PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST A T 79TH STREET, NEW YOR K, NY 10024
Number 3646, 37 pp., 14 figures, 5 tables June 25, 2009
Pacific Flying Foxes (Mammalia: Chiroptera):
Two New Species of Pteropus from Samoa,
Probably Extinct
KRISTOFER M. HELGEN,1 LAUREN E. HELGEN,2 AND DON E. WILSON3
ABSTRACT
Two new species of flying foxes (genus Pteropus) from the Samoan archipelago are described on
the basis of modern museum specimens collected in the mid-19th century. A medium-sized species
(P. allenorum, n. sp.) is introduced from the island of Upolu (Independent Samoa), based on a
specimen collected in 1856 and deposited in the Academy of Natural Sciences of Philadelphia. It
has not been collected again, and we regard it as almost certainly extinct. This species is smaller
bodied and has much smaller teeth than both extant congeners recorded in the contemporary
fauna of Samoa (Pteropus samoensis and P. tonganus). The closest relative of this new species may
be Pteropus fundatus of northern Vanuatu. The disjunct historical distribution of these two small-
toothed flying foxes (in Vanuatu and Samoa) suggests that similar species may have been more
extensively distributed in the remote Pacific in the recent past. Another species, a very large flying
fox with large teeth (P. coxi, n. sp.), is described from two skulls collected in Samoa in 1839?1841
during the U.S. Exploring Expedition; it too has not been collected since. This robust species
resembles Pteropus samoensis and Pteropus anetianus of Vanuatu in craniodental conformation but
is larger than other Polynesian Pteropus, and in some features it is ecomorphologically convergent
on the Pacific monkey-faced bats (the pteropodid genera Pteralopex and Mirimiri). On the basis of
eyewitness reports from the early 1980s, it is possible that this species survived until recent decades,
or is still extant. These two new Samoan species join Pteropus tokudae of Guam, P. pilosus of
Palau, P. subniger of the Mascarenes, and P. brunneus of coastal north-eastern Australia as flying
Copyright E American Museum of Natural History 2009 ISSN 0003-0082
1 Division of Vertebrate Zoology (Mammalogy), American Museum of Natural History; Division of Mammals, National
Museum of Natural History, Smithsonian Institution, Washington, D.C. 20013-7012; Vertebrate Zoology, Bernice P.
Bishop Museum, Honolulu, Hawaii (helgenk@si.edu).
2 Division of Mammals and Department of Entomology, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. (helgenl@si.edu).
3 Division of Mammals, National Museum of Natural History, Smithsonian Institution, Washington, D.C.
(wilsond@si.edu).
foxes with limited insular distributions that survived at least until the 19th century but are now
most likely extinct.
INTRODUCTION
Capable of flight, bats are the only mam-
mals to have naturally colonized many of the
remote oceanic islands and archipelagos of the
Pacific?eastern Melanesia, Polynesia, and
Micronesia, including the islands of Vanua-
tu, Fiji, New Caledonia, Samoa, Tonga,
Wallis and Futuna, Niue?, the Cook Islands,
New Zealand, Lord Howe Island, Guam, the
Marianas, Palau, and the Carolines (Flannery,
1995). Some insect-eating bats are found in
modern and subfossil insular faunas through-
out the region, including representatives of the
emballonurid genus Emballonura (Micronesia,
Vanuatu, Fiji, Samoa, Tonga); the hipposider-
id Hipposideros (Vanuatu); the vespertilionids
Miniopterus (Vanuatu, New Caledonia), Myo-
tis (Vanuatu, possibly Samoa), Nyctophilus
(New Caledonia, formerly Lord Howe, possi-
bly Fiji), and Chalinolobus (Lord Howe, New
Zealand); the molossid Chaerephon (Vanuatu,
Fiji, formerly Tonga); and the monogeneric
family Mystacinidae (Mystacina), today en-
demic to New Zealand (Hill and Daniel, 1985;
Flannery, 1995; Hand et al., 1998; Parnaby,
2002a; Helgen and Flannery, 2002; Simmons,
2005). Most remote Pacific bat species, how-
ever, are fruit- and nectar-feeding species
classified in the family Pteropodidae (see
Andersen, 1912). The genus Pteropus (the
??flying foxes??) is represented by a diverse
complement of species in eastern Melanesia,
Micronesia, and Polynesia (extending as far
east as the Cook Islands). Also occurring in
the region are two older generic lineages
endemic to the remote Pacific?Notopteris
(two species of blossom bats, known from
the modern faunas of New Caledonia, Vanua-
tu, and Fiji, as well as the subfossil record of
Tonga) and Mirimiri (one species of ??monkey-
faced bat??, recorded only from montane
forests on the island of Taveuni in Fiji) (Hill
and Beckon, 1978; Flannery, 1995; Parnaby,
2002b; Helgen, 2005; Palmeirim et al., 2007).
In the 18th and 19th centuries, European
explorers of Pacific archipelagos encountered
a number of endemic vertebrate species?
especially birds, but also lizards and bats,
which became rapidly extinct soon after their
discovery by science. Setting aside New
Zealand and the Hawaiian Islands, both of
which are well known as epicenters of
historical (as well as prehistoric) avian extinc-
tion (Tennyson and Martinson, 2006; Worthy
and Holdaway, 2002; Pratt, 1994; Ziegler,
2002), some of the better documented exam-
ples of Polynesian extinctions prior to the 20th
century include the Tahitian Sandpiper
(Prosobonia leucoptera) and Raiatea Parakeet
(Cyanoramphus ulietanus) of the Society
Islands, which were last recorded before
1800; the Mysterious Starling (Aplonis mavor-
nata) of the Cook Islands, the Tongan Giant
Skink (Tachygia microlepis), and the Tahiti
Parakeet (Cyanoramphus zealandicus), all of
which were last recorded in the first half of the
19th century; and the Samoan Wood-Rail
(Pareudiastes pacificus), last recorded in the
latter half of the 19th century (Flannery and
Schouten, 2001). There are many other avian
examples (Steadman, 2006b). The arrival of
European impacts in the Pacific during the
1700s and 1800s clearly fostered or accelerated
an astonishing extinction pulse in these insular
vertebrate faunas?a pulse that has carried on
unabated and today threatens the survival of
many critically endangered species throughout
the region (Flannery, 1995; Stattersfield et al.,
1998; Steadman, 2006b).
The purpose of the present paper is to bring
to light two additional examples of Polynesian
vertebrate species?both previously unnamed
flying foxes?that have not been recorded
since the 19th century. Both new species
originate from the archipelago of Samoa in
West Polynesia (figs. 1, 2), where only two
species of Pteropus (P. tonganus and P.
samoensis) are known in the contemporary
fauna (e.g., Andersen, 1912; Cox, 1983;
Wilson and Engbring, 1992; Flannery, 1995;
Banack, 1996, 1998; Brooke, 2001). One of the
new species is recorded by a single museum
specimen collected on the island of Upolu in
1856. Deposited in the collections of the
Academy of Natural Sciences of Philadel-
phia, this specimen has been overlooked by
systematists since the time of its collection.
(Although identified as ??Pteropus pselaphon???
in the Academy?s accession catalog and as
2 AMERICAN MUSEUM NOVITATES NO. 3646
??Pteropus tonganus?? on its skull box, it bears
no particular morphological resemblance to
either of these species.) The other species is
represented by two skulls in the United States
National Museum in Washington, D.C.,
collected in Samoa during the 1838?1842
U.S. Exploring Expedition to the Pacific (see
Wilkes, 1844; Peale, 1848; Cassin, 1858;
Philbrick, 2003). These have been identified
and cataloged as Pteropus samoensis since
their collection, and they even apparently
formed part of the hypodigm for the original
taxonomic description of P. samoensis by
Peale (1848). However, we argue that these
two skulls represent a species distinct from
(and, presumably, formerly sympatric with) P.
samoensis, with a considerably more robust
skull and teeth than any extant Pteropus from
Polynesia. We describe both of these over-
looked Samoan bat taxa as new species, and
designate a lectotype for Pteropus samoensis
Peale, 1848, to preserve the traditional usage
of this epithet (i.e., for the shorter faced and
more cranially robust of the two Pteropus
species known to survive in Samoa today;
Andersen, 1912; Wilson and Engbring, 1992).
MATERIALS AND METHODS
Specimens discussed herein are deposited in
the collections of the American Museum of
Natural History, New York (AMNH);
Australian Museum, Sydney (AM); Academy
of Natural Sciences, Philadelphia (ANSP);
Natural History Museum, London (BMNH);
Museum of Comparative Zoology at Harvard
University, Cambridge, Mass. (MCZ); Muse?um
National d?Histoire Naturelle, Paris (MNHN);
United States National Museum of Natural
History, Smithsonian Institution, Washington,
D.C. (USNM); Museum fu?r Naturkunde,
Humboldt Universita?t, Berlin (ZMB); and
Zoological Museum of the University of
Copenhagen, Copenhagen (ZMUC).
Terminology for cranial and dental features
follows Giannini et al. (2006) and Giannini
and Simmons (2007). All measurements of
length are in millimeters. Standard external
measurements for most museum specimens
were recorded by the original collectors in the
field; in other cases forearm lengths were
measured from dry skins or from specimens
preserved in alcohol. Craniodental and exter-
Fig. 1. Map of the southwest Pacific region. Adapted from Steadman (2006b).
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 3
nal variables were measured with hand-held
callipers to the nearest 0.1 mm. Single-tooth
measurements are measured on the crown.
For wing measurements (chord or straight-
line lengths in the case of curved phalangeal
measurements), digit is abbreviated as ??D??,
metacarpal as ??M??, and phalanx as ??P??; thus,
D2P2 refers to the second phalanx of the
second digit, and so forth.
Cranial measurements are abbreviated
(and, where necessary, defined) as follows:
CBL, condylobasal length; ONL, orbitonasal
length, here defined as the distance from the
anterior edge of the orbit to the midpoint of
the premaxillae; ZYG, greatest bizygomatic
width; MTR, alveolar length of maxillary
toothrow, C1?M2; CC, external, alveolar
distance across upper canines; M1M1, exter-
nal, alveolar distance across upper first
molars, M1?M1; M2M2, external, alveolar
distance across upper second molars, M2?M2;
BBC, breadth of braincase at zygomata; ML,
greatest length of mandible; CHM, height of
mandible to coronoid process; LTR, alveolar
length of mandibular toothrow, c1?m3; LM1,
length of first upper molar; WM1, width of
first upper molar. Unless otherwise noted, all
craniodental measurements are based only on
adult skulls, identified as those in which the
mature dentition is fully in place and the
basioccipital-basisphenoid (basilar) suture is
completely fused. Because our measured
samples of Pteropus species featured in this
paper do not exhibit statistically significant
intersexual metric differences in these sampled
craniodental variables (i.e., t-test comparisons,
p . 0.05), we have pooled adult males, adult
females, and unsexed adult skulls in our
univariate tabulations and morphometric
analyses. We calculated standard descriptive
statistics (mean, standard deviation, and
observed range) for the samples of popula-
tions and species listed in table 1. Our plot-
tings of specimen scores in multivariate
analyses provide visual patterns that reflect
similarity or contrast in the combination of all
cranial and dental dimensions among samples
of different species. Principal components
analyses and discriminant function analyses
were computed using the combination of
Fig. 2. A map of Samoa, showing political boundaries and principal islands, with inset showing the
location of Apia on the island of Upolu, the type locality of Pteropus allenorum. Adapted from
Steadman (2006b).
4 AMERICAN MUSEUM NOVITATES NO. 3646
cranial and dental measurements indicated in
table 1, with the exception of CBL (unavail-
able for either available skull of P. coxi) and
BBC (measurable in only one of the two skulls
of P. coxi), omitted to allow for the inclusion
of these critical samples. All measurement
values were transformed to natural logarithms
prior to analysis. Principal components were
TABLE 1
Forearm and Selected Cranial Measurements of the Four Flying Foxes of Samoa
Values for Fijian P. samoensis and P. anetianus of Vanuatu are shown for comparison (provided for each sample
are mean 6 SD, range of measurements, and sample size) Measurements for Samoan and Fijian populations
based on adult specimens at AM, AMNH, ANSP, BMNH, USNM, and ZMUC. Cranial measurements for P.
anetianus are based on specimens at AMNH, BMNH, and USNM; external measurements for P. anetianus are
compiled from Andersen (1912), Felten and Kock (1972), and Flannery (1995) where explicit measurements
were provided by those sources. Asterisked values refer to an estimated measurement from an incomplete skull.
P. allenorum P. tonganus P. coxi P. samoensis P. samoensis P. anetianus
Samoa subadult Samoa adults Samoa adults Samoa adults Fiji adults Vanuatu adults
Forearm 116 138 6 5.6 ? 140 6 6.0 128 6 4.2 126 6 6.3
? 130?146 ? 128?154 122?134 117?135
n 5 1 n 5 11 ? n 5 15 n 5 10 n 5 17
CBL 50.0 62.0 6 1.29 ? 58.1 6 1.50 58.7 6 0.97 58.9 6 14.5
? 59.7?64.1 ? 55.7?60.3 57.1?59.8 55.1?60.7
n 5 1 n 5 16 ? n 5 17 n 5 7 n 5 17
ONL 16.5 21.5 6 0.89 19.9 18.4 6 0.77 18.8 6 1.03 18.0 6 0.80
? 19.5?22.7 19.6?20.3 16.6?20.1 17.6?20.7 16.4?19.4
n 5 1 n 5 19 n 5 2 n 5 23 n 5 11 n 5 21
ZYG 26.0 34.3 6 1.58 37.5* 33.9 6 1.40 34.3 6 1.10 35.5 6 1.28
? 31.8?37.1 37*?38 31.8?37.4 32.0?35.7 31.7?37.7
n 5 1 n 5 19 n 5 2 n 5 19 n 5 10 n 5 19
BBC 20.6 21.9 6 0.62 23.6 21.7 6 0.57 22.1 6 0.51 22.1 6 0.84
? 20.5?23.0 ? 20.6?22.9 21.3?23.2 20.4?23.4
n 5 1 n 5 19 n 5 1 n 5 23 n 5 10 n 5 19
MTR 18.4 23.6 6 0.60 23.5 21.7 6 0.61 21.7 6 0.85 22.1 6 0.57
? 22.7?24.9 23.4?23.6 20.4?22.6 20.0?22.8 20.3?22.8
n 5 1 n 5 19 n 5 2 n 5 23 n 5 11 n 5 20
CC 9.7 11.7 6 0.53 12.8 11.8 6 0.39 11.6 6 0.61 11.9 6 0.67
? 10.8?12.7 12.7?12.9 10.8?12.5 10.0?12.2 9.9?13.0
n 5 1 n 5 19 n 5 2 n 5 23 n 5 11 n 5 21
M2M2 13.0 15.0 6 0.69 15.4 14.1 6 0.62 14.4 6 0.55 14.6 6 0.62
? 13.9?16.4 15.2?15.7 13.1?15.6 13.7?15.6 13.0?15.4
n 5 1 n 5 19 n 5 2 n 5 19 n 5 9 n 5 19
ML 38.7 50.5 6 1.37 50.6 47.2 6 1.18 46.3 6 1.19 47.7 6 1.31
? 48.3?51.3 50.6?50.7 44.0?48.9 43.2?47.6 43.3?49.9
n 5 1 n 5 19 n 5 2 n 5 23 n 5 11 n 5 21
MH 17.7 24.0 6 1.07 28 25.8 6 1.40 24.0 6 0.91 27.2 6 1.15
? 21.1?25.3 27.7?28.3 21.5?27.8 22.7?25.3 24.4?28.5
n 5 1 n 5 19 n 5 2 n 5 23 n 5 11 n 5 21
LTR 26.7 6 0.77 27.2 24.9 6 0.70 24.8 6 0.84 25.5 6 0.84
? 25.9?28.2 26.6?27.8 22.5?25.9 22.8?25.4 23.2?26.9
n 5 1 n 5 19 n 5 2 n 5 23 n 5 11 n 5 20
LM1 4.2 5.05 6 0.21 5.92 5.41 6 0.23 4.90 6 0.19 5.5 6 0.34
? 4.84?5.50 5.89?5.94 4.98?5.85 4.76?5.17 4.77?6.06
n 5 1 n 5 17 n 5 2 n 5 22 n 5 4 n 5 22
WM1 1.9 2.99 6 0.21 3.15 2.80 6 0.13 2.59 6 0.09 3.0 6 0.23
? 2.64?3.43 3.12?3.17 2.45?3.08 2.47?2.67 2.53?3.34
n 5 1 n 5 17 n 5 2 n 5 22 n 5 4 n 5 22
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 5
extracted from a covariance matrix, and
canonical variates were extracted from the
discriminant function analyses. The software
program Statistica 8.0 (Statsoft, Tulsa, Okla.)
was used for all analytical procedures.
SYSTEMATICS
Pteropus allenorum, new species
HOLOTYPE: The holotype of Pteropus alle-
norum is ANSP 1234, a skin in alcohol (fig. 3)
with the cranium and mandible removed and
cleaned (fig. 7), collected at Apia (13u499S,
171u449W), a harbor and settlement on the
island of Upolu (and today the capital of
Independent, or Western, Samoa; fig. 2),
apparently by H.C. Caldwell in April 1856
(see below), and donated to the museum by
W.S.W. Ruschenberger. Judging from its
craniodental development (fig. 7; see Helgen,
2004a; Giannini et al., 2006), this unsexed
specimen is a nearly mature subadult.
A faded tag bearing scripted ink writing
accompanies the skin in alcohol, which we
take to be the original or at least the oldest tag
associated with it. Both this tag and another?
affixed to the specimen, less faded, and
bearing writing in pencil?give the provenance
of the specimen as ??Apia, Upolu??. Written in
ink on the mandible is the faded annotation
??Apia??. The accession catalog, skull box, and
a penciled label inside the skull box give the
locality as ??Navigator Islands, Apia, Apola??.
(The ??Navigator Islands?? is a 19th-century
appellation for the Pacific archipelago today
known as Samoa, incorporating the modern-
day political boundaries of both Independent
Samoa and American Samoa; see Wilkes,
1844; Keesing, 1934.) The relatively large
and high island of Upolu (area 1100 km2,
maximum elevation ca. 1100 m), home to the
harbor of Apia, is the second largest island in
the Samoan archipelago (after the adjacent
island of Savai9i, with area 1820 km2 and
maximum elevation ca. 1850 m).
The faded tag in the alcohol jar with the
holotype also bears the date ??April 1856??. We
consider this most likely to be the specimen?s
date of collection, rather than the date of
accession at ANSP. (Other specimens listed on
the same page of the ANSP mammal accession
catalog list a ??Date of Presentation?? to the
museum a decade later, in 1865 or 1866.) Tags
associated with the jar of alcohol and the skull
bear the name ??Dr. W.S.W. Ruschenberger??
or ??W.S.W.R.??, who is listed in the ANSP
accession catalog under the column of
??Donor??, rather than ??Collector??.
Amongst ??Donations to the Museum??
received in 1857, the Proceedings of the
Academy of Natural Sciences of Philadelphia
gives the following entries, listed consecutively
(Anonymous, 1858: i):
A collection of Echinodermata, Acelephae,
and Mollusca in alcohol, from the Navigator
Fig. 3. The fragmentary holotype skin of Pteropus allenorum (ANSP 1234, preserved in alcohol).
6 AMERICAN MUSEUM NOVITATES NO. 3646
Islands. Presented by Drs. W. S. W.
Ruschenberger, and Henry Clay Caldwell,
U.S.N[avy].
A specimen of Pteropus from the same
locality.
Based on its unique and concordant label
data (noting its collection in Samoa in 1856
and presentation to the museum by
Ruschenberger) we strongly suspect that this
latter specimen mentioned amongst the
Proceedings donations in 1857 is the holotype
of P. allenorum, and not any other specimen of
Pteropus currently in the collections (or listed
in the catalogs) at ANSP.
Fowler (1901) discussed and described
ichthyological collections from Samoa like-
wise donated to ANSP by Ruschenberger and
Caldwell, of which he noted: ??The following
specimens were collected many years ago by
Dr. H.C. Caldwell, by whom they were
presented to the Academy.?? Fowler also
described the new taxon Mugil caldwelli,
noting that he ??named this species for Dr.
Caldwell, who collected the type.?? We have
been unable to discover if Ruschenberger
actually took part in the collecting efforts in
Samoa in 1856, or if he was simply a financial
sponsor of these exploratory efforts. It is clear,
however, that Caldwell was directly responsi-
ble for the collection of some zoological
specimens during this voyage to Samoa. We
suspect based on the evidence at hand that he
(or his assistants and colleagues during his
visit to Apia) was the collector of the holotype
of Pteropus allenorum.
The holotype is the only specimen of
Pteropus allenorum known to us.
DIAGNOSIS: Pteropus allenorum is a rela-
tively small to medium-sized (figs. 4?8) mem-
ber of the genus Pteropus (forearm 116 mm in
the young holotype), probably with a brown
head, tinged with russet; a golden-brown
mantle, dusky brown back, and warm brown
limbs and wing membranes; very small cheek-
teeth (with an upper cheektooth size gradient
such that P3 . P4 , M1, according to overall
bulk), but proportionally large canines and
incisors; a moderately elongate rostrum; and a
relatively gracile mandible.
DISTRIBUTION: Pteropus allenorum is re-
corded historically only from the Samoan
island of Upolu. We speculate that, like most
native elements in the Samoan avifauna
(Steadman, 2006b), the actual historic or
prehistoric distribution of this species was
not limited solely to this single island, but
probably extended to Savai9i and to other
islands of Samoa, if not to adjacent archipel-
agos (even if Upolu truly was its last place of
occurrence). Further excavations of subfossil
material in Samoa and further study of
subfossil material from the adjacent archipel-
agos of Tonga and Fiji may help to clarify the
past distribution of this species. It is not yet
reported from the subfossil record of Tonga,
the only Polynesian archipelago where the
chiropteran subfossil record has been studied
in some detail (Koopman and Steadman,
1995), although in light of the elucidation of
this species in the historical fauna of Samoa,
closer study of Pteropus osteological material
from Tongan excavations is certainly warrant-
ed. Although likely extinct, we suggest that P.
allenorum should be sought after during future
biotic inventory efforts in Samoa on the
chance that an overlooked extant population
survives somewhere in the archipelago (see
Discussion, below).
ETYMOLOGY: We have chosen the specific
epithet allenorum to honor the name of Allen,
in the plural. The epithet simultaneously
acknowledges Harrison Allen (1841?1897), a
zoologist, anthropologist, and physician
(Hrdlicka, 1914), who assembled much of the
ANSP chiropteran collection in the late 19th
century, and Allen Drew, who kindly hosted
the Helgens during a visit to Philadelphia in
2006, during which the holotype of allenorum
was first examined. We suggest ??Small Samoan
Flying Fox?? as an appropriate common name.
DESCRIPTION: As noted above, the only
available specimen of Pteropus allenorum is
represented by a skin stored in alcohol with an
accompanying skull that, although broken,
preserves most cranial features. The skin is
fragmentary and fragile but includes the head
skin (separated from the rest of the body),
most of the dorsal skin of the body, the limbs
and wing membranes (somewhat decayed and
partly discolored), and some other small
clumps of fur. Because it has been preserved
in alcohol for more than 150 years and its
overall state of preservation is poor, it is
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 7
difficult to characterize the original external
appearance of this specimen. Based on the
single specimen available, we think that the
best that can be stated is that the holotype
probably had a brown head tinged with russet,
a golden-brown mantle, a dusky brown back,
and warm brown limbs and wing membranes.
We suggest that the general appearance of P.
allenorum was probably that of a rather
??furry?? flying fox, similar in pelage quality
to P. samoensis (see photograph in Flannery,
1995), in which the fur is longish and not
strongly adpressed, as opposed to many flying
foxes (such as P. tonganus), in which the fur
tends to be shorter, sleeker, and often clearly
adpressed dorsally. The lengths of the hairs in
the fur on the mid-back reach to about 25 mm.
The fur is sparser and paler on the front of the
face, and there is no darkened eye-ring
encircling the eye. The forearm in the holotype
measures 116 mm, and we expect that fully
grown adults would have a forearm length of
ca. 116?125 mm, smaller than either extant
Samoan congener, Pteropus samoensis and P.
tonganus (see table 1).
The holotype cranium is broken behind the
orbits, but most of it is preserved in two intact
pieces, which we carefully reconstructed to
prepare an image of the overall skull (fig. 7).
Compared to other Pteropus, the rostrum is of
??moderate length?? sensu Andersen (1912).
Despite its youth, the postorbital processes
are rather well developed. The back of the
palate forms a broad ??U?? shape. We estimate
the condylobasal length of the (nearly mature
but broken) holotype skull of allenorum to be
50 mm and the zygomatic width to be ca. 26
mm; measurements of full-grown adults would
Fig. 4. Bivariate ecomorphological contrasts in Samoan Pteropus. A plot of rostral (orbitonasal) length
versus zygomatic width discriminates the four Pteropus species recorded from Samoa. Sample represents all
adult specimens (or nearly adult in the case of the unique holotype of P. allenorum) from the Samoan
archipelago. Open triangles indicate P. samoensis; closed diamonds, P. tonganus; closed square, P. allenorum;
open circles, P. coxi.
8 AMERICAN MUSEUM NOVITATES NO. 3646
thus somewhat exceed these values, particu-
larly in zygomatic width.
A striking feature of P. allenorum is the
small size of the teeth relative to the size of the
skull, even when perfectly unworn, as in the
holotype. As indicated above, the very small
skull and teeth of P. allenorum allow for its
instant discrimination against the sympatric
large-toothed forms P. tonganus and P.
samoensis (figs. 4?8; table 1). The upper den-
tition is largely complete in the holotype (right
C1 is loose from the jaw but preserved in the
box; right P1 is missing, represented only by
an empty alveolus). The upper incisors are
proportionally very broad (fig. 7). The canines
are long and narrow, with a moderately
developed posterior cingulum. P1 is present.
The cheekteeth posterior to P1 are relatively
very small and narrow. The length of the
maxillary toothrow (C1?M2) measures 18.0
mm, markedly smaller than in Samoan con-
geners (table 1; fig. 5). The soft palate is not
preserved.
The holotype mandible is complete and
gracile in overall appearance. It has compar-
atively weak posterior processes (fig. 7), re-
sembling P. fundatus of Vanuatu (Felten and
Kock, 1972), yet it is more gracile in the
reduced size of the coronoid and angular
processes despite its slightly larger overall size
compared to that species. The incisors and
canines and most of the right premolars and
molars have been dislodged from the mandible
but are preserved separately in the skull box
(we reinserted these to provide the dorsal view
of the mandible and its dentition, fig. 7). The
lower canines and incisors are much larger
(both in absolute and relative terms) than the
corresponding teeth in P. fundatus. The
posterior premolars and molars are small
and rather narrow, but p1 is especially well
developed.
Fig. 5. Bivariate ecomorphological contrasts in Samoan Pteropus (continued). A plot of maxillary
toothrow length versus the distance across the upper canines also discriminates the four Pteropus species
recorded from Samoa. Sample and symbols as for figure 4.
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 9
Overall, the skull of P. allenorum requires
closest comparison with P. fundatus, a flying
fox of similar cranial and dental size, also
endemic to a remote Pacific archipelago
(Vanuatu). The premolars and molars of P.
allenorum match those of P. fundatus closely in
both shape and absolute dimensions (table 2).
More striking dental contrasts between P.
allenorum and P. fundatus lie in the compar-
ative size of the anterior dentition. In P.
allenorum the incisors are much broadened
and the canines vertically and anteroposteri-
orly relatively more elongate than in P.
fundatus (fig. 7). We suggest that this juxta-
position of relatively less massive cheekteeth
but more massive incisors and canines in P.
allenorum relative to P. fundatus probably
reflects salient differences in feeding mode and
ecology between these two ecomorphologi-
cally distinctive taxa. P. allenorum also can be
distinguished immediately from P. fundatus by
its darker coloration (P. fundatus is a pale
flying fox; fig. 9) and its larger body and skull
size (the forearm measures 95?102 mm [n 5
21] and condylobasal length measures 44?49
mm [n 5 17] in adult P. fundatus; Felten and
Kock, 1972; Flannery, 1995; compare to P.
allenorum in table 1). Given their morpholog-
ical similarity and geographic proximity in the
remote Pacific, P. allenorum and P. fundatus
may be close relatives within the genus
Pteropus. However, we stress that similarities
between them may not necessarily reflect a
close phylogenetic relationship, but perhaps
instead a shared pattern of ecomorphological
convergence on isolated Pacific archipelagos.
A study drawing on molecular sequence data
from a wide taxonomic and geographic sample
Fig. 6. Maxillary toothrows of the four Pteropus species of Samoa, to scale. A, Pteropus allenorum
(ANSP 1234, unsexed subadult); B, Pteropus samoensis (ANSP 1867, unsexed adult); C, Pteropus coxi
(USNM 3791, adult, probably male); D, Pteropus tonganus (AMNH 68738, adult male). Scale bar 5 5 mm.
10 AMERICAN MUSEUM NOVITATES NO. 3646
of Pteropus is needed to distinguish between
these possible scenarios (see Giannini et al.,
2008).
The dentition of Pteropus allenorum is
considerably less reduced than that of
Pteropus scapulatus (of Australia and southern
New Guinea) and the presumed phylogenetic
allies of that species (P. woodfordi and P.
mahaganus of the Solomon Islands and P.
gilliardorum of the Bismarck Archipelago),
which have relatively small and nearly fea-
tureless cheekteeth and together are thought
to constitute a distinctive Australian and
Pacific group of specialist nectar-feeding
flying foxes (Thomas, 1888; Andersen, 1912;
Sanborn, 1931; Van Deusen, 1969; Flannery,
1995; Bonaccorso, 1998; Helgen, 2004a; al-
though see Giannini et al., 2008). However,
Pteropus allenorum shares with these species a
similarly gracile mandible, relatively large
canines, small cheekteeth, and similar body
size (e.g., fig. 7), attributes suggestive of
similarities in lifestyle, although likely conver-
gently derived.
Like P. allenorum, Pteropus vetulus Jouan,
1863 (a New Caledonian endemic), is a small
Pacific Pteropus with very small cheekteeth,
broad upper and lower incisors, and a
conspicuously large p1. Like P. allenorum, it
is also a rather darkly colored Pteropus with
Fig. 7. Skulls of smaller-toothed, medium-sized Pteropus species of the southwest Pacific region. A,
Partially cleaned skull of Pteropus fundatus, endemic to Vanuatu (AM M26897, adult male, Mota); B,
reconstructed skull of Pteropus allenorum, endemic Samoa (ANSP 1234, unsexed subadult, Upolu); C, skull
of Pteropus mahaganus, endemic to the Solomon Archipelago (AM M6280, adult male, Bougainville). Scale
bar 5 10 mm.
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 11
thick and furry rather than adpressed dorsal
pelage (see photograph published by Flan-
nery, 1995: 299). Pteropus allenorum is distin-
guished from P. vetulus by its larger skull and
body size (the forearm measures 92?112 mm
in adult P. vetulus); dorsoventrally longer
canines (short and deep in P. vetulus); simpler
upper first molar (labial edge subdivided into
two distinct cusps in P. vetulus); proportion-
ally more elongate and evenly sloping rostrum
(blunter, with more precipitous slope down-
ward from the braincase in lateral profile in P.
vetulus); upper cheektooth size (bulk) gradient
such that P3 . P4 , M1 (P3 , P4 . M1 in P.
vetulus); less pronounced occipital cresting
(strongly marked, with a subrectangular pos-
terior braincase conformation in P. vetulus);
and more gracile mandible with a less devel-
oped angular process and lower coronoid
process (slightly more robust with broader
and higher respective processes in P. vetulus).
Selected wing and leg measurements in the
holotype of P. allenorum are as follows (in
mm, measured from wet skin): pollex (with
claw) 48.5, (without claw) 42.6; D2M 58.7;
D2P1 17.5; D2P2 12.5; D3M 78.4; D3P1 57.3;
D3P2 67.1; D4M 77.9; D4P1 50.6; D4P2 45.5;
D5M 85.0; D5P1 37.7; D5P2 38; tibia ca. 52;
hindfoot (with claws) 30, (without claws) 28.
NATURAL HISTORY: That Pteropus allenor-
um has such a markedly reduced dentition
relative to its extant and extinct Samoan
congeners undoubtedly reflects differences in
its overall diet relative to those species. It may
Fig. 8. Multivariate morphometric comparisons in Samoan Pteropus (principal components analysis),
drawing from 10 log-transformed craniodental measurements, divide the four Pteropus species recorded
from Samoa into four discrete quadrant clusters. Sample and symbols as for figure 4. In this case, overall
size can be visualized on the first component (increasing from right to left), while the loadings on the second
component serve as an indication of general ??robustness?? (increasing from top to bottom). See text and
table 3. Parenthetical numbers on the axes indicate the proportion of variance for each principal component
(see table 3).
12 AMERICAN MUSEUM NOVITATES NO. 3646
TABLE 2
Measurements of Premolars and Molars in Pteropus allenorum and P. fundatus, Two Relatively Small
Pacific Pteropus
L indicates length; W, width
P. allenorum P. fundatus
ANSP AM AM AM
1234 M26898 M26897 M26896
? male male female
P3 L 3.4 3.3 3.5 3.2
P3 W 2.1 2.1 2.1 1.9
P4 L 3.2 3.4 3.3 3.3
P4 W 2.1 2.3 2.5 2.1
M1 L 4.2 4.2 4.3 4.3
M1 W 1.9 2.2 2.2 2.0
M2 L 1.9 1.6 1.9 1.6
M2 W 1.3 1.2 1.4 1.3
p1 L 1.9 1.5 1.8 1.7
p1 W 1.6 1.6 1.7 1.5
p3 L 3.4 3.3 3.5 3.1
p3 W 1.7 1.9 1.8 1.7
p4 L 3.5 3.3 3.4 3.3
p4 W 1.9 2.0 1.9 1.7
m1 L 3.6 3.5 3.4 3.6
m1 W 1.9 2.0 1.8 1.9
m2 L 2.7 2.6 2.9 2.6
m2 W 1.8 1.7 1.8 1.7
m3 L 1.4 1.5 1.7 1.4
m3 W 1.2 1.1 1.3 1.0
TABLE 3
Principal Components Analysis Comparing 36 Pteropus Skulls from Samoa
Factor loadings, eigenvalues, and percentage of variance for the first three principal components in a
principal components analysis are included. Principal components are extracted from a covariance matrix of
10 log-transformed cranial and dental variables; see figure 8, where the first two components are plotted.
PC1 PC2 PC3
ONL 20.7125 0.6602 0.0844
ZYG 20.8246 20.2427 0.0664
MTR 20.9207 0.2991 0.0855
CC 20.7654 20.4451 0.1307
M2M2 20.7184 0.3431 0.1674
ML 20.9378 0.2100 0.1338
MH 20.5859 20.7038 0.3107
LTR 20.9449 0.1669 0.1222
M1L 20.4139 20.8000 20.2295
M1W 20.8795 20.0120 20.4397
Eigenvalues 0.0298 0.0123 0.0031
% variance 58.9999 24.4082 6.0630
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 13
have specialized as a nectar-feeder or by eating
inflorescences or smaller fruits and nuts than
those typically utilized by these larger bats, as
the smaller-toothed species P. mahaganus and
P. fundatus (fig. 7) do in the Solomon archi-
pelago and Vanuatu, respectively (Bonaccor-
so, 1998; Flannery, 1995). However, no notes
about ecological attributes or context of
collection are associated with the holotype of
Pteropus allenorum, such that nothing is firmly
recorded of its basic biology.
The morphological resemblance between
P. allenorum and P. fundatus of northern
Vanuatu (fig. 7) may indicate that similar
small-bodied flying foxes were formerly more
widespread throughout the Pacific theatre. If
so, their discovery in the modern faunas or
subfossil records of Fiji, Tonga, and on other
islands of the Vanuatu and Samoan archipel-
agos might be expected in the future. For
example, the current geographic restriction of
P. fundatus to the small outlying Banks and
Torres island groups of Vanuatu (Flannery,
1995) seems likely to be a relictual distribution
indicative of ??pseudoendemism?? in that ar-
chipelago (Steadman, 1997, 2006a). In this
vein, we note with interest that Hickey (2007)
discussed a small, dark-colored, inflorescence-
eating Pteropus observed on the large island of
Malekula in Vanuatu. Hickey (2007) specu-
lated that if not a juvenile P. tonganus, this
might be a small flying fox related to P.
fundatus. The chiropteran faunas of Vanuatu
and Fiji undoubtedly remain incompletely
inventoried, so future discoveries are certainly
to be expected (Helgen and Flannery, 2002;
Helgen, 2004c, 2005).
Pteropus coxi, new species
HOLOTYPE: The holotype of Pteropus coxi
is USNM 3791, skull of an adult, probably
male, from the ??Samoan Is.??, collected during
the U.S. Exploring Expedition of 1838?1842.
The skin of this specimen bore the number
3953 but is now apparently lost (L.K. Gordon,
in litt.).
PARATYPE: USNM 3790, skull of an adult,
probably female, from ??Samoan Is.??, also col-
lected during the U.S. Exploring Expedition.
The skin of this specimen bore the number
Fig. 9. Skins of the endemic Pteropus of Vanuatu. A, P. fundatus, AM M26896, adult female, Mota. B,
P. anetianus, USNM 278062, adult female, Espiritu Santo.
14 AMERICAN MUSEUM NOVITATES NO. 3646
3952 but, like the holotype skin, is now
apparently lost (L.K. Gordon, in litt.).
DIAGNOSIS: Pteropus coxi is a large flying
fox similar in cranial conformation to P.
samoensis and P. anetianus?with a robust
skull featuring a rostrum of moderate length,
a mandible with heavily developed posterior
processes, and large molars?but considerably
larger than P. samoensis and P. anetianus in
cranial size, with larger canines and cheekteeth
(figs. 4?6, 8, 10, 14). Apart from P. tubercu-
latus of the Santa Cruz Islands (an outlying
island group between the Solomon archipela-
go and Vanuatu), P. coxi is the only species in
the genus in which the very large upper
canines sometimes (in the case of P. coxi, in
one of the two available specimens) bear a
secondary, posterior cusp.
DISTRIBUTION: The U.S. Exploring Expe-
dition visited the Samoan archipelago from 7
October to 10 November in 1839, exploring
the islands of Tutuila, Upolu, Savai9i, and the
Manua group; some members of the expedi-
tion also returned to Upolu for visits in
following years (one ship, the Porpoise re-
turned 4-8 September 1840 to Upolu; two
ships, the Peacock and Flying Fish, returned 6
February to 6 March 1841; M. Grunes, in
litt.). Pteropus coxi is known only by the
holotype and paratype, which were collected
at some time during the U.S. Exploring
Expedition, but localized only generally to
the ??Samoan Is.[lands]?? or ??Samoan Isle?? on
their accompanying labels. We regard P. coxi
as a Samoan endemic, although the precise
insular extent of its historical distribution
Fig. 10. Skulls of the four larger Pteropus species of Vanuatu and Polynesia. A, P. tonganus (AMNH
68738, adult male, ??Samoa??); B, P. samoensis (USNM 4465, unsexed adult, ??Samoan Archipelago??); C, P.
coxi (USNM 3953/3791, probably male, ??Samoan Archipelago??); D, P. anetianus (USNM 278062, adult
female, Espiritu Santo, Vanuatu). Scale bar 5 10 mm.
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 15
remains unknown. Future subfossil excava-
tions in Samoa may clarify its prehistoric or
historical distribution.
ETYMOLOGY: The specific epithet honors
Dr. Paul A. Cox of the Institute for
Ethnomedicine in Jackson Hole, Wyoming,
in recognition of his research on flying foxes,
tropical ecology, and biodiversity conserva-
tion in Pacific archipelagos in general and in
Samoa in particular. We suggest ??Large
Samoan Flying Fox?? as an appropriate
common name for this species.
DESCRIPTION: Nothing is known with cer-
tainty about the external appearance of P.
coxi, because the skins originally associated
with both skulls have been lost and cannot be
traced (L.K. Gordon, in litt.). However, a few
clues are available that may help to envision
its appearance. First, the skull of P. coxi is
larger and more robust than its sympatric
congeners on Samoa, an indication that it was
probably a heavier bat than P. samoensis and
P. tonganus, with a somewhat longer forearm.
Another clue to its appearance may be found
in the original description of Pteropus samoen-
sis by Peale (1848). The U.S. Exploring
Expedition collected P. samoensis, P. tonga-
nus, and P. coxi in Samoa in 1839-1841, but
Peale (1848) implicitly referred all of these to
P. samoensis in his original description of P.
samoensis, noting:
The species was first discovered on the island
of Tutuila, and subsequently in all the
islands of the Samoan Group; we obtained
numerous specimens, and found the young
animals somewhat-lighter coloured than the
old ones, but in other respects there is but
little variation in colour or size.
In preparing his description of P. samoensis,
we assume from his comments that Peale had
all or most Samoan Pteropus specimens from
the U.S. Exploring Expedition at hand (of
these, those that are specifically localized
indicate their collection on Tutuila, Upolu,
and Olusinga [5 Olosega] Islands; table 4).
The skins of both specimens of P. coxi (in
addition to the still-available skulls) were
apparently still extant in 1848 and were
apparently referred to P. samoensis. As such,
we suggest that these specimens can be
assumed to have formed part of the original
hypodigm of Pteropus samoensis (as can
TABLE 4
Specimens of Pteropus from Samoa at USNM Collected during the U.S. Exploring Expedition (all of which
apparently comprise the original hypodigm or syntype series of Pteropus samoensis)
Pteropus samoensis Peale, 1858
USNM 3788/3949 ??Samoan Archipelago??, adult, skin (3949) and skull (3788)
USNM 3789 ??Samoan Arch.??, adult female, skull, accompanying skin
(USNM 3950) missing
USNM 3947 ??Samoan Archip.??, adult, skin
USNM 4465 ??Samoan Ids??, adult, skull
USNM 22562 Tutuila Island, adult, skull, accompanying skin (8594) missing
USNM 37860/8597 Tutuila Island, adult, skin (8597) and skull (37860). Lectotype
of Pteropus samoensis Peale, 1858,designated herein (see
figs. 12, 13)
USNM 37861/8593 Olusinga (Olosega) Island, adult male, skin (8593) and skull
(37861)
USNM 37862/8596 Tutuila Island, adult female, skin (8596) and skull (37862)
USNM 37878/8595 Upolu Island, adult, skin (8595) and skull (37878)
Pteropus coxi, n. sp.
USNM 3790 ??Samoan Isle??, skull, accompanying skin (3952) missing
USNM 3791 ??Samoan Isle??, skull, accompanying skin (3953) missing
Pteropus tonganus Quoy and Gaimard, 1830
USNM 3954 ??Samoan Island??, adult male, skin with skull in situ
USNM 3955 ??Samoan Isle??, adult, skin without skull
USNM 38681/3956 ??Samoan Isle??, subadult, skin (38681) and skull (3956)
16 AMERICAN MUSEUM NOVITATES NO. 3646
several specimens of P. tonganus collected in
Samoa during the expedition; see below,
where we designate a lectotype for P. samoen-
sis to preserve its traditional taxonomic
association). Peale?s (1848: 20?21) statement
about variation (reiterated in the observations
of Cassin [1858] a decade later) might allow us
to infer that the skins of P. coxi were not
especially different in overall pelage patterning
and form from his series of true P. samoensis,
which he characterized as ??head ? tawny, the
front gray; ears small, rounded, black; neck of
the old animals rufus [sic], in the younger
animals tawny, body and throat reddish-
brown; hair erect, and somewhat woolly, most
smooth on the back; wing-membranes black;
irides brown.?? Although P. coxi is undoubt-
edly a larger bat than P. samoensis, Peale?s
(1848) statement about the lack of noticeable
size variation in his series also suggests that
the skins of P. coxi were probably not
markedly larger in apparent size as compared
to true P. samoensis (and P. tonganus).
The skull of P. coxi is larger and more
robustly constructed than that of P. samoen-
sis, with a broader rostrum and palate; a
longer maxillary toothrow featuring more
massive individual teeth; broader and thicker
zygomata and postorbital processes; and a
dentary with a deeper ramus, more massive
coronoid, condylar, and angular processes,
and more heavily sculpted surfaces for mus-
cular attachments. The canines are more
massive in general (vertically taller, broader,
more anteroposteriorly elongate), and the
upper canines bear more prominent internal
ridging. In the holotype, the right upper
canine bears a groove separating a small
secondary cusp from the main body of the
canine, and a similar, incipient secondary cusp
can be seen on the left upper canine. The
paratype, a somewhat younger animal judging
from toothwear, also with massive canines,
does not have these same secondary cusps.
This trait may be individual in its variability,
but it could also be a sexual difference. Males
and females differ in canine size and robusti-
city in many species of pteropodids, including
many if not most species of Pteropus
(Andersen, 1912: 75: ??canines in males of
nearly all species longer and heavier than in
females??). Individual premolars and molars
are both wider and longer in P. coxi than in P.
samoensis (e.g., table 1). P1 (upper) is very
small but persists in the adult dentition. The
first lower premolar (p1) is large. In general
cranial robustness, particularly in the confor-
mation of the mandible, P. coxi resembles P.
anetianus of Vanuatu, with which it is most
closely allied in our morphometric compari-
sons (e.g., fig. 14), but the two species differ
both in cranial and dental size (greater in P.
coxi) as well as in the proportional length of
the rostrum (longer in P. coxi). In its extreme
cranial robustness, large teeth, and incipient
(if variable) secondary cusps in the upper
canines, P. coxi might be regarded as ecomor-
phologically convergent to some extent on the
cranially robust and large-toothed Pacific
??monkey-faced bats?? of the Solomon archi-
pelago (genus Pteralopex) and Fiji (genus
Mirimiri), which occur in the broader region
but not in Samoa, and to which flying foxes
are not immediately related (Hill and Beckon,
1978; Flannery, 1995; Parnaby, 2002b;
Helgen, 2005). (These large bats are thought
to feed on nuts, hard, thick-skinned fruits, and
perhaps tree exudates; Flannery, 1995; Fisher
and Tasker, 1997; Helgen, 2005.)
NATURAL HISTORY: Nothing is known of
the habits or biology of P. coxi, the largest of
Polynesian flying foxes, and we assume that
the species is now extinct, perhaps for longer
than a century, as the only known specimens
were collected in 1839?1841.
However, published observations from the
early 1980s by the botanist Paul Cox, discuss-
ing a flying fox of unusual size in Samoa,
could indicate that P. coxi survived until
recently. In his paper on the bats of Samoa,
Cox (1983) noted an encounter with ??Pteropus
samoensis??:
I will never forget the first time I saw one of
these giant bats in the rainforest. One day,
while climbing a tree, I saw what appeared to
be an eagle flying away from a liana flower.
The bat I saw in my field glasses appeared to
have a wingspan of five feet or more and
lacked the white fur on the back of the neck
that characterizes the locally common flying
fox, P. tonganus. This large bat was black and
its behavior was completely unusual. I later
thoroughly enjoyed watching them soar,
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 17
Fig. 11. Color variation in the extant Pteropus spp. of Samoa. A, P. tonganus (USNM 566603, adult
female, ??Samoa??), dorsal coloration pattern; B, P. samoensis (USNM 338624, adult male, Tutuila), dark
phase (most common); C, P. samoensis (USNM 565827, adult female, Tutuila), white phase (very rare); D, P.
samoensis (USNM 3947, unsexed young adult, ??Samoan Archipelago??), pale, straw-colored phase (less
common in Samoa, more common in Fiji).
18 AMERICAN MUSEUM NOVITATES NO. 3646
eagle-like, high above the forest in midday
sun.
In this paper, Cox (1983) concluded that P.
samoensis is perhaps the largest species of
Pteropus anywhere?a strange claim, consid-
ering that P. samoensis is a medium-sized
member of the genus, and even sympatric
Pteropus tonganus is slightly heavier (despite
equivalent forearm lengths in Samoa [table 1],
body mass averages 383 g in three adult
Samoan P. samoensis and 410 g in nine adult
Samoan P. tonganus at USNM). After an
attempt to collect verifying voucher speci-
mens, and further reflection, Cox (1984a)
noted:
Two specimens [later] shot by hunters were
confirmed as P[teropus] samoensis by Dr. Karl
Koopman at the American Museum of
Natural History. These had wingspans of
only 3.5 feet, the size considered typical for
the species. The larger bats that I observed
appeared to behave like normal P. samoensis,
however their size raises the possibility that
my observations include a second, but
undescribed endangered species. These find-
ings emphasize the paucity of available
information and the urgency of a thorough
investigation.
Understandably, most subsequent authors
have attributed Cox?s observations of giant
black bats to somewhat exaggerated or
fantastical descriptions of Pteropus samoensis
(e.g., Wilson and Engbring, 1992), a conclu-
sion that Cox apparently later accepted (Cox,
1999). However, we raise the possibility
(however slight) that Cox instead encountered
the species that we describe here as P. coxi, a
very large species of Samoan flying fox that is
probably closely related to P. samoensis, with
which it must occur (or have occurred)
sympatrically. If Cox truly observed P. coxi
in the forests of Upolu in 1981, then he has
provided the only known description of its
external features?i.e., that it is ??black??, with
a wingspan of ca. ??5 feet?? (5 1.5 m) (Cox,
1983; later revised to ??4 feet?? [5 1.2 m] by
Cox, 1999). Certainly neither P. samoensis nor
P. tonganus achieves such a large body size in
Samoa (Wilson and Engbring, 1992). That
Fig. 12. Skull of USNM 8597/37860, lectotype of
Pteropus samoensis Peale, 1848. Scale bar 5 10 mm.
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 19
Cox?s clearest observations of this large bat
took place in upland forests during the day
(Cox, 1983, 1984a), when P. samoensis also
flies (e.g., Andersen, 1912; Wilson and
Engbring, 1992; Thomson et al., 1998, 2002),
provides the only ecological information that
might conceivably refer to P. coxi.
Cox?s (1984a) prescient call regarding the
??urgency of a thorough investigation?? into the
possibility that more than two extant species
of Pteropus persist in Samoan forests has gone
unheralded. This urgency has increased with
the passage of time and with the discoveries
reported herein. To us, Cox?s account raises
the slight possibility that populations of P.
coxi could persist in Samoa, especially in
remote montane habitats on the high forested
islands of Upolu and Savai9i. This possibility
should be pursued in further biological explo-
rations in the archipelago and in interviews
with local communities throughout Samoa.
Should P. coxi persist in Samoa as a rare
species, some field studies that have attempted
to document the biology of ??Pteropus samoen-
sis?? in recent years (e.g., Cox, 1983; Wilson
and Engbring, 1992; Banack, 1996, 1998;
Brooke, 2001) could conceivably be based on
studies of more than one biological species
(i.e., P. samoensis and P. coxi), even after
distinctions between P. samoensis and P.
tonganus, sometimes confused in the past,
were made clear to Samoan fieldworkers
(Wilson and Engbring, 1992). Although it is
tempting to associate this newly elucidated
taxon with Cox?s observations in the early
1980s, we suggest that these are best viewed
Fig. 13. Skin of USNM 8597/37860, lectotype of Pteropus samoensis Peale, 1848. A, dorsal view. B,
ventral view.
20 AMERICAN MUSEUM NOVITATES NO. 3646
with skepticism in the absence of any addi-
tional evidence for the recent survival of P.
coxi. A dark brown color phase is the most
common coloration pattern in Samoan popu-
lations of Pteropus samoensis (fig. 11), and the
wingspan of flying foxes is difficult to estimate
from any distance. As such, Cox?s (1983, 1984)
observation of very large, blackish bats in
flight during the day could easily be based on
sightings of P. samoensis. Furthermore, while
20th-century specimens of P. samoensis and P.
tonganus from Samoa are not uncommon in
museums (e.g., AM, AMNH, ANSP, USNM,
ZMUC), we know of no museum specimens of
P. coxi collected since the type series was
taken more than 160 years ago. Regardless of
whether more than two species of Pteropus
survive today in Samoa, it is clear to us from
our own experiences and from our close
reading of relevant literature that it is indeed
only P. samoensis and P. tonganus that have
been regularly encountered by field biologists
working in Samoa during the past 15 years
(e.g., Wilson and Engbring, 1992; Craig and
Syron, 1992; Elmqvist et al., 1992, 1994; Craig
et al., 1994a, 1994b; Morrell and Craig, 1995;
Grant and Banack, 1995, 1999; Banack, 1996,
1998; Pierson et al., 1996; Grant et al., 1997;
Miller and Wilson, 1997; Richmond et al.,
1998; Thomson et al., 1998, 2002; Brooke et
al., 2000; Nelson et al., 2000; Webb et al.,
2000; Brooke, 2001; Banack and Grant, 2002,
2003).
A LECTOTYPE FOR PTEROPUS
SAMOENSIS PEALE, 1848
As discussed above, Peale (1848) did not
designate a type specimen for Pteropus sa-
moensis in the original description of that
Fig. 14. Bivariate plot of the two canonical variates in a discriminant function analysis contrasting
samples identified as Pteropus samoensis (Fiji and Samoa, triangles), Pteropus anetianus (Vanuatu, closed
circles), and Pteropus coxi (Samoa, open circles). Parenthetical numbers on the axes indicate the proportion
of variance for each variate (see table 5).
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 21
species (and did not discriminate specimens of
P. tonganus and P. coxi from the remainder of
the Samoan Pteropus specimens referred to P.
samoensis). Accordingly, all Samoan Pteropus
collected during the U.S. Exploring Expedi-
tion must be regarded as syntypes of P.
samoensis, including those specimens included
here in the type series of P. coxi. Lyon and
Osgood (1909: 258) (followed by Poole and
Schantz, 1942: 142) regarded the series of
specimens of P. samoensis from Tutuila as
??typical??, based on Peale?s statement that
??this species was first discovered on Tutuila,
and subsequently on all the islands of the
Samoan group; we obtained numerous speci-
mens.?? This act by Lyon and Osgood formally
restricted the type locality of P. samoensis to
Tutuila, but no lectotype was designated.
The syntype series of P. samoensis is
composite (table 4), comprising specimens of
three biological species (nine specimens of the
taxon traditionally identified as P. samoensis,
three specimens of P. tonganus, and two
specimens of P. coxi; table 4). Selection of a
lectotype for P. samoensis is required to
preserve what can be regarded as the tradi-
tional association of the name, as established
most authoritatively by Andersen?s (1912)
description and overview of P. samoensis.
(Unfortunately, in preparing his account,
Andersen did not have access to the entire
syntype series collected by the U.S. Exploring
Expedition, and in particular he did not
examine the type series of P. coxi.)
Included within the syntype series of P.
samoensis at USNM are three specimens
marked as coming from Tutuila, which has
been regarded as the type locality following
Lyon and Osgood (1909) and Poole and
Schantz (1942). Each of these three specimens
represents the biological species corresponding
to Andersen?s understanding and usage of the
name Pteropus samoensis Peale, 1848. The first
of the three specimens listed by Lyon and
Osgood (1909: 142), an unsexed adult, USNM
8594/22562 (the latter number misquoted as
25562 by Lyon and Osgood), was at some
point ??turned over to the Dep[artment of]
Comp[arative] Anat[omy]??, and it now con-
sists only of a skull with the basicranial region
broken. The second, an adult female, USNM
8596/37862, consists of a study skin and
accompanying skull with the back of the
braincase sawn off and the left upper molar
row smashed. The third, USNM 8597/37860,
an unsexed adult, comprises a study skin and
skull, both in reasonably good condition. It is
this last specimen, figured here (figs. 12, 13),
that we choose as the lectotype of Pteropus
samoensis Peale, 1848.
THE SYNONYMY OF PTEROPUS
SAMOENSIS PEALE, 1848
In the context of designating a lectotype for
P. samoensis, it is profitable to briefly review
the nomenclatural synonymy of the species. So
far as we can ascertain, the first name ever
applied to a flying fox from Samoa appears
in the narrative account of the U.S. Explor-
ing Expedition published by Wilkes (1844).
Wilkes (1844: 128) wrote, ??there are no traces
among these islands of any native quadruped,
nor any other of the mammalia, except a
species of bat (Pteropus ruficollis), which is
very destructive to the bread-fruit.?? This name
(ruficollis), introduced without description,
must be regarded as a nomen nudum, and
can be taken to apply to all flying foxes that
Wilkes and his crew may have seen in Samoa
(at least four species at the time of their
voyage) or collected in Samoa (at least three
TABLE 5
Underlying Statistics for Discriminant
Function Analysis
Correlations, canonical correlations, eigenvalue,
cumulative proportion of variance contrast eight
log-transformed measurements for adult skulls
identified as Pteropus samoensis (25 skulls), P.
anetianus (15 skulls), and P. coxi (2 skulls) (see
fig. 14, where specimen scores are plotted)
CV1 CV2
ONL 0.2255 20.6424
ZYG 20.5093 20.5915
MTR 20.1682 20.7728
CC 20.1113 20.5118
M2M2 20.0032 20.1207
ML 20.2695 20.7578
MH 20.6815 20.2677
LTR 20.2631 20.7630
Canonical correlation 0.7628 0.5841
Eigenvalue 1.3915 0.5178
Cumulative proportion of variance 0.7288 1.0000
22 AMERICAN MUSEUM NOVITATES NO. 3646
species; table 4). Wilkes? nomen nudum is
probably a lapsus for Pteropus rubricollis E.
Geoffroy, 1810 (a synonym of P. subniger
(Kerr, 1792), an extinct flying fox endemic to
the Mascarenes), a taxonomic epithet widely
applied to flying foxes in the early to middle
19th century (see Andersen, 1912). Andersen
(1912) and subsequent systematic workers
(e.g., Wilson and Engbring, 1992; Simmons,
2005) overlooked Wilkes? usage of ruficollis
and attributed the original use of the name to
Nicoll (1908), also as a nomen nudum, in his
book Three Voyages of a Naturalist. Nicoll
(1908) also employed another lapsus, rufficol-
lis, in the index of his book, and yet another,
fuscicollis, in an earlier publication (Nicoll,
1904). It is clear to us that in each case Nicoll?s
names were used for Samoan flying foxes in
the same general sense as employed by Wilkes
(1844). Both of these additional nomina nuda
can be regarded simply as incorrect subse-
quent spellings of Wilkes? original nomen
nudum, ruficollis; all of these are partial
synonyms of Pteropus samoensis Peale, 1848.
A scientific name that deserves careful
consideration in light of the discoveries report-
ed in this paper is Pteropus whitmeei Alston,
1874. Alston (1874: 96) discussed a small
collection of bats received from the Rev. S.J.
Whitmee in Samoa, which consisted of four
specimens?one individual of Emballonura
semicaudata, two individuals of Pteropus ton-
ganus (reported at the time as ??P. flavicollis??),
and one specimen that Alston judged to be an
undescribed species, introduced under the new
name Pteropus whitmeei. The collector of the
holotype of whitmeei indicated that this latter
species was ??very common in Samoa?? and that
he ??once saw a number together ? estimated
at over a thousand?? in an inland crater on
Savai9i (Whitmee, 1874b). Alston?s description
of whitmeei is cursory and based on a single
specimen preserved intact (i.e., with skull in
situ) in alcohol (such that only color, external
measurements, and the comparative size of the
first upper premolar are discussed). This type
specimen was deposited in the personal collec-
tion of ??the Rev. Canon Tristam?? (Alston,
1874: 96), and we do not know its current
whereabouts. This specimen was not available
to Andersen (1912), who, while based in
London, attempted to examine all megachir-
opteran type specimens for his signal treatise on
pteropodid systematics, and this specimen is
apparently not in the BMNH today. Andersen
(1912) argued that the type of whitmeei must be
an incompletely grown specimen of P. samoen-
sis, and we agree. Alston noted that whitmeei
??is most nearly allied to P. vitiensis Gray
(5 Pteropus samoensis nawaiensis Gray, 1870,
the Fijian subspecies), of which it may probably
be regarded as the Samoan representative??,
and he distinguished it from typical P. samoen-
sis (which he knew only on the basis of Peale?s
original description) by its smaller size alone.
Alston?s description of the pelage and the
excellent accompanying color plate of the type
specimen match the coloration of many skins
of Pteropus samoensis samoensis at USNM and
elsewhere. Its forearm length (113 mm) falls
within the range of subadult P. samoensis. This
forearm length also compares favorably with
the nearly mature holotype of Pteropus allenor-
um (116 mm). It is not possible that whitmeei is
an earlier name for P. allenorum, however, as
Alston mentioned that the first upper premolar
in whitmeei is heavier than in P. s. nawaiensis,
itself a much larger-toothed bat than P.
allenorum. We suggest that Pteropus whitmeei
is unlikely to be an earlier name for P. coxi
because of the very small size of the holotype,
which instead matches subadult P. samoensis in
size and corresponds precisely in its coloration
to the most common color variant of P.
samoensis. As long as its holotype remains
unavailable to systematists, the name whitmeei
should rest within the synonymy of P. samoen-
sis, where Andersen (1912) and all subsequent
authors have placed it, and where we judge that
it is best arranged on the basis of information
provided in the original description (Alston,
1874). Whitmee?s (1874b) comments about the
abundance of ??P. whitmeei?? further suggest
that he was referring to one (or both) of the
more commonly collected bats of Samoa (i.e.,
Pteropus samoensis and/or P. tonganus), and
probably indicate his inability to distinguish
the two (or more) Pteropus species extant in the
archipelago at that time. Other flying fox
specimens sent to BMNH by Whitmee from
Samoa represent both P. samoensis and P.
tonganus (Andersen, 1912: 188, 287).
Apart from whitmeei and ruficollis and its
various lapsed nomenclatural manifestations,
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 23
the synonymy of Pteropus samoensis also
includes the names nawaiensis and vitiensis,
taxonomic names applied to Fijian popula-
tions of P. samoensis by Gray (1870). We
follow all recent authors in recognizing the
Fijian P. s. nawaiensis as a subspecies of P.
samoensis (e.g., Wodzicki and Felten, 1975;
Wilson and Engbring, 1992; Flannery, 1995;
Ingleby and Colgan, 2003; Simmons, 2005).
Pteropus s. nawaiensis differs from the nomi-
nate subspecies of Samoa in its smaller body
size (in average terms; see table 1; body weight
averages 353 g in seven adults from Fiji [AM]
versus 383 g in three adults from Samoa
[USNM]), but without concomitant differenc-
es in cranial size (table 1), and in its less
variable coloration?most Fijian animals are
dark brown with a paler, straw-colored
mantle, in contrast to the greater chromatic
variability evident in museum samples of P.
samoensis (fig. 11).
The synonymy of Pteropus samoensis (in-
cluding synonyms from Fiji) can thus be
delineated as follows:
Pteropus ruficollis Wilkes, 1844: 128. Type locality
??Samoa??. Part; nomen nudum.
Pteropus samoe?nsis Peale, 1848: 20. Type locality ??Tutuila
? and, all islands of the Samoan Group.?? Part; type
series including specimens referred here to Pteropus
tonganus Quoy and Gaimard, 1830, and Pteropus coxi,
n. sp. Type locality restricted to Tutuila (American
Samoa) by Lyon and Osgood (1909: 142). Clarification
of composite type series and lectotype designation
provided above.
Pteropus whitmeei Alston, 1874: 96. Type locality
??Samoa??.
Pteropus nawaiensis Gray, 1870: 107. Type locality
??Nauai?? and ??Ovalau?? (Fiji). Lectotype designation
and restriction of type locality to Nauai by Andersen
(1912: 283?284).
Pteropus vitiensis Gray, 1870: 109. Type locality ??Ovalau??
(Fiji).
Pteropus fuscicollis Nicoll, 1904: 413. Part; nomen nudum
and incorrect subsequent spelling of P. ruficollis
Wilkes, 1844: 128.
Pteropus rufficollis Nicoll, 1908: 245. Part; nomen nudum
and incorrect subsequent spelling of P. ruficollis
Wilkes, 1844: 128.
DISCUSSION
RELATIONSHIPS AND BIOGEOGRAPHY
In his classic monograph on the Pteropo-
didae, Andersen (1912) subdivided the large
genus Pteropus into 17 species groups. No
single reviewer has reviewed the content of the
entire genus in detail since, although some
authors have provided important partial revi-
sions (e.g., Musser et al., 1982; Bergmans,
1990, 2001; Koopman, 1994; Giannini et al.,
2008). It is becoming clear, especially from
molecular studies, that species groups of
Pteropus as currently constituted will require
substantial revision and rearrangement before
these groupings effectively portray evolution-
ary relationships (Kirsch et al., 1995; Colgan
and da Costa, 2002; Giannini and Simmons,
2003, 2005; Giannini et al., 2008), and for now
we suggest these should be employed only as
??groupings of convenience??.
Andersen (1912) united P. samoensis (in-
cluding the Fijian subspecies P. s. nawaiensis,
then recognized as a distinct species) and P.
anetianus of Vanuatu into a unique species
group, designated as ??the Pteropus samoe?nsis
Group.?? These species share a skull that is not
particularly large, but that is extremely robust,
with a relatively short rostrum, heavy mandi-
ble, and large teeth. Based on these same
morphological features, we tentatively add P.
coxi to this species group, which may consti-
tute an older Pacific lineage in the genus
(Ingleby and Colgan, 2003; Giannini et al.,
2008).
Andersen (1912) classified the other com-
mon species of Samoa, P. tonganus, in the
mariannus species group, which includes P.
mariannus from the Mariana Islands, P.
pelewensis from the Palau, P. ualanus from
Kosrae, and P. yapensis from Yap, a group of
closely related allopatric species of the remote
central Pacific (Simmons, 2005). This group of
species is united by a rather uniform color
pattern that includes a pale yellowish mantle
contrasting with a blackish back, and dark
brown underparts with scattered silver hairs.
More recently, especially through genetic
studies (Colgan and da Costa, 2002; Ingleby
and Colgan, 2003), it has become clear that P.
tonganus is most closely related to P. con-
spicillatus of the Moluccas, New Guinea, and
Australia, a species that it closely resembles
morphologically, but that Andersen (1912)
instead placed within a separate species group
(the conspicillatus group). As noted, ongoing
studies such as these provide an indication
that previously recognized species groupings
24 AMERICAN MUSEUM NOVITATES NO. 3646
within Pteropus are united out of geographic
convenience or primarily by features that are
variably plesiomorphic in nature or conver-
gently derived. These will eventually require
comprehensive revision, ideally drawing from
united anatomical and molecular comparisons
(see Giannini et al., 2008).
If we were to place P. allenorum within the
species group framework advocated by
Andersen (1912) and Koopman (1994), it
would be within the chrysoproctus species
group (cf. Simmons, 2005). Other members
of this group include P. chrysoproctus from
the Moluccas, P. rayneri from the Solomon
Islands, and P. fundatus from Vanuatu. These
species share a dentition characterized by
relatively small cheekteeth, with somewhat
more robust incisors and canines. However,
based on our own preliminary examinations,
we suggest that the chrysoproctus group (with
or without inclusion of P. allenorum) is
unlikely to be monophyletic, and we advocate
further study of the immediate relationships of
P. allenorum in particular.
ECOLOGY AND EXTINCTION
With the advent in recent decades of
rigorous zooarchaeological excavations in
Pacific archipelagos, and the resulting system-
atic studies of osteological remains, it is now
well established that both insular extirpations
and global extinctions of vertebrate species
have been widespread and severe throughout
the region since the first arrivals of human
settlers to many island groups in recent
millennia (e.g., Steadman, 1993, 1995, 1997,
2006a, 2006b; Steadman and Kirch, 1990;
Pregill, 1993; Pregill and Dye, 1989; Balouet
and Buffetout, 1987; Mead et al., 2002;
Molnar et al., 2002; Helgen, 2004b). Docu-
mented prehistoric vertebrate extinctions in
the Pacific primarily concern birds, lizards,
and land crocodiles; relatively few examples
concerning mammals are known (Flannery,
1995). Mammalian examples of Holocene
extinction events documented with recourse
to insular subfossil records (each extinction
possibly dating to either prehistoric times or to
recent centuries) thus far include rodent
extinctions in the Bismarck Archipelago and
Solomon Islands (Flannery and Wickler, 1990;
Flannery and White, 1991), bat extinctions in
Hawaii and on Lord Howe Island (Ziegler,
2002; McKean, 1973), insular extirpations of
bats in Tonga and the Cook Islands (Hill,
1979; Wodzicki and Felten, 1981; Tiraa, 1992;
Flannery, 1995; Koopman and Steadman,
1995; Weisler et al., 2006), and insular
extirpations of marsupials and rodents anthro-
pogenically transported to islands in Northern
Melanesia and West Polynesia (Flannery et
al., 1988; White et al., 2000). Closer study of
zooarcheological material already excavated
from various Pacific islands will probably
document additional examples of mammalian
extinction and insular extirpation in the
broader region, particularly for bats (see
Steadman, 2006b: 68; Steadman, in litt.), as
undoubtedly will future zooarchaeological
fieldwork. In West Polynesia, detailed studies
of subfossil vertebrates of Fiji and Tonga have
been published (Molnar et al., 2002; Pregill,
1993; Pregill and Dye, 1989; Steadman, 2006b
and references therein), but excavations in
Samoa have not yet revealed an extensive
Holocene vertebrate record. Prehistoric verte-
brate extinctions in that archipelago, if they
have occurred, remain to be documented
(Green and Davidson, 1969, 1974; Jennings
et al., 1976; Nagaoka, 1993; Steadman and
Pregill, 2004; Steadman, 2006b).
Other Pacific vertebrate extinctions, not
clearly associated with the initial colonization
of Pacific islands by humans, have happened
more recently?within the past 200 years. This
is demonstrated by the discovery (and depo-
sition of representative specimens in museum
collections) of a number of locally endemic
species, mainly birds, that were encountered
during the period of early, pioneering biolog-
ical exploration of the Pacific by European
sailors and scientists (primarily between the
late 17th and early 20th centuries), but that are
no longer to be found in the archipelagos
where they were collected. Avian examples of
historical extinctions are recorded especially
from New Zealand (Tennyson and Martinson,
2006; Worthy and Holdaway, 2002) and
Hawaii (Pratt, 1994; Ziegler, 2002), but also
from the Solomon Islands, New Caledonia,
the Society Islands, Guam, and the Micro-
nesian island of Kosrae (Mayr and Diamond,
2001; Steadman, 2006b and references there-
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 25
in). As far as we are aware, there is only one
previously documented case of an historical
vertebrate extinction in Samoa. This concerns
the large flightless gallinule Pareudiastes paci-
ficus, a Samoan endemic that occurred histor-
ically on Savai9i (Hartlaub and Finsch, 1871),
probably also on Upolu (Whitmee, 1874a),
and perhaps on other islands in American
Samoa (Armstrong, 1932). Known by a
handful of specimens collected between 1869
and 1873 on Savai9i, it probably became
extinct in the late 19th or early 20th century
(Hartlaub and Finsch, 1871; Stair, 1897;
Olson, 1975; DuPont, 1976; Reed, 1980;
Merlin and Juvik, 1983; Pratt et al., 1987;
Taylor, 1998), although some authorities are
convinced that it may still survive in upland
Savai9i as a critically endangered species
(Bellingham and Davis, 1988; Stattersfield et
al., 1998). Whether extinct or critically endan-
gered, as a Samoan endemic known with
certainty only from a small number of 19th-
century specimens, and not definitively re-
corded during the 20th century, Pareudiastes
pacificus offers a close analogy to the two
flying foxes described in this paper. We
suggest that Pteropus allenorum and P. coxi
are probably best regarded as extinct, as
neither species has been definitively encoun-
tered in the past 150 years. However, as with
Pareudiastes pacificus, it would not be sur-
prising if exhaustive searching in the archipel-
ago, perhaps especially on remote islets (cf.
Freifeld et al., 2001) and in upland habitats,
revealed that a population of one or both
species of these large bats still persists.
Pteropus allenorum and P. coxi coexisted
into modern times with at least three other bat
species definitively recorded in the contempo-
rary fauna of Samoa?the small, insectivorous
emballonurid Emballonura semicaudata (mass
averaging 7 g) and the large fruit-eating
pteropodids Pteropus samoensis and Pteropus
tonganus. Pteropus allenorum and P. coxi are
the only endemic chiropteran species known in
the Samoan fauna; the ranges of these other
bats extend (or extended) beyond Samoa at
least to Fiji and Tonga (although Pteropus
samoensis is extinct in Tonga today; Koopman
and Steadman, 1995), and two of them (E.
semicaudata and P. tonganus) are compara-
tively widespread in the broader Pacific region
(Flannery, 1995; Helgen and Flannery, 2002).
Additionally, a species of the vespertilionid
genus Myotis may also have been present in
the 19th-century fauna of Samoa, although
this is disputed (see below).
Specimens retained in museum collections
today reveal that naturalists who visited
Samoa between 1839 and 1856 encountered
four species of Pteropus in the archipelago: P.
samoensis (many specimens, 1839 and later),
P. tonganus (three specimens, 1839?1841), P.
coxi (two specimens, 1839?1841), and P.
allenorum (one specimen, 1856). Members of
the U.S. Exploring Expedition visited Samoa
for about two months over the years 1839?
1841, collecting at least nine specimens of P.
samoensis, but only two specimens of P. coxi
and three of P. tonganus (table 4). This
provides a possible indication that P. samoen-
sis was more common than P. tonganus in
Samoa 170 years ago, although interspecific
differences in behavior (Pteropus samoensis is
diurnal and less skittish than P. tonganus) may
also explain these differences. In any case,
Pteropus tonganus is today the most common
pteropodid in Samoa, P. samoensis is much
less common, and P. allenorum and P. coxi are
probably extinct (Wilson and Engbring, 1992).
With these disruptions over the past century
and a half in taxonomic (and trophic) diversity
(and possibly in relative abundance) in the
flying fox component of Samoa, we speculate
that the tropical ecology of Samoan forests
has probably changed in tandem. Flying foxes
have been characterized as ??strong interac-
tors?? in remote insular tropical forest ecosys-
tems (Cox et al., 1991) because they are
important pollinators and seed dispersers in
these vertebrate-depauperate environments
(Cox, 1982, 1984b; Marshall, 1983; Fujita
and Tuttle, 1991; Whittaker and Jones, 1994;
Banack, 1998; Shilton et al., 1999; Cox and
Elmqvist, 2000; Webb et al., 2000; Nelson et
al., 2000, 2005; Elmqvist et al., 2002; Meehan
et al., 2002, 2005; McConkey and Drake,
2002, 2006, 2007; Parsons et al., 2007). The
extinction of Samoa?s largest and smallest
flying foxes may have had particularly acute
effects on forest ecology, with the remaining,
intermediate-sized Pteropus species unlikely to
replicate all of their ecological interactions.
Many Samoan plants may have depended on
26 AMERICAN MUSEUM NOVITATES NO. 3646
Pteropus coxi and P. allenorum as key
pollinators and seed dispersers, and would
have coevolved with them. We encourage
future studies contrasting cranial and dental
morphologies of the four modern Pteropus of
Samoa in greater detail. In light of previous
studies of chiropteran craniodental features
and their ecological correlates, research along
these lines may further illuminate ecological
differentiation in these four presumably sym-
patric congeners (cf. Aguirre et al., 2002;
Campbell et al., 2007; Dumont, 1997, 2004,
2006; Dumont and O?Neal, 2004; Freeman,
1988, 1995; Steadman, 1997).
Ultimately, recourse to extensive subfossil
deposits will provide the only means to
definitively characterize the complete prehu-
man and pre-European content of the Samoan
bat fauna, should such deposits prove eventu-
ally to be available (Steadman, 2006b). The
elucidation of P. allenorum and P. coxi
demonstrates that the large-bodied bat fauna
of Samoa was twice as diverse in historical
times (comprising four species larger than 200
g) than previously suspected. Other lines of
evidence lead us to suspect that the small-
bodied chiropteran fauna of Samoa may also
have been more species-rich than indicated by
the single insectivorous species known from
the archipelago today (E. semicaudata). The
molossid Chaerephon bregullae and the small
pteropodid Notopteris macdonaldi are both
known from the modern faunas of Vanuatu
and Fiji, to the west of Samoa, and from the
subfossil fauna of Tonga, to the south
(Koopman and Steadman, 1995). Other spe-
cies of Chaerephon are represented in the
modern oceanic insular faunas of Northern
Melanesia (Flannery, 1995), and a second
species of Notopteris (N. neocaledonica) occurs
in the modern oceanic fauna of New Cale-
donia. Clearly, representatives of Chaerephon
and Notopteris (like Pteropus and Embal-
lonura) crossed extensive oceanic barriers
repeatedly during their histories of dispersal
throughout the Pacific; their scattered distri-
bution across multiple remote insular archi-
pelagos suggests to us that Chaerephon and
Notopteris could conceivably have colonized
and occurred in Samoa in the past. Both
Chaerephon and Notopteris (like Emballonura)
are reliant on caves as roosting sites, which
may render them more susceptible to decline
and extinction in the face of intensive exploi-
tation or cave disturbances than forest-roost-
ing bats (Flannery, 1995). As in Tonga, these
elements of the fauna might have become
extinct in Samoa either before or soon after
European exploration and impacts in the
region began, before any modern museum
specimens were (or, perhaps, could be)
obtained. Alternatively, Chaerephon and
Notopteris could occur today in remote cave
systems in Samoa, remaining undetected to
date. Similarly, the vespertilionid genus
Nyctophilus is represented in the modern
faunas of Northern Melanesia (Flannery,
1995), New Caledonia (Parnaby, 2002a), and
Vanuatu (P. Bouchet, personal commun.;
Steadman, 2006b), and persisted into the
19th century in Fiji, at least if the record
presented by Dobson (1878: 174?175) is to be
accepted (e.g., see Miller, 1907; Parnaby,
2002a). On remote Pacific islands, species of
Nyctophilus are associated with montane
forest habitats (Bonaccorso, 1998; Parnaby,
2002a; P. Bouchet, personal commun.), and
we suggest that the mountains of Samoa, little
surveyed for small bats, might well harbor a
representative of the genus, or have done so
into the recent past. Montane forests in Fiji
are the place to search for the continued
occurrence of the genus in that archipelago, as
well.
Another indication of a potentially richer
Samoan ??microchiropteran?? fauna was pro-
vided by Dobson (1878), who described the
vespertilionid taxon Myotis insularum (initial-
ly under the name combination Vespertilio
insularum), the holotype of which was said to
have been collected in the ??Navigators?
Islands?? (i.e., Samoa). According to Dobson
(1878: 313), the holotype was collected by a
??Mr. Schmeltz??. This was most likely
Johannes Dietrich Eduard Schmeltz, custodi-
an of the Godeffroy Museum in Hamburg,
which sold natural history specimens collected
throughout the Pacific region to many
European museums during the 19th century
(see Evenhuis, 2007). Our understanding is
that Schmeltz organized Godeffroy specimens
in Hamburg, but was not a field collector
himself. If the holotype of M. insularum
originated from Samoa, it was likely received
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 27
from Dr. Eduard Gra?ffe, a Godeffroy repre-
sentative based in Apia for a decade beginning
in 1861, who was apparently responsible for
the collection of most Godeffroy specimens
from Samoa (Hoffman, 1999; Evenhuis,
2007). The collectors Andrew Garrett and
Franz Hu?bner also collected in Samoa for the
Godeffroy trading company and museum
(Evenhuis, 2007: 20).
Because no further specimens of Myotis
have been obtained in Samoa, most authors
have regarded the occurrence as probably
erroneous (Tate, 1941; Findley, 1972; Cox,
1983; Koopman, 1994; Helgen and Flannery,
2002). Nevertheless, insularum has never been
firmly equated with any other species in the
genus; its taxonomic status remains indeter-
minate (Simmons, 2005). The discovery of P.
allenorum and P. coxi amongst Samoan
museum material collected in the mid-19th
century suggests to us that the status of M.
insularum as a rare or now-extinct Samoan bat
deserves consideration and renewed study.
The natural presence of Myotis in Samoa
would hardly be surprising, as Myotis is the
most geographically widespread chiropteran
genus on earth (Findley, 1972; Ruedi and
Mayer, 2001). Like all chiropteran genera
present in the modern and subfossil faunas
of Fiji, Samoa, or Tonga apart from the
endemic montane genus Mirimiri (i.e.,
Pteropus, Notopteris, Emballonura, and
Chaerephon), Myotis also occurs in the rela-
tively remote archipelago of Vanuatu
(Medway and Marshall, 1975; Hill, 1983;
Flannery, 1995; Helgen and Flannery, 2002).
Comprehensive systematic study of Old World
Myotis will be requisite to establish whether
the holotype of insularum is truly morpholog-
ically or genetically distinctive relative to
phenetically similar Myotis taxa (cf. Findley,
1972), as would be expected if the holotype
originated (and not as a vagrant) from such a
remote archipelago. In the meantime, we
concur with Simmons (2005) that M. insular-
um should be provisionally ranked as a
Samoan taxon in need of clarifying study. In
this light we note that Kra?mer (1903) listed
two native names applied to small insectivo-
rous bats in Samoa?pe?ape?avai, said to apply
to ??Emballonura semicaudata??, and apa?auvai,
said to apply to ??Emballonura fuliginosa?? (a
synonym of E. semicaudata). We presume this
to be an error of interpretation on Kra?mer?s
part, and that both of these Samoan names
(if indeed they are truly different names) apply
to Emballonura semicaudata. We mention
Kra?mer?s account simply to reference his
apparent impression that more than one kind
of small insectivorous bat was traditionally
recognized by Samoans in historical times (see
also Cox, 1983: 519).
The case of Myotis insularum warns that
careful consideration is needed before accept-
ing the geographic provenance of specimens
from possibly ??unlikely?? (or subsequently
unverified) localities, perhaps especially when
associated label or catalog data provide a
general locality only (such as the ??Navigators?
Islands?? in the case of insularum). Notable
errors have been made in the past in attribut-
ing bats to Pacific archipelagos on the basis of
incorrect locality data. Emballonura semicau-
data was mistakenly recorded from the
Marshall Islands by Tate and Archbold
(1939) (see Sanborn, 1953; Lemke, 1986).
The syntypes of Phyllorhina taitensis
Fitzinger, 1861, a synonym of the horseshoe-
bat Hipposideros speoris (Schneider, 1800),
endemic to the Indian subcontinent, were
supposedly collected in Tahiti, far outside
the natural distribution of Hipposideros, and
they are undoubtedly incorrectly localized
(Dobson, 1877). An early record of Notop-
teris from Ponape? in the Caroline Islands,
based on two immature specimens at the
Naturalis Museum in Leiden (Jentink, 1887,
1888), is probably erroneous (Helgen, in
Simmons, 2005). A particularly relevant case
of misattribution is that of Pteropus laniger, a
species described by Harrison Allen in 1890
(originally as Pteropus lanigera [sic]) on the
basis of two specimens, located today at
USNM and MCZ (Allen, 1890; Helgen and
McFadden, 2001). The type series of P.
laniger, purchased from Ward?s Natural
Science Establishment in Rochester, New
York (a well-known supplier of natural
history specimens), was supposedly collected
in the ??Samoa Islands??, but Andersen (1912)
established that the lectotype of laniger
(designated by Helgen, in Helgen and
McFadden, 2001: 98?99) is morphologically
indistinguishable from Pteropus insularis
28 AMERICAN MUSEUM NOVITATES NO. 3646
Hombron and Jacquinot, 1842, a small flying
fox known only from Micronesia. Indeed,
Allen (1890) based his description more on
geography than anything else; he recognized
that his new taxon laniger was more or less
identical to Pteropus phaeocephalus (a syno-
nym of P. insularis), and he did not offer any
features to distinguish the two nominal forms.
We have independently verified Andersen?s
conclusions regarding laniger based on direct
comparisons of the lectotype of laniger and
series of insularis (at USNM) and through
multivariate craniometric analyses involving
different insular populations of P. insularis,
based on specimens at AMNH, ANSP,
BMNH, MNHN, USNM, and ZMB (our
results, not shown here, suggest to us that the
type series of laniger likely originated from
islands in the Chuuk group of the Carolines).
Certainly, misattribution of the type locality in
the description of Pteropus laniger should give
pause for thought in a paper in which we
describe two species of Pteropus, both credited
to Samoa, on the basis of incomplete 19th
century museum specimens. In contrast to the
situation with laniger, however, for both new
species described here (P. allenorum and P.
coxi), our type material is associated with firm
sources (ANSP associates working on Upolu
in the case of allenorum, and the U.S.
Exploring Expedition in the case of coxi),
rather than natural history dealers. Further-
more, in contrast to Allen?s description of
laniger, we describe P. allenorum and P. coxi
because both taxa are distinguishable from all
previously described Pteropus species, from
Polynesia and beyond.
In summary, based on the recorded histor-
ical distributions of bat genera across remote
Pacific archipelagos, we can easily envision a
prehuman bat fauna in Samoa?s tropical
rainforests that might have included represen-
tatives of Emballonura, Chaerephon, Myotis,
and Nyctophilus as insectivores, Notopteris
and Pteropus allenorum as nectarivores or
flower and smaller fruit feeders, and Pteropus
samoensis, P. coxi, and P. tonganus as large
frugivores, with the latter group of three
species perhaps dominating in inland, mon-
tane, and coastal landscapes, respectively. Of
these, only Emballonura and the four Pteropus
are confirmed members of Samoa?s Holocene
fauna, and only Emballonura and two of the
Pteropus (P. samoensis and P. tonganus)
persist today.
We interpret currently available evidence
from museum specimens and published ac-
counts to suggest that, apart from Pteropus
allenorum and Pteropus coxi, at least five
taxonomically valid species of bats have
become extinct during the past 200 years.
Four of these are species of Pteropus: Pteropus
subniger (Kerr, 1792) of Re?union and
Mauritius in the Mascarene archipelago, last
recorded in the latter part of the 19th century
(Cheke and Dahl, 1981; Moutou, 1982;
Bergmans, 1990); P. pilosus Andersen, 1908,
of Palau, last recorded in the mid-19th century
(Andersen, 1912; Flannery, 1995); P. brunneus
Dobson, 1878, of coastal northern Australia
(recorded only from Percy Island), still known
only by the holotype (Dobson, 1878;
Andersen, 1912); and P. tokudae Tate, 1934,
of Guam, last recorded in the late 1960s or
early 1970s (Perez, 1973a, 1973b; Wiles, 1987;
Flannery, 1995). Apart from these flying
foxes, the mystacinid Mystacina robusta
Dwyer, 1962, of New Zealand, last recorded
in 1967 (Hill and Daniel, 1985; Flannery,
1987, 1995), is the only other bat that can be
definitively regarded as becoming extinct in
recent centuries, although it is likely that
Nyctophilus howensis McKean, 1973, an ex-
tinct bat recorded only by a subfossil skull
from Lord Howe Island, also survived into
historical times (to the late 19th century;
McKean, 1973). Some bats often considered
to be extinct are actually taxonomically
invalid or problematic (e.g., Acerodon lucifer,
Pipistrellus sturdeei; see Simmons, 2005). In
the western Pacific, this category includes
??Pteropus loochooensis?? of Japan, which we
regard as a synonym of P. mariannus based on
independent study (K.M. Helgen, personal
obs.), and Nyctimene sanctacrucis Troughton,
1931 (of the Santa Cruz Islands), which may
be indistinguishable from populations of
Nyctimene major elsewhere in the Solomon
Archipelago (K.M. Helgen, personal obs.).
Other bats often considered to be extinct have
been recently rediscovered as living animals
(e.g., Dobsonia chapmani, Myotis planiceps; see
Simmons, 2005). Several mormoopid and
phyllostomid species from the West Indies
2009 HELGEN ET AL.: TWO NEW SPECIES OF PTEROPUS 29
also became extinct during the Holocene and
may have persisted into historical times, but as
far as we know these taxa remain known
solely from subfossil remains rather than
historical museum specimens (Simmons,
2005; see also Tejedor et al., 2004, 2005).
Thus, of the seven to eight bat species that
probably became extinct within the past 200
years, six are species of Pteropus, and all but
one of these extinctions (P. subniger of the
Mascarenes) took place in the Pacific region.
These extinctions are not well understood, and
they have been tentatively linked to overhunt-
ing and habitat destruction especially
(Flannery and Schouten, 2001). As oceanic
island specialists, many species of Pteropus are
especially vulnerable to natural population
perturbations, such as tropical storms and
epidemic disease (Flannery, 1989; Elmqvist et
al., 1994; Pierson et al., 1996; McConkey et
al., 2004; Esselstyn et al., 2006), and human
impacts such as hunting and deforestation for
agriculture can severely amplify the effects of
these natural impacts (Wiles and Payne, 1986;
Mickleburgh et al., 1992; Craig et al., 1994b;
Brooke and Tschapka, 2002; Struebig et al.,
2007). Any of these factors may have been
involved in the decline to extinction of these
various insular flying foxes. A more extensive
(and speculative) review of possible factors
underlying the extinction of P. allenorum and
P. coxi is beyond the scope of the present
paper, but the elucidation of these new
Samoan species points to the need for further
study in this direction. In any case, the
apparent disappearance of these two
Pteropus species parallels the extinction of at
least one flightless bird, Pareudiastes pacificus,
and the decline and extinction of many
elements in the Samoan land snail fauna
(Cowie, 2001; Cowie and Cook, 2001; Cowie
and Robinson, 2003), declines that have
progressed in the face of ongoing deforesta-
tion, growing human populations, and intro-
ductions of non-native species in the Samoan
fauna and flora over the past century (Evans
et al., 1992; Whistler, 1992; Mueller-Dombois
and Fosberg, 1998; Stattersfield et al., 1998;
Cox, 1999). These anthropogenic perturba-
tions underscore the ecological fragility of
many oceanic insular ecosystems, and they
remind us of the role of conservation biology
in documenting and understanding extinctions
so that further losses can be prevented.
ACKNOWLEDGMENTS
Reports like ours are possible only because
critical specimens have been preserved and
curated since their deposition, decades and
even centuries ago, in natural history museums
around the world. We thank Darrin Lunde,
Eileen Westwig, and Nancy Simmons
(AMNH), Sandy Ingleby and Tish Ennis
(AM), Paula Jenkins, Daphne Hills, and
Richard Harbord (BMNH), Robert Asher,
Irene Thomas, and Peter Giere (ZMB), Hans
Baagoe (ZMUC), and Geraldine Veron and
Jacques Cuisin (MNHN) for access to speci-
mens and other kindnesses during visits to
museums. We thank Wim Bergmans, Norberto
Giannini, and Don Drake for insightful reviews
of our manuscript; Rusty Russell, Marissa
Grunes, David Steadman, Tim Flannery,
Sandra Banack, Paul Cox, Adam Miles,
Alfred Gardner, Storrs Olson, Dan Polhemus,
Philippe Bouchet, F. Russell Cole, and Allen
Allison for valuable discussion; and Suzanne
Cole for assistance at the Library of Congress.
We especially thank Ned Gilmore for facilitat-
ing our work in the collection and archives of
ANSP, Linda Gordon and Helen Kafka for
assistance at the Smithsonian Institution, and
Allen and Heather Drew for their hospitality
during our visit to Philadelphia. This research
was completed during a postdoctoral fellow-
ship to K.M.H. funded by the Smithsonian
Institution, travel funding from the South
Australian Museum, and graduate fellowships
to K.M.H. from the Australian-American
Fulbright Foundation, National Science
Foundation, American Society of Mamma-
logists, and the University of Adelaide.
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