The Avifauna of Kosrae, Micronesia: History, Status, and ... etal... · cally low hu man pop u la...
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91 Pacific Science (2016), vol. 70, no. 1:91–127 doi:10.2984/70.1.8 © 2016 by University of Hawai‘i Press All rights reserved Kosrae, the easternmost high island of the Caroline islands of Micronesia (5° 19 ′ N, 162° 59 ′ E) and one of four states that compose the Federated States of Micronesia, is a 110 km 2 volcanic island surrounded by fringing coral reefs and a few small satellite islands (Figure 1). The mountainous interior is steep, rugged, and uninhabited, rising up to 630 m above sea level at Mount Finkol. Precipitation is high, averag ing 4,842 mm /yr (Western Regional Climate Center 2004). The vegetation is dom- inated by tropical rainforest in the mountains and mangroves along the coast (e.g., Maxwell 1982, Cole et al. 1999). Because of its histori- cally low human population (6,616 humans in 2010 with a density of 54.7 /km 2 ; Gorenflo 1993, Office of SBOC 2013), which is concen- trated along the coast, the environment of Kosrae remains relatively pristine. Segal (1989) provided an excellent treatise on the island’s history, culture, government, and economy, but some of his information on its natural his- tory is inaccurate. Kosrae is seldom visited by birders and or- nithologists because it is small, isolated, and The Avifauna of Kosrae, Micronesia: History, Status, and Taxonomy 1 Floyd E. Hayes, 2,5 H. Douglas Pratt, 3 and Carlos J. Cianchini 4 Abstract: Kosrae, the easternmost high island of Micronesia, is a 110 km 2 vol- canic island rising up to 630 m above sea level. It is seldom visited by birders and ornithologists because it is small, isolated, and lacks any previously recognized extant endemic bird species. We review the history of research on the island’s avifauna and summarize the status of each species, including documentation for six new species: Northern Pintail ( Anas acuta), Gray Plover ( Pluvialis squatarola), Common Snipe ( Gallinago gallinago ), Bristle-thighed Curlew ( Numenius tahitien- sis), Red-necked Stint ( Calidris ruficollis), and Whiskered Tern ( Chlidonias hy- brida). We discuss previously undescribed vocalizations of endemic taxa and provide online reference to recordings. We also present supporting evidence for the recognition of two taxa as full biological species: Kosrae Fruit Dove ( Ptili- nopus hernsheimi; formerly part of the P. porphyraceus complex) and Kosrae White- eye ( Zosterops cinereus, as a split from Gray-brown White-eye, Z. ponapensis). The avifauna of Kosrae includes 53 nat urally occurring species of birds of which 13 breeding residents are extant (2 endemic species, 4 endemic subspecies) and 2 are extinct (both endemic species), 21 are boreal migrants from breeding pop- ulations in the temperate Northern Hemisphere (including 11 exclusively Pale- arctic migrants and 3 exclusively Nearctic migrants), 5 are austral migrants from breeding populations in the temperate Southern Hemisphere, and 12 are visi- tors from breeding populations on tropical islands elsewhere in the Pacific. Two additional species have been introduced; one has a self-sustaining feral popula- tion and the other is extirpated. Because of the island’s low human population and relatively pristine environment, resident breeding birds are thriving with no serious threats to their sur vival at present, except for overhunting of the Mi- cronesian Imperial Pigeon ( Ducula oceanica oceanica). 1 Manuscript accepted 4 September 2015. 2 Department of Biology, Pacific Union College, 1 Angwin Avenue, Angwin, California 94508. 3 North Carolina Museum of Natural Sciences, 11 West Jones Street, Raleigh, North Carolina 27601 (e-mail: [email protected]; website: www.hdouglaspratt .com). 4 P.O. Box 495, Kosrae, Federated States of Microne- sia 96944 (e-mail: [email protected]). 5 Corresponding author (e-mail: [email protected]).
Transcript of The Avifauna of Kosrae, Micronesia: History, Status, and ... etal... · cally low hu man pop u la...
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Pacifi c Science (2016), vol. 70, no. 1:91–127 doi:10.2984/70.1.8© 2016 by University of Hawai‘i PressAll rights re served
K osrae, the east ern most high island of the Car o line is lands of Micronesia (5 ° 19 ′ N, 162 ° 59 ′ E) and one of four states that com pose the Federated States of Micronesia, is a 110 km 2 vol ca nic island surrounded by fring ing coral
reefs and a few small sat el lite is lands ( Figure 1 ). The moun tain ous in te rior is steep, rug ged, and un in hab it ed, ris ing up to 630 m above sea level at Mount Finkol. Precipitation is high, av er ag ing 4,842 mm / yr (Western Regional Climate Center 2004). The veg e ta tion is dom-i nated by trop i cal rainforest in the moun tains and man groves along the coast (e.g., Maxwell 1982, Cole et al. 1999). Because of its his tor i-cally low hu man pop u la tion (6,616 hu mans in 2010 with a den sity of 54.7 / km 2 ; Gorenfl o 1993, Offi ce of SBOC 2013), which is con cen-trated along the coast, the en vi ron ment of Kosrae re mains rel a tively pris tine. Segal (1989) pro vided an ex cel lent trea tise on the island ’ s his to ry, cul ture, gov ern ment, and econ o my, but some of his in for ma tion on its nat u ral his-tory is in ac cu rate.
Kosrae is sel dom vis ited by bird ers and or-ni thol o gists be cause it is small, iso lat ed, and
The Avifauna of Kosrae, Micronesia: History, Status, and Taxonomy 1
Floyd E. Hayes , 2,5 H. Douglas Pratt , 3 and Carlos J. Cianchini 4
Abstract: Kosrae, the east ern most high island of Micronesia, is a 110 km 2 vol-ca nic island ris ing up to 630 m above sea lev el. It is sel dom vis ited by bird ers and or ni thol o gists be cause it is small, iso lat ed, and lacks any pre vi ously rec og nized ex tant en demic bird spe cies. We re view the his tory of re search on the island ’ s avi fauna and sum ma rize the sta tus of each spe cies, in clud ing doc u men ta tion for six new spe cies: Northern Pintail ( Anas acuta ), Gray Plover ( Pluvialis squatarola ), Common Snipe ( Gallinago gallinago ), Bristle - thighed Curlew ( Numenius tahitien-sis ), Red - necked Stint ( Calidris rufi collis ), and Whiskered Tern ( Chlidonias hy-brida ). We dis cuss pre vi ously undescribed vo cal i za tions of en demic taxa and pro vide online ref er ence to re cord ings. We also pres ent supporting ev i dence for the rec og ni tion of two taxa as full bi o log i cal spe cies: Kosrae Fruit Dove ( Ptili-nopus hernsheimi; for merly part of the P. porphyraceus com plex) and Kosrae White - eye ( Zosterops cinereus, as a split from Gray - brown White - eye, Z. ponapensis ). The avi fauna of Kosrae includes 53 nat u rally oc cur ring spe cies of birds of which 13 breed ing res i dents are ex tant (2 en demic spe cies, 4 en demic sub spe cies) and 2 are ex tinct (both en demic spe cies), 21 are bo real mi grants from breed ing pop-u la tions in the tem per ate Northern Hemisphere (in clud ing 11 ex clu sively Pa le-arc tic mi grants and 3 ex clu sively Ne arc tic mi grants), 5 are aus tral mi grants from breed ing pop u la tions in the tem per ate Southern Hemisphere, and 12 are vis i-tors from breed ing pop u la tions on trop i cal is lands else where in the Pacifi c. Two ad di tional spe cies have been in tro duced; one has a self - sus tain ing fe ral pop u la-tion and the other is ex tir pat ed. Because of the island ’ s low hu man pop u la tion and rel a tively pris tine en vi ron ment, res i dent breed ing birds are thriv ing with no se ri ous threats to their sur vival at pres ent, ex cept for over hunt ing of the Mi-cro ne sian Imperial Pigeon ( Ducula oceanica oceanica ).
1 Manuscript ac cepted 4 Sep tem ber 2015. 2 Department of Biology, Pacifi c Union College , 1
Angwin Avenue, Angwin, California 94508. 3 North Carolina Museum of Natural Sciences, 11
West Jones Street, Raleigh, North Carolina 27601 (e - mail: dpratt14 @ nc.rr.com; website: www.hdoug laspratt .com ).
4 P.O. Box 495, Kosrae, Federated States of Microne-sia 96944 (e - mail: cjcianchini @ ya hoo.com).
5 Corresponding au thor (e - mail: fhayes @ puc.edu).
92 PACIFIC SCIENCE · Jan u ary 2016
un til now lacked any pre vi ously rec og nized ex tant en demic bird spe cies. The avail able in-for ma tion on the avi fauna of Kosrae is widely scattered in the tech ni cal lit er a ture and has been dom i nated by re ports of new or un usual spe cies, which of ten neglected to re port de-tails of pre vi ously recorded spe cies. The only sum ma ries of the avi fauna since Baker ’ s (1951) com pre hen sive re view of the birds of Micro-nesia in clude fi ve check lists with a few sym-bols denoting the sta tus or a lit er a ture source for the oc cur rence of each spe cies on Kosrae (Owen 1977, Pyle and Engbring 1985, Pratt et al. 1987, Engbring et al. 1990, Wiles 2005). As a con se quence, it is cum ber some for re-search ers to ex tract in for ma tion on the sta tus of the island ’ s birds. The pur pose of this pa-per is to pro vide a de tailed sum mary of or ni-tho log i cal his tory and over view of what is
known of the avi fauna of Kosrae, in clud ing doc u men ta tion for six new spe cies for the is-land. We also dis cuss pre vi ously undescribed vo cal i za tions of en demic spe cies and sub spe-cies, and we pres ent supporting ev i dence for the rec og ni tion of three taxa as full bi o log i-cal spe cies.
meth ods
We reviewed the per ti nent lit er a ture on the birds of Kosrae, gleaning data on col lec tors and ob serv ers, the ranges of dates in which they vis it ed, their ob ser va tions of birds, re-ports of spec i mens col lect ed, the mu se ums in which spec i mens were de pos it ed, and the Kosraean names of birds. We also reviewed the avail able online cat a logs of spec i mens of var i ous mu se ums, in clud ing those in the
F igure 1. Map of Kosrae, Federated States of Micronesia.
Avifauna of Kosrae · Hayes et al. 93
ORNIS 2 da ta base (ornis2.ornisnet.org); how-ev er, not all mu se ums po ten tially hosting spec i mens from Kosrae have posted their cat-a logs online. We also searched for ar chived re cord ings of bird vo cal i za tions from Kosrae in the Macaulay Library of the Cornell Uni-versity Laboratory of Ornithology (macaulay-library.org) and the online source xeno - canto ( www.xeno-canto.org ), spon sored by the Xeno - canto Foundation with sup port from the Naturalis Biodiversity Center. Our online searches yielded pre vi ously un pub lished data for both skin and acous tic spec i mens.
We com piled a list of bird spe cies reported from Kosrae. We con sid ered a spe cies to be documented if its oc cur rence was based on ver i fi able phys i cal ev i dence such as a spec i-men, pho to graph, or sound re cord ing. Un-documented sight re cords were eval u ated based on the in for ma tion avail able to us. Each spe cies was placed into one of three catego-ries: (1) nat u rally oc cur ring spe cies, (2) in tro-duced spe cies, and (3) hy po thet i cal spe cies whose pres ence on Kosrae has been in ad e-quately documented. Terms for cur rent abun dance in clude “ com mon ” (recorded dai ly), “ un com mon ” (recorded less than daily with fi ve or more re cords), and “ rare ” (less than fi ve re cords). Status terms in clude “ ex-tinct ” (no lon ger liv ing), “ mi grant ” (an nual mi gra tory move ments, non breed ing), “ res i-dent ” (breed ing), and “ vis i tor ” (non breed ing va grant from is lands in trop i cal Pacifi c). Mi-gratory birds are de fi ned by the bio geo graph-i cal realm or re gion in which they breed (Hayes 1995): “ aus tral mi grant ” (tem per ate Southern Hemisphere), “ Aus tra lian mi grant ” (Australia or New Zealand), “ bo real mi grant ” (tem per ate Northern Hemisphere), “ Ne arc tic mi grant ” (tem per ate North America), and “ Pa le arc tic mi grant ” (tem per ate Eurasia). In the case of sea bird re cords with published co-or di na tes, dis tances from Kosrae were recal-culated us ing Google Earth.
A sum mary of the avi fau na, in clud ing En-glish, sci en tifi c, and Kosraean names; abun-dance; sta tus; and type of doc u men ta tion, is pro vided in Table 1 . Unless oth er wise spec i-fi ed, tax on omy and En glish names fol low Gill and Donsker (2015). Our tax o nomic in no va-tions are based on cri te ria for spe cies lim its
among in su lar pop u la tions pro posed by Pratt (2010), with a fo cus on po ten tial iso lat ing mech a nisms, geo graphic dis tance (i.e., de gree of iso la tion), qua si - clinal var i a tion, and com-par i sons with closely re lated spe cies. Popula-tions that have ob vi ous po ten tial iso lat ing mech a nisms in two or more qual i ta tively dif-fer ent categories are con sid ered spe cies un less proven oth er wise. This view con sid ers ge netic data in for ma tive but not es sen tial to tax o-nomic de ci sions and fol lows Gill ’ s (2014) re-cent pro posal to shift the bur den of proof in spe cies - level tax on omy of birds. All tax o nomic com ments herein were con trib uted by H.D.P.
Acronyms for mu se ums and sound ar chives are as fol low: AMNH = Amer i can Museum of Natural History, New York, New York; BPBM = Bernice Pauahi Bishop Museum, Ho-nolulu, Hawai ‘ i; DMNH = Del a ware Mu-seum of Natural History, Wilmington, Del a ware; FMNH = Field Museum of Na-tural History, Chicago, Il li nois; ML = Ma-caulay Library, Cornell Lab of Ornithology, Ithaca, New York; MCZ = Museum of Com-parative Zoology, Harvard University, Cam-bridge, Mas sa chu setts; MNHN = Mus é um National d ’ Histoire Naturelle Zoologie, Par is, France; MTD = Senckenberg Naturhistorische Sammlungen Dresden, Dresden, Germany; NSMT = National Museum of Nature and Sci-ence, Tokyo, Japan; RMNH = Nationaal Na-tuurhistorisch Museum, Leiden, Netherlands; SMF = Senckenberg Naturmuseum Frank-furt, Frankfurt, Germany; USNM = United States National Museum, Washington, DC; UWBM = Burke Museum, University of Washington, Seattle, Washington; XC = Xeno - canto Foundation ( www.xenocanto.org ); YIO = Yamashina Institute of Ornithology, Abiko, Japan; YPM = Peabody Museum of Natural History, Yale University, New Haven, Con-necticut; ZIN = Zoological Museum of the Zoological Institute of the Rus sian Academy of Sciences, St. Petersburg, Russia.
re sults
Ornithological History
The fi rst sci en tifi c stud ies of Kosrae ’ s avi fauna were conducted by French bi ol o gists Ren é
TA
BL
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Sum
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the
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and
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Bir
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tlet
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s in
quie
tus
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kie
f,f Kal
kecf
,g Kal
kafd K
alka
cfe
CR
, E2
SpM
i cro
ne si
an M
yzom
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omel
a ru
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raSc
husc
h,f S
ruus
r,g S
rusr
,d Sri
hsre
CR
, E2
SpK
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hite
-eye
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tero
ps c
iner
eus
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ahm
,f Tuw
ram
,g Tru
m,d T
uhra
me
CR
, E1
SpM
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ne si
an S
tarl
ing
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onis
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caU
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,g We,
d Wac
eC
R, E
2Sp
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rae
Star
ling
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onis
cor
vi n
aK
ako-
wec
gX
R, E
1Sp
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e-fa
ced
Parr
otfi n
chE
ryth
rura
tri
chro
aSc
hisc
h na
weh
,f Sru
usr
pecl
ang,
g Sru
sr-e
n-ek
u,d
Srih
sr in
eku
eC
R, E
2Sp
a Abu
ndan
ce: C
= c
om m
on, U
= u
n com
mon
, R =
rare
, X =
ex t
inct
.b S
tatu
s: A
M =
aus t
ral m
i gra
nt, B
M =
bo r
eal m
i gra
nt, E
1 =
en de
mic
spe
cies
, E2
= en
dem
ic s
ub sp
e cie
s, N
M =
Ne a
rc ti
c m
i gra
nt, I
= in
tro d
uced
, PM
= P
a le a
rc ti
c m
i gra
nt, R
= re
s i de
nt,
V =
vis
i tor.
c Doc
umen
tati
on: P
h =
pho t
o gra
ph, S
i = si
ght,
Sp
= sp
ec i m
en.
d San
tos
et a
l. (1
970)
, Seg
al (1
985,
198
9), E
ngbr
ing
et a
l. (1
990)
.e Y
i and
Lee
(197
6).
f Cou
ltas
(193
1).
g Hill
(195
7).
96 PACIFIC SCIENCE · Jan u ary 2016
Primev è re Lesson and Prosper Garnot, who vis ited Kosrae dur ing 5 – 15 June 1824 with a French ex pe di tion aboard the cor vette La Co-quille, re cord ing at least 10 spe cies (Lesson 1829 – 1830 a, b, 1831). Lesson de scribed two new spe cies from Kosrae (Lesson and Garnot 1826, Lesson 1827). Oustalet (1895) suspected that none of Lesson ’ s spec i mens from Kosrae were pre served, which was the case with one of the new spe cies that he de scribed, but three spec i mens of two spe cies from Kosrae are listed online in the MNHN, and at least two spec i mens from Kosrae are in the RMNH (Hoek Ostende et al. 1997, Dekker and Quaisser 2006).
Freidrich Heinrich von Kittlitz, a Rus sian bi ol o gist, vis ited Kosrae from 8 De cem ber 1827 to 1 Jan u ary 1828 with a Rus sian ex pe-di tion aboard the La Seniavine (Kittlitz 1832 – 1833, 1835, 1836, 1858). Kittlitz recorded 19 spe cies, of which 17 were col lected and de pos-ited in the ZIN. Some spec i mens were sold or traded to the SMF (Steinbacher 1954) and the RMNH (Mees 1964, Dekker and Quaisser 2006). Six new spe cies were de scribed from Kosrae, two of which were never col lected again.
Friedrich Hermann Otto Finsch, a Ger-man bi ol o gist, col lected birds on Kosrae from 20 to 29 Feb ru ary 1880 (Finsch 1880 a, 1881). Finsch ob served 18 spe cies, of which 14 were col lected and a new sub spe cies was de-scribed. Unfortunately, he did not state where his spec i mens were de pos it ed, but none are listed online at the SMF and the MTD, and none of the other po ten tial mu se ums in Ger-many and Holland have an online cat a log.
The online cat a log of the FMNH lists four spec i mens from the Louis B. Bishop Collec-tion that were obtained on Kosrae from 28 Jan u ary to 7 Feb ru ary 1899. It is un cer tain whether Bishop col lected the spec i mens him-self or obtained them from an other col lec tor.
Charles Haskins Townsend, an Amer i can bi ol o gist, col lected birds on Kosrae dur ing an ex pe di tion of the U.S. Fish Commission aboard the Albatross dur ing 7 – 9 Feb ru ary 1900. The spec i mens were de pos ited in the USNM (Townsend and Wetmore 1919), but a few were traded or sold to the MCZ (online cat a log).
Ernst Gotthilf Sarfert, a Ger man eth nol-o gist, vis ited Kosrae dur ing Feb ru ary to May 1910 as a mem ber of the South Sea Expedition spon sored by the Hamburg Scientifi c Foun-dation. Sarfert (1919) published an ec dotal ac-counts about the island ’ s most com mon bird spe cies and wrote ex ten sively about the meth-ods used to hunt birds.
Jap a nese bi ol o gists col lected birds on Kos-rae in the early 1900s (Taka - Tsukasa and Ku-roda 1915 a, b; Momiyama 1920, 1922; Kuroda 1922; Hachisuka et al. 1932, 1942), but no in-for ma tion was published on the col lec tors and dates of col lec tion. The online cat a log of the YIO lists a spec i men col lected by (Keisuke ? ) Kobayashi on 4 April 1915, an other spec i men by Toji Mishima on 23 Jan u ary 1929, and 69 spec i mens by Hyojiro Orii dur ing 2 – 9 Au gust 1931 (Orii ’ s name ap pears on only one la bel, but the hand writ ing ap pears to be the same on all other la bels). Two of Orii ’ s spec i mens were obtained by the FMNH and one by the YPM (online cat a logs). Six spec i mens at the NSMT were obtained by an un known col lec tor, pos-si bly Hyojiro Orii, dur ing 2 – 16 April 1931 and on 10 Sep tem ber 1931 (online cat a log).
William Coultas, an Amer i can bi ol o gist, vis ited Kosrae as a mem ber of the Whitney South Sea Expeditions of the AMNH from 15 Jan u ary to 11 June 1931 (Coultas 1931). He col lected 426 spec i mens of 22 spe cies for the mu se um, a few of which were traded or sold to the DMNH. Coultas pro vided the fi rst rel-a tively com plete list of Kosraean bird names. Some of his spec i mens from Kosrae were re-ported by oth ers (Bogert 1937, Mayr and Amadon 1941, Ripley and Birckhead 1942, Stickney 1943).
After World War II, nearly all bird ob ser-va tions were made by Amer i cans. Biologist Jo-seph T. Marshall Jr. vis ited Kosrae dur ing 23 – 27 No vem ber 1955 while study ing rats for the Pacifi c Islands Rat Ecology Project. He col lected four spe cies of birds for the USNM (Marshall 1957), but only two spec i mens of one spe cies are in the online cat a log.
Margaret Hill, a teach er, briefl y reported her ob ser va tions of birds and in ter views with lo cal in for mants from July 1956 to March 1957 (Hill 1957), al though a few of the spe cies she reported were al most cer tainly misidenti-
Avifauna of Kosrae · Hayes et al. 97
fi ed. She also in cluded the Kosraean names of birds.
John H. Brandt col lected two nests from Kosrae on 10 May 1959, which were de pos ited in the USNM (online cat a log). Brandt (1962) published a de tailed sum mary of nests and eggs col lected on Chuuk, but never published any thing on Kosrae.
Harvey Gordon Segal, an other teacher and ed u ca tion co or di na tor, re sided on Kosrae from 1964 to 1968 be fore mov ing per ma-nently to Pohnpei. Segal authored two books on the birds of Micronesia (Santos et al. 1970, Segal 1985) and a com pen dium on Kosrae (Se-gal 1989) that in cluded in for ma tion, al though some times in ac cu rate, on the island ’ s or ni tho-log i cal his tory and birdlife, in clud ing Kos-raean names and folk lore.
C. John Ralph vis ited Kosrae in Sep tem ber 1979 (pre cise dates not giv en) and col lected 22 spec i mens of six spe cies of birds that were de-pos ited at the USNM (online cat a log).
Floyd E. Hayes, a teach er, re sided in Taf-unsak from 27 Au gust 1981 to 1 Jan u ary 1982 and from 15 Jan u ary to 21 May 1982. He also trav eled by ship from Kosrae to Pohnpei dur-ing 1 – 4 Jan u ary 1982. Hayes published de tails for 11 new spe cies of birds for Kosrae (Hayes 1985) and data on mi gra tory shore bird pop u-la tions (Hayes 1986).
During the sum mer of 1983, a U.S. Fish and Wildlife Service (USFWS) team, in clud-ing Philip Ashman, John Engbring, Da vid Jickling, James Moore, and Pe ter Pyle, con-ducted a sur vey of the na tive for est birds of Kosrae from 30 June to 29 July 1983 (Pyle 1983, Pyle and Engbring 1987, Engbring et al. 1990). Using the var i able cir cu lar plot method for 8 min ute counts, they es ti mated the den-si ties and to tal pop u la tion sizes of birds on Kosrae based on 313 sta tions for a to tal of 26.35 km 2 . Their fi eld work added two new spe cies of birds for the island.
Mi chael Lauret, a phy si cian, vis ited the is-land from 28 Jan u ary to 8 March 1988 and re-ported three new spe cies of birds (Lauret 1990). Ornithologists H. Douglas Pratt and Robert L. Pyle briefl y vis ited Kosrae dur ing 24 – 26 Feb ru ary 1988 and ob served a few note wor thy birds (Lauret 1990, Wiles et al. 2000). Pratt recorded vo cal i za tions of eight
spe cies of birds, which were de pos ited at the ML.
During the First Joint U.S. – U.S.S.R. Cen-tral Pacifi c Expedition in 1988, Angela K. Kepler, Cameron B. Kepler, Da vid H. Ellis, and Jeffrey S. Hatfi eld stud ied sea birds from a ship while cruis ing from Hawai ‘ i to Singa-pore, pass ing south of Kosrae on 12 Oc to ber 1988 (Kepler et al. 1992). In an ex traor di-nary day, they added six new sea birds to the avi fauna of Kosrae, show ing how lit tle is known about the move ments of sea birds in Micronesia.
Visits by bird ers and or ni thol o gists have in creased in re cent years. Between 8 Feb ru ary and 5 March 1989, three spec i mens of birds from Kosrae were col lected by S. E. Shubel, Berlin Sigrah, and D. Tahlavan and were de-pos ited in the UWBM (online cat a log). Todd Mark vis ited dur ing 7 – 10 Oc to ber 2008 and recorded the vo cal i za tions of fi ve spe cies of birds. Carlos Cianchini has re sided on the is-land since 2008 and obtained pho to graphs of six new spe cies, one reported in Pratt et al. (2010) and the oth ers reported here in. Glenn McKinlay, a New Zealand bird er, vis ited Kos-rae dur ing 30 April to 3 May 2010 and photo-graphed a new bird (Buden and McKinlay 2010). W. Ross Silcock led a group of bird ers in clud ing Al Reyer, Bill Scheible, and Ken Lowder to Kosrae dur ing 6 – 10 June 2011 and conducted a pe lagic trip ap prox i ma tely 6 km out from Utwe on 7 June 2011 (reported here in). Pablo Oleiro, a grad u ate stu dent of Dylan Kesler at the University of Missouri, Co lum bia, co or di nated a bird sur vey in early 2012, but the re sults have yet to be published. Floyd E. Hayes briefl y returned to Kosrae dur ing 18 – 28 June 2013 and 17 – 27 June 2014 and photographed two new birds for the is-land (reported here in). Grady L. Pettigrew taught at the College of Micronesia in Kosrae from 9 Sep tem ber 2013 un til his un timely death on 17 Au gust 2014 and found a new bird (reported here in).
In re cent de cades, the avi fauna of Kosrae has been listed in sev eral sim ple check lists (Owen 1977, Pyle and Engbring 1985, Pratt et al. 1987, Engbring et al. 1990, Wiles 2005) that usu ally cite only the fi rst source for new re cords since Baker (1951). The fol low ing
98 PACIFIC SCIENCE · Jan u ary 2016
spe cies ac counts doc u ment sub se quent re-cords for mi gra tory vis i tors to re veal pat terns of oc cur rence.
Species Accounts
Northern Shoveler ( Anas clypeata )
Rare bo real mi grant. Hayes (1985) found eight fe males or im ma tures at Okat Harbor on 13 Oc to ber 1981. C.J.C. photographed fi ve of six fe males or im ma tures fl y ing over Lelu Harbor to ward Tofol on 4 Oc to ber 2014 and photographed three at Tofol on 11 Oc to ber 2014 ( Figure 2 ), pro vid ing the fi rst docu-mented re cord for Kosrae. This spe cies is a reg u lar vis i tor to most of the island groups of Micronesia (Wiles 2005).
Northern Pintail ( Anas acuta )
Uncommon bo real mi grant. Segal (1989: 220) listed this spe cies among three duck spe cies that are “ oc ca sional vis i tors ” but pro-vided no fur ther de tails. In late De cem ber 2009 (ex act date un known), Maxson Nithan reported to C.J.C. that two ducks had been found in Utwe. One of them, a fe male, died but was photographed ( Figure 3 A ) and the other disappeared the next day. On 13 Jan u-ary 2011, C.J.C. photographed two males (Figure 3 B ) on the grounds of the ath letic com plex in Lelu and was told by oth ers that
fi ve had been pres ent the week be fore. He photographed three males nearby at Foko Lik, Lelu, on 18 Jan u ary (Figure 3 C ) and 2 Feb ru-ary 2011. On 26 No vem ber and 1 De cem ber 2011, C.J.C. photographed two fe male - plum-aged in di vid u als near Tofol. The date is early enough that these birds could have been eclipse - plum aged males rather than fe males. C.J.C. and Grady Pettigrew ob served a fe male at Tofol on 14 March 2014. These re cords, in four suc ces sive win ters, in di cate that the Northern Pintail may be an an nual vis i tor to Kosrae in very low num bers. It is the most fre-quent win ter ing duck through out Micronesia (Wiles 2005).
Feral Chicken ( Gallus gal lus × sonnerati )
Introduced res i dent. Ornithologists have gen er ally regarded the Red Junglefowl ( G. gal lus ) as hav ing established fe ral breed ing pop u la tions on Kosrae and other is lands groups of Micronesia (Baker 1951, Owen 1977, Pyle and Engbring 1985, Pratt et al. 1987, Wiles 2005). Eriksson et al. (2008) re cently con fi rmed the hy brid or i gin of the do mes tic chick en, which com pli cates the tax on omy of fe ral pop u la tions through out the trop i cal Pa-cifi c. On Kosrae, Kittlitz (1836:284) stated that it “ runs wild ” (trans lated from Ger man). Finsch (1881:114) stated that they were “ the off spring of imported birds, which have run wild and have returned to the orig i nal plum-age. ” However, a half - cen tury lat er, Coultas
F igure 2. First photo documented re cord of Northern Shoveler ( Anas clypeata ) for Kosrae at Tofol on 12 Oc to-ber 2014. Photo by C.J.C.
F igure 3. First documented re cords of Northern Pintail ( Anas acuta ) for Kosrae at: ( A ) Utwe in late De cem ber 2009; ( B ) Lelu on 15 Jan u ary 2011; and ( C ) Lelu on 17 Jan u ary. Photos by C.J.C.
Avifauna of Kosrae · Hayes et al. 99
(1931:241) reported that it was “ in ter bred so badly with do mes tic strains ” that he was un-able to fi nd a “ good ex am ple ” and that the “ few that are found in a ‘ wild state ’ re main so close to hu man hab i ta tion that it is use less to con sider [it] a pure strain. ” He also “ never once en coun tered this bird in the moun tains or high moun tain val leys. ” Bequaert (1939, 1941) found the par a sitic fl y Ornithoctona pli-cata (Hippoboscidae) on one or more spec i-mens col lected by Coultas. During 1981 – 1982, F.E.H. never saw one > 100 m from a hu man home, al though he saw one shot by Oliod Moura in a man grove swamp > 100 m from the nearest house in Walung on 29 De cem ber 1981. Nevertheless, Engbring et al. (1990) found a few wild birds scattered rel a tively uni-formly in man groves and rainforest through-out the un in hab ited in te rior of Kosrae with a den sity of 2 / km 2 , in di cat ing that a self - sus tain ing fe ral pop u la tion still existed in July 1983. Since 2008, C.J.C. and F.E.H. have heard in di vid u als in un in hab ited ar eas.
breed ing: No da ta. spec i mens: AMNH (Peterson and Brisbin
2005), NSMT (online), YIO (online), ZIN (Kittlitz 1836).
Kermadec Petrel ( Pterodroma neglecta )
Rare aus tral mi grant off shore. Kepler et al. (1992) ob served two in di vid u als (pale and dark morphs) ap prox i ma tely 147 km (orig i nally re-ported as 157 km) south - south east of Kosrae (04 ° 03 ′ N, 163 ° 30 ′ E) on 12 Oc to ber 1988. On 7 June 2011, W. Ross Silcock et al. ob-served two all - dark, small tubenoses that might have been this spe cies sev eral ki lo me-ters off shore from Utwe. Elsewhere in Micro-nesia, it has been recorded as a vis i tor in Chuuk and the Marshall Islands (Wiles 2005).
Wedge - tailed Shearwater ( Puffi nus paci-fi cus )
Common vis i tor off shore. First reported with out de tails by Hachisuka et al. (1932), pre-sum ably based on a male dark - morph spec i-men obtained on 8 Au gust 1931 (YIO 00708). F.E.H. ob served 50 in di vid u als from a fi sh ing boat a few ki lo me ters west of Tafunsak on 8
Oc to ber 1981. All were dark - morph ex cept for a sin gle pale - morph in di vid u al. In the same ar ea, F.E.H. saw a doz en, in clud ing a sin gle pale - morph in di vid u al, on 12 Oc to ber 1981. The shear wa ters were feed ing with large fl ocks of noddies ( Anous spp.). On 2 Jan u ary 1982, F.E.H. ob served 63, in clud ing 15 pale - morph in di vid u als, at sea be tween about 85 km west - northwest of Kosrae and 50 km south - south east of Pingelap, mostly near the lat ter (Hayes 1985). Kepler et al. (1992) ob-served 29 dark - morph in di vid u als at sea south of Kosrae on 12 Oc to ber 1988. On 7 June 2011, W. Ross Silcock et al. ob served one Wedge - tailed Shearwater and prob a bly oth ers too dis tant to iden tify sev eral ki lo me ters off Utwe. On 21 Feb ru ary 2012, a pale - morph Wedge - tailed Shearwater was found on land in Utwe and brought alive to C.J.C., who pho-tographed it. This is the most com mon and wide spread shear wa ter through out Microne-sia (Wiles 2005).
spec i men: YIO (online).
Flesh - footed Shearwater ( Puffi nus carneipes )
Rare aus tral mi grant off shore. Kepler et al. (1992) ob served one in di vid ual about 167 km south - south east of Kosrae (03 ° 59 ′ N, 163 ° 46 ′ E) on 12 Oc to ber 1988. It has also been re-corded in the Marshall Islands (Wiles 2005).
Christ mas Shearwater ( Puffi nus nativitatis )
Rare vis i tor off shore. Pyle and Engbring (1987) ob served a sin gle in di vid ual about 12 km east of Lelu on 28 July 1982. On 7 June 2011, W. Ross Silcock et al. saw a small, all - dark shear wa ter that was prob a bly this spe-cies. It also vis its the Commonwealth of the Northern Marianas, Marshall Islands, and Wake (Wiles 2005).
Tropical Shearwater ( Puffi nus bailloni di-chrous )
Uncommon res i dent. Lesson ’ s (1829 – 1830 a: 432) “ p é trel d é sol é ” and Kittlitz ’ s (1858:358) “ Schwarzlicher Sturmvogel ” pre-sum ably re fer to this spe cies. Coultas (1931) obtained a spec i men that was trapped in a
100 PACIFIC SCIENCE · Jan u ary 2016
chicken coop on 25 April 1931 (Baker 1951). F.E.H. ob served fi ve in di vid u als from a fi sh-ing boat a few ki lo me ters west of Tafunsak on 8 Oc to ber 1981 and two more in the same area on 12 Oc to ber 1981. The USFWS team heard one fl y ing over the Yela River at night on 8 July 1983 (Pyle 1983, Engbring et al. 1990) and ob served four to six at sea about 15 km south east of Lelu on 28 July 1983 (Pyle and Engbring 1987, Engbring et al. 1990). Kepler et al. (1992) did not ob serve any dur ing their cruise through Micronesia in Oc to ber 1988.
breed ing: Coultas (1931) es ti mated that two or three dozen nested on two steep cliffs in the cen ter of the island, where na tives used to tie ropes of grass and lower them selves over the edge to cap ture the birds for food, but he and his as sis tants were un able to cap ture any dur ing an at tempt with ropes. Pyle (1983) and Engbring et al. (1990) suggested that one call-ing above the Yela River at night on 8 July 1983 was returning to its nesting site in the moun tains, but no nests were lo cat ed. It is un-known whether any still nest on Kosrae. This spe cies is recorded as a res i dent or vis i tor in most island groups of Micronesia (Wiles 2005).
spec i men: AMNH (Baker 1951).
Bulwer ’ s Petrel ( Bulweria bulwerii )
Rare vis i tor off shore. Kepler et al. (1992) ob served sev eral in di vid u als > 100 km south of Kosrae on 12 Oc to ber 1988. It has been re-corded in most of the island groups of Micro-nesia (Wiles 2005).
Wilson ’ s Storm Petrel ( Oceanites oceanicus )
Rare aus tral mi grant off shore. Kepler et al. (1992) ob served one in di vid ual ap prox i ma tely 145 km (orig i nally reported as 157 km) south - south east of Kosrae (04 ° 03 ′ N, 163 ° 31 ′ E) and ei ther a Wilson ’ s Storm Petrel or a Band - rumped Storm Petrel ( Oceanodroma castro ) ap-prox i ma tely 165 km south - south east of Kosrae (03 ° 59 ′ N, 163 ° 45 ′ E) on 12 Oc to ber 1988. Elsewhere in Micronesia, it has been re-corded as a vis i tor only in the Marshall Is-lands (Wiles 2005).
Tris tram ’ s Storm Petrel ( Oceanodroma tris-trami )
Rare vis i tor off shore. Hayes (1985) ob-served two large, fork - tailed storm pe trels at sea ap prox i ma tely 88 km (orig i nally reported as 84 km) west - northwest of Kosrae (5 ° 36 ′ N, 162 ° 10 ′ E) on 2 Jan u ary 1982 (and up to four dur ing sub se quent days in Pohnpei state). They were iden ti fi ed as “ most like ly ” Tris-tram ’ s Storm Petrel be cause no white was ob-served on the pri mary shafts, which is typ i cal of Matsudaira ’ s Storm Petrel. Subsequent au-thor i ties regarded them as Matsudaira ’ s Storm Petrel be cause at the time Tris tram ’ s Storm Petrel was not known to fol low ships and white in the pri mary shafts of Matsudaira ’ s Storm Petrel was con sid ered less con spic u-ous than the paler rump of Tris tram ’ s Storm Petrel (Pyle and Engbring 1985, Pratt et al. 1987, Wiles 2005). However, Tris tram ’ s Storm Petrel is now known to fol low ships (Yoshiki Watabe, kantorilode.weebly.com / dark - rum ped - storm - pe trels.html) and a re view of nu-mer ous online pho tos of both spe cies re veals that the white pri mary shafts of Matsudaira ’ s Storm Petrel are a dis tinc tive fi eld mark contrasting with the brown up per wing bar, whereas the slightly paler rump of Tris tram ’ s Storm Petrel is more eas ily overlooked as a fi eld mark. Based on the ob served fi eld marks, we now be lieve these birds were Tris-tram ’ s Storm Petrels. Matsudaira ’ s Storm Petrel has been recorded as a vis i tor in Micronesia east to Pohnpei (Wiles 2005) whereas Tris tram ’ s Storm Petrel has been documented in the Marianas, in clud ing Guam (D. Vice pers. comm.; de tails to be published else where), and by a fe male spec i-men found dead by Nancy Vander Velde on Majuro, Marshall Islands, on 7 April 2007 (BPBM 185080).
Red - tailed Tropicbird ( Phaethon rubri-cauda )
Rare vis i tor. Finsch ’ s (1880 a, b, 1881) ob ser-va tions of Red - billed Tropicbird ( P. aethereus ) at Kosrae dur ing Feb ru ary 1880 have been re-garded by oth ers as Red - tailed Tropicbirds
Avifauna of Kosrae · Hayes et al. 101
(Amerson 1969, Wiles 2005; more de tails pro-vided in Red - billed Tropicbird ac count within Hypothetical Species section). Lauret (1990) ob served sin gle in di vid u als on three oc ca sions near Okat Harbor in Feb ru ary 1988 and re-ported that HDP and R. L. Pyle ob served a fl ock of six in the same lo cal ity later that month; the birds appeared to be en gaged in court ship fl ights and vo cal ized loud ly, sug-gesting pos si ble breed ing, but none has been con fi rmed. This spe cies has been recorded from most island groups in Micronesia (Baker 1951, Wiles 2005) and has been expanding its breed ing range in re cent de cades (H.D.P., pers. obs.).
White - tailed Tropicbird ( Phaethon lepturus dorotheae )
Common res i dent. It oc curs in small groups through out the island and at sea, of ten fl y ing at great heights above the moun tains. During 1981 – 1982, F.E.H. ob served up to a dozen at a time at Okat and Utwe Harbors. On 2 Jan u ary 1982, F.E.H. ob served four far out at sea be tween Kosrae and Pingelap; the fi rst was recorded about 91 km west - northwest of Kosrae. Engbring et al. (1990) reported a mean den sity of 18 / km 2 with an es ti mated to-tal pop u la tion of 1,828; den si ties were high-est in man groves and neg a tively cor re lated with el e va tion, with none oc cur ring above 400 m ( Table 2 ), and were higher in Walung (34 / km 2 ) than in other re gions. Engbring et al. (1990) ob served 20 – 25 at a time soar ing above Okat Valley in Tafunsak dur ing July 1983. However, C.J.C. has not ob served more than fi ve at a time since 2008, suggesting that the pop u la tion has de clined.
breed ing: Finsch (1880 a, 1881) cap tured one on its nest in late Feb ru ary 1880. This spe cies is res i dent in most island groups of Mi-cronesia (Wiles 2005).
spec i mens: AMNH (Baker 1951), YIO (on-line).
Asian Cattle Egret ( Bubulcus coromandus )
Rare vis i tor. Lauret (1990) saw a sin gle in-di vid ual fl y over head and land in a pas ture
near two cows on 30 Jan u ary 1988, but did not re port the ex act lo cal i ty. The spe cies has in-creased in num bers and range in Micronesia in re cent de cades and is now an an nual vis i tor in large num bers in the west ern part of the re-gion (Wiles 2005, H.D.P., pers. obs.). Mi cro-ne sian cat tle egrets be long to the Indo - Pacifi c spe cies re cently split (Rasmussen and Ander-ton 2005) from the Af ro - Amer i can B. ibis, which is in tro duced in the Ha wai ian Islands (Pyle and Pyle 2009). The two are eas ily dis-tin guished in al ter nate (breed ing) plum age be cause B. coromandus has the en tire head and neck col ored ru fous - buff, but birds in ba sic plum age are prob a bly not dis tin guish able in the fi eld, even though B. coromandus has lon-ger legs, so Lauret ’ s sighting is iden ti fi ed on geo graphic like li hood rather than ac tual ob-served char ac ters. See Pratt (2014) for a dis-cus sion of En glish names of cat tle egrets.
Pacifi c Reef Heron ( Egretta sa cra )
Common res i dent. Micronesia is inhabited by the nom i nate sub spe cies E. s. sa cra (Baker 1951). It for ages along the coastal tidal fl ats, beaches, and man groves, but it also wan ders in land along streams and ditches. Coultas (1931) noted in di vid u als at the deep end of a cave (100 m from the en trance) on the north side of the island. Bequaert (1939, 1941) found the par a sitic fl y Ornithoctona plicata (Hippo-boscidae) on one or more spec i mens col lected by Coultas. During 1981 – 1982, F.E.H. ’ s max-i mum count was 23 at Okat Harbor on 18 April 1982. Mayr and Amadon (1941) stud ied plum age var i a tion among spec i mens, in clud-ing 49 from Micronesia (26 from Kosrae), and reported that 54 % in Micronesia were dark - morph, 40 % were white - morph, and 6 % were pied - morph. However, they pointed out that white - morph ju ve niles of ten have dark feath-ers, ren der ing it dif fi cult to iden tify gen u ine pied - morph in di vid u als, and that white - morph in di vid u als may have been pref er en-tially col lect ed. In Kosrae, Coultas (1931) reported that dark - morph and white - morph in di vid u als were equally com mon with a few pied - morph in di vid u als. Engbring et al. (1990) ob served ap prox i ma tely equal num bers of
102 PACIFIC SCIENCE · Jan u ary 2016
dark - and white - morph in di vid u als and did not fi nd any pied - morph in di vid u als. F.E.H. recorded the plum age of each sighting dur ing 1981 – 1982 ( n = 325) and 2013 ( n = 64) and found that most were dark - morph (64 % vs. 67 % ) with smaller per cent ages of white - morph (29 % vs. 19 % ) and pied - morph (7 % vs. 14 % , some prob a bly were white - morph ju-ve niles with scattered dark feath ers), suggest-ing that the pro por tion of morphs remained rel a tively sta ble dur ing three de cades. One bi-lat er ally asym met ri cal in di vid ual in 1983 – 1982 was com pletely white on one side and dark on the oth er.
breed ing: Segal (1989) stated that there were sev eral rook er ies around the island, but F.E.H. never saw any; al though, in June 2013, he noted her ons roosting over night in the man groves at Yen - Yen Island in Lelu. This spe cies oc curs through out Micronesia ex cept Wake (Wiles 2005).
spec i mens: AMNH (Mayr and Amadon 1941, Baker 1951), USNM (Townsend and Wetmore 1919, Baker 1951), YIO (online), ZIN (Kittlitz 1836).
Great Frigatebird ( Fregata mi nor )
Rare vis i tor. Finsch (1880 a, 1881) ob served one in late Feb ru ary 1880. Hill (1957) re-ported that it oc ca sion ally fl ies over head or chases noddies ( Anous spp.), but did not state whether she ac tu ally saw one. F.E.H. ob served sin gle in di vid u als, both fe males with a white head and chest, at Okat Harbor on 31 Oc to-ber 1981 and at Malem on 31 De cem ber 1981. This spe cies is a res i dent or vis i tor in most is-land groups of Micronesia, in clud ing nearby Pohnpei (Wiles 2005).
Lesser Frigatebird ( Fregata ariel )
Rare vis i tor. Philip Ashman (in Pyle 1983, Pyle and Engbring 1987) ob served a male and a fe male near Malem on 12 July 1983. C.J.C. photographed a dis tant fl ock of nine frigate-birds at Lelu on 11 April 2015, but they were too dis tant to iden ti fy. Two weeks lat er, C.J.C. photographed a male and a fe male at Okat Harbor on 25 April 2015, pro vid ing the fi rst documented re cord for Kosrae ( Figure 4 ).
Nonmangrove (Mostly Tropical Rainforest) Areas
SpeciesMan-groves
0–100m
100–200m
200–400m
400–600 m
0–630m Total
Feral Chicken 1 2 2 2 1 2 2White- tailed Tropicbird 82 4 8 9 0 30 66Brown Noddy 886 275 38 49 27 164 274Black Noddy 129 72 10 10 0 42 55White Tern 366 153 136 67 22 123 160Purple- capped Fruit Dove 44 72 90 106 42 83 77Micronesian Imperial- Pigeon
7 57 77 96 28 69 60
Island Swiftlet 118 332 344 276 381 322 290Micronesian Myzomela 3,975 1,325 665 550 495 973 1,433Kosrae White- eye 1,098 2,062 2,000 1,897 1,350 1,981 1,846Micronesian Starling 637 946 1,247 958 513 989 935Blue- faced Parrotfi nch 0 180 84 50 244 131 111
Source: Engbring et al. (1990).
TABLE 2
Densities (Individuals / Km2) of Bird Species in Mangroves and at Dif fer ent Elevations in Nonmangrove Areas (Mostly Tropical Rainforest) of Kosrae in July 1983
Avifauna of Kosrae · Hayes et al. 103
F igure 4. First photo documented re cord of Lesser Frig-atebird ( Fregata ariel ) for Kosrae at Okat Harbor on 25 April 2015. Photo by C.J.C.
This spe cies is a vis i tor in sev eral island groups of Micronesia (Wiles 2005), with the nearest known breed ing site at Helen Atoll, south-west ern Palau (Pratt and Etpison 2008).
Brown Booby ( Sula leucogaster )
Uncommon vis i tor. Coultas (1931:240) col-lected a spec i men on 19 April 1931 (date from Baker 1951) and noted in di vid u als roosting on a small is let at Walung, but reported that, “ ac-cording to the na tives, this spe cies has never been known to nest. ” During 1981 – 1982, F.E.H. recorded up to three in di vid u als, in-clud ing at least one im ma ture, on seven oc ca-sions from 8 Oc to ber 1981 to 16 May 1982. On 29 De cem ber 1981, F.E.H. found two adults sit ting among nesting Brown Noddies ( Anous stolidus ) in man groves be tween Okat Harbor and Walung, but they were not nest-ing. C.J.C. photographed an im ma ture just south of Okat Harbor on 10 No vem ber 2010 and saw an in di vid ual of un known age off Utwe on 28 De cem ber 2013. F.E.H. photo-graphed an adult off Utwe on 18 June 2014. This spe cies is a res i dent or vis i tor in all island groups of Micronesia (Wiles 2005).
spec i men: AMNH (Baker 1951).
Kosrae Crake ( Porzana monasa )
Extinct res i dent. Kittlitz (1836) col lected two spec i mens in De cem ber 1827, which he
ini tially iden ti fi ed as the Spotless Crake ( P. tabuensis; Kittlitz 1836) and later de scribed, al-though with some un cer tain ty, as a new spe-cies (Kittlitz 1858). They were found “ in the damp places in the mid dle of the for ests ” (trans lated from French; Kittlitz 1836:286) or “ on the ground in those places in the for est that are al ways deeply shaded and damp ” (trans lated from Ger man; Kittlitz 1858:31). Finsch (1880 a, 1881) searched for it un suc-cess fully in 1880, suggesting it may have al-ready been ex tinct by then; he ex am ined the spec i mens and be lieved they represented P. tabuensis . Hartlaub (1893) redescribed the spec i mens, dem on strat ing that they repre-sented a valid spe cies. Coultas (1931) in cluded a map of Kosrae that shows where he searched, re veal ing that he trav eled ex ten sively through low land and mon tane for ests. Coultas (1931:247) wrote that “ it is a god of the bush and much rev er enced by the peo ple. Several old sters seemed to re mem ber their fore fa thers speak ing of the bird, but none of them ad mit-ted hav ing seen it, ex cept an el derly dea con, a staunch pil lar of the church, who claimed to have had the bird pointed out to him some 20 years be fore my vis it. ” Kittlitz (1836, 1858) re-ported that its na tive name is setamanot. Coultas (1931:247) stated that its na tive name, nay tai mai not, means “ to land in the ta ro, ” suggesting that it lived in the swamps in which taro was cul ti vat ed. According to Hill (1957), King John, an 80 yr old ex - king, re mem bered that a Ger man sci en tist had col lected one in a cave in the moun tains about 50 yr ear li er, but none had been found since; it may have been a fuzzy mem ory of Finsch ’ s un suc cess ful search for it 77 yr ear li er. Extensive fi eld work at all el e va tions by the USFWS team failed to fi nd any in July 1983 (Engbring et al. 1990), yet Segal insisted that hunt ers have seen “ a ‘ chick en - like bird, ’ red dish brown in col or, that runs rather than fl ys [ sic ] when sight ed ” (Segal 1985:16), and later re it er ated that “ once in a while pig hunt ers re port a run ning for est bird high up in the moun tains ” (Segal 1989:219 – 220). Steadman (1986) published X - rays of the two spec i mens, which revealed that it was prob a bly fl ight less. Its ex tinc tion is usu ally at trib uted to pre da tion by in tro duced rats, which Kittlitz (1836:284) reported as
104 PACIFIC SCIENCE · Jan u ary 2016
abun dant and caus ing “ great dam age ” to ba-nana and sugar cane; in the words of Coultas (1931:247), “ in as much as the island is over-run with rats, one couldn ’ t ex pect to fi nd many ground birds. ”
breed ing: No da ta. spec i mens: ZIN (Kittlitz 1836, 1858;
Steadman 1986).
Watercock ( Gallicrex cinerea )
Rare vis i tor. Glenn McKinlay photo-graphed an in di vid u al, pos si bly a ba sic - plum-aged male, at Utwe dur ing 1 – 2 May 2010, pro vid ing the fi rst documented re cord for Oceania (Buden and McKinlay 2010). A pre-vi ous sight re cord from Palau was regarded as hy po thet i cal by Pratt et al. (1987).
Pacifi c Golden Plover ( Pluvialis fulva )
Common bo real mi grant. This is the third most com mon shore bird on Kosrae (Hayes 1986), for ag ing at sandy beaches, man grove swamps, grassy ar eas, and other open hab i tats. During 1981 – 1982, the max i mum count was 35 birds on 9 March 1982, with slightly lower num bers dur ing fall (up to 30) and win ter (up to 30; Hayes 1986). A few lin ger dur ing the sum mer. The USFWS team saw up to 12 at Okat Harbor on 1 July 1983 (J. Engbring in Hayes 1986) and F.E.H. saw up to 13 at Okat Harbor on 27 June 2014. This spe cies oc curs through out Micronesia (Wiles 2005).
spec i mens: AMNH (Stickney 1943, Baker 1951), FMNH (online), ZIN (Kittlitz 1836).
vo cal i za tions: ML (online).
Gray Plover ( Pluvialis squatarola )
Rare bo real mi grant. C.J.C. photographed an adult molting into ba sic plum age at Kos-rae Nautilus Resort, Lelu, on 1 – 3 No vem ber 2014 ( Figure 5 ), pro vid ing the fi rst re cord for Kosrae. It was dis tin guished from a Pacifi c Golden Plover by its larger size, grayer plum-age, and white tail when seen in fl ight. This spe cies oc curs through out Micronesia ex cept Wake (Wiles 2005).
Lesser Sand Plover ( Charadrius mongolus )
Rare Pa le arc tic mi grant. First recorded with out de tails by Hachisuka et al. (1932), pre-sum ably based on an adult fe male in prebasic molt col lected on 2 Au gust 1931 (YIO 00314). Hayes (1986) found two ba sic - plum aged birds at Okat Harbor on 3 Oc to ber 1981 and an al-ter nate - plum aged adult at Okat Harbor on 6 April 1982. F.E.H. photographed an al ter-nate - plum aged adult male of the nom i nate race at Okat Harbor on 20 June 2013. This spe cies oc curs through out Micronesia ex cept Wake (Wiles 2005).
spec i men: YIO (online).
Common Snipe ( Gallinago gallinago )
Rare Pa le arc tic mi grant. Grady Pettigrew and C.J.C. en coun tered a group of three snipes at Tofol on 29 March and four on 4 April 2014, representing the fi rst Gallinago re-ported on the island. On the lat ter oc ca sion, C.J.C. obtained a blurry photo of one fl y ing away ( Figure 6 ) that revealed a prominent, sharply de fi ned, white trailing edge to the sec-ondaries, which is a di ag nos tic fi eld char ac ter for Common Snipe (Hayman et al. 1986, Pratt et al. 1987, Message and Taylor 2005), pro vid-ing the fi rst re cord for Kosrae. On 4 No vem-ber 2014, C.J.C. found three more snipe at Tofol and recorded the calls of one on a vid eo; the calls, which sound some what like tear ing cloth, re sem bled those of Common Snipe (XC215455). Although a rare va grant in
F igure 5. First documented re cord of Gray Plover ( Plu-vialis squatarola ) for Kosrae at Kosrae Nautilus Resort, Lelu, on 1 No vem ber 2014. Photo by C.J.C.
Avifauna of Kosrae · Hayes et al. 105
F igure 6. First documented re cord of Common Snipe ( Gallinago gallinago ) for Kosrae at Tofol on 4 April 2014. Photo by C.J.C.
F igure 7. First photo documented re cord of Bar - tailed Godwit ( Limosa lapponica ) for Kosrae at Walung on 27 June 2013. Photo by F.E.H. Micronesia, this spe cies has been recorded
in most island groups (Wiles 2005, Vander-Werf 2006, Pratt et al. 2010).
Black - tailed Godwit ( Limosa limosa )
Rare Pa le arc tic mi grant. Hayes (1985, 1986) found a ba sic - plum aged in di vid ual at Okat Harbor on 13 De cem ber 1981. The US-FWS team saw and photographed an other at Okat Harbor dur ing 1 – 17 July 1983 (Pyle 1983, J. Engbring in Hayes 1986, Pyle and Engbring 1987; photo published in the lat ter). This spe cies oc curs through out Micronesia ex cept Wake (Wiles 2005).
Bar - tailed Godwit ( Limosa lapponica )
Uncommon Pa le arc tic mi grant. Hayes (1985, 1986) found two at Walung on 8 Sep-tem ber 1981 and reg u larly saw a few through-out the win ter of 1981 – 1982 be tween Okat Harbor and Walung, with up to 13 at Okat Harbor on 17 Oc to ber 1981 and smaller num-bers dur ing win ter (up to 6) and spring (up to 2). Pratt and Pyle ob served one at Okat Har-bor on 21 Feb ru ary 1988 (Wiles et al. 2000). A few lin ger dur ing the sum mer. The US-FWS team ob served three at Okat Harbor in July 1983 (Pyle 1983, J. Engbring in Hayes 1986, Pyle and Engbring 1987). F.E.H. pho-tographed a ba sic - plum aged adult at Walung on 27 June 2013 ( Figure 7 ), pro vid ing the fi rst
doc u men ta tion of the spe cies for Kosrae. C.J.C. photographed one of un known age near Treelodge Hotel, Putuk Te, Lelu, on 10 De cem ber 2014. This spe cies oc curs through-out Micronesia ex cept Wake (Wiles 2005).
Whimbrel ( Numenius phaeopus )
Common Pa le arc tic mi grant. Flocks of the East Asian sub spe cies N. p. variegatus (Baker 1951) for age among rocks and seagrasses on tidal fl ats dur ing low tide. Coultas (1931) noted fl ocks of 20 or more roosting at night on tiny is lets at Walung. During 1981 – 1982, Hayes (1986) ob served a max i mum of 100 (27 Sep-tem ber 1981) roosting on the branches of Pan-danus ( Pandanus sp.) and Coconut Palms ( Cocos nucifera ) on tiny Kiul Island at Okat Harbor; smaller num bers oc curred dur ing win ter (up to 35) and spring (up to 45). A few lin ger dur ing the sum mer, with up to 11 ob-served dur ing July 1983 (Pyle 1983, J. Eng-bring in Hayes 1986) and up to two seen by F.E.H. dur ing June 2013 and June 2014. This spe cies oc curs through out Micronesia ex cept Wake (Wiles 2005).
spec i mens: FMNH (online), ZIN (Kittlitz 1836).
Bristle - thighed Curlew ( Numenius tahiti-ensis )
106 PACIFIC SCIENCE · Jan u ary 2016
Rare Ne arc tic mi grant. Hill (1957) re-ported that it was “ com monly seen ” dur ing 1956 – 1957, but un doubt edly she misidentifi ed the much more com mon Whimbrel, which she did not re port. Segal (1985, 1989) listed it for Kosrae but no de tails were giv en. On 24 June 2014, F.E.H. photographed a sol i tary in-di vid ual at the northwest tip of Utwe - Ma ( Figure 8 ), pro vid ing the fi rst re cord for Kos-rae. It was dis tin guished from the sim i lar Whimbrel by its buffy or ange rump. It has been recorded in most of the island groups of Micronesia, in clud ing Pohnpei and the Mar-shall Islands (Wiles 2005), thus its oc cur rence on Kosrae was expected.
Gray - tailed Tattler ( Tringa brevipes )
Uncommon Pa le arc tic mi grant. This spe-cies is dif fi cult to dis tin guish from the more com mon Wandering Tattler ( T. incana ) by
plum age, but is eas ier by voice. Hayes (1986) es ti mated that 5 % of the tat tlers on Kosrae were of this spe cies. They pre ferred the tidal fl ats at low tide and of ten fl ocked to geth er. During 1981 – 1982, it was more com mon in fall, with a high count of 35 at Tafunsak on 2 Oc to ber 1981 (Hayes 1986), than dur ing spring, with a high count of 5 at Okat Harbor on 13 May 1982. The USFWS team ob served a sin gle in di vid ual on the old run way at Lelu in July 1983 (J. Engbring in Hayes 1986). F.E.H. failed to fi nd any dur ing June 2013 and June 2014. This spe cies oc curs through out Micronesia (Wiles 2005).
spec i mens: AMNH (Stickney 1943, Baker 1951), ZIN (Kittlitz 1836).
Wandering Tattler ( Tringa incana )
Common Ne arc tic mi grant. This is the most com mon shore bird on Kosrae (Hayes 1986), for ag ing among rocks and seagrasses on tidal fl ats, sandy beaches, man groves, and oc ca sion ally in land along moun tain streams. During 1981 – 1982, max i mum counts were higher dur ing fall (135 at Okat Harbor on 4 Oc to ber 1981) than dur ing win ter (100) or spring (80; Hayes 1986), and large fl ocks roosted at night on the run way un der con-struc tion at Okat Harbor. A few lin ger dur-ing the sum mer, with up to 10 per day seen by F.E.H. in June 2013 and smaller num bers dur-ing July 1983 (J. Engbring in Hayes 1986). Both tat tler spe cies are wide spread through-out Micronesia, but the Gray - tailed Tattler is more com mon west ward and Wandering Tat-tler is more com mon east ward.
spec i mens: AMNH (Stickney 1943, Baker 1951), FMNH (online), MNHN (online), USNM (Townsend and Wetmore 1919), YIO (online).
vo cal i za tions: ML (online).
Common Sandpiper ( Actitis hypoleucos )
Rare Pa le arc tic mi grant. Separating this spe cies from a ba sic plum aged Spotted Sand-piper ( A. macularius ), a Ne arc tic mi grant, is dif fi cult. Hayes (1985) saw sin gle birds, pos si-bly the same in di vid u al, on four oc ca sions from 15 Jan u ary to 1 April 1982 at Lelu and
F igure 8. First documented re cord of Bristle - thighed Curlew ( Numenius tahitiensis ) for Kosrae at Utwe - Ma on 24 June 2014. Photo by F.E.H.
Avifauna of Kosrae · Hayes et al. 107
Figure 9. First documented re cord of Common Sand-piper (Actitis hypoleucos) at Okat Harbor on 21 Feb ru ary 2014. Photo by C.J.C.
Figure 10. First photo documented re cord of Sanderling (Calidris alba) for Kosrae at Okat Harbor on 13 No vem-ber 2013. Photo by C.J.C.
Tofol, but did not pos i tively iden tify the spe-cies; how ev er, the lack of spot ting by 1 April, at a time when it is of ten ac quired by Spotted Sandpiper (es pe cially adults; Pyle 2008), is sug ges tive of the Common Sandpiper. C.J.C. photographed an im ma ture at Okat Harbor on 21 Feb ru ary 2014 ( Figure 9 ), pro vid ing the fi rst documented re cord for Kosrae. It was dis-tin guished from a Spotted Sandpiper by its tail extending well be yond its rel a tively shorter wings, shorter wingstripe, and less barred wing co verts. This spe cies has been recorded in most island groups of Micronesia (Wiles 2005), in clud ing re cent con fi r ma tion in Pohn-pei (Pratt et al. 2010). The Spotted Sandpiper has only been recorded from the Marshall Is-lands (Wiles 2005).
Ruddy Turnstone ( Arenaria interpres )
Common bo real mi grant. This is the sec-ond most com mon shore bird on Kosrae (Hayes 1986), for ag ing along the coasts and oc ca sion ally oc cur ring on roads dur ing high tide. During 1981 – 1982, the highest counts oc curred dur ing late win ter (160 at Okat Har-bor on 27 Feb ru ary 1982) and spring (120 at Okat Harbor on 9 March 1982), with smaller num bers dur ing fall (up to 85; Hayes 1986). A few lin ger dur ing sum mer. The USFWS team ob served up to eight at Tafunsak and six at Okat Harbor on 17 July 1983 (J. Engbring in Hayes 1986), and F.E.H. saw six at Okat Har-bor on 20 June 2013. This spe cies oc curs through out Micronesia (Wiles 2005).
spec i mens: AMNH (Stickney 1943, Baker 1951), NSMT (online), YIO (online), ZIN (Kittlitz 1836).
Sanderling ( Calidris alba )
Rare bo real mi grant. Hayes (1985, 1986) ob served ba sic - plum aged in di vid u als at Okat Harbor on 14 No vem ber 1981 and at Tafun-sak on 18 De cem ber 1981. The USFWS team ob served an al ter nate - plum aged in di vid ual at Okat Harbor on 1 July 1983 (Pyle and Eng-bring 1987) and pos si bly the same in di vid ual at Tafunsak on 3 July 1983 (J. Engbring in Hayes 1986; Pyle 1983 reported the date of 5 July 1983). C.J.C. photographed an al ter nate - plum aged in di vid ual at Okat Harbor on 13 No vem ber 2013 ( Figure 10 ), pro vid ing the fi rst doc u men ta tion of the spe cies for Kosrae. This spe cies oc curs through out Micronesia (Wiles 2005).
Red - necked Stint ( Calidris rufi collis )
Rare Pa le arc tic mi grant. C.J.C. photo-graphed a worn adult Calidris sp. molting into ba sic plum age ( Figures 11 A, 11 B ) at To-fol on 21 No vem ber 2011. Although the char-ac ters of the bird are con sis tent with this spe cies, distinguishing it from other sim i lar con ge ners is very dif fi cult (e.g., Message and Taylor 2005, O ’ Brien et al. 2006, Chandler 2009). The bird ap pears too pale, es pe cially on the head and neck, for ei ther Semipalmated Sandpiper ( C. pusilla ) or Little Stint ( C. minuta ), but its iden tity could not be con fi rmed. C.J.C.
108 PACIFIC SCIENCE · Jan u ary 2016
photographed a more de fi n i tively iden ti fi ed ju-ve nile at Okat Harbor on 30 Oc to ber 2013 (Figure 11 C ), which lin gered un til 13 No-vem ber 2013, pro vid ing the fi rst documented re cord for Kosrae. It was dis tin guished from Semipalmated Sandpiper by its fi ner bill tip, less scaly back, whiter tertial edges, and lon-ger wing pro jec tion, and from Little Stint by its thicker bill tip, lack of man tle lines, duller tertial edges, and shorter wing pro jec tion. The Red - necked Stint is a reg u lar mi grant in west ern Micronesia east to Chuuk and a va-grant in the Marshall Islands, Little Stint is a va grant in the Commonwealth of the North-ern Marianas and Marshall Islands (Wiles 2005), and Semipalmated Sandpiper is un re-corded in Micronesia but is a va grant in the Ha wai ian Islands (Pyle and Pyle 2009).
Sharp - tailed Sandpiper ( Calidris acuminata )
Uncommon Pa le arc tic mi grant. Hayes (1985, 1986) saw up to two at Okat Harbor on fi ve oc ca sions from 27 Sep tem ber to 31 Oc to-ber 1981, and also sin gle in di vid u als at Malem on 12 No vem ber 1981 and at Tofol on 22 No-
vem ber 1981. C.J.C. photographed an im ma-ture among a fl ock of fi ve at Tofol dur ing 23 – 24 No vem ber 2011 ( Figure 12 ), pro vid ing the fi rst doc u men ta tion of the spe cies for Kos-rae. C.J.C. photographed four more at Okat Harbor on 23 No vem ber 2013. This spe cies oc curs through out Micronesia (Wiles 2005).
Brown Noddy ( Anous stolidus pileatus )
Common res i dent. Coultas (1931:242) con-sid ered it com mon, es pe cially in the south, and reported it oc cur ring “ high up on the ridges of the tallest moun tains. ” During 1981 – 1982, F.E.H. ob served large fl ocks of > 1,000 in di vid u als, of ten vis i ble from shore, fol low-ing schools of fi sh at sea. Fisherman rou tinely followed such fl ocks to lo cate fi sh. Engbring et al. (1990) found Brown Noddies abun dant in all ter res trial hab i tats, with an av er age den-sity of 274 / km 2 and an es ti mated to tal pop u-la tion of 27,243; den si ties were highest in man groves and neg a tively cor re lated with el-e va tion, but some oc curred above 400 m (Ta-ble 2), and den si ties were higher at Walung (472 / km 2 ) than in other ar eas. During June 2013 and June 2014, F.E.H. never saw more than a few dozen fl y ing to or from their roosts de spite even ing searches from the west ern, north east ern, and south east ern coasts, and never saw any large for ag ing fl ocks from shore, suggesting ei ther a sea sonal de par ture
Figure 11. First documented re cords of Red-necked Stint (Calidris rufi collis) for Kosrae: (A, B) a pre sumed adult at Tofol on 21 No vem ber 2011 and (C) a ju ve nile at Okat Harbor on 30 Oc to ber 2013. Photos by C.J.C.
Figure 12. First photo documented re cord of Sharp-tailed Sandpiper (Calidris acuminata) for Kosrae at Tofol on 23 No vem ber 2011. Photo by C.J.C.
Avifauna of Kosrae · Hayes et al. 109
of most noddies from the island or a dra matic de cline in the pop u la tion. This spe cies oc curs through out Micronesia (Wiles 2005).
breed ing: Finsch (1880 a , 1881) ob served nests among epi phytic ferns in tall trees. Coultas (1931) suspected that Brown Noddies nested year round. F.E.H. found ap prox i ma-tely 50 nests in man groves near Molsron Mwot, be tween Okat Harbor and Walung on 29 De cem ber 1981. When camping at higher el e va tions in July 1983, Engbring et al. (1990) of ten heard adults and their young call ing at night. C.J.C. ob served a mixed col ony of nod-dies nesting an nu ally since 2012 at Inya Mu-tunenea, a man grove la goon north of Lelu; nest con struc tion be gins in early No vem ber and the last chicks are fl edged in May. C.J.C. also ob served about 50 nests (3 photographed) at Molsron Mwot on 5 March 2011, and about 20 nesting in trees be side the Kosrae Island Resource Management Authority in Tofol in early 2011.
spec i mens: AMNH (Baker 1951), MCZ (online), USNM (Townsend and Wetmore 1919), YIO (online), ZIN (Kittlitz 1836).
vo cal i za tions: ML (online).
Black Noddy ( Anous minutus marcusi )
Common res i dent. Coultas (1931:242 – 243) stated that it fa vored roosting on “ low shrubs and trees of out ly ing reefs and iso lated small is lands. ” Engbring et al. (1990) reported a mean den sity of 55 / km 2 and an es ti mated to-tal pop u la tion of 5,461; den si ties were highest in man groves and neg a tively cor re lated with el e va tion (Table 2). Engbring et al. (1990:80) ob served “ lim it ed, scattered num bers ” in the for ested in te rior and recorded at least one pair “ perched in a tree high in the in te ri or, ” but none were seen above 400 m (Table 2). This spe cies has been recorded from all island groups in Micronesia (Wiles 2005).
breed ing: Finsch (1880 a , 1881) reported large col o nies nesting in man groves in late Feb ru ary 1880. Nehrkorn (1899) reported that eggs had been col lected at Kosrae. F.E.H. lo cated two nests on the branches of tall trees about 75 m above sea level along the Mutunte River in Tafunsak on 26 De cem ber 1982. In July 1983, Engbring et al. (1990) found a col-
ony of sev eral hun dred to a few thou sand nests on a small is let at Walung, a sec ond col ony of a few hun dred nests on a man grove island near Utwe, and a third col ony of a few hun dred nests in man groves at Lelu Harbor, but F.E.H. did not ob serve any ev i dence of nesting in June 2013. C.J.C. ob served a mixed col ony of noddies nesting an nu ally at Inya Mutunenea, a man grove la goon north of Lelu, since 2012; nest con struc tion be gins in early No vem ber and the last chicks are fl edged in May.
spec i mens: AMNH (Baker 1951), YIO (on-line), ZIN (Kittlitz 1836).
Vocalizations: ML (online).
White Tern ( Gygis alba pacifi ca )
Common res i dent. F.E.H. reg u larly ob-served small num bers at lower el e va tions, along the shore, and for ag ing at sea, where they of ten ac com pa nied fi sh ing fl ocks of nod-dies, in clud ing 5 seen from a fi sh ing boat a few ki lo me ters west of Tafunsak on 8 Oc to-ber 1981 and 10 more in the same area on 12 Oc to ber 1981. On 2 Jan u ary 1982, F.E.H. ob-served six far out to sea be tween Kosrae and Pingelap, mostly near the lat ter. Engbring et al. (1990) reported a mean den sity of 160 / km 2 with an es ti mated to tal pop u la tion of 15,860; den si ties were highest in man groves and neg a tively cor re lated with el e va tion, al-though some were ob served above 400 m (Table 2). Engbring et al. (1990) noted 24 dur-ing an off shore fi sh ing trip south east of Lelu on 28 Ju ly. Kepler et al. (1992) ob served an un-spec i fi ed num ber of in di vid u als > 100 km south of Kosrae on 12 Oc to ber 1988. This spe cies oc curs through out Micronesia (Wiles 2005).
breed ing: A fl edg ling was found on the ground and photographed by C.J.C. at Lelu Island on 20 Feb ru ary 2011.
spec i mens: AMNH (Baker 1948, 1951), NSMT (online), YIO (online), ZIN (Kittlitz 1836).
Laughing Gull ( Leucophaeus atricilla )
Rare Ne arc tic mi grant. C.J.C. photo-graphed a ba sic - plum aged adult on a con crete plat form be side the cause way connecting Lelu
110 PACIFIC SCIENCE · Jan u ary 2016
Island with the main island on 4 Jan u ary 2008; it lin gered through Feb ru ary 2008 (last date un known) and was seen twice at Okat Harbor (photo published; Pratt et al. 2010). C.J.C. photographed a fi rst - cy cle im ma ture at Okat Harbor on 25 March 2011 and an other fi rst - cy cle im ma ture at Okat Harbor on 24 Feb ru-ary 2014. This spe cies has been recorded as a va grant in Palau, Commonwealth of the Northern Marianas, Pohnpei, and Marshall Islands (Wiles 2005, Pratt et al. 2010).
Greater Crested Tern ( Thalasseus bergii )
Uncommon vis i tor. Hayes (1985) ob served two at Tafunsak and Okat Harbor from 31 Au gust to 13 Sep tem ber 1981 and up to seven (six adults and an im ma ture) along the west coast from 5 De cem ber 1981 to 5 May 1982. F.E.H. saw one at Walung on 26 June 2013, and C.J.C. saw two at Okat Harbor on 18 Feb-ru ary 2014. This spe cies has been recorded from all island groups in Micronesia ex cept Wake (Wiles 2005).
Little Tern ( Sternula albifrons )
Uncommon Pa le arc tic mi grant. Hayes (1985) ob served up to fi ve at Okat Harbor from 3 Oc to ber 1981 to 5 May 1982. Pratt and Pyle ob served fi ve at Okat Harbor on 24 Feb ru ary 1988 (Wiles et al. 2000). This spe-cies oc curs through out Micronesia ex cept Wake (Wiles 2005).
Sooty Tern ( Onychoprion fuscatus )
Rare vis i tor. Hayes (1985) heard and saw three fl y ing over Okat Harbor on 20 April 1982 and one over Okat Harbor on 13 May 1982. Kepler et al. (1992) ob served a fl ock of about 200 for ag ing ap prox i ma tely 145 km (orig i nally reported as 157 km) south - south-east of Kosrae (04 ° 03 ′ N, 163 ° 31 ′ E) on 12 Oc to ber 1988. This spe cies oc curs through-out Micronesia (Wiles 2005).
Whiskered Tern ( Chlidonias hybrida )
Rare Pa le arc tic mi grant. On 25 June 2013, F.E.H. ob served an al ter nate - plum aged adult
cir cling Utwe - Ma, but did not see it well enough to ver ify its iden tity un til the fol low-ing day, 26 June 2013, when he ob served and photographed it for ag ing in Utwe Harbor ( Figure 13 ). It was dis tin guished from a Com-mon Tern ( Sterna hirundo ) by its all - red bill, darker gray un der parts, and shorter tail with white outer tail feath ers. The Whiskered Tern was un known in Micronesia un til a spec i men was taken on Yap in 1976 (Clapp and Lay-bourne 1983). Subsequently, it has been reported in Palau, the Marianas, Yap, and Pohnpei (Wiles 2005, Pratt et al. 2010), and it is now a reg u lar vis i tor to west ern Micronesia in sub stan tial num bers (Pratt and Etpison 2008). This re cord rep re sents the fi rst for Kosrae and the east ern most in Micronesia, suggesting an on go ing ex pan sion of this spe cies in the re gion.
South Polar Skua ( Stercorarius maccormicki )
Rare aus tral mi grant off shore. Kepler et al. (1992) ob served two pale - morph adults ap-prox i ma tely 145 km south - south east of Kos-rae (04 ° 03 ′ N, 163 ° 31 ′ E) on 12 Oc to ber 1988. Elsewhere in Micronesia it has been reported at Palau and Pohnpei (Wiles 2005).
Parasitic Jaeger ( Stercorarius parasiticus )
Figure 13. First documented re cord of Whiskered Tern (Chlidonias hybrida) for Kosrae at Utwe on 26 June 2013. Photo by F.E.H.
Avifauna of Kosrae · Hayes et al. 111
Rare bo real mi grant off shore. On 12 Oc-to ber 1981, F.E.H. briefl y ob served two jae-ger - like birds, prob a bly of this spe cies, from a fi sh ing boat a few ki lo me ters west of Kos-rae, but did not re port them among other new bird re cords be cause of un cer tainty of iden ti-fi ca tion (Hayes 1985). Kepler et al. (1992) ob-served two un iden ti fi ed jae gers ap prox i ma tely 145 km south - south east of Kosrae (04 ° 03 ′ N, 163 ° 31 ′ E), and a ju ve nile Parasitic Jaeger ap-prox i ma tely 120 km south - south west of Kos-rae (04 ° 22 ′ N, 162 ° 21 ′ E) on 12 Oc to ber 1988. Elsewhere in Micronesia this spe cies has been reported only for Pohnpei (Wiles 2005).
Rock Dove ( Columba livia )
Extirpated in tro duc tion. Santos et al. (1970:33) suggested that the Jap a nese may have in tro duced car rier pi geons to carry mes-sages, “ but we have no re cord of them now. ” Hayes (1985) found three in di vid u als at Lelu on 16 De cem ber 1981, which were still pres-ent when he left the island in May 1982. They were ob vi ously in tro duced, ei ther from a vis-it ing ship or as es caped pets. However, none were seen by the USFWS team in July 1983 (Engbring et al. 1990), in di cat ing that they failed to es tab lish a self - sus tain ing fe ral pop-u la tion. Elsewhere in Micronesia, this spe cies failed to es tab lish itself in Chuuk, Pohnpei, and the Marshall Islands, where birds have been detected in the past (Wiles 2005), but ap par ently a self - sus tain ing pop u la tion oc curs on Wake (Jones 1995).
Kosrae Fruit Dove ( Ptilinopus hernsheimi )
Common res i dent. This en demic spe cies was de scribed from Kosrae by Finsch (1880 a ). A spec i men taken in 1859 at Ebon in the Mar-shall Islands was orig i nally de scribed as a new spe cies (Pe ters and Griscom 1928) but later syn on y mized with the Kosrae form by Ripley and Birckhead (1942), who suspected its lo cal ity was mislabeled. Later au thors as-sumed the lo cal ity was cor rect; if so, the lack of sub se quent re cords from the Marshall Is-lands in di cates that any pre vi ous pop u la tion is now ex tir pated (Baker 1951, Amerson 1969, Spennemann 2006). The calls of this spe cies
are a con stant sound in the rainforest, but the birds are hard to see ex cept in fl ight. Engbring et al. (1990) reported a mean den sity of 77 / km 2 with an es ti mated pop u la tion of 8,672 on Kos-rae; den si ties were higher in rainforest than in man groves and in creased with el e va tion up to 400 m, but ta pered off at the highest el e va-tions (Table 2). Because fe males do not vo cal-ize and most in di vid u als were detected by sound, the den sity and to tal pop u la tion could be twice as high (Engbring et al. 1990). It re-mains com mon on Kosrae through 2015 (C.J.C. and F.E.H., pers. obs.). F.E.H. ob-served one at 589 m near the sum mit of Mt. Mutunte on 21 June 2013.
breed ing: No da ta. spec i mens: AMNH (Ripley and Birckhead
1942, Baker 1951), DMNH (online), FMNH (online), MCZ (online), NSMT (online); USNM (Townsend and Wetmore 1919, Baker 1951), YIO (online).
vo cal i za tions: ML (online), XC (online). tax o nomic notes: The Kosrae Fruit Dove
was long regarded as a com po nent of a large poly typic Crimson - crowned Fruit Dove P. porphyraceus com plex that in cluded the Mi-cro ne sian forms hernsheimi (Kosrae), ponapen-sis (Chuuk and Pohnpei), and pelewensis (Palau), along with three Poly ne sian sub spe-cies (Mayr 1945, Baker 1951). Pratt et al. (1980), followed by all sub se quent au thors, be gan the breakup of this com plex by split-ting the Palau Fruit Dove ( P. pelewensis ) based on strik ing plum age and vo cal dif fer ences. Pratt et al. (1987) suggested that the remain-ing Mi cro ne sian and Poly ne sian sub spe cies might best be con sid ered two spe cies, but a thor ough anal y sis of the com plex had to await the col lec tion of ad e quate com par a tive vo cal ma te ri al. Vocal dif fer ences now sug gest that the P. porphyraceus com plex may com prise more than the two spe cies pre vi ously hy-poth e sized (Pratt et al. 1987). Unfortunately, past de scrip tions, such as that of Pratt et al. (1987), tended to con fl ate the songs of dif fer-ent re gional groups and un der stated what was con sid ered at the time to be in tra spe cifi c var i a tion.
The Mi cro ne sian and Poly ne sian groups are widely sep a rated geo graph i cally and dif fer in col or a tion and voice. Mi cro ne sian sub spe cies
112 PACIFIC SCIENCE · Jan u ary 2016
have a bold yel low band at the tip of the tail, whereas P. porphyraceus fasciatus (Sa moan Ar-chipelago) has an in con spic u ous yel low ish - gray band, and the nom i nate sub spe cies (Fiji, most of Tonga, Niue) has a pale gray band, with P. porphyraceus graeffei (Wallis and Fu-tuna and Niuafo ‘ ou, Tonga) in ter me di ate. Mi cro ne sian forms have a deep pur ple, al most black, belly patch that can be brown ish - red in many in di vid u als of P. hernsheimi (Baker 1951). The belly patch is paler in nom i nate P. por-phyraceus and dull red in Sa moan birds. The head and breast are strongly tinged yel low in Mi cro ne sian birds, but are green ish gray in Poly ne sian ones, and the fron tal cap is more pur ple and less rosy than in Mi cro ne sian birds.
The typ i cal song of all Poly ne sian forms has a halt ing or stum bling rhythm as if the bird stops and starts again: hup - brrOOAH, a - hup - brrOOAH, hoo - HOO - hoo hoo doo, hoo - HOO - hoo hoo, hoo - HOO - hoo hoo, hoo - HOO - hoo hoop - hoop - hoop, hoo - HOO - hoop hoop (from ML 139953). Very rare ly, they ut ter a var i ant, as in the fi rst song on ML 94984 from Vava ‘ u, Tonga, with rhythm more evenly paced. Fruit doves on Chuuk and Pohnpei ut ter a hooting song that has an ini tial syn co pated phrase fol-lowed by a long rhyth mic se ries of 12 – 20 nearly iden ti cal sin gle - note hoots that slow to-ward the end: hooo, hup - hoo, whoop hoo hoo hoo hoo, etc. Occasionally, the sin gle hoots may be slightly two - syl la bled as tawoo tawoo tawoo, for ex am ple (sev eral ex am ples in ML 55697), but at a dis tance this struc ture is not ap par ent. The pace of the long se ries may vary from song to song, even when uttered by the same in di vid u al. As fi rst noted by Engbring et al. (1990), the Kosrae Fruit Dove ’ s higher - pitched song dou bles the notes in the long se ries, with the sec ond note em pha sized and sharply clipped, like a hic cup: hooo, hup - HOOO, a - HOOT a - HOOT a - HOOT a - HOOT, etc. Among roughly 40 songs of Kosrae birds, we found one (ML 55624, sec ond song) in which the fi rst syl la ble of the “ hic cup ” was muted or de leted so that the song some what re sem bled that of ponapensis, but the over all rhythm and other hernsheimi char ac ter is tics remained the same. We found no dou ble - note songs among 10 recorded ex am ples from Chuuk and 40 from Pohnpei (to tals are ap prox i ma tions be-
cause whether to count a par tic u lar re cord ing, based on its qual i ty, is an ar bi trary de ci sion of the re search er).
Given the wide geo graphic sep a ra tion of the Poly ne sian sub spe cies and their po ten tial iso lat ing mech a nisms of voice and col or a tion, they clearly war rant rec og ni tion as a sep a rate spe cies by Pratt ’ s (2010) cri te ria, and can re-tain the En glish name Crimson - crowned Fruit Dove. Pratt and Mittermeier (2016) dis-cuss geo graphic var i a tion among the three nom i nal Polynesian sub spe cies of Crimson - crowned Fruit Dove, which may com prise only two valid sub spe cies and a hy brid pop u-la tion.
Molecular stud ies also sup port the sep a ra-tion of Poly ne sian P. porphyraceus at the spe-cies lev el. Cibois et al. (2014) found the broadly de fi ned P. porphyraceus to be poly phy let ic, with Mi cro ne sian P. ponapensis closer to the Mel a ne sian Red - bel lied Fruit Dove P. greyi than to Poly ne sian P. porphyraceus . Unfortu-nately, Cibois et al. (2014) in cluded no sam-ples from Kosrae, so for now, the sta tus of P. hernsheimi must be based on mor pho log i-cal and be hav ioral cri te ria. We orig i nally re-garded the color dif fer ences be tween P. hern-sheimi and P. ponapensis as mat ters of de gree rather than kind, typ i cal of dif fer ences among sub spe cies, but del Hoyo and Collar (2014), us ing a scor ing sys tem pro moted by To bias et al. (2010) for de ter min ing pu ta tive spe cies lim its, rec og nized P. hernsheimi as a full en demic spe cies, based on col or a tion and mea sure ments alone. Now that strik ing vo-cal dif fer ences be tween them have been well documented, we agree that P. hernsheimi and P. ponapensis are best regarded as sep a rate bi-o log i cal spe cies.
En glish names in this com plex have led to some past con fu sion. Owen (1977, pers. comm.) coined the name “ pur ple - capped ” be-cause he con sid ered “ crim son - crowned ” to be in ac cu rate for Mi cro ne sian rep re sen ta tives. Pratt et al. (1987) used Purple - capped for the whole porphyraceus com plex, but most other sources have used Crimson - crowned. With this re vi sion, Purple - capped Fruit Dove can be used for P. ponapensis and Crimson - crowned Fruit Dove can be re stricted to the Poly ne sian forms.
Avifauna of Kosrae · Hayes et al. 113
Mi cro ne sian Imperial Pigeon ( Ducula oce-anica oceanica )
Uncommon res i dent. The nom i nate sub-spe cies, which oc curs in Kosrae and in Jaluit and Elmore in the Marshall Islands (Baker 1951), was fi rst de scribed from Kosrae by Lesson and Garnot (1826). According to Sarfert (1919) its num bers had sharply de-clined by 1910 due to hunt ing with guns and it had be come rare along the coast. Coultas (1931) stated that the pi geons con gre gated in tall trees along the coast in the even ings to roost and scattered at dawn to for age in the moun tains. Coultas (1931) ob served that the island res i dents had be come pro fi cient at im i-tat ing their calls and kill ing them with stones and clubs. He felt that there were not enough Jap a nese hunt ers to have re duced their num-bers, but lat er, according to Segal (1989), the pi geons were nearly ex ter mi nated by Jap a-nese hunt ers dur ing World War II, when many hu mans were starv ing. Bequaert (1939, 1941) found the par a sitic fl y Ornithoctona plicata (Hippoboscidae) on one or more spec-i mens col lected by Coultas. During 1981 – 1982, F.E.H. sel dom saw this bird, but of ten heard its deep low moan and coarse cooing, with most sightings oc cur ring away from the coast. Native hunt ers seem to have lit tle trou-ble fi nd ing this spe cies; dur ing 1981 – 1982, F.E.H. met hunt ers who had shot up to three birds within a few hours. Engbring et al. (1990) reported its mean den sity as 60 / km 2 with an es ti mated to tal pop u la tion of 7,474; den si ties were highest at in ter me di ate el e va-tions, low est in man groves (Table 2), and var-ied re gion al ly, rang ing from 46 / km 2 in Tafunsak to 84 / km 2 in Walung. During sev-eral long hikes in the moun tains in 2013 and 2014, F.E.H. heard only a few in di vid u als, suggesting the pop u la tion has de clined. The spe cies is heavily hunted with an open sea son lacking a bag limit in Jan u ary. However, poach ers take many dur ing other months and sell each for $ 10 on the black mar ket. This spe cies ex hib its an ir reg u lar, pos si bly relic-tual (Baker 1951), dis tri bu tion on both high is lands and atolls through out the Car o line Islands (Wiles 2005). Todd Mark obtained the fi rst re cord ings from Kosrae of both the
barking “ song ” (XC75516) and the low moan-ing call (XC75517).
breed ing: Finsch (1880 a, 1881) col lected molting im ma tures and molting adults in late Feb ru ary and suggested that they breed year - round. C.J.C. photographed a re cently fl edged ju ve nile at Pukusrik Wan on 11 Sep tem ber 2008.
spec i mens: AMNH (Baker 1951), DMNH (online), FMNH (online), MCZ (online), MNHN (online), RMNH (Hoek Ostende et al. 1997), SMF (online), USNM (Townsend and Wetmore 1919, Baker 1951), YIO (online), ZIN (Kittlitz 1836).
vo cal i za tions: XC (online).
Pacifi c Long - tailed Cuckoo ( Urodynamis taitensis )
Uncommon aus tral mi grant. Kittlitz (1836) ob served one but failed to col lect it in De cem-ber 1827. Coultas (1931) reported that the spe cies ar rived around 1 Feb ru ary and oc-curred ev ery where on the island. He col-lected three spec i mens, on 15 March, 5 April, and 15 April 1931 (Bogert 1937). During 1981 – 1982, F.E.H. saw only one bird, fl y ing over the main road at Tofol on 14 March 1982. The USFWS team ob served an im ma ture in rainforest near Tafunsak on 5 Ju ly, a sec ond in man groves at Walung on 12 Ju ly, and a third in rainforest near Okat Harbor on 23 July 1983 (Pyle 1983, Pyle and Engbring 1987, Engbring et al. 1990). C.J.C. has oc ca sion ally ob served and heard up to three an nu ally since 2007 at Pukusruk Wan, north of Lelu, and photographed one on 26 Oc to ber 2009. This spe cies, whose mi gra tory pat tern has been re-cently stud ied (Gill and Hauber 2012), oc curs through out Micronesia (Wiles 2005).
spec i mens: AMNH (Bogert 1937, Baker 1951).
Island Swiftlet ( Aerodramus inquietus inqui-etus )
Common res i dent. This spe cies was fi rst de scribed from Kosrae by Kittlitz (1836). The nom i nate sub spe cies A. i. inquietus is en demic to Kosrae (Baker 1951); else where in Microne-sia, this spe cies oc curs on Chuuk and Pohnpei
114 PACIFIC SCIENCE · Jan u ary 2016
(Baker 1951, Wiles 2005). Large fl ocks for-age above the for est can opy through out Kosrae. Coultas (1931) con sid ered it the most com mon bird on the island and reported it from the tops of the highest peaks. F.E.H. ob-served a few at 589 m near the sum mit of Mt. Mutunte on 21 April 2013. Engbring et al. (1990) reported a mean den sity of 290 / km 2 with an es ti mated to tal pop u la tion of 27,863; den si ties were highest at the highest el e va-tions, low est in man groves (Table 2), and var-ied re gion ally with much higher den si ties at Tafunsak (549 / km 2 ) than in Malem (162 / km 2 ) or Walung (147 / km 2 ). As pointed out by Gary Wiles (pers. comm.), the var i able cir cu lar plot method may be un suit able for study ing this spe cies due to its swift and cir cling fl ight; the es ti mate of nearly 28,000 birds may be too high, re quir ing the equiv a lent of 28 col o nies with 1,000 birds each, yet only a few col o nies with none ex ceed ing 1,000 birds are known. Nevertheless, the spe cies is still abun dant.
breed ing: Coultas (1931) reported roosting and nesting in caves scattered through out the island. Engbring et al. (1990) vis ited the Wiya Bird Cave at Tafunsak on 1 July 1983 and ob-served many birds on nests with beg ging off-spring and an egg that fell to the ground. C.J.C. ob served nests at a cave in Yela, south of Okat Harbor, on 15 Jan u ary 2010. F.E.H. saw sev eral hun dred ac tive nests at the Wiya Bird Cave, Tafunsak, on 20 June 2013 and 25 June 2014. On 19 June 2014, F.E.H. ex am ined 11 nests, each made of moss (most of which was still green), in a small man - made Jap a nese cave be side the mu nic i pal wa ter sup ply in Malem; four nests had one egg, one nest had a chick that had just hatched (egg shell still pres ent), one nest had a quar ter - grown chick, and fi ve nests were emp ty.
spec i mens: AMNH (Baker 1951), USNM (Baker 1951), YIO (online), ZIN (Kittlitz 1836).
vo cal i za tions: XC (online). Todd Mark recorded the ca coph ony of swift lets (XC75520) in side Wiya Bird Cave on 10 Oc to ber 2008, pro vid ing the fi rst re cord ings of this spe cies ’ voice (other than clicks used for echo lo ca tion recorded at Chuuk). Because the cave is shal-low enough for light to pen e trate through out, Kosrae swift lets do not echolocate there. The
calls are high - pitched, fairly loud squeaky notes uttered in a pat tern less jum ble, and ap-par ently only within the con fi nes of nesting caves, even though they may not be re lated to nav i ga tion. Mark recorded a sim i lar vo cal i za-tion (XC100258) in a cave on Pohnpei, but swift lets for ag ing in the open are ut terly si lent (Pratt et al. 1987) on that island and at Chuuk. On the lat ter, the only sounds recorded have been echo lo ca tion clicks (ML 5509) and squeaks of nes tlings (ML 5508). Possible in-ter is land dif fer ences in the vo cal i za tions of swift lets should be stud ied.
Mi cro ne sian Myzomela ( Myzomela rubra-tra rubratra )
Common res i dent. This spe cies was fi rst de scribed from Kosrae by Lesson (1827). The nom i nate sub spe cies is en demic to Kosrae (Baker 1951). It is com mon in ev ery ter res trial hab i tat, in clud ing man groves. Coultas (1931) stated that it did not oc cur at higher el e va-tions, but Engbring et al. (1990) of ten re-corded it above 400 m, and F.E.H. ob served sev eral at 589 m near the sum mit of Mt. Mu-tunte on 21 April 2013. During 1981 – 1982, F.E.H. of ten ob served chil dren shoot ing them with home made sling shots and once saw a bird killed by a thrown rock. Engbring et al. (1990) reported a mean den sity of 1,433 / km 2 with an es ti mated to tal pop u la tion of 136,358; den si ties were highest in man groves (higher in Sonneratia than Rhizophora trees) and in versely cor re lated with el e va tion (Table 2). This spe-cies oc curs in all island groups in Micronesia ex cept the Marshall Islands and Wake (Wiles 2005).
ab nor mal plum age: C.J.C. photographed a hypomelanic in di vid ual with gray replacing black plum age at Tafayet, Lelu, on 1 Au gust 2014.
breed ing: Finsch (1880 a, 1881) found an un fi n ished nest in a man grove in late Feb ru-ary. Coultas (1931) noted fe males feed ing young long af ter they could fl y, and also noted adult males cop u lat ing with ju ve nile fe males. Baker (1951), based on an ex am i na tion of molt in spec i mens, con cluded that nesting ac tiv i ties oc cur from De cem ber to March and molt be-gins in Jan u ary and peaks in March. In April
Avifauna of Kosrae · Hayes et al. 115
1982, F.E.H. ob served a nest with chicks at Tafunsak.
spec i mens: AMNH (Baker 1951), NSMT (online), RMNH (Dekker and Quaisser 2006), USNM (Townsend and Wetmore 1919), YIO (Momiyama 1922; online), ZIN (Kittlitz 1836).
vo cal i za tions: ML (online), XC (online). Myzomela taxa are note wor thy for sing ing im-pres sively com plex dawn songs quite dis tinct from their day time ut ter ances, and the Kos-rae rep re sen ta tive is an ex cel lent ex am ple. By itself, it per forms a dawn cho rus that sounds like many spe cies sing ing to geth er. Singing be gins be fore sun rise when it is still too dark to see the birds and con tin ues for about 25 min un til enough light is pres ent for vis i bil i ty. It then ceases quite sud den ly, as if some one has thrown a switch (ML 55634, 55659). The dawn song is a con tin u ous se ries of burry whis tles and jerky back - and - forth phrases, many seg-ments of which lend them selves to trans lit er-a tion as cheeky cheeky can ’ t beat the heat; too sweet to eat; or she eats a piz za . It is gen er ally sim i lar to the dawn song of M. r. dichromata on Pohn-pei (ML 55674), or M. r. kurodai on Yap (ML 140151), to se lect two ex am ples, but with a quicker pace and fewer slurred notes. Individ-ual var i a tion is great, and some dawn songs are more lilt ing and less jer ky. In con trast, the day-light vo cal i za tions of Mi cro ne sian Myzomelas are rather un in ter est ing, but highly var ied, with short buzzy calls, sharp tseep! notes that may be re peated as a se quence, and raspy scolds. They also ut ter a rapid trill or whinny (ML 55614) whose func tion is un known, but seems to be an alarm call. The day time song is short and highly var i able, usu ally in clud ing one em phatic note and two notes slurred to-gether as seee - oo SEET or schweer cheap - beer .
tax o nomic notes: The nom i nate and east-ern most sub spe cies, which is en demic to Kosrae, dif fers from other sub spe cies in that adult sex ual di mor phism is much re duced, and ju ve nile males are dark sooty gray with a tri-an gu lar patch of red - tipped feath ers in the cen ter of the back. Other sub spe cies of M. ru-bratra oc cur in Pohnpei, Chuuk, Yap, Palau, and the Marianas (Baker 1951).
Populations of Mayr ’ s (1945) orig i nal M. cardinalis com plex are dis trib uted among three
widely sep a rated geo graphic cen ters: one in east ern Melanesia, one in Samoa, and one in Micronesia. On the ba sis of re gion - wide dif-fer ences in col or a tion and vo cal i za tions, Pratt et al. (1987) sep a rated the M. rubratra group (Micronesia) from the M. cardinalis group (east ern Melanesia and Samoa). See Pratt and Mittermeier (2016) for fur ther de tails of the lat ter and Higgins et al. (2008) for il lus tra-tions by H.D.P. of the en tire M. cardinalis com plex.
Kosrae White - eye ( Zosterops cinereus )
Common res i dent. This en demic spe cies was fi rst de scribed from Kosrae by Kittlitz (1832 – 1833). It is com mon, for ag ing in small fl ocks in vir tu ally ev ery ter res trial hab i tat, in-clud ing man groves. F.E.H. ob served it at 589 m near the sum mit of Mt. Mutunte on 21 April 2013. Engbring et al. (1990) reported a mean den sity of 1,846 / km 2 with an es ti mated to tal pop u la tion of 203,375; den si ties were higher in the rainforest than in man groves and were in versely cor re lated with el e va tion (Table 2). Engbring et al. (1990) noted that fl ocks typ i cally in cluded four to eight in di vid-u als and that early in the morn ing large num-bers fl ew from the low lands to ward the in te ri or, but they were un cer tain whether it was a daily or sea sonal pat tern.
breed ing: No da ta. F.E.H. photographed a ju ve nile on 22 June 2013 ( Figure 14 ) with an en larged rec tus, pure yel low bill, green - tinged plum age (pos si bly an ar ti fact of light ing), and mostly yel low legs with a few dark blotches. A pre sum ably older im ma ture photographed by C.J.C. on 6 No vem ber 2009 has a yel low bill with dark blotches and adult plum age col or. No dis tinc tive ju ve nile col or a tion has been pre vi ously de scribed for the Kosrae White - eye. Distinct ju ve nile plum ages are rare ly, if ev er, found among white - eyes (van Balen 2008).
spec i mens: NSMT (online), SMF (Stein-bacher 1954, but not listed online), USNM (Townsend and Wetmore 1919), YIO (Taka - Tsukasa and Kuroda 1915 a; online), YPM (on-line), ZIN (Kittlitz 1836).
vo cal i za tions: ML (online), XC (online). The vo cal rep er toire of the Kosrae White - eye
116 PACIFIC SCIENCE · Jan u ary 2016
is de scribed here for the fi rst time and com-pared to that of the Gray - brown White - eye ( Z. ponapensis ) of Pohnpei (ML num bers are cur rently listed un der Gray - brown White - eye). Call notes are of two types. Isolated for-ag ing birds or pairs ut ter a na sal, reedy tschew (ML 55625) that any one fa mil iar with the ge-nus would rec og nize as that of a Zosterops, but dis tinc tive in its own right. Similar calls of Gray - brown White - eyes are higher pitched and less reedy. Roaming fl ocks of fi ve to seven Kosrae White - eyes of ten ut ter a dif fer ent se-ries of calls as they fl y from perch to perch, but do not call in fl ight. These notes are con-trastingly loud and sharp, with the sound qual ity of a child ’ s squeak - toy (ML 55626, 55645). Observers in North America might com pare them to calls of the Brown - headed Nuthatch ( Sitta pusilla ), but more vig or ous. Kosrae White - eyes also ut ter iso lated sin gle notes that seem to be alarm calls, or short bursts that may rep re sent a true song. Appar-ently the Kosrae White - eye has no dawn song. In two dawn re cord ing ses sions in places where the white - eye is com mon dur ing the day (see Micronesian Myzomela ac count), H.D.P. could de tect no in put from the Kos-rae White - eye. However, two days later on Pohnpei, he dis cov ered that the Gray - brown White - eye has a dis tinc tive com plex dawn song (ML 55690).
tax o nomic notes: This spe cies was long con sid ered the nom i nate sub spe cies of the Dusky White - eye com plex along with Z. c. fi nschii of Palau and Z. c. ponapensis of Pohn-pei (Baker 1951) (see Figure 15 ). Pratt et al. (1980) split the Palau form, the Dusky White -
eye ( Z. fi nschii ), on the ba sis of strik ing dif-fer ences in col or a tion, vo cal i za tions, and be hav ior, leav ing the Kosrae and Pohnpei birds as sub spe cies of the Gray - brown White - eye (Pratt et al. 1987, Wiles 2005), a de fault tax on omy that can be eval u ated now that the vo cal dif fer ences be tween the two forms have been documented. The two dif fer ob vi ously in col or, with the Kosrae bird lacking any brown in the oth er wise gray plum age while Pohnpei birds are strongly tinged ol ive - brown dor sally and on the fl anks (Figure 15). The white eye - ring of the Kosrae White - eye is but a sin gle row of feath ers and con se quently is of ten not clearly vis i ble in study skins, which may ex-plain Baker ’ s (1951) misstating that it lacks an eye - ring. Pohnpei birds also have smaller bills (Baker 1951), per haps resulting from char ac-ter dis place ment be cause Pohnpei has two other white - eye spe cies. Clearly, the two forms have adapted to com pletely dif fer ent avian com mu ni ties and prob a bly dif fer in their for ag ing ecol o gy, which has not been stud ied.
Given all the dif fer ences in vo cal i za tions, col or a tion, mor phol o gy, and very likely ecol-o gy, con tinu ing to con sider Z. cinereus con-spe cifi c with Z. ponapensis is clearly un ten a ble; there fore, we treat the two forms as sin gle is-land en demic spe cies con fi ned to Kosrae and Pohnpei, re spec tive ly.
Mi cro ne sian Starling ( Aplonis opaca opaca )
Common res i dent. The nom i nate sub spe-cies was de scribed from Kosrae by Kittlitz (1832 – 1833) and is en demic to Kosrae (Baker 1951). Coultas (1931) noted that this bird was the sec ond most com mon landbird spe cies and that fl ocks oc curred through out the island at all el e va tions. F.E.H. ob served it at 589 m near the sum mit of Mt. Mutunte on 21 April 2013. Coultas (1931) reported that im ma ture - plum-aged birds outnumbered adults by a ra tio of 5:1, and that youn ger birds fl ock to geth er. The high ra tio of im ma tures to adults also oc curs else where (Engbring et al. 1990) and sug gests ei ther a high rate of re pro duc tion or an un usu-ally long re ten tion of juvenal plum age (Jen-kins 1983). Engbring et al. (1990) reported a mean den sity of 835 / km 2 with an es ti mated
Figure 14. Adult (A) and ju ve nile (B) Kosrae White-eyes (Zosterops cinereus) at Malem on 19 June 2014 and at Ta-funsak on 22 June 2013, re spec tive ly. The ju ve nile has a swol len rec tis, pure yel low bill (black in adult), green-tinged plum age (gray in adult), and yel low legs with dark blotches (darker in adult). Photos by F.E.H.
Avifauna of Kosrae · Hayes et al. 117
to tal pop u la tion of 101,955; den si ties were higher in rainforest than in man groves and were highest at low and in ter me di ate el e va-tions (Table 2). Seven ad di tional sub spe cies are found in Micronesia west to Palau and north to Pagan in the Marianas; it is ab sent in the Marshall Islands and Wake (Baker 1951, Wiles 2005).
breed ing: No da ta. spec i mens: AMNH (Baker 1951), MCZ
(online), RMNH (Dekker and Quaisser 2006), USNM (Townsend and Wetmore 1919, Baker 1951), YIO (Momiyama 1920, 1922; Taka - Tsukasa and Yamashina 1931; online), ZIN (Kittlitz 1836).
vo cal i za tions: ML (online).
Kosrae Starling ( Aplonis cor vi na )
Extinct res i dent. Kittlitz (1832 – 1833, 1835) de scribed this en demic spe cies on Kosrae based on an un known num ber of spec i mens. Hume and Peterson (2003) pointed out that
the cor rect pub li ca tion date of the de scrip tion is 1833. Kittlitz (1858) men tioned only three spec i mens de pos ited in the Zoological Mu-seum of the Zoological Institute of the Rus-sian Academy of Sciences in St. Petersburg, which were col lected on 15, 21, and 29 De-cem ber 1827 (Hume 2002). However, two ad-di tional spec i mens, col lected by Kittlitz and sub se quently traded or sold, were later dis cov-ered in the Nationaal Natuurhistorisch Mu-seum in Leiden (Mees 1964). A sixth spec i men in the Senckenberg Naturmuseum Frankfurt in Frankfurt (Hartert 1891) was destroyed dur ing World War II (Steinbacher 1954, Hume 2002). The existing spec i mens in clude two ju ve niles, a sub adult, and two adults. Ac-cording to Kittlitz (1835:7), this spe cies was sol i tary, lived “ deep in the wooded re gion in the cen tre of the island and is just about the only bird oc ca sion ally to be seen here. ” Kittlitz (1835) also noted that it for aged on large in sects, small liz ards, and fruit. Subse-quent col lec tors, es pe cially Coultas (1931),
Figure 15. Three Mi cro ne sian spe cies of Zosterops for merly con sid ered con spe cifi c: Dusky White-eye (Z. fi nschii) of Palau, Gray-brown White-eye (Z. ponapensis) of Pohnpei, and Kosrae White-eye (Z. cinereus) of Kosrae. Painting by H.D.P.
118 PACIFIC SCIENCE · Jan u ary 2016
searched ex ten sively for this spe cies with out suc cess. Hill (1957) claimed that an in for mant and two oth ers in Utwe were “ fa mil iar with it ” and had re cently seen one in the “ Finkol Mountains ” above Utwe dur ing 1955 or 1956. The USFWS team failed to fi nd it in July 1983 de spite ex ten sive searches.
Hume (2002) reviewed four hy poth e ses for its ex tinc tion: (1) com pet i tive ex clu sion by the Mi cro ne sian Starling, which ar rived later and outcompeted it at lower el e va tions, forc ing it to higher el e va tions in the in te ri or; (2) pre da-tion by in tro duced rats, which Kittlitz (1836:284) reported as abun dant and caus ing “ great dam age ” to ba nana and sugar cane; (3) an in tro duced avian dis ease, as in the Ha wai-ian Islands; and (4) hu man dis tur bance and hunt ing. Hume (2002:153) suggested that “ a com bi na tion of events was to blame, ” and we con cur. There is strong ev i dence that early avian col o nists on is lands are forced to spe-cial ize and be come vul ner a ble to ex tinc tion by in ter spe cifi c com pe ti tion from later col o-nists, which tend to be highly com pet i tive gen er al ists (Ricklefs and Bermingham 1999, 2002). We sus pect that the ap par ently dis tinc-tive hab i tat, di e tary, and mor pho log i cal spe-cial i za tions of the Kosrae Starling (Hume 2002) were a di rect out come of com pe ti tion with a later ar riv ing con ge ner, the Mi cro ne-sian Starling, thus greatly re duc ing its pop u la-tion and range, and ren der ing it vul ner a ble to ex tinc tion by one or more of the other fac tors.
breed ing: The two ju ve nile spec i mens were re cently fl edged, ap par ently by late No-vem ber (Hume 2002).
spec i mens: RMNH (Mees 1964), ZIN (Kittlitz 1836).
Blue - faced Parrotfi nch ( Erythrura trichroa trichroa )
Common res i dent. The nom i nate sub spe-cies was fi rst de scribed from Kosrae by Kittlitz (1835) and is en demic to Kosrae (Baker 1951). Coultas (1931:246) regarded it as com mon, oc cur ring in fl ocks through out the island and even at “ quite high el e va tions. ” F.E.H. ob served it only in open ar eas at lower el e va tions at Okat Harbor, Tafunsak, and To-fol in 1981 – 1982 and 2013 – 2014. Engbring
et al. (1990) reported a mean den sity of 111 / km 2 with an es ti mated to tal pop u la tion of 10,893; den si ties were highest at the highest el e va tions, none were in man groves (Table 2), and re gional var i a tion oc curred with the high-est den si ties in Malem (177 / km 2 ) and the low-est in Walung (62 / km 2 ). Engbring et al. (1990) sel dom saw fl ocks larger than four in di vid u-als. It is much more com mon on Kosrae than Pohnpei (Engbring et al. 1990). Other Mi cro-ne sian sub spe cies in habit Chuuk and Pohn-pei ( E. t. clara ) and Palau ( E. t. pelewensis; Baker 1951, Wiles 2005).
breed ing: No da ta. spec i mens: SMF (Steinbacher 1954, but not
listed online), YIO (Taka - Tsukasa and Ya-mashina 1931; online), ZIN (Kittlitz 1836).
vo cal i za tions: ML (online). The me tal lic or in sect - like call notes of this spe cies are well known, but the full song is not, nor has it been pre vi ously reported in Micronesia. H.D.P. re-corded it in a man grove for est at the Pacifi c Tree Lodge on the north east ern coast of Kos-rae on 23 Feb ru ary 1988 (ML 55637 0:44ff). It matches de scrip tions given by Goodwin (1982) and Clement et al. (1993) as com pris ing three dis tinct el e ments uttered re peat edly in se quence. One el e ment is the three - note var-i ant of the me tal lic call tee - dee - deet, an other is a shrill drawn - out descending note tseeeooo, and the other a se ries of upslurred whis tles seet - seet - seet, etc. The en tire song would be: tseeeoo, tee - dee - deet tee - dee - deet tseeeooo seet - seet - seet - seet - seet - seet - seet - seet tseeeoo tee - dee - deet tseeeoo tee - dee - deet tseeeoo tee - dee - deet seet - seet - seet - seet - seet - seet tseeeoo tee - dee - deet tseeeoo .
Hypothetical Species
Eastern Spot - billed Duck ( Anas zonorhyncha )
Possibly a rare vis i tor. Santos et al. (1970) listed this spe cies for Kosrae but pro vided no fur ther de tails. In Micronesia, it has been re-corded only in the Marianas (Wiles 2005).
Pacifi c Black Duck ( Anas superciliosa )
Possibly a rare vis i tor. Segal (1989:220) listed this spe cies among three duck spe cies that are “ oc ca sional vis i tors ” but pro vided no
Avifauna of Kosrae · Hayes et al. 119
fur ther de tails. It breeds at Chuuk and Palau, but these pop u la tions are ap par ently sed en-tary, as are other pop u la tions found across the trop i cal Pacifi c and south to New Zealand (Pratt et al. 1987).
Red - billed Tropicbird ( Phaethon aethereus )
Possibly a rare vis i tor. Finsch (1880 a, b, 1881) reported ob serv ing both the White - tailed Tropicbird and Red - billed Tropicbird in the Marshall Islands and in Kosrae dur ing Feb ru ary 1880. His de scrip tion of the lat ter from Kosrae is un equiv o cal: “ ob served sev eral dis tinc tive (red beak, white tail!) in sight of is-land at sea ” (trans lated from Ger man; Finsch 1880 a: 310). Baker (1951) ac cepted Finsch ’ s re-cords, but Amerson (1969) regarded his ob ser-va tions of Red - billed Tropicbirds from the Marshall Islands (Finsch 1880 b ) as mis taken iden ti fi ca tions of Red - tailed Tropicbirds. Finsch was well aware of the ex is tence of the Red - tailed Tropicbird, which he ob served at Pohnpei only a few days af ter leav ing Kos-rae (Finsch 1880 a ). However, Wiles (2005) pointed out that Finsch (1880 c ) saw more Red - billed Tropicbirds than White - tailed Tropicbirds and did not see any Red - tailed Tropicbirds dur ing his voy age from Hawai ‘ i to the Marshall Islands. Although the Red - billed Tropicbird is much less com mon than the Red - tailed Tropicbird in Hawai ‘ i, it has been recorded at least 19 times as far west as Johnston Atoll (VanderWerf and Young 2007), which is > 5,000 km west of their near-est col o nies in Mexico. Kosrae is an ad di tional 3,250 km west of Johnston Atoll. Given the un cer tainty of Finsch ’ s ob ser va tions and ab-sence of sub se quent re cords in Micronesia, it has been regarded as hy po thet i cal (or even de-let ed) for Micronesia by sub se quent au thor i-ties (Owen 1977, Pyle and Engbring 1985, Pratt et al. 1987, Wiles 2005).
Masked Booby ( Sula dactylatra )
Possibly a rare vis i tor. Hill (1957) reported that a boo by, “ ap par ent ly ” this spe cies, was “ not com mon ” dur ing 1956 – 1957. It has been recorded in all of the island groups of Micro-nesia ex cept Pohnpei and Kosrae (Wiles
2005), so its oc cur rence on Kosrae is plau si-ble but re quires fur ther doc u men ta tion.
Greater Sand Plover ( Charadrius le-schenaultii )
Possibly a rare Pa le arc tic mi grant. We have been un able to fi nd a spec i men or de tailed published re cord from Kosrae. Lesson men-tioned “ Oulan ” in his brief de scrip tion of the spe cies from In dia (Lesson 1826) and in a sub-se quent trea tise of or ni thol ogy (Lesson 1831), resulting in sub se quent au thor i ties list ing it for Kosrae up through Wiles (2005). How-ever, Oustalet (1895:48 – 49) pointed out that “ Oulan, ” which was ital i cized un like other lo-cal ity names in Lesson (1826, 1831), repre-sented a vul gar name for the spe cies in In dia rather than a lo cal i ty, and was misinterpreted for sim i larly spelled “ Oualan, ” the name for Kosrae used else where by Lesson (1827, 1829 – 1830 a, b, 1831) and Lesson and Garrot (1826). Unfortunately the mis in ter pre ta tion was per-pet u ated for more than a cen tu ry. Because this spe cies has been recorded from all ma jor island groups west of Kosrae (Wiles 2005), its even tual doc u men ta tion in Kosrae is ex-pected.
Swinhoe ’ s Snipe ( Gallinago megala )
Possibly a rare Pa le arc tic mi grant. Identi-fi cation of snipes to spe cies is no to ri ously dif-fi cult (Message and Taylor 2005). Among a group of four snipes at Tofol on 4 April 2014, C.J.C. obtained a blurry photo of one fl y ing away ( Figure 16 ) that clearly lacked the white trailing edge of the Common Snipe ( G. gal-linago ). It appeared too dark for Latham ’ s Snipe ( G. hardwickii ), which has been docu-mented by a spec i men from the Marshall Is-lands (Amerson 1969) with un con fi rmed sight re cords from Yap and Rota (Pratt et al. 2010) and an un pub lished pho to graphic re cord from Guam (D. Vice, pers. comm.). It re sem bled a Swinhoe ’ s Snipe, which is a fairly com mon win ter vis i tor in Palau, Yap, and the Marianas, and a rare vis i tor to Chuuk (Wiles 2005), but it also could have been a Pin - tailed Snipe ( G. stenura ), only re cently con fi rmed for the Mar-ianas (D. Vice, pers. comm.; de tails to be
120 PACIFIC SCIENCE · Jan u ary 2016
published else where), al though the legs ap pear to be too short, barely pro trud ing be yond the tail (Figure 16). We regard this identifi cation as hy po thet i cal un til bet ter doc u men ta tion is avail able.
Greater / Lesser Yellowlegs ( Tringa melano-leuca / fl avipes )
Possibly a rare Ne arc tic mi grant. Santos et al. (1970) listed the Greater Yellowlegs and Segal (1989:220) in cluded the “ Yellow Legs ” among sev eral shore bird spe cies that are “ less com mon but yearly vis i tors, ” but no fur ther de tails were pro vid ed. The Greater Yellowlegs has been recorded as a va grant in the Com-monwealth of the Northern Marianas, Mar-shall Islands, and Wake, and the Lesser Yellowlegs has reached the Marshall Islands (Wiles 2005).
Red - necked Phalarope ( Phalaropus lobatus )
Possibly a rare bo real mi grant. Hill (1957) reported it from Kosrae but did not state whether she had ac tu ally seen any. She stated that an in for mant “ seems to be cer tain ” of its iden tity based on a pic ture (con fu sion with the Sanderling is pos si ble) and a de scrip tion of its hab its. In Micronesia, it is known only from the north ern Mariana Islands (Wiles 2005) and Palau (Pratt et al. 2010).
Cuckoo sp. ( Cuculus optatus )
Possibly a rare Pa le arc tic mi grant. C.J.C. very briefl y ob served a gray, cuck oo - like bird with a long, barred tail, but was un able to ob-tain any pho tos, at Okat Harbor on 13 Oc to-ber 2014. In Micronesia, two spe cies of this ge nus have been recorded from Palau and Yap, with un con fi rmed re ports from the Com-monwealth of the Northern Marianas and Pohnpei (Wiles 2005).
Short - eared Owl ( Asio fl ammeus )
Possibly a rare vis i tor. Ridgway (1881) briefl y de scribed an unsexed adult spec i men (USNM 66235) col lected at “ Strong ’ s Island, ” which he be lieved was in the West Indies. Kelso (1938) pointed out that Strong ’ s Island was an old name for Kosrae and sub se quent au thor i ties up through Wiles (2005) con sid-ered the spec i men to be a valid re cord from Kosrae. According to the la bel, the un dated spec i men was col lected by “ Gulick ” of “ Wes-leyan University. ” Presumably it was col-lected by either Luther H. Gulick, John T. Gulick, or Theodore W. Gulick, all bro th ers who were American missionaries (John later became an accomplished biologist, specializ-ing in mollusks and evolutionary theory). Luther and John briefl y visited Kosrae dur-ing 22–30 August 1852, and John returned during 6–15 October 1852 (Jewett 1895, Gulick 1932). Kelso (1938) and R. B. Clapp (in Engbring et al. 1990) mea sured the spec-i men ’ s wings, which were 20 – 40 mm shorter than those of con ti nen tal pop u la tions and typ i cal of shorter - winged in su lar forms. Baker (1951) in cluded it within the en demic sub spe cies A. f. ponapensis of nearby Pohnpei but did not dis cuss its or i gin. R. B. Clapp (in Engbring et al. 1990) and H.D.P. found that the al leged color dis tinc tions of A. f. ponapen-sis in Mayr ’ s (1933) de scrip tion could be matched by spec i mens of nom i nate A. f. fl ammeus of Asia, thus its smaller size is its only mor pho log i cal dis tinc tion as an en demic sub spe cies. Engbring et al. (1990) suggested that the Kosrae spec i men may have ar rived on Kosrae as a va grant from Pohnpei or that it was ac tu ally col lected on Pohnpei but
Figure 16. First pos si ble re cord of Swinhoe’s Snipe (Gal-linago megala) for Kosrae at Tofol on 4 No vem ber 2014. Photo by C.J.C.
Avifauna of Kosrae · Hayes et al. 121
mislabeled as orig i nat ing from Kosrae. Al-ternatively, the spec i men may have repre-sented a res i dent pop u la tion on Kosrae that was sub se quently ex tir pat ed, but the island prob a bly lacked suf fi cient grass land hab i tat to sup port a res i dent pop u la tion. Luther re sided on Pohnpei from 1852 – 1859, but vis ited Kosrae only briefl y with John in 1852 (Jewett 1895). After his fi rst visit to Kosrae, John wrote, “ I was sur prised not to see more wild birds in my ram bles on Kusaie. We heard the cooing of pi geons and the songs of two or three other kinds of small birds ” (Gulick 1932:79). Had an owl been en coun tered dur-ing ei ther of John ’ s vis its, he prob a bly would have writ ten about it. Theodore collected birds on Pohnpei (and possibly Kosrae as well) and may have collected the owl speci-men. He wrote, “ My recreation has been gar-dening, paddling canoes, shooting birds, and the like. I have made a collection of Ponape [Pohnpei] birds, most of which are quite dif-fer ent from ours at the Sandwich [Hawaiian] Islands ” (Chamberlain 1860:9). We do not know the fate of his bird collection from Pohnpei, but we suggest it was eventually de-posited at Wesleyan University and included the owl specimen, which was mislabeled as originating from Kosrae. Occasional re cords of Short - eared Owl from the Mariana Islands (Wiles 2005), Yap (Pratt et al. 2010), Marshall Islands (Spennemann 2004), and Wake (Jones 1995) are prob a bly mi grants of the lon ger - winged nom i nate race from Asia.
spec i men: USNM (Ridgway 1881, Kelso 1938, Engbring et al. 1990).
Sacred Kingfi sher ( Todiramphus sanctus )
Possibly a rare Aus tra lian mi grant. Lauret (1990) briefl y ob served a king fi sher at Malem on 7 Feb ru ary 1988, but was un able to view its un der parts. It was most likely this highly mi-gra tory spe cies, which has also been ten ta tively reported from Kapingamarangi Atoll in south-ern Pohnpei State (Buden 1998) and Nauru (Buden 2008), and con fi rmed on Kwajalein Atoll in the Marshall Islands (Schipper 1985).
Carolinian Reed Warbler ( Acrocephalus syr-inx )
This spe cies was con sid ered by many au-thors to be an ex tir pated res i dent un til Eng-bring et al. (1990) fi rst questioned the ve rac ity of his tor i cal re ports. Hartlaub (1852:131) listed “ Tatare syr inx ” for “ Ualan ” (Kosrae) and “ Lu-gunor, ” which he at trib uted to Kittlitz with-out cit ing a ref er ence. Kittlitz (1835:6) de scribed the spe cies from the is lands of “ Ul-cei ” and “ Lugunor ” ; the for mer re fers to Woleai in Yap and the lat ter is an atoll in Chuuk. Hartlaub ev i dently con fused “ Ulcei ” with “ Ualan, ” which led to con fu sion in the sub se quent lit er a ture. Kittlitz (1858) discussed “ Sylvia syr inx ” (a syn o nym of this spe cies) and then “ Lamprothornis ” (a syn o nym of the Mi-cro ne sian Starling) on p. 92, but did not men-tion “ Ualan ” un til p. 93 as a lo cal ity for the lat ter spe cies. Finsch (1881:108) stat ed, “ I sus-pect Calamoherpe syr inx [an other syn o nym] may still be found in Kushai, al though it is not men tioned by Kittlitz and es caped my no tice. ” Despite Finsch ’ s state ment that nei ther he nor Kittlitz had found it on Kosrae, Baker (1951) cited Hartlaub (1852), Kittlitz (1858), and Finsch (1880 a, 1881) among other au thors for its oc cur rence on Kosrae, even though none had ever reported ob serv ing or collecting it on Kosrae. Elsewhere it oc curs in Yap, Chuuk, and Pohnpei (Wiles 2005).
dis cus sion
Thus far, 53 nat u rally oc cur ring spe cies of birds have been re li ably reported from Kosrae. Of these, 39 have been prop erly documented by ei ther a spec i men de pos ited in a per ma nent mu seum col lec tion (24 spe cies) or a published pho to graph (15 spe cies), and 14 are based on one or more sight re cords (Table 1). Twelve po ten tially nat u rally oc cur ring spe cies have been reported on the ba sis of in suf fi ciently sub stan ti ated sight re cords or er ro ne ous in ter-pre ta tion of the lit er a ture and are there fore con sid ered hy po thet i cal. Two ad di tional spe-cies have been in tro duced (one represented by spec i mens, the other by sight re cords), of which one has a self - sus tain ing fe ral pop u la-tion and the other is ex tir pat ed.
Of the 15 nat u rally oc cur ring breed ing res-i dents, 13 spe cies are ex tant (in clud ing two en demic spe cies and four en demic sub spe cies)
122 PACIFIC SCIENCE · Jan u ary 2016
and two spe cies (both en dem ic) are ex tinct. Of the 38 non breed ing res i dents, 21 are bo real mi grants (in clud ing 11 ex clu sively Pa le arc tic mi grants and 3 ex clu sively Ne arc tic mi grants), 5 are aus tral mi grants from breed ing pop u la-tions in the tem per ate Southern Hemisphere, and 12 are vis i tors that breed on trop i cal is-lands else where in the Pacifi c (Table 1).
Further fi eld work will un doubt edly add new spe cies to the island ’ s avi fauna and ad di-tional re cords of the rarer spe cies. All re cords of new spe cies and those of spe cies repre-sented only by sight re cords should be prop-erly documented by a published pho to graph or sound re cord ing, or a spec i men de pos ited in a per ma nent mu seum col lec tion.
Although data on the pop u la tions of mi gra-tory shore birds and res i dent landbirds have been published (Hayes 1986, Engbring et al. 1990), ad di tional mon i tor ing of these pop u la-tions and those of other spe cies is needed to de tect any fu ture changes in pop u la tion lev-els. Much more in for ma tion is needed on the breed ing sea son, breed ing bi ol o gy, for ag ing be hav ior, and hab i tat re quire ments of Kosrae ’ s res i dent spe cies. Because of the island ’ s his tor-i cally low hu man pop u la tion den sity (Goren-fl o 1993) and rel a tively pris tine hab i tats (Wortel 2010), most of the breed ing birds ap-pear to be thriv ing with no se ri ous threats. The Mi cro ne sian Imperial Pigeon, which is still heavily hunted, is the only ex cep tion. Vigilance is warranted and steps should be taken to mit i gate any fu ture threats to the is-land ’ s avi fau na, such as hab i tat de struc tion and in tro duced spe cies.
ac knowl edg ments
We thank the fol low ing in di vid u als for their as sis tance: Oliod Moura and Larry Neth for assisting F.E.H. in the field; Roger Clapp and John Engbring for ad vice dur ing prep a ra tion of an ear lier draft; Mary LeCroy for send ing Coultas ’ s un pub lished man u script; Todd Mark, Grady Pettigrew, and Ross Silcock for shar ing their field ob ser va tions, pho to graphs, and sound re cord ings; Barbara Roth and Ste-fan Kreft for trans la tion of Ger man text; Saya Uehara for trans la tion of Jap a nese text; Eric
VanderWerf, Gary Wiles, Dan Vice, and Pe-ter Pyle for help with iden ti fy ing some of the pho to graphs; Greg Budney, Matt Medler, and Jessie Barry of the Macaulay Library for as-sisting H.D.P. in ar chiv ing his re cord ings; Rich ard Roe for reviewing our ac counts of ex-tinct spe cies; Molly Hagemann and Nancy Vander Velde for pro vid ing in for ma tion on the Tris tram ’ s Storm Petrel spec i men and the Gulick family; and Gary Wiles and an anon y mous ref eree for thor oughly review-ing the sub mit ted man u script.
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