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OCCASIONAL PAPERS
THE MUSEUM
TEXAS TECH UNIVERSITY
NUMBER 81 14 JANUARY 1983
PUBLICATIONS OF THE MUSEUM
TEXAS TECH UNIVERSITY
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A NEW SPECIES OF TUBE-NOSED FRUIT BAT
(NYCTIMENE) FROM THE BISMARCK ARCHIPELAGO,
PAPUA NEW GUINEA
JAMES DALE SMITH AND CRAIG S. HOOD
Tube-nosed fruit bats of the genus Nyctimene (Chiroptera,
Megachiroptera, Pteropodidae), and its sister-taxon Paranycti-
mene, are unique among the various taxa that comprise the fam-
ily Pteropodidae, being distinguished by their peculiar tubular
nostrils. The nine currently recognized species of Nyctimene
occur in Indo-Australia from Timor, through Sulawesi and the
Moluccas, New Guinea and tropical Australia, and the Bismarck
and Solomon Islands to Santa Cruz Island.
In the summer of 1979, the Taylor South Seas Expedition from
the Natural History Museum of Los Angeles County (LACM), led
by one of us (Smith), visited the Bismarck Islands of New Ireland
and New Britain for the purpose of surveying the bat fauna of
this poorly known region. Preliminary results of this expedition
were reported by Smith and Hood (1981). A more extensive report
of collections made, plus the results of a second expedition (1981)
to the Bismarcks (including Manus and Duke of York islands), is
in progress. In addition to the capture of many species new to the
fauna of the Bismarcks, we encountered an undescribed species of
Nyctimene, which is diagnosed and discussed below.
Nyctimene masalai, new species
Demonic Tube-nosed Fruit Bat
Holotype.—LACM 65798, adult male (preserved in alcohol,
skull removed) collected on 9 July 1979 by James Dale Smith
2 OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY
(original number 4522) at Ralum, 10 m.. New Ireland Island,
New Ireland Prov., Papua New Guinea (lat. 3° 33' S, long. 152°
22'
E).
Paratype.—'LACM 65799, young adult male (preserved in alco-
hol, skull removed) collected on 19 July 1979 by James Dale
Smith (original number 4878) at 2 km. NW Hilalon, sea level,
New Ireland Island, New Ireland Prov., Papua New Guinea (lat.
3° 51' S, long. 152° 39' E).
Distribution.—Bismarck Archipelago, Papua New Guinea,
known only from New Ireland Island.
Diagnosis.—Size moderately large (see measurements below).
Color mottled, dark reddish brown above; black, middorsal strip
(5 mm. wide) from back of crown to base of tail; grayish white
with yellowish brown wash below; spots on wings, ears, and nar-
ial tubes whitish. Cranium narrow and elongate (rectangular
rather than squarish); rostrum relatively long and narrow; frontal
sinuses inflated and parallel to each other (not converging in
supraorbital region); maxillary toothrows straight, converging
anteriorly; bony palate moderately domed (not flat), deepest ante-
rior to second upper premolars; postdental palate not markedly
pandurate; pterygoid wings low. Dentition with marked reduction
of coronal cusps; canines relatively short and broad-based, upper
pair lacking labial cusps; P3 and p3 unicuspid and nearly subcon-
ical.
Measurements. —Selected cranial and external measurements (in
mm.) of holotype and paratype, respectively: condylobasal length,
30.7, 29.7; zygomatic breadth, 20.4, 20.9; mastoid breadth, 13.7,
13.5; interorbital breadth, 6.2, 5.7; breadth across canines, 5.6, 5.8;
length of maxillary toothrow, 10.9, 10.4; breadth across upper
molars (MI-MI), 8.3, 9.0; length of mandibular toothrow, 12.5,
12.1; length of mandible, 24.0, 23.4; height of coronoid process,
13.3, 13.7; length of head and body, 103, 94; length of tail, 22, 21;
length of hind foot, 14, 13; length of ear, 14, 14; length of fore-
arm, 67.5, 63.5; weight (grams), 52.3, 45.2.
Description.—Head long and narrow; face deep; ears broad,
bluntly pointed; narial tubes long, directed anteriorly; flight
membranes dark brown; wing attached to dorsal surface of foot at
base of third toe; large white blotches on dorsal surface of fore-
arm and all digits of wing (spots on membrane between fingers
pale yellowish brown); leading edge of ear pinna and narial tube
with white spots.
Pelage and Coloration.—Dorsal pelage long (8-9 mm.) and lax;
extending along pectoral limb to proximal third of forearm and
SMITH AND HOOD—NEW SPECIES OF NYCTIMENE
FIG. 1.—Dorsal and ventral views of skulls of Bismarck Nyctimene. A, /V. ulbi-
venter (LACM 65786); B, N. vizcaccia (LACM 65787); C, N. cyclotis (BBM-NG
28398); D, N. masalai (LACM 65798, holotype); E, N. major (LACM 65802). See
text for discussion.
OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY
IG. 3.—Upper (A) and lower (B) dentition of Nyctimene masalai (LACM 65798,
otype).
g. 3B); internal cusp absent (again, probably fused with main
ip); short labial and lingual ridges extending rearward from
itral cusp, then dropping abruptly to shelflike heel of tooth;
rd lower premolar (p4) slightly smaller than p3; anterior,
SMITH AND HOOD—NEW SPECIES OF NYCTIMENE 7
external cusp high; internal cusp not distinct; these two cusps
connected by a ridge that arcs around anterior margin of tooth;
posteriorly, internal and external ridges drop rather abruptly to
heel of tooth.
Molars. First and only upper molar (Ml) subequal in size to
P4; cusp positions and ridges similar to those of P4 but much
lower; tooth nearly flat as viewed in profile. First lower molar
(ml) similar to p4 in size and coronal morphology; anterior por-
tion of tooth only slightly higher than heel; second lower molar
(m2) slighly shorter than ml; no distinct cusps apparent, crown
flat.
Soft
palate.—Entire
length
of
soft palate covered with
20-21
palatal ridges, eight or nine of which are interdental; first ridge
short, extending straight across palate between first upper premo-
lars; second through fifth or sixth ridges similar in shape with a
lateral branch extending anteriorly from toothrow at about 45°
angle, then bending sharply to cross midline at perpendicular
angle; next few ridges with indentation medially, may connect at
midline; all aforementioned ridges rather thick, rounded, and
separated by deep grooves, their surface wrinkled. Posterior to
interdental ridges is a series of more widely spaced, delicate ridges
clothed by many sharply pointed, toothlike papillae, anteriormost
of which randomly and irregularly traverse soft palate; many do
not reach midline; last eight or nine postdental ridges more or
less regular in form, spacing, and nearly all extend, unbroken,
across soft palate.
Etymology.—The epithet for the new species, masalai, is taken
from the Tolai language and means forest demon or devil.
COMPARISON AND DISCUSSION
The most recent critical review of the tube-nosed fruit bats is
that by Andersen (1912fr:681-722, 828). He treated 13 species of
Nyctimene (papuanus, albiventer, minutus, varius, cyclotis,
cephalotes, geminus, major, scitulus, lullulae, aello, robinsoni,
and certans). The holotypes of these, except tor cephalotes (not
examined by Andersen and apparently lost), are housed in the
collection of the British Museum (Natural History). All except N.
cephalotes (Pallas, 1767), N. albiventer (Gray, 1862), N. major
(Dobson, 1877), N. aello (Thomas, 1900), N. robinsoni Thomas,
1904, N. major lullulae Thomas, 1904, and N. certans Andersen,
1912a were described by Andersen (1910). Since the review by
Andersen (19126), seven additional taxa of Nyctimene have been
8 OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY
described: N. vizcaccia Thomas, 1914, N. draconilla Thomas,
1922a, N. celaeno Thomas, 1922&, N. sanctacrucis Troughton,
1931, N. bougainville Troughton, 1936, N. malaitensis Phillips,
1968, and N. albiventer minor Phillips, 1968. Paranyctimene rap-
tor
Tate,
1942,
also
was described
after
Andersen
(1912&).
Tate
(1942: 341-343) discussed some, but not all, of these taxa in his
account of pteropodids contained in the Archbold collections.
Likewise, Laurie and Hill (1954: 46-48) treated most, but not all,
of these bats. They assigned the various taxa of Nyctimene known
to occur in the geographic area covered by them to species and
subspecies, but did not discuss or otherwise justify these assign-
ments. More recently, several other workers (Phillips, 1968:817-
825; McKean, 1972:16-20; and Koopman, 1979:12) have remarked
on regionally limited taxa of Nyctimene. To date, however, the
genus remains unreviewed in its entirety.
Like many bats from the Indo-Australian region, most species
of Nyctimene are represented only by the holotype or extremely
small series (one or two in most cases) from scattered localities,
many of which are islands. In addition, much of the crucial cran-
ial material is damaged or has heavily worn dentition, thereby
limiting critical comparisons and analyses of characters. These
troublesome factors are further aggravated by a rather high degree
of individual variability that is apparently common among pte-
ropodid bats. In the past, species of Nyctimene have been based
mostly on coloration and overall size. In preparing the descrip-
tion on N. masalai, we have examined and directly compared it
with the 15 holotypes of Nyctimene in the British Museum (Nat-
ural History), holotypes of N. malaitensis and N. albiventer
minor (Bernice P. Bishop Museum, Honolulu), and representative
series of all other species of Nyctimene except N. sanctacrucis. In
addition, we have attempted to identify and use characteristics in
the diagnosis and description that seem applicable to all taxa of
Nyctimene.
Nyctimene masalai is easily distinguished from N. m. major,
with which it is sympatric, on the basis of large overall size of the
latter (Figs. IE and 2A). The cranium of major is flatter in pro-
file, much more massive, and the dentition is characteristically
more cuspidate than that of masalai (Fig. IE). Nyctimene masalai
approaches N. m. scitulus in overall size, but remains distinct
because of the qualitative characters mentioned above. This is
also the case with N. geminus, which currently is regarded as a
geographic race of major. Nyctimene lullulae (also regarded as a
SMITH AND HOOD—NEW SPECIES OF NYCTIMENE 9
subspecies of major) is slightly smaller than masalai, but again, is
distinguished by cranial and dental features typical of major.
Nyctimene robinsoni, N. aello, and N. celaeno (regarded as a race
of aello by Laurie and Hill, 1954) occupy geographic ranges that
are allopatric to that of N. masalai. Nyctimene robinsoni, an
apparently close relative of N. major, is distinguished from masa-
lai by the same general suite of characters that separates masalai
from major. Large size, highly cuspidate dentition, and a unique
broad middorsal stripe easily separate N. aello and N. celaeno
from N. masalai.
Nyctimene masalai is slightly larger in overall size compared to
N. cyclotis (Figs. 1C and 2C). It is conceivable that these two
occur sympatrically {cyclotis having been recently reported from
New Britain Island by Smith and Hood, 1981), but as yet they
remain allopatric. While similar in size, cyclotis is readily identi-
fied by its generally dark, extremely long and wooly pelage, as
well as uniquely cuspidate postcanine dentition (Fig. 1C). The
premolars and molars of cyclotis are round as opposed to rectan-
gular and the premolars have three strong cusps; the palate is
markedly arcuate. The cranium of cyclotis tends to be more
squarish than rectangular. The large round palatal fenestrations
shown in Fig. 1C are not wholly artifactual, but are frequently
encountered in specimens of cyclotis, and are often asymmetrical.
They do not occur in all specimens, but their form, position, and
incidence of occurrence seem consistent enough to consider them
a feature of the species.
The two remaining species that occur in sympatry with N.
masalai are N. albiventer and N. cephalotes. They are similar in
size, but both are smaller than masalai. The Bismarck Archipe-
lago (including the Admiralty Islands) is the only geographic
area in which albiventer and cephalotes are known to occur sym-
patrically. Nyctimene albiventer, in its current context, is a wide-
ranging species that occurs from the northern Moluccas through
New Guinea, the Bismarcks and the Admiralties, to the Solomon
Islands. Laurie and Hill (1954) placed N. papuanus, N. draco-
nilla, and N. bougainville as subspecies of N. albiventer. With
the exception of bougainville (see below), these associations seem
to be correct, although Koopman (1979:6) regarded draconilla as a
distinct species (the specimens referred to draconilla by Greig-
Smith, 1975, and mentioned by Koopman, 1979, as partial justifi-
cation of this arrangement, are Paranyctimene raptor). The name
cephalotes was first introduced into the Bismarck area when
10 OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY
Laurie and Hill (1954) relegated N. vizcaccia Thomas, 1914, to
subspedfic status under N. cephalotes. Formerly, cephalotes was
applied to bats that occurred generally west of New Guinea
(Timor, Peleng Island, off the east coast of Sulawesi, and the
Moluccas); one specimen from Numfoor Island, Geelvinck Bay,
Irian Jaya, was also assigned to cephalotes (Laurie and Hill,
1954; Koopman, 1979).
In the preparation of this description of masalai and the devel-
opment of comparative criteria for species of Nyctimene, we
stumbled inadvertently onto a problem concerning the identity of
albiventer and cephalotes. Neither albiventer nor cephalotes is
especially well defined in the literature and, as noted above, the
holotype of the latter appears to be lost. We arrived at our under-
standing of cephalotes by first defining the nature of albiventer.
For this, we used the holotypes of albiventer, papuanus, and dra-
conilla as well as considerable comparative material from the
mainland of New Guinea. We regard albiventer to be a moder-
ately small species with a narrow, brownish black middorsal strip.
The dorsal pelage is not mottled and the venter is generally uni-
formly whitish or yellowish white. The cranium is squarish with
an extremely short rostrum and globose braincase (Figs. 1A and
2E), and the palate and maxillary toothrow are broad and arcuate
rather than narrow, straight-sided, and convergent anteriorly. The
second upper premolar is bicuspidate with a strong external cusp
and a lower, usually prominent, internal cusp (Fig. 1A), but tooth
wear may obliterate the internal cusp. The second lower premolar
also is cuspidate with a strong external cusp, usually flanked by a
short anterior and posterior loph, and a prominent internal cusp.
There is some individual variation in the distinctness of the
internal cusp throughout the geographic range of the species and
wear quickly obliterates its appearance. However, this tooth and
its upper counterpart are always broad and round rather than
long and narrow. All of these features easily distinguish albiventer
from masalai. They do not, however, characterize tube-nosed fruit
bats from the Solomon Islands that have been previously assigned
to N. albiventer bougainville.
With albiventer so defined, we are left with one remaining spe-
cies in the Bismarcks—supposedly N. cephalotes. This bat agrees
in size and general external appearance with albiventer, but its
dorsal pelage is usually mottled and the venter is often darker. In
describing N. bougainville, Troughton (1936) made similar obser-
vations in his comparison with N. papuanus, and these appear to
SMITH AND HOOD—NEW SPECIES OF NYCTIMENE 11
have influenced Pohle's (1953) association of bougainville with
albiventer. Subsequent authors have followed this assignment.
More importantly, the cranium and dentition of the remaining
taxon differ considerably from those of the form that we regard as
albiventer. The cranium is rectangular with a relatively longer
rostrum than in albiventer, and the braincase is elongate, not glo-
bose (Fig. 1B). The second upper and lower premolars lack a dis-
tinct internal cusp. Often there is a marked, flangelike ridge
sweeping in a graceful and gentle curve from the posterior inter-
nal margin of the cingulum upward to the apex of the prominent
and narrow external cusp (Fig. 1B). This is especially apparent on
unworn teeth. The teeth are usually longer and somewhat nar-
rower than those of albiventer, although this tendency may be
obscured by wear and erosion. The preceding features characterize
specimens formerly referred to N. albiventer bougainville from the
Solomon Islands, the holotype of N. vizcaccia, and a larger series
of topotypes (Ruk, Rooke, or Umboi Island) in the Bernice P.
Bishop Museum.
Finally, we compared the Bismarck and Solomon specimens
with those referred to cephalotes from Peleng Island and the
Moluccas. The latter agree in external appearance and general
shape of the cranium. The crania of Bismarck and Solomon spec-
imens tend to be less rectangular than either those of cephalotes
or masalai. Specimens of cephalotes from Peleng Island and the
Moluccas are larger in overall size, and the upper and lower pre-
molars have a moderately prominent internal cusp. This is also
true of the specimen from Numfoor. On the lower premolar, this
cusp may be reduced to a promontory or shoulder on the internal
ridge that extends from the posterior cingulum to the apex of the
external cusp. Thus, given these differences, we regard Nyctimene
vizcaccia Thomas (1914) to be a valid species, separate and distinct
from cephalotes and masalai, and occupying a geographic range
in the Bismarck and Solomon Islands. All three taxa appear to be
allied and may ultimately be regarded as members of a
"cephalotes-group." Furthermore, we regard N. bougainville from
the Solomon Islands, heretofore assigned to N. albiventer, as a
junior synonym of N. vizcaccia and as a valid subspecies of that
species, Nyctimene vizcaccia bougainville, new combination.
There seems to be little evidence to warrant recognition of the
subspecies minor from Fauro, Choiseui, and Santa Ysabel islands.
Although slightly smaller in overall size, representative specimens
are not markedly removed from the range of variation in N. v.
12 OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY
bougamville, and we therefore regard N. albiventer minor as a
junior synonym of N. v. bougainville.
Nyctimene masalai differs from supposed cephalotes from
Peleng Island and the Moluccas by being larger in overall size,
having a somewhat broader, yet rectangular cranium, and a rela-
tively longer rostrum. The dentition of masalai is, perhaps, the
most reduced of any species of Nyctimene in terms of coronal
cuspidation (Fig. 3). Nyctimene malaitensis Phillips (1968) is
known only from the type specimen from Malaita Island,
Solomon Islands. It appears to be a species distinct from N. v.
bougainville which is smaller in all respects. Nyctimene malaiten-
sis does approach masalai in size, but the cranium is less rectan-
gular, the rostrum is shorter and broader, and the palate is flat,
not domed as in masalai. The dentition of the holotype of malai-
tensis is badly worn. The foundations of the teeth are broad and
rounded, and those of the upper and lower second premolars
appear to have supported internal cusps.
SUMMARY
A new species, Nyctimene masalai, is described from New Ire-
land Island, Bismarck Archipelago, Papua New Guinea. The new
species is compared with all other species of Nyctimene except N.
sanctacrucis. In these comparisons, useful characteristics for iden-
tifying the species of Nyctimene are presented, and Nyctimene
albiventer and N. cephalotes are discussed in detail. As a result,
Nyctimene vizcaccia is raised to species rank. Nyctimene bougain-
ville, from the Solomon Islands (previously assigned to N. albi-
venter), is placed as a junior synonym of N. vizcaccia, and
Solomon representatives are assigned to Nyctimene vizcaccia bou-
gainville. Nyctimene albiventer minor, also from the Solomons, is
put into the synonymy oi N. vizcaccia bougainville.
ACKNOWLEDGMENTS
We thank John E. Hill for reviewing the manuscript and making material in
his care available to us. John Wright, Natural History Museum of Los Angeles
County, and William Presch III, Dept. Biology, California State Univ., Fullerton,
also commented on the manuscript. We thank the curators of the following insti-
tutions for allowing us access to critical material: K. F. Koopman, American
Museum of Natural History, New York; B. Marlowe, Australian Museum, Sydney;
A, Ziegler, Bernice P. Bishop Museum, Honolulu; W. Z. Lidicker, Jr., Museum of
Vertebrate Zoology, Univ. California (Berkeley); H. Felten and D. Kock, Natur-
Museum Senckenberg, Frankfurt; D. E. Wilson, United States National Museum,
Washington; H. Hackethal, Museum fur Naturkunde, der Humboldt Universitat
SMITH AND HOOD—NEW SPECIES OF NYCTIMENE 13
zu Berlin, Berlin, DDR. Also, we would like to extend our thanks to M. Raga
(former First Assistant Director), N. Kwapena (current First Assistant Director), G.
Maynes, F. Kimbag, and C. Umkau, Division of Wildlife, Department of Lands
and Environment, Papua New Guinea, for granting permission to conduct field
research in the Bismarck Archipelago and for providing invaluable assistance and
companionship in the field. Lance Hill and J. Pernetta, Dept. Biology, Univ.
Papua New Guinea, provided institutional affiliation for our field project, and A.
Norrie graciously provided us with shelter and a base of operation during our stay
on New Britain in 1979 and 1981. Susan Smith prepared the illustrations for the
paper. Last, but not least, we wish to thank Mrs. Reese Taylor for her continued
financial and moral support without which none of this work would have been
possible.
LITERATURE CITED
ANDERSEN,
K,
1910.
Ten
new
fruit-bats
of
the
genera
Nyctimene,
Cynopterus,
andEonycteris. Ann. Mag. Nat. Hist., ser. 8, 6: 621-625.
———. 1912a. A new Nyctimene from New Guinea. Ann. Mag. Nat. Hist., ser.
8, 9:95-96.
———. 1912&. Catalogue of the Chiroptera in the collection of the British
Museum. British Mus. (Nat. Hist.), 2nd ed., l:ci+854 pp.
DOBSON, G. E. 1877. On a collection of Chiroptera from Duke-of-York Island
and the adjacent parts of New Ireland and New Britain. Proc. Zool. Soc.
London, pp. 114-127.
GRAY,
J.
E. 1862.
Description
of
some
new
species
of
Mammalia.
Proc.
Zool.
Soc. London, pp. 261-263.
GREIG-SMITH, P. W. 1975. Notes on a collection of bats and their ectoparasites
from the Sepik District, Papua New Guinea. Sci. New Guinea, 3:117-
122.
KOOPMAN,
K.
F.
1979.
Zoogeography
of
mammals from islands
off
the north-
eastern coast of New Guinea. Amer. Mus. Novit., 2690:1-17.
LAURIE,
E. M.
0,
AND
J.
E.
HILL.
1954.
List
of
land mammals
of
New Guinea,
Celebes, and adjacent islands, 1758-1952. British Mus. (NaL Hist.), iv+175
pp.
MCKEAN, J. L. 1972. Notes on some collections of bats (order Chiroptera) from
Papua New Guinea and Bougainville Island. Tech. Paper Div. Wildlife
Res., C.S.I.R.O., Australia, 26:1-35.
PALLAS,
P.
S.
1767.
Spicilegia
Zoologica
quibus
novae
imprimis
el
obscurae
animalium species inconibus descriptionibus atque commentarius illus-
trantur. 3. Berolini.
PHILLIPS,
C.
J.
1968. Systemalics
of
megachiropteran bats in the
Solomon
Islands. Univ. Kansas PubL, Mus. Nat. Hist., 16:777-837.
POHLE, H. 1953. Uber die Fledertiere von Bougainville. Z. Saugetierk., 17:127-
137.
SMITH, J. D., AND C. S. HOOD. 1981. Preliminary notes on bats from the Bis-
marck Archipelago (Mammalia: Chiroptera). Sci. New Guinea, 8:81-121.
TATE, G. H. H. 1942. Pteropodidae (Chiroptera) of the Archbold Collections.
Bull. Amer. Mus. Nat. Hist., 80:331-347.
THOMAS,
0.
1900.
Description
of
a new
fruit-bat
from New
Guinea.
Ann.
Mag. Nat. Hist., ser. 7, 5:216-217.
14 OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY
———. 1904. New bats and rodents from West Africa, the Malay Peninsula, and
Papuasia. Ann. Mag. Nat. Hist., ser. 7, 14:196-202.
———. 1914. On mammals from Manus Island, Admiralty group, and Ruk
Island. Bismarck Archipelago. Ann. Mag. Nat. Hist., ser. 8, 13:434-439.
———. 1922a. On mammals from New Guinea obtained by the Dutch scientific
expeditions of recent years. Nova Guinea, 13:723-740.
———. 1922&. New mammals from New Guinea and neighboring islands. Ann.
Mag. Nat. Hist.. ser. 9:261-265.
TROUGHTON,
E. LEG.
1931.
Three
new bats
of
the
genera
Pteropus,
Nyctimene,
and Chaerephon from Melanesia. Proc. Linnaean Soc. New South
Wales, 56:204-209.
———. 1936. The mammalian fauna of Bougainville Island, Solomon group.
Res. Australian Mus., 19:341-353.
Addresses
of
authors;
JAMES DALE
SMITH,
Dept.
of
Biological
Sciences.
Califor-
nia State University, Fullerton, California 92634 and Section of Mammals, Natural
History Museum of Los Angeles County, 900 Exposition Blvd., Los Angeles, Cali-
fornia
90007;
CRAIG
S.
HOOD,
The
Museum
and
Dept.
of
Biological
Sciences,
Texas Tech University, Lubbock, Texas 79409. Submitted 20 June, accepted 2
August 1982.
