmer, Ditlev Thaanum, Alexander Wetmore Theodore T. Dranga, Austin Jones, Frank R. Lawrence, H.C. Reno,* Lorrin A. Thurston, Gerrit P. Wilder William G. Anderson, George Kaufman Victor Pietschmann Paul S. Galtsoff,* N.S. Castle, John F. Reppun, Leon Amboy, Anatolio Polo BBS - Bureau of Biological Survey; BPBM - Bernice P. Bishop Museum; BSFW - Bureau of Sport Fisheries and Wildlife; CC - Clare- mont College; HDFG - Hawaiian Division of Fish and Game; HSB - Honolulu Star Bulletin; NAS - National Audubon Society; POBSP - Pacific Ocean Biological Survey Program; POFI - Pacific Ocean Fisheries Investigation; UF - University of Florida; UH - University of Hawaii; USCG - U.S. Coast Guard; USFW - U.S. Fish and Wildlife Service; USN - U.S. Navy; VU - Vienna University. *Biologist-in-charge. Appendix Table 1. Date of Visit 1949 1950 1951 1954 1955 1956 1957 1958 May 25-27 June 14 May 1 Nov. 5 Dec. 25-26 Jan. 22-2 Mar. 24-26 May 9 Dec. 10 Dec. if DEC 7 Jan. 24 Jan. 15 Apr. 14 May 14 Oct. 18 Dec. ok Jan. 14 Apr. 2 May 16 June (continued ) Vessel Navy PBY USFW Hugh M. Smith USFW Hugh M. Smith Aerial survey W Li USFW John R. Manning USFW John Manning USFW John Manning Ip Ip Aerial survey 269 Personnel Alfred M. Bailey HDFG: Vernon E. Brock; UH: William A. Gosline; POFI personnel POFI personnel BSFW: Philip Dumont, Johnson A. Neff 1 "W 1 " 1! POFI: Tamio Otsu, Walter Matsumoto POFI: Daniel T. Yamashita, Herbert S. Shippen POFI: E.C. Jones, T.S. Hida BSFW: John W. Aldrich, Karl W. Kenyon, Dale W. Rice, Chandler S. Robbins BSFW: Karl W. Kenyon, Dale W. Rice BSFW: Dale W.. Rice 270 Appendix Table 1. Date of Visit 1961 12 Mar. 162 Tew. 26 Feb.- 8 Mar. 1963 18-23, 27 June 1964 13-14 Mar. 16-20 Aug. 16-17 Sept. 1965 15-19 Mar. 21-22 Mar. 1966 1 Apr. 20-26 Sept. 25-27 Sept. 1967 16-17 Mar. (continued) Vessel USCGC Planetree Aerial survey USS Moctobi (ATF-105 ) USS Tawakoni (ATF-114) USCGC Planetree (WAGL-307 ) USNS Shearwater (T-AG-177 USCGC Basswood USNS Shearwater (T-AG-177 ) USCGC Blackhaw (WAGL-390) USCGC Buttonwood (WAGL-30 USCGC Ironwood (WAGL-297) Navy helicopter Personnel HDFG: Kramer David H. Woodside, Raymond J. UF: Archie F. Carr POBSP: A. Binion Amerson, Jr., F. Allen Blagden, Robert W. McFarlane, Fred C. Sibley, William O. Wirtz II* POBSP: A. Binion Amerson, Jr., Fred C. Sibley* POBSP: A. Binion Amerson, Jr.,* George S. Wislocki; BSFW: Eugene Kridler, Edward O'Neal; HDFG: Ronald L. Walker; UH: Loren W. Kroenke POBSP: Kenneth EF. Amerman,* Allen Anderson, Peter Marshall, Richard W. Merrill, J. Douglas Whitman, Paul W. Woodward; UH: Allen L. Young POBSP: Robert R. Fleet, Charles R. Long; BSFW: Eugene Kridler;* HDFG: Ronald L. Walker; UH: John Beardsley POBSP; Kenneth E. Amerman, Roger B. Clapp, J. Vincent Hoeman, Dennis L. Stadel, William O. Wirtz II* POBSP: Winston E. Banko; BSFW: Kridler,* Chandler Robbins; HDFG: Ronald L. Walker Eugene BSFW: Eugene Kridler;* HDFG: Nelson Rice, Ronald L. Walker; UH: Andrew J. Berger BSFW: Karl W. Kenyon, Eugene Kridler;* HDFG: Ronald L. Walker; CC: Sherwin Carlquist; HSB: Warren Roll POBSP: T. James Lewis;* USN: George Young BSFW: Van T. Harris, Chandler Robbins* e71 Appendix Table 1. (continued) Date of Visit Vessel Personnel 1967 21-23 Mar. USCGC Basswood POBSP: C. Douglas Hackman; BSFW: Eugene Kridler;* HDFG: Ernest F. Kosaka; UH: John Maciolek, Richard Wass 28, 30 May- LT 2081, POBSP: David L. Burckhalter, Robert 1 June LT 2086, L. DeLong,* Dennis L. Stadel, F. Chris- LT 2087 tian Thompson, Robert Tuxson; USN: Ronald Amerson 3-9 July ? BSFW: Eugene Kridler, 7? 28-30 Aug. LT 2081, POBSP: Roger B. Clapp, Charles A. Ely,* LT 2086, David I. Hoff; USN: Wrangel LT 2087 2/-29 Sept. USCGC Buttonwood BSFW: Robert Ballou, Eugene Kridler,* John L. Sincock; UDFG: Ronald L. Walker 1968 22-24 Mar. USCGC Ironwood POBSP: Roger B. Clapp; BSFW: Karl W. (WAGL-297 ) Kenyon, Eugene Kridler,* John L. Sincock; HDFG: Ernest F. Kosaka; USCG: Douglas Keran 8 Aug. ? BSFW: Eugene Kridler, ? 1969 10-12 Feb. Navy helicopter BSFW: Eugene Kridler, David L. Olsen; USN: Ronald Amerson By Mars USCGC Buttonwood BSFW: Karl W. Kenyon, Eugene Kridler, 2 Apr. David L. Olsen, John L. Sincock; NAS: George Laycock 26-31 May Navy helicopter BSFW: Hugene Kridler; BPBM: Donald All- red; USN: Ronald Amerson 5 June Mahi BSFW: David L. Olsen,* John L. Sincock; HDFG: Ernest F. Kosaka; UH: Karl Bathen, Thomas Clark, James McVay, William Patz- ert; BPBM: Douglas Yen; Ronald Kent 28 July ? BSFW: ? 10-19 Sept. USCGC Buttonwood BSFW: Eugene Kridler,* David L. Olsen, John L. Sincock; UH: John Maciolek Ce Appendix Table 2. Publications and manuscripts on collections and studies made on Pearl and Hermes Reef * PROTOZOA Cushman, 1925 Records 25 species of Foraminifera from Tanager Expedition PORIFERA Galtsoff, 1933 Records 1 sponge species from 1930 COELENTERA TA Galtsoff, 1933 Records 1 jellyfish species and 12 coral species from 1930 MOLLUSCA Pilsbry, 1917 Records 1 mollusk species collected by Munter Pilsbry, 1920 Lists 3 mollusk species collected by Munter Pilsbry, 1927 Lists 1 barnacle from Tanager collection Bartsch, 1931 Describes a new pearl oyster from Galtsoff's collection Galtsoff, 1933 Lists 10 mollusk species from 1930 Schilder, 1933 Reports 14 species of Cypraeacea col- 3 lected by Pietschmann in February 1928 Dall, Bartsch, and Rehder, Records 8 species of mollusks 4 1938 ECHINODERMATA Clark, 1925 Reports 10 species of echinoderms from Tanager collection Fisher, 1925 Lists 3 species of starfish collected by Tanager Expedition Holly, 1932 Reports 13 echinoderms from collections made in 1927-28 *Authors are in chronological order. Appendix Table 2. (continued) 273 ECHINODERMATA (continued) Galtsoff, 1933 Clark, 1949 Lists 4 species of sea urchins, 3 starfish, 1 brittle star, and 9 holo- thurian species from 1930 Lists 33-34 brittle stars (Ophiuroidea) collected by Albatross and Tanager Ex- peditions ANNELIDA Treadwell, 1925 Holly, 1935 Hartman, 1966 Reports 1 annelid from Tanager collection Lists 15 Polychaeta from Pietschmann's 1928 collection Summarizes earlier polychaetous annélids; lists 11 species ARTHROPODA 1. Crustacea Edmondson, 1925 Edmondson, 1930 Galtsoff, 1933 Edmondson, 1935 2. Chilopoda Attems, 1938 3. Arachnida Brennan, 1965 Kohls, Sonnenshine and Clifford, 1965 Kohls, 1966 Garrett and Haramoto, 1967 Reports 33 species of decapods from Tanager collection Describes 1 new species of crab taken in 1927 by T.T. Dranga Mentions collection of over 50 species taken in 1930; lists only 5 species Describes 1 new crustacea species Records 1 myriapod species collected by Tanager Expedition Records 1 chigger from POBSP material Records 1 tick species from POBSP material Records 1 ixodid species from POBSP material Summarizes 2 earlier records of acarina eT Appendix Table 2. (continued) ARTHROPODA (continued) 3. Arachnida (continued) Kohls and Clifford, 1967 Amerson, 1968 Brennan and Amerson, 1971 Goff, 1971 Bryan, et al., 1926 Wheeler, 1934 Lopes, 1938 Zimmerman, 1940 Munro, 1942 Usinger, 1942 Thompson, 1948 Zimmerman, 1948a Zimmerman, 1948b ieimeranaiae 1948e Zimmerman, 1948d Zimmerman, 1957 Zimmerman, 1958 Beardsley, 1966 Mentions occurrence of ixodid species from POBSP data Records 2 tick species from POBSP col- lections Records 1 chigger collected by J.L. Gressitt in December 1970. Records 3 chiggers collected by J.L. Gressitt in December 1970 INSECTA Lists ca. 24 insects from Tanager col- lection Lists 1 ant species Describes new sarcophagid from Tanager material Reports a neuropteran Mentions 2 insects on ship while off- shore in 1891 Records 1 new Nysius species from Tanager collection Records 2 Mallophaga species from Tanager collection Lists 1 Orthoptera and 2 Dermoptera Lists 1 Orthoptera species Lists 3 Hemiptera species Lists 1 Homoptera species Lists 1 Neuroptera species Lists 1 Lepidoptera species Lists 52 insect species a Appendix Table 2. (continued) Maa, 1968 Amerson and Emerson, 1971 1. Pisces Brooks, 1860 Fowler and Ball, 1925 Fowler, 1927 Pietschmann, 1930 Fowler, 1931 Schilder, 1932 Galtsoff, 1933 Pietschmann, 1938 POFI (1950-56) mss. Ikehara, 1953 Jones and Reintjes, 1953 Gosline and Brock, 1965 2. Reptilia | Morrell, 1832 Paty, 1857 Brooks, 1860 Bailey, 1918 275 INSECTA (continued) Records 2 hippoboscid flies from POBSP collection Records 2 Mallophaga species from POBSP material CHORDATA First record of fish in 1859 Reports 16 fish species taken by Tanager Expedition Lists 1 fish species from Tanager Ex- pedition Lists 9 fish species collected by Dranga in 1927 Adds 3 fish species, collected by Dranga, April 1927 Discusses hemirhampids collected in 1928 Lists 31 fish species collected in 1930 Reports fishes collected in 1927-28 Consists of 5 trip reports listing fish collections and observations Records 5 species of bait fish from 1950 and 1951 POFI surveys Records 4 fish species Includes fish from earlier trips First record of sea turtles Notes presence of turtles Records presence of turtles Reports sleeping turtles 276 Appendix Table 2. (continued) CHORDATA (continued) 2. Reptilia (continued) Galtsoff, 1933 POFI (1950-56) mss. Bailey, 1956 HDFG (1961-64) mss. POBSP (1963-68) mss. BSFW (1964-69) mss. Carr, 1964 Kemble, 1966 3. Mammalia Baieary | USS Atkinson and Bryan, 1913 Bailey, 1918 Wetmore, ms. Gregory, 1924 Wetmore, 1925 Bryan et al., 1926 Galtsoff, 1933 Allen, 1942 Records sea turtles in 1930 Consists of 5 trip reports listing turtle observations Records many large sea turtles at North Island seen by Willett in 1913 Includes 2 trip reports of Refuge surveys Includes 12 trip reports from island surveys Includes 15 trip reports of Refuge surveys Notes presence of sea turtle Reports turtle observations by Osbun in 1850 Mentions presence of seals in May 1857 Notes estimate by Frear in 1912 Reports less than 60 seals and a school of porpoise observed in 1913 1923 personal journal notes presence of seals and rabbits Mentions Tanager personnel killed all but one of the rabbits seen in May 1923 Notes presence of seals in 1923 Notes presence of number of rabbits on Southeast in 1923 Counts 68 seals Reports seal colony found by Tanager Expedition Eq(T( Appendix Table 2. (continued) CHORDATA (continued) 3. Mammalia (continued) POFI (1950-56) mss. Consists of 5 trip reports listing mammal observations Bailey, 1956 Records ca. 50 seals seen by Willett in 1913 King, 1956 Summarizes previous information on Hawaiian Monk Seals Kenyon and Rice, 1959 Reports observations of Hawaiian Monk Seals from 1956 and 1957 aerial surveys Svihla, 1959 Summarizes previous seal data Rice, 1960a Gives results of 1956 and 1957 aerial Surveys of Hawaiian Monk Seals Rice, 1960b Reports observations of dolphin in March 1958 HDFG (1961-64) mss. Includes 2 trip reports of Refuge surveys POBSP (1963-68) mss. Includes 12 trip reports from island surveys BSFW (1964-69) mss. Includes 15 trip reports of Refuge surveys Ralee, 1964 Mentions presence of seals Kemble, 1966 Reports seal observations by Osbun in 1850 Tomich, 1969 Records Hawaiian Monk Seal and bottlenose dolphin; excellent bibliography Laycock, 1970a Notes presence of seals in March 1969 at Southeast Laycock, 1970b Notes rabbit data in April 1923 as re- ported by Wetmore Kenyon, ms. Discusses man's influence on seals 278 Appendix Table 2. (continued) CHORDATA (continued) 4, Aves Elschner, 1915 Willett, ms. Willett, 1919 Wetmore, ms. Galtsoff, 1933 Munro, 1942 Fisher and Baldwin, 1945 Munro, 1945 Fisher, 1946 Baldwin, 1947 Munro, 1947 POFI (1950-56) mss. Bailey, 1956 Aldrich, Robbins, and Rice, ms. Rice, ms. a Richardson, 1957 Rice, ms. b Records 5 bird species in September 1914 1912-13 bird observations Records 1 storm petrel species in 1912-13 Records 21 bird species during 1923 Tanager Expedition Records 6 bird species, summer 1930 Records 4 bird species seen offshore 6-7 July 1891 Records introduction and fate of Laysan Rail Records severe storm of 1930 or 1931 which killed many seabirds Discusses unsuccessful introduction of Laysan Rail Discusses Laysan Rail introduction Speculates on destruction of bird life on low islands by tidal waves 5 trip reports listing bird observations Records 11 species seen by Willett on North Island, March 1913 Records about 40,000 of both albatrosses on 9 December 1956 Memo to Dr. J.W. Aldrich records 12 species of birds from aerial survey of 14 October 1957 Records 8 breeding seabird species Records 43% nest destruction of Black- footed Albatross and 6% of Laysan Alba- tross by January 1958 tidal wave Appendix Table 2. (continued ) e719 CHORDATA (continued ) 4, Aves (continued) HDFG (1961-64) mss. Rice and Kenyon, 1962 POBSP (1963-68) mss. BSFW (1964-69) mss. Pettingill, 1964 Clapp and Woodward, 1968 Sibley and McFarlane, 1968 Laycock, 1970a 1. Vascular Plants Kemble, 1966 Brooks, 1860 Bitter, 1900 Bailey, 1918 Gregory, 1924 Christophersen and Caum, 1931 Galtsoff, 1933 FLORA Includes 2 trip reports of Refuge surveys Aerial surveys of populations of alba- trosses during 1956-57 and 1957-58 seasons Includes 12 trip reports from island surveys Includes 15 trip reports of Refuge surveys Mentions 17,800 breeding pairs of Laysan Albatross observed by Rice and Kenyon Lists 17 new bird species records for the atoll from POBSP data Records 4 gull species from POBSP data Notes presence of 5 bird species on Southeast in March 1969 First record of plants Second record of plants Reports 2 plant species from photographs by F.G.E. Walker in 1899(7) Notes presence of grass in 1913 Lists plants introduced by Tanager Ex- pedition in 1923 Reports 11 species of vascular plants from Tanager Expedition Lists 2 plant species from 1930 280 Appendix Table 2. (continued) FLORA (continued) 1. Vascular Plants (continued) HDFG (1961-64) mss. POBSP (1963-68) mss. Iamoureux, 1964 Laycock, 1970a St. John, 1970 2. Algae Galtsoff, 1933 Howe, 1934 POFI (1950-56) mss. Taylor, 1964 Tsuda, 1966 Elschner, 1915 Galtsoff, 1933 Thorp, 1936 Bryan, 1937 Bryan, 1942 Bryan, 1954 Kroenke and Wollard, 1965 Notes 4 plant species Includes botanical notes in 12 trip reports Mentions 11 plant species taken by Tanager Expedition Mustard sprayed with herbicide Describes 1 new species collected in 1923 Records 2 algae species from 1930 Reports 12 algae collected by Galtsoff in 1930 Reports 4. algae species in 1956 Lists 1 algae species collected by Galtsoff in 1930 Reports 39 species of marine benthic algae taken by POBSP in June 1956, August and September 1964, and earlier records GEOPHYS ICAL Describes reef, lagoon, and North Island Records hydrographic data from 1930 Records of sediments from Galtsoff's 1930 collection General description of atoll General description of atoll General description of atoll Gives gravity observations made in March 1964 281 Appendix Table 2. (continued) GEOPHYSICAL (continued ) Standen, 1967 Describes formation of sand islands (POBSP) Gross et al., 1969 Analysis of Gemini VII photographs Appendix Table 3. Annotated list of vascular plants from Pearl and Hermes Reef. Specimens are found in the herbaria of the Na- tional Museum of Natural History (USNM), the Bernice P. Bishop Museum (BPBM), and the University of Hawaii (UH) Gramineae *Cynodon dactylon (L.) Pers. Specimens only from Southeast Island. Lamoureux s.n. (UH); Sibley 10 (USNM); Young 115 (UH). Eragrostis variabilis (Gaud.) Steud. Specimens from Grass, North and South- east Islands; known also from Seal Island. Caum 38, 45 (BPBM); Lamoureux s.n. (UH); Long 2272, 2285, 2313 (UH); Sibley 5 (USNM); Young 105, 122 (UH). Lepturus repens (Forst.) R. Br. Specimens from Grass, Little North, North, and Southeast Islands; known from Seal Island. Caum 39, 46 (BPBM) as L. repens var. subulatus Fosb.; Lamoureux s.n. (UH); Long 2270 (appears to be var. subulatus Fosb.), 2273, 2274, 2277, 2279, 2292, 2293, 2302, 2311, 2312 (UH); Young 110, 128 (UH). *Setaria verticellata (L.) Beauv. Specimens only from Southeast Island; known also from Grass Island. Lamoureux s.n. (BPBM); Long 2269 (UH); Sibley 12 (USNM); Young 109 (UH). Liliaceae *Allium sp. Found growing on Southeast Island refuseheap, eradicated March 1963. No specimens collected. Palmae *Cocos sp. Planted in 1928-29 on Southeast Island, all dead or dying in 1930 (Galtsoff, 1933: 14). No specimen record. *Pritchardia pacifica Wendl. Planted in 1923 on Southeast Island by Tanager Expedition personnel (Gregory, 1924: 21). Wo specimen record. *Presumably exotic. 282 Appendix Table 3. (continued) Casuarinaceae *Casuarina equisetifolia L. Specimen only from Southeast Island; planted in 1963 by U.S. Navy. Young 120 (UH). Amaranthaceae Achyranthes splendens var. reflexa Hbd. Specimens from Grass and North Islands; known from Seal Island. Caum 50 (BPBM); Lamoureux s.n. (UH); Long 2298 (UH); Wilder 3 (BPBM). Nyctaginaceae Boerhavia repens L. Specimens from Grass, Little North, North and Southeast Islands; known from Seal Island. Caum 40, 41, 47, 48 (BPBM); Galtsoff s.n. (USNM); Lamoureux s.n. (UH); Long 2271, 2291, 2295, 2306, 2307, 2310 (UH); Sibley 3 (USNM); Young 98, 130 (UH). Aizoaceae Sesuvium portulacastrum L. Specimens from Seal and Southeast Islands. Caum 3, 55 (BPBM); Galtsoff s.n. (USNM); Lamoureux s.n. (UH); Long 2276 (UH); Sibley 7 (USNM); Young 100, 103 (UH). Portulacaceae Portulaca lutea Sol. Specimens from North and Southeast Islands. Sibley 9 (USNM); Long 2283 (UH); Young 104, 108, 123 (UH). Capparidaceae Capparis sandwichiana Dc. Specimens only from Seal Island. Wilder 2 (BPBM); Caum 54 (BPBM); Lamoureux s.n. (UH). Cruciferae *Brassica campestris L. Specimens from only Southeast Island; known also at North Island. Sibley 2 (USNM); Young 106, 107, 111, 116 (UH). *Coronopus a (L.) J.E. Smith. Specimen only from Southeast Island. Sibley 8 (USNM). Lepidium bidentatum var o-waihense (C. and S.) Fosb. Specimens from Grass, North, and Southeast Islands; known from Seal Island. Caum 51 (BPBM); Lamoureux s.n. (UH); Long 2286, 2289, 2299 (UH); Sibley 14 ee Young 99, 124 (UH). Zygophyllaceae Tribulus cistoides L. Specimens from Grass, Little North, North, Seal and Southeast Islands. Caum 44 (BPBM); Lamoureux s.n. (UH); Long 2268, 2282, 2296, 2305 (UH); Sibley 1 (USNM); Wilder 1 (BPBM); Young 102, 126 (UH). 283 Malvaceae *Malvastrum coromandelianum (L.) Garcke. Specimen only from Southeast Island. Young 114 (UH). *Hibiscus tiliaceus L. Planted in 1923 on Southeast Island by Tanager Ex- pedition personnel (Gregory, 1924: 21). No specimen record. Boraginaceae Tournefortia argentea L. f. Specimens from Grass, Little North, and North Islands. Long 2297, 2300 (UH); Young 118, 129 (UH). Solanaceae Solanum nelsoni Dunal. Specimens from Grass, North, and Southeast Islands; known also from Seal Island. Caum 49 (BPBM); Lamoureux s.n. (UH); Long 2275, 2301 (UH); Sibley 4 (USNM); Wilder 5 (BPBM); Young 101, 125 (UH). Solanum nigrum L. Specimens only from Southeast Island; known also from Grass Island. Sibley 11 (USNM); Young 112 (UH). ‘Cucurbitaceae Sicyos caumii St. John. Specimens from Seal (type locality), North and Southeast Islands. Caum 42, 53 (BPBM); Galtsoff (USNM); Lamoureux s.n. (UH); Long 2280, 2303 (UH); Sibley 6 (USNM) as Cucumis sativus L.; Wilder 4 (BPBM); Young 117, 119, 121 (UH). Goodeniaceae Scaevola taccada (Gaertn.) Roxb. Specimens from Grass, North, and Southeast Islands; known from Seal Island. Caum 52 (BPBM); Galtsoff s.n. (USNM); Lamoureux s.n. (UH); Long 2265, 2281, 2287, 2304 (UH); Young 97, 127 (UH). Compositae *Sonchus oleraceus L. Specimens only from Southeast Island. Long 2290 (UH); Young 113 (UH). 284 Appendix Table 4a. Movements of Black-footed Albatross from Pearl and Hermes Reef Original Banding Data Recapture Data Band Number Date Age-Sex* Where Recaptured Date Age-Sex* Grass Island 737-257 24 06-26-63 N-U Ocean shores beach, 09-??-65 U-U east of Olympia, (found Washington dead ) 1767-42371 03-19-65 N-U At Sea, 33°23'N, 01-21-67 U-U 122°18'W (dead) 767 -4.2398 03-19-65 N-U At Sea, ca. 37°30'N 07-22-67 U -U 767 -4.24.23 03-19-65 N-U Near Chirikof I., 08-22-67 iMG Alaska (ca. 56'N, 156°W) Kittery Island 7137-25459 06-26-63 L-U At Sea 15 mi. S.W. O4-25-64 U-U of Point Sur, California 7737-25483 06-26-63 L-U At Sea, 59°18'N, 08-20-65 U-U 143°20'W in the (dead) Gulf of Alaska 737 -25496 06-26-63 L-U Off Nemuro Hok, 08-07 -67 U-wU Japan (ca. 43°10'N, (found 145°hO'R) dead) 1737-25945 06-26-63 L-U 15 mi. W. of Cres- 07-25-65 U-U cent City, Cali- fornia 1737-25951 06-26-63 L-U Pacific Coast of 08-06-66 U-U Hokkaido, Japan (dead) (42°16'N, 142°27'E) North Island 737-25 080 06-23 -63 L-U At Sea, 15 mi. W. of 07-09-65 U-U Crescent City, Cali- fornia *A = adult; I = immature; L = local; N - nestling; S - subadult; U = unknown. Appendix Table Ha. Original Banding Data Age-Sex Band Number North Island (3254.10 737 -25406 767 -41723 Seal Island 7737-25514 Date 06-24 -63 06-24 -63 03-12-65 06-26-63 Southeast Island 7377-31030 737-31216 737-31250 (e-31392 737-31463 737 -31464 737-31614 1737-31634 737-31666 02-26-63 02-27-63 02-27 -63 02-27-63 02-27-63 02-27-63 02-27 -63 02-27-63 02-27-63 *Recaptured by BSFW- (continued) 285 Recapture Data Where Recaptured Kure Atoll Trig I., French Frigate Shoals Kure Atoll At Sea, 32°25'N, 137°30'W Sand I., Midway Atoll At Sea, 31°20'N, 173°00'W At Sea, ca. 46°00'N, 147°10'W Near Waldport, Ore- gon (ca. 44°20'N, 124°00'w) Kure Atoll Kure Atoll Kure Atoll Kure Atoll Kure Atoll Date 01-15-69 03 -22-66% 03-03-69 03-09-64 02-25 -67* 05-12-66 07-05-69 06-28-69 12-13-65 12-13-65 01-24-65 12-14-65 01-14-69 Age-Se 286 Appendix Table 4a. (continued) Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-sex Southeast Island 737-31540 02-28-63 A-U At Sea, ca. 55°50'N, 07-21-67 A-U 154°20'W 737-31557 02-28-63 A-U Near Kaines I., 10-12-68 A-U British Columbia (ea. 50°20'N; 128° 00'W) 1737-31589 02-28-63 A-U Near Shumagin I., 08-2? -67 A-U Alaska (dead) 1737-31598 02-28-63 A-U Tillamook, Oregon o4-27-69 A-U (enki 5c20iN, (found 123°50'W) dead) 737-31818 02-28-63 A-U At Sea, ca. 55°50'N, 07-20-67* A-U 154°20'W 1737-32046 02-28-63 A-U On beach 12 mi. S. 07-04-66 A-U of Gray's Harbour, (dead) Washington 737-32068 02-28-63 A -U At Sea, 54°30'N, 10-03-63 A-U 133°50' SSW of For- rester Is., Alaska 1737-32151 02-28-63 A-U At Sea, 48°15'N, O4-29-65 A-U 126°30'W (dead) 1737-32257 02-28-63 A-U At Sea, 48°15'N, O4 -29-65 A-U 126°30'W 1737-32247 03-01-63 A-U Kure Atoll 02-06-65 A-U Kure Atoll 01-17-66 A-U Kure Atoll 11-29-68 A-U 1737-32248 03-01-63 A-U At Sea, 59°38'N 07-05 -65 A-U 1737-32287 03-01-63 A-U At Sea, 46°31'N, 05-01-63 A-U 131°57'W (dead) *Recaptured by BSFW. Appendix Table 4a. (continued) Original Banding Data Band Number Date Southeast Island 7737-34035 1737-34307 7737-34316 1737-32461 1737-32463 737 -32499 1737-34235 737-34738 (ai—327e3 7137-32540 737 -33087 737 -33260 1737-33300 737 -34992 03-01-63 03-02-63 03-02-63 03-03-63 03 -03 -63 - 03-03-63 03-03-63 03-03-63 03-04-63 03-04-63 03-04-63 03-04-63 03-04-63 03-04-63 Age-Sex A A 287 Recapture Data Where Recaptured Date Age-Sex Kure Atoll 11-08-68 A-U At Sea, 23°20'N, 04-22-66 A-U 176°40'W, 280 Mi. SE of Midway Atoll S. of Port Oxford, 06-03-69 A -U Oregon (ca. 42°30'N, (found 124°20'w) dead) Kure Atoll 01-24-65 A-U Kure Atoll 12-10-65 A-U (with egg) Kure Atoll 01-14-69 A-U At Sea, ca: 25°N, 05-16-66 A-U 155 At Sea, 4o°ho'n, 07-20-66 A-U 23 mi. NE of Hachihote, Japan 59°00'N, 140°50'E, 06-77-63 A-U near Amka, U.S.S.R. (dead) Kure Atoll 01=25 -65 A-U At Sea, ca. 27°10'N, 05-05-68 A -U 170°10'W Kure Atoll 02-07-66 A-U 20imi. S. of Ti- 06-26-64 U-U juana, Mexico At Sea, 31°27'N, O4-2h -65 U-wU 166°08'E Cape Lookout, Ore- 08-01-69 A-U gon (ca. 45°20'N, 124°00'w) 288 Appendix Table 4a. (continued) Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex Southeast Island 1737-32522 03-04-63 I-vU At Sea, ca. 30°N, 02-06-66 U-U 140°W 1737-32667 03-06-63 A-U Kure Atoll 12-07-65 A-U Kure Atoll 01-14-69 A-U 7737-32686 03-06-63 A-U At Sea, 24°30'N, 11-25 -64 A-U 151°25 'W 737 -35199 03-06-63 A-U At Sea, 57°54'N, 09-15 -63 A-U 180°54', near the (dead) Pribilof I. 1737-32711 03-07-63 A-U 15 mi. SE of Cres- 07-15-67 A-U cent City, Cali- fornia 1737-35298 03-07-63 A-U Kure Atoll 12-06-65 A-U Kure Atoll 12-01-68 A-U (with egg) 1737-35445 03 -07 -63 A-U Beach at Ilwaco, 07-14-63 U-U Washington ca. (found 46°10'N, 124°00'w dead) 1737-32806 03-08-63 L-U East I., French 06-10-69 A-U Frigate Shoals 1737-32860 03-08-63 L-U MtuSea sms 50 N, 05-16-67 U-U 174°30'W (dead) 737 -32867 03-08-63 Ib = WY At Sea, ca. 35°N, 06-25 -64 Uy =U) 140°W 737 -32966 03-08-63 N-U On beach, Santa Mon- 07-07-65 Ua ica, California (found (BiessIes 03-08-63 L-U At Sea, 22°2k'N, 03-24-64 U-U 25) (05)! (dead) — Appendix Table 4a. (continued) Original Banding Data Age-Sex Band Number Date Southeast Island 737-33796 03-08-63 i = 737-33990 03-08-63 agen 737-33991 03-08-63 L-U 737-37637 06-20-63 aw 1737-39019 06-20-63 L - U 737-39178 06-23 -63 N-wvU Appendix Table 4b. Hermes Reef Original Banding Data Band Number Date Age-sex At Sea, ca. 43°N, 136°W 44 -711603 09-05 -45* At Sea, ca. 50°, 145°W 509-03279 05-14-54 527-65504 07-04-54 *Banded by Yocom (see Yocom, 1965: 187-188). 7377-31376. 289 Recapture Data Where Recaptured Date Age-Sex Sultan Lake, Ore- 04-30-69 U-U gon (ca. Lheoo'n, (found 124°00'W) dead) Terra de Mar, Pa- 05-29-65 U-wvU cific City, Oregon At Sea, 43°00'N, 08-01-67 U-wU 142°20'W (dead) Kure Atoll 03-16-66 A-U (dead) Ford Ord, Califor- 07-19-64 U-U nia ca. 36°30'N, (found 121°h0'w dead) Kure Atoll 12-10-68 Bs 1) (with egg) Movements of Black-footed Albatross to Pearl and Recapture Data Where Recaptured Southeast I. Southeast I. Southeast I. Band Date Age-Sex 02-27-63 A-U 02-28-63 A-U 03-13-64 A-U replaced on 02-27-63 with 290 Appendix Table 4b. Original Banding Data Band Number Date (continued ) Age-Sex French Frigate Shoals, Hast I. 737-37004 Kure Atoll 737-45767 737-93980 737-937 06 737-93740 1737-92652 737-92654 Midway Atoll, Eastern I. 587-58199 597 -69479 587 -72834 597-27231 597-27234 597 -27286 597-5 2847 597 -52912 697 -73944 697 -72553 06-08-63 11-16-63 01-17-64 01-21-64 02-12-64 01-31-65 01-31-65 06-03 -57+ 07-01-59 04 -06-60+ O4-06-60+ 04-06 -60+ 04-06 -60+ 04-06 -60+ O4-06-60+ 02-22-63+ 02-25 -63+ *Recaptured by BSFW. +Banded by BSFW. L U eS Cf «4 (=| Recapture Data Where Recaptured Southeast Southeast Southeast Eastern I. Atoll Southeast North I. Southeast Southeast Southeast Southeast Southeast Southeast Southeast Southeast Southeast Southeast Southeast Southeast LT. Date 03-21-67* 03-13-64 03-13-64 02-17-64 03-13-64 03-17-65 03-21-65 03-21-65 02-27-63 03-03-63 03-07 -63 03-04-63 02-26-63 03-13-64 03-06-63 03-01-63 03-16-65 03-16-65 Age-Sex eq eG € Gq U U U U U U U U Appendix Table 4b. Original Banding Data Age-Sex Band Number Date Midway Atoll, Sand I. DoH (250) DSM Zi Ls) 587-27157 5ST =z 0 Appendix Table 5. Original Banding Data Age-Sex Band Number Grass Island 737 -26473 71737-29019 71737-26942 O4-06-60+ O4 -07 -60+ O4-10-60+ O4 -10-60+ Date 06 -26-63 06-26-63 06-27 -63 Kittery Island 7737-24258 06-26-63 Little North Island 1737-38558 North Island 737-4797 737-24827 737 -24833 06-24-63 06-23 -63 06-23-63 06-23-63 +Banded by BSFW. (continued) 291 Recapture Data Where Recaptured Southeast I. Southeast I. Southeast I. Southeast I. Date 03-01-63 02-28-63 03-06-63 02-27-63 Age-Sex A-U A-U A-U A-U Movements of Laysan Albatross from Pearl and Hermes Reef Recapture Data Where Recaptured Kure Atoll Kure Atoll Kure Atoll Kure Atoll Kure Atoll Kure Atoll Kure Atoll At Sea, 29°45'N, On 50! E Date 05-14-69 05-20-69 O4-17-66 03-25-69 03-25-69 05 -06-69 02-25 -69 02-19-64 Age-sex A-U A-U U-U (dead) A-U A-U A-U A-U U-U 292 Appendix Table 5. (continued) Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex North Island 1737-24841 06-23 -63 N-U AGASea, 137720" N, 12-20-65* U -U AGN GLO Ms 1737-24905 06-23 -63 L-U Eastern I., Midway 04-01-68 U-U Atoll (found dead ) 1737-24153 06-24-63 L-U At Sea, 42°52'N, 05-13-64 U-wU 153° 2k 'E 757-17198 03-17-65 N-U Kure Atoll 03-10-69 A-U Seal Island 1737-24310 06-26-63 N-U Kure Atoll 02-24-69 A-U 737 -24365 06-26-63 L-U At Sea, ca. 36°N, 05 -13-64 U-U 142°R Southeast Island 737 -36039 03-06-63 N-U At Sea, ca. 37°N, 12-27-64 U-U 151°E 1737-36223 03 -06-63 N-U Kure Atoll 04-17-66 U-wU (dead) 1737-36227 03-06-63 N-U Kure Atoll o4-01-69 A-U 1737-36277 03-06-63 N-U Kure Atoll 02-25-69 A-U 1737-36527 03 -07 -63 N-U At Sea, 30°00'N, 01-04-64 U-U 150°43'E 1737-37637 06-20-63 N-U Kure Atoll 03-16-66 U-UvU 737 -37986 06-21-63 N-U Kure Atoll O4-11-69 A-U 757-18422 03-15 -65 N-U Kure Atoll 04-01-69 A-wU 1757-19488 03-15 -65 N -U Kure Atoll 02-24 -69 A-U Kure Atoll *Recaptured by BSFW. Appendix Table 5. (continued) Original Banding Data Band Number Date Age-Sex Southeast Island 757-19562 03-15-65 N - U 757 -20484. 03-16-65 N-U 71757-20828 03-15 -65 N-U 737-21168 03-16-65 N-U 1757-21409 03-16-65 N-U 1757-22320 03-16-65 N-U Appendix Table 6a. Movements Original Banding Data Band Number Date Southeast Island 723 -60810 7113-99525 Appendix Table 6b. 03-01-63 03-16-65 Age-Sex A-U A -U Original Banding Data Band Number Kure Atoll 793 -20559 1935-21293 793 -20381 *Recaptured Date 11-23 -64 03-10-65 10-21-64 by BSFW. Age-Sex KN 293 Recapture Data Where Recaptured Date Age-Sex Kure Atoll 02-24 -69 A-U Kure Atoll 05-05 -69 N - U Laysan I. 03-27-66 U-U to 03-30-66%* Kure Atoll O4-14 -69 A-U Laysan I. 03-27-66 U-U to 03-30-66% Kure Atoll 02-25 -69 A-U Recapture Data Where Recaptured Laysan IL. Kure Atoll Recapture Data Where Recaptured Seal I. Seal I. Kure Atoll Southeast I. of Bonin Petrels from Pearl and Hermes Reef Date — 09-19-64 09-10-66 Movements of Bonin Petrels to Pearl and Hermes Reef Date 03-19-65 03-18-65 11-23-65 03-16-65 Age-sex AN - U N10) Age-Sex A-U A-U A -U A-U 294 Appendix Table 7a. Movements of Blue-faced Boobies from Pearl and Hermes Reef Original Banding Data Recapture Data Band Number Date Age-sSex Where Recaptured Date Age-Sex Kittery Island 1737-26522 06-26-63 N -U Kure Atoll 06-29-67 A=-U 7137-26549 06-26-63 N-U Laysan I. 09-16-64 A-F Johnston Atoll 05-20-65 A-U Little North Island 558-83563 06-23 -63 N-U Laysan I. 03-09-65 A-F 587-91109 08-29-67 N-U Kure Atoll 07-25 -68 s-U North Island 558-83080 06-23 -63 A-U Whale-Skate I., 08-16-65 A-F French Frigate Shoals Whale-Skate I., 06-26-66 A-F French Frigate Shoals Whale-Skate I., 06-03 -67 A-F French Frigate Shoals 558-83100 06-23 -63 A-U Tistanski hs 03-13-65 A-F 558-83580 06-24 -63 N -U Lisianski IT. 09-02-67 A-F (in roost- ing club) 5558-83595 06-23-63 A-U Kure Atoll 11-10-63 U-U Seal Island 737 -26562 06-26-63 N-U Laysan I. 295 Appendix Table 7b. Movements of Blue-faced Boobies to Pearl and Hermes Reef Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex French Frigate Shoals 71757-26182 06-11-66 A-U Southeast I. 05 -28-67 A-U (found dead) Kure Atoll 697 -70633 08-06-62+ I -U Kure Atoll 10-09-63 Ss -U Southeast I. 03-13-64 Ss -U Kure Atoll 10-26-64 A-U Kure Atoll 11-24-65 A-F Kure Atoll 07-01-66 A-F (with nest- ling) Kure Atoll 05 -31-67 A-F (with 2 eggs) Kure Atoll 03-28-68 A-F (with egg) Kure Atoll 03-01-69 A-F (with 2 eggs) 1737-99590 09-24-66 N -U Southeast I. 05 -30-67 s-U Kure Atoll 11-30-68 A-U Kure Atoll 04-03-69 A-F +Banded by BSFW. 296 Appendix Table 8a. Movements of Red-footed Boobies from Pearl and Hermes Reef Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex Grass Island 587-80011 08-18-64 A-U Johnston Atoll 04-26-65 A-wU (found dead ) North Island 587 -80060 08-20-64 N-U East I., French 06-17-66 Ss -U Frigate Shoals 587-91131 08-29-67 Ss -U Kure Atoll 07-20-68 s-U Southeast Island 1737-30079 02-26-63 A-U Johnston Atoll 03-09-65 A -U Southeast I. 09-25 -66 1737-30093 02-26-63 S-U Kure Atoll 11-11-64 A-U Kure Atoll 06-30-68 A-U 1737-30100 02-26-63 A-U Kure Atoll 01-29-65 A-U Kure Atoll 06-09-67 A-U East I., French 06-07-68 A-U Frigate Shoals Kure Atoll 07-24-68 A-U 1737-30151 02-27-63 A -U Johnston Atoll 02-27 -66 A-F 737-30156 02-27-63 I-U Johnston Atoll 03-26-65 A-U 7737-30281 02-28-63 A-U Kure Atoll 06-07 -66 A-U 737 -38050 06-19-63 N - U Lisianski TI. 06-17-66 U -U 7137-38051 O6=10=63) 9 =. U) Johnston Atoll 03-26-65 me Lisianski I. 09-01-67 A-U 737-38053 06-21-63 A-U Kure Atoll 07-15 -66 A-U Appendix Table 8a. Original Banding Data Band Number 757 -89355 757 -89390 757-43 066 587-85172 767 -4.0621 Appendix Table 8b. Date 08-17-64 08-17-64 09-25 -66 08-28-67 08-08-68 Original Banding Data Band Number Date (continued ) Age-Sex (A)- U A-U Ss =U a) (A)- U Age-Sex French Frigate Shoals, East I. 1757-26100 06-10-66 A = UI French Frigate Shoals, Trig I. 1757-27618 08-13 -66 Johnston Atoll 737 -43576 7137-44695 Kure Atoll 737 -45360 737 -45332 02-22-66 02-23-66 10-14-63 10-14-63 *Recaptured by BSFW. S = 297 Recapture Data Where Recaptured Date Age-Sex Johnston Atoll 05-04-65 (I)- U Lisianski I. 03-12-65 A-U Lisianski I. 09-04-67 s-U Johnston Atoll 03-19-69 sS-U Johnston Atoll 06-23-69 s-U Movements of Red-footed Boobies to Pearl and Hermes Reef Recapture Data Where Recaptured Date Age-Sex Southeast I. 03 -22-67 A-U (on egg) Southeast I. 05 -30-67 A-U Southeast I. 02-10-69* A-U Southeast I. 05 -31-67 A-U (with chick) Southeast I. 05-31-67 A-U Eastern I., Midway 12-27-67 U-wU North I. 08-20-64 s-U Kure Atoll 01-29-65 A-U Kure Atoll 10-03 -66 A-U 298 Appendix Table 8b. (continued) Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex Kure Atoll 1737-45332 10-14-63 A-U Southeast I. 08-28-67 A-U 1737-95942 12-12-64 A-U Southeast I. 02-10-69* A-U 767 -45935 07-09-66 A-U Southeast I. 03-23-68 A-U Laysan I. 1757-28129 10-21-66 A-U Southeast I. 08-28-67 A-U Midway Atoll, Eastern I. 767 -49147 06-22-66 A-U Southeast I. 03 -21-67* A-U Appendix Table 9a. Movements of Brown Boobies from Pearl and Hermes Reef Original Banding Data Recapture Data : Band Number Date Age-Sex Where Recaptured Date Age-Sex Southeast Island 1737-30106 02-26-63 A-U Southeast I. 06-19-63 A-U (with newly hatched young) Laysan I. 03-07-65 A-U Seal I., Pearl and 03-18-65 A-M Hermes Reef 1737-37516 06-19-63 N-U Kure Atoll 05 -20-66 A-M 737-37552 06-19-63+ N-U Funafuti I., Ellice 02-06-67 U-U TS ie (found dead ) 1737-37557 06-21-63 A-U Kure Atoll Ol. -20-66 A-M *Recaptured by BSFW. +Also banded with #737-37564 on 06-21-63. 299 Appendix Table 9a. (continued) Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-sex TESTO 06-21-63 A-U Southeast I. 09-20-66 A-M (with nest- ling) 1757-89307 08-17-64 s-U Wake I. 12-31-66 U-U 7157-89344 08-17-64 N-U Kure Atoll 06-04-67 A-M 587-85182 08-28-67 A-M Kure Atoll 06-03-69 A-M (with nest- (with 2 ling) eggs) Appendix Table 9b. Movements of Brown Boobies to Pearl and Hermes Reef Original Banding Data Recapture Data Band Number Date Age-sex Where Recaptured Date Age-Sex Kure Atoll 1737-98173 11-08-64 A-F Seal I. 03-18-65 A-F Southeast I. 03 -21-67* A-M Wake Island 7157-89729 04-27-65 A-U Southeast I. 09-26-66 A-M Appendix Table 10. Movements of Great Frigatebirds from Pearl and Hermes Reef Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date renee Grass Island 1737-30334 03 -05 -63 s -U Kure Atoll 05 -20-66 Ss -U 737-30348 03-05 -63 A-M Kure Atoll 05-22-66 A+-M 7377-30351 03-05 -63 s -U Kure Atoll 05 -22-66 s -U *Recaptured by BSFW. 300 Appendix Table 10. (continued) Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex Grass Island 1737-30359 03-05 -63 s-U Kure Atoll 06-20-66 s-U 737-26702 06-26-63 A-M Kure Atoll 07-31-68 A-M 1737-26707 06-26-63 A-M Kure Atoll 06-10-66 A-M 737 -26708 06-26-63 A-M Kure Atoll 06-11-69 A-M 737-26717 06-26-63 N-U Kure Atoll 05 -07-66 Ss -U 1737-26723 06-26-63 N-U Kure Atoll 05-15-66 Ss -U 1737-26739 06-26-63 N-U Kure Atoll 06-04-67 Ss -U (dead) 737-26741 06-26-63 N-U Johnston Atoll 01-29-69 Ss -U (dead) 737-26751 06 -26 -63 A-F Kure Atoll 06-17-69 A-F 1737-26752 06-26-63 A-M Kure Atoll 05 -25 -66 A-M 1737-26759 06-26-63 N-U Kure Atoll 06-05 -67 A-F Kure Atoll 06-17-69 A-F 1737-37396 06-26-63 N-U Kure Atoll 06-15 -66 s-U 587-80008 08-18-64 N-U Kure Atoll 07-20-68 Ss -F 587-80013 08-18-64 N-U Kure Atoll 07-02-67 Ss -U 587-80031 08-18-64 N-U Kure Atoll O4 -20-66 Ss -U 587-80040 08-18-64 L-U Manicomi I., off- 12-16-65 U-U shore Samar I., Philippine Is. (10°59'N, 125 °37'E) 587-80044 08-18-64 N-U Kure Atoll 04-20-66 Ss -U Kure Atoll 06-25 -67 S-F 301 Appendix Table 10. (continued) Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex North Island 7737-37138 06-23 -63 A-M Kure Atoll 05 -06-66 A-M 1737-37145 06-23-63 A-F Tubabao I., off- 12-11-65 A-F shore Samar I., Philippine Is. (1°03'N,125°42'E) 737 -37163 06-23 -63 N-U Kure Atoll 05 -12-66 s -U 737 -37164 06-23 -63 N-U Kure Atoll 05 -20-66 Ss -U 71737-37171 06-23 -63 A-M Kure Atoll 12-15-65 A-M 737-37178 06-23 -63 N-U Whale-Skate I., 09-29-65 I-vU French Frigate Shoals 1737-37196 06-24 -63 A-F Kure Atoll 05-26-68 A-F (dead) 587-80059 08-20-64 A-F Kure Atoll 07-10-66 A-F 587-80094 08-20-64 N-U Kure Atoll 05 -20-66 s-U 587-80100 08-18-64 N-U Johnston Atoll 03-11-67 I-U 587-80104 08-20-64 N-U Kure Atoll 06-04-67 Ss -U (dead) 587-80106 08-20-64 N - U Kure Atoll 04-23 -66 s-U 587-80108 08-20-64 N-U Kure Atoll 06-05 -67 Ss -U 587-80114 08-20-64 N-U Kure Atoll 06-05 -66 Ss -U 587-80116 08-20-64 N-U Kure Atoll 05-23-66 s-U 71767-42210 03-17-65 A-M Kure Atoll 05-14-69 A-M 587-91136 08-29-67 Ss -U Talibon, Bohol I., 01-77-68 U-U Philippine Is. Southeast Island 1737-30034 02-26-63 A-M Kure Atoll 05-31-68 A-M (with egg) ee eee ee 302 Appendix Table 10. (continued) Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date | Age-Sex Southeast Island | 1737-30253 02-28-63 S-U Kure Atoll 06-08-67 S-M Kure Atoll 05 -31-68 U-U (dead) 1737-30321 03 -02-63 A-M Kure Atoll 06-10-66 A-M 7137-37493 06-19-63 N-U Kure Atoll 06-20-66 S-U 1737-37123 06-21-63 A-M Kure Atoll 08-09-64 A-M Kure Atoll 05 -23-66 A-M 737-37124 06-21-63 A-U Johnston Atoll 10-04-66 A-F Southeast I. 05-30-67 A-F (wit egg) Southeast I. 07-03 -67 A-F (with week old chick) 737-30119 06-26-63 A-F Kure Atoll o4-21-64 A-F (dead ) 757-89301 08-17-64 N-U Kure Atoll 05 -06-66 Ss -U 757 -89354 08-17-64 N-U Kure Atoll 05-10-66 Ss -U 757-43 048 09-25 -66 N-U Kure Atoll 07-28-68 S-F 757-43056 09-25 -66 N-U Lisianski I. 09-02-67 s -U 1757-43070 09-25 -66 A-F Kure Atoll 06-27-68 A -F 587 -83832 05 -30-67 A-M Kure Atoll 06-09-67 A-M (with egg) 303 Appendix Table 11. Movements of Ruddy Turnstones to Pearl and Hermes Reef Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex Alaska, Pribilof Is., St. George 722-10282 08-04-66 A-U St. George, Pribi- 08-01-67 A-U lof I., Alaska Southeast I. 08-28-67 A-U (found dead ) 7222-14020 08-18-66 I-F Southeast I. 05 -28-67 U-wU (recov- ered) 1103 -02294 08-18-67 A-U Southeast I. 08-28-67 A-U (found dead ) Appendix Table 12a. Movements of Sooty Terns from Pearl and Hermes Reef Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex Southeast Island 1723-64552 06-20-63 A-U Kure Atoll 06-14-67 A-U (dead) 723 -64928 06-21-63 A-U Eastern I., Mid- 10-20-65 A-U way Atoll 723 -64.994 06-21-63 A-U Eastern I., Mid- 08-15 -64 A-U way Atoll 7153-43213 08-17 -64 I-vU Johnston Atoll 08-21-68 A-U 753-43868 08-17-64 A -U Lisianski I. 06-18-66 A-U 1753-44069 08-18-64 A-U Johnston Atoll O4 -23 -66 A-U 754-4797 08-18-64 A-U Tistanska i. 05-30-67 A-U (nesting) 753-44.982 08-18-64 A= U Kure Atoll 05 -20-66 304 Appendix Table 12b. Movements of Sooty Terns to Pearl and Hermes Reef Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex Johnston Atoll 1753-16049 08-06 -63 A-U Southeast I. 06-01-67 A-U (with egg) 1753-24199 09-15-63 A-U Southeast I. 08-28 -67 A-U 7153-24357 09-16-63 A-U Southeast I. 06-01-67 A-U (with egg) 7153-24662 09-19-63 A-U Southeast I. 08-17-64 A-F (collected) 7153-73704 03-09-64 A-U Southeast I. 06-01-67 A-U (with egg) 843 -99581 08-21-65 A-U Southeast I. 06-01-67 A-U (with egg) 933 -4.1833 08-21-66 A-U Southeast I. 06-01-67 A-U (with chick) Kure Atoll 813-91771 08-28-64 A-U Kure Atoll 06-03 -66 A-U (with egg) Southeast I. 06-01-67 A-U 813-91969 08-30-64 A-U Southeast I. 06-01-67 A-U (with egg) 793-20138 09-07 -64 A-U Southeast I. 05-30-67 A-U (with egg) 903 -085 19 06-22-66 A eG Southeast I. 05 -30-67 A =i (breeding) 305 Appendix Table 12b. (continued) Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-sex Kure Atoll 903 -14367 08-20-66 A-U Southeast I. 05 -28-67 A-U (with egg) Lisianski Island 943 -04396 06-17-66 A= & Southeast I. 05 -29-69* A-U Midway Atoll, Eastern Island 863 -07439 07-22-65 A-=—Ur-...-Southeastel. 03-22-67 A-U Appendix Table 13a. Movements of Black Noddies from Pearl and Hermes Reef Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex Southeast Island 632-20082 06-18-63 A-U Whale-Skate I., 08-30-65 A-U French Frigate Shoals 632-20751 06-21-63 A-U Whale-Skate I., 06-26-66 A-U French Frigate Shoals 65 2-4.2283 08-20-64 N-U Whale-Skate I., 08-13-65 A-U French Frigate Shoals 632-20573 06-18-63 A-U Ingisalayarsiiss IE 09-03-67 A-U 632-20761 06-21-63 A-U Lisianski I. 09-04-67 A-U 632-20731 06-21-63 A-U Kure Atoll 01-25-65 A-U (dead) 652-42330 08-18-64 A-U Sand I., Midway 03 -05 -66 U -U Atoll 65 2-4.2340 08-17-64 A-U Kure Atoll 05-11-66 A-U *Recaptured by BSFW. aa hn 306 Appendix Table 13a. (continued) Original Banding Data Recapture Data Band Number Date Age-Sex Where Recaptured Date Age-Sex Southeast Island 632-20558 06-18-63 A-U Kure Atoll 07-07-67 A-U Kure Atoll 07-30-68 A -U (dead) Appendix Table 13b. Movements of Black Noddies to Pearl and Hermes Reef Original Banding Data Recapture Data Band Number Date Age-sex Where Recaptured Date Age-Sex French Frigate Shoals, Whale-Skate Island 712-58386 06-26-66 A-U Southeast I. 05 -30-67 A-U Laysan Island 1723-60435 02-12-63 A-U Southeast I. 05-30-67 A-U Midway Atoll, Sand Island 662-05446 02-20-64 U-U Southeast I. 08-18-64 A-U *Banded by BSFW. x U.S. GOVERNMENT PRINTING OFFICE; 1974 O - 561-843 1+ fie OT 4 er ee Toms” 2 m “wow 2 Nee mm SSP 2 Ne ee ee z w = = u? = 7) ; NLILSNI NVINOSHLIWS, Sa Iyvyag EVER AR i ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWS SaI¥ \ le, = = Dy: : = =a = z = 9 Lip, B 2 U7G.5 Nw: aN Gee z of 77 = a Pp 7* = AS z = < \S 2 , GE = z z = Fie ll = Pe ale : = RARI ES SMITHSONIAN INSTITUTION (pO REESNT UNVINOSHETNS "3a 1YVdail LIBRARI ES SMITHSONIAN _ fy. ‘3 @ 2 é us a fea tity, oa 5 a b) 3 [ad 4 fa a & Ga! ce és = ao = . aj er 2 =) 2 a = h TALULSNI LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLINS S31 r = Ly eS = S = NS S i D 5 2 5 phy te 5 an ae > re = E GG = - ca Ee ea E YG a E 2 = as - Gy = = me = i Z ; EN tee : 5 : “INSTITUTION NOILNALILSNI NVINOSHLINS S3JIYVYEIT LIBRARIES SMITHSONIAN ID 7) z s (a7) z on z ae “ = = <= a ES << = =f Ws = < =] z = = =U? ay ode ANS x Zz Er Oo T= ro) Se LI ro) Ve ane k= oO § n 22) 7) 72) DY yj D WS ba) a ©} an (e) 2G O GF a5 je) ne Z = z Ee =a. = ye = i 3 2 z pate = z NVINOSHLINS SAIYVYGIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWS Salk |NLILSNI = Zp) = 7) S 7) = = RS uw = uw a uw 2 2 WX = < a “S = g Sl NSN x = = o = S = a =>) ys > =) > =) > 2 S = "se E — c x = a) ne” =e a = o — a = z o Zz Oo z a Zz om LALILSNI NVINOSHLINS, S3 luvyygia_LIBRARI ES SMITHSONIAN NOILNLILSNI_ NVINOSHLINS, SI : Oy : = | P = = @ = Z ty, S = z = Ls = VW 2 B SIN 8 g = a4 yg 2 : = z = 2 |, — baa > = —v a 5 x 8 S a = JRARIES SMITHSONIAN INSTITUTION NOILALILSNI_ NVINOSHLIWS LIBRARIES SMITHSONIAN a S Zz = me) = = a g & uw a < = = s Pa mS. = We = c ce “¢ pea 3S fo) = co = az 2 z 2 = g .NLILSNI NVINOSHLINS SSIYVYUGIT LIBRARIES SMITHSONIAN _INSTITUTION NOILNLILSNI NVINOSHLINS S31 S z S = z | 5 = E ig ty = = B : ee S P es] — = 4 | — E “Ty - FE 5 : Md z 3 IRARIES SMITHSONIAN NOILNLILSNI NVINOSHLINS S31YVUdIT_LIBRAR NVINOSHLINS S3IY¥VYaIT LIBRARIES SMITHSONIAN NVINOSHLINS S31uvyalt we wn z z z Zz | 4 = SS ass < INN = = z Zz z IW Xe = 6 fe) (2) 9 S Re 2 2 Z Z PY 2 = = E E | ae > = = = = f - 5 “ LA ie) n a” JLOLILSNI NVINOSHLINS S31uNVaaIT cp MITHSONIAN _ - NVINOSHLIWS Say i bs ae tp) = 2 = siete A i 2 See wu = rw] Oo uJ z AG uJ GN g = ipl jp 3 5 ZN 28 = : =i Z = 2K = = FQ, a 5 ee ws = A nee NS DSSS are fe each 0 SNS Shae ya Nas Swosne i _ SMITHSONIAN \MBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI_NVINOSHLINS Sa1uvugi7_LIBRARIES $3 Sie INS Sai NS 2) = a a ae) = = py, = = < We z 5 Nu it $735 = 3 = 3 SES Do WwW B Vf a B eB B 2 SAO" 2 FS ENS eel EA ily See Nas zt E = = 7. 2 =) ee S ae dae i ANLILSNI_NVINOSHLINS SAIUVYIT LIBRARIES SMITHSONIAN INSTITUTION NOILOLILSNI_NVINOSHLINS a
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