HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

OCCASIONAL PAPERS

of the

MUSEUM OF NATURAL HISTORY
l^yj^ CQlJSfi^yniversity of Kansas
LIBfCawrence, Kansas

^NUMBER 19, PAGES 1-47. NOVEMBER 9, 1973

Jj[|5^ffS^ATICS OF THE GENUS RHOGEESSA
T^n^OPTERA: VESPERTILIONIDAE )

By
Richard K. LaVal'

The several species of iXeotropical bats of the vespertihonid
genera Rhogeessa and Baeodon inchide some of the smallest of
living mammals, although one species approaches in size the com-
mon big brown bat, Eptesinis fusciis, of North America. Most of
the species occur at low ele\ati()ns, in habitats ranging from desert
scrub to rain forest, and appear to be insecti\'orous. They range
from the lowlands of northern Mexico to southern Brasil, but are
poorly represented in collections from South America south of
Venezuela and Colombia.

Allen (1S66) described the genus Rho<i,ecssa including two spe-
cies, R. parvula and R. tiiniida. He implied close relationships of
Rho(geessa with Nycficejus [:=Nijcticeius], Nyctinoinus [=Ta-
darida], and with the Noctilionidae. Dobson (1878) referred Rho-
geessa to a subgenus of the genus Vespenigo [^=Vespertilio]. How-
ever, Thomas ( 1892 ) retained Rhogeessa as a full genus, and stated
that it was most closely related to Nycticejus [=^Nycticeius]. Sub-
sequently, Miller (1897) and others have followed Thomas. Miller
( 1906) erected a new genus, Baeodon, for Rhogeessa alleni Thomas.
Although Simpson (1945) included Baeodon in Rhogeessa, other
recent authors, including Tate (1942), Hall and Kelson (1959),
and Koopman and Cockrum ( 1967 ) ha\'e retained them as separate
genera.

The eight nominal species of Rhogeessa were described in the
following sequence: R. pawtda (Tres Marias Islands, Nayarit) and

1 Adjunct Professor, Department of Systematics and Ecology, and Research
Associate, Museum of Natural History, Unixersity of Kansas. (Present address:
Organization for Tropical Studies, Unixersidad de Costa Rica, Ciudad Uni-
versitaria, Costa Rica, C.A.)

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

R. tumida (Veracruz) by H. Allen in 1866; K. allen'i (Jalisco) by
Thomas in 1892; R. minutilla (Margarita Island) and R. g,racilis
(Puebla) by Miller in 1897; R. io (Venezuela) and R. velilla (Ecua-
dor) by Thomas in 1903, and R. bombijx (Colombia) by Thomas
in 1913. Hall (1952) reduced R. tiimida to a subspecies of R.
parvula, an arrangement followed by Hall and Kelson (1959) and
several authors prior to 1959. No new taxa were described in the
genus Rhogeessa until Goodwin's (1958) revision, in which he
named three subspecies, R. tumida major (Oaxaca), R. tumida
riparia (Venezuela), and R. parvula aeneus (Yucatan). At the
same time he included R. bombijx as a subspecies of R. tumida,
and reduced R. minutilla, R. io, and R. velilla to subspecies of R.
parvula.

The distributional ranges Goodwin gave for the seven subspecies
he recognized suggest a montage of isolated subspecies, some sepa-
rated by gaps of over 1500 miles. On the basis of my preliminary
examination of specimens in tlie Museum of Natural History at the
University of Kansas, I concluded that his taxonomic arrangement
failed to elucidate the actual intra- and interspecific relationships
within the genus. Comments made by Alvarez and Avifia (1965)
and by Jones, et al. ( 1971 ) also suggest the need for a revision of
the genus Rhogeessa. Therefore, with the 650 specimens now avail-
able, I have undertaken a new revision, employing multivariate
statistical analyses.

MATERIALS AND METHODS

A total of 653 specimens of Rhogeessa was examined. Included
were 8 alleni (2 bacula), 350 R. tumida (43 bacula), 160 R. parvula
(28 bacula), 112 R. mimitilla (11 bacula), 9 R. gracilis (4 bacula)
and 14 (1 baculum) of an unnamed species. Although a substantial
number of juvenal specimens was encountered, no more than one
per locality was included in the total number of specimens exam-
ined. Because many important specimens are preseived in alcohol,
skulls from these specimens were removed and cleaned.

I examined the following holotypes: R. gracilis: R. tumida: R.
tumida major: R. tumida riparia; R. parvula; R. parvula aeneus;
R. minutilla. Holotypes of R. alleni, R. bombyx, R. io, and R. velilla
were examined for me by J. E. Hill of the British Museum of Natural
History. C. O. Handley of the National Museum of Natural History
generously made available to me his notes on the holotypes of
R. io, as compared with various specimens in the NMNH. Topo-
types of R. velilla and R. parvida were examined by me.

I recorded all available label and field note information, plus
three external measurements, length of fur, nine cranial measure-
ments, three bacular measurements, color of dorsum and of venter,
presence or absence of sagittal crest, age (adults: phalangeal

SYSTEMATICS OF THE GENUS RHOGEESSA 3

epiphyses ossified; juveniles: epiphyses not ossified) and toothwear
( 1 — none; 2 — present but scareely noticeable; 3 — easily noticeable
on canines and molars, but little cusp detail obscured; 4 — heavy,
much cusp detail obscured). Measurements were taken following
in general the methods outlined by Handley ( 1959 ) , with excep-
tions or additions as noted below: forearm (FA); third metacarpal
(SMC); tibia (from center of knee joint to point of attachment of
calcar to ankle); fur (greatest length in center of dorsum); greatest
length of skull (GLS — including incisors); depth of braincase
(DB); postorbital width (POWO; mastoid width (MW); width
across second upper molars (M2-M2); width of second upper molar
(M2 — as measured with ocular micrometer); maxillary toothrow
(MAX); mandibular toothrow (MAND); measurements of l^acular
length, depth, and width h>llow LaVal (1973, Fig. 1).

A set of nine specimens ( Table 1 ) was chosen as color standards
and matched to named colors from Ridgway (1912). Unfortunately
certain specimens, notably those from northwestern Venezuela, fell
outside the limits of variation delimited by the color standards;
these were compared directly with Ridgway (1912).

Table 1. Colors from Ridgway (1912) applied to specimens used as color

standards (tip color only).

Color

Specimen no.

no.

Dorsum

Venter

TCWC 19756

1

Fuscous-Black

Buffy Brown

KU 102619

2

Hair Brown

Cartridge Buff

KU 29886

3

Taw n\ -Olive

Light Ochraceous

-Buff

KU 107494

4

Buffv Brown

Cartridge Buff

KU 105565

5

Buckthorn Brown

Light Ochraceous

-Buff

KU 90754

6

Pinkish Cinnamon

Cinnamon Buff

UNM 27545

7

Warm Buff

Light Ochraceous

-Buff

KU 97050

8

ca. Light Ochraceous-Buff

Pinkish Buff

KU 61171

9

Dresden Brown

Light Ochraceous

-Buff

Karyotypes were not examined by me, but I have seen partial
results of a study of karyotypic variation in the genus now being
conducted by R. J. Baker (pers. com.) and his students. Baker and
Patton ( 1967 ) documented the differences in karyotypes between
R. tumicia from Chiapas and R. powula from Sonora and Nayarit.
Specimens identified by Baker and Patton as R. (iraciUs are not of
that species (Baker, pers. com.) and are almost certainly R. parvulo,
collected at the opposite end of the long cline beginning in Sonora.

Scanning electron micrographs were made of hairs from the
mid-dorsal region of specimens of each species. Because the appear-
ance of Rhogeessa hairs varies substantially depending on the angle
of view and the portion of the hair photographed, all photographs
were made of the mid-section of each hair; the hair was oriented

4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

so that the side of the scales bearing the V-shaped depression in
the rim would faee the film. Polaroid negatives (4 X 5 in) were
exposed for 70 seconds on a ETEC Autoscan scanning electron
microscope, made available to me by the Department of Ento-
mology, Kansas State University, Manhattan.

Penes were removed from 89 of the males examined, and cleared
and stained to reveal the bacula, using Anderson's ( 1960 ) method.
Bacula were measured with an ocular micrometer.

All data recorded were punched on IBM cards, and statistical
analyses performed on the GE 635 computer at the University of
Kansas Computation Center. Although f-tests performed between
the sexes indicated a significant level of sexual dimorphism for
some measurements at some localities, the small sample sizes
prevalent in this genus made separation of the sexes impractical.
Samples usually contained a fairly well-balanced proportion of tlie
two sexes, which would tend to offset any possible bias due to
sexual dimorphism.

Gabriel's Sum of Squares Simultaneous Testing Procedure (SS-
STP) was performed, first with all species in the same analysis,
and then each species was analyzed separately. An SS-STP program
available at the University of Kansas Museum of Natural History
(as modified by Gary Powers) gives 13 standard statistics, calcu-
lates and lists non-significant subsets, and prints these in graphic
form, with means in descending sequence (Figs. 10, 11, 13). The
SS-STP procedure, as related to systematics, is discussed by Smith
(1972), who lists some of the more important literature references.

The University of Kansas Numerical Taxonomy Program (NT-
SYS) developed by E. J. Rohlf, J. Kishpaugh, and R. Bartcher was
utilized to give the following statistical information: 1) standard-
ized data correlations between localities and a two-dimensional
phenogram based on these; 2) cophenetic correlation, in which
cophenetic values are plotted on scatter diagrams against the cor-
relation coefficients used in the phenogram; 3) standardized data
distances between localities, and a two-dimensional phenogram
based on these; 4) a second cophenetic correlation, plotting co-
phenetic values against distance coefficients; 5) a principal com-
ponent analysis in which the first five principal components are
calculated; 6) a Varimax Factor Rotation by tlie Kaiser Method;
and 7) the variance-covariance matrices projected onto tlie principal
components (both rotated and non-rotated), and two-dimensional
scattergrams plotting each of the principal components against
each of the others.

A stepwise discriminant analysis was also carried out, using the
University of California BMD07M program; this program identi-
fies each individual specimen based on tlie criteria deri\'ed from
the analysis. It computes canonical correlations and coefficients

SYSTEMATICS OF THE GENUS RHOGEESSA 5

for canonical variables, and plots the first two canonical variables
on a two-dimensional scattergrani.

All of the multivariate analyses were computed using the same
nine characters, all of which are skin and skull measurements made
by myself. These were chosen for several reasons, one of which
dealt with problems resulting from missing data. Discriminant
function analysis indicated that all contributed significantly to
variation observed among the samples analyzed. Other characters
examined either did not contribute to variation among samples, or
contributed so strongly to singling out one or more species or
samples that I felt they would tend to obscure the close relation-
ships which are typical of species within this genus. Further, I
wanted to see if a multivariate analysis based on a few measure-
ments made of morphological characters, which I would expect
to be highly correlated, would support suspected relationships based
on other kinds of characters (such as distribution of hair on uro-
patagium, size of i3, presence of cingual cusps, bacula, hair, etc.).
Blackith and Reyment ( 1971 ) stated that "Where the organisms
are fairly closely related, and all the measurements arc quantita-
tive, .... Satisfactory analyses have been made with as few as
three to six characters, although ten might be regarded as more
optimal. ..."

In the analyses performcxl by the NT-SYS program, only sample
means were utilized. To lu>lp assure that means used would be
representative of the geographic area wliich they represented,
several small samples often were pooled (for example, several
samples of one or two specimens each from various localities on
the Caribbean slope of Honduras were combined). In other cases,
large samples from a single locality were utilized. The areas from
which each of the samples was taken, along with the abbre\'iations
used in this paper, are as follow: Tamaulipas, coast (Tamp 1);
Tamaulipas, interior (Tamp 2); San Louis Potosi (SLP); Veraciaiz,
northern (VC-N); Veracruz, central (VC-C); Veracruz, southern
(VC-S); Campeche (Camp); Yucatan and northern Quintana Roo
(Yuc); Chiapas, Pacific Coast (Chiapas); Oaxaca, Pacific slope
(Oax); Cuerrero (Guer); Michoacan (Mich); Colima (Colima);
Jalisco (Jal); Nayarit (Nay); Sinaloa, southern (Sin-S); Sinaloa,
central (Sin-C); Sonora (Son); Guatemala, Pacific Coast (Gnat);
El Salvador (El Salv); Honduras, Caribbean slope (Hond); Nica-
ragua, Caribbean slope (Nic); Costa Rica, Interior (CR); Panama,
Pacific slope (Pan); Colombia, north (Colomb); Venezuela, north-
west (Venez-NW, 1-5); Venezuela, coast (Venez-C, 1-2); Vene-
zuela, interior (Venez-int); Venezuela, northeast (Venez-NE);
Trinidad (Trin); Guyana (Guyana). All specimens of R. (gracilis,
R. allcui. and the unnanuxl species are pooled into single samples.

Fewer samples, each representing more indi\'idual samples com-
bined into larger pooled samples, were used in the stepwise dis-

6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

criminant analysis, because the kind of output generated by that
program would be unintelligible if a very large number of indi-
vidual samples were chosen. These samples, along with their ab-
breviations, are as follows: Southwestern Mexico — Oaxaca and
Guerrero (SWMEX); Michoacan (MICH); West-central Mexico-
Jalisco, Colima, Nayarit, southern Sinaloa (WCMEX); Northwest-
ern Mexico — northern Sinaloa and Sonora (NWMEX); Eastern
Mexico — Tamaulipas, San Luis Potosi, and Veracruz (EMEX);
Northern Yucatan Peninsula (YUC); Central America — Tabasco
through Panama, excluding Caribbean slope from Nicaragua to
Panama (CENTAM); Caribbean slope from Nicaragua to Panama
(NICPAN); Northern Colombia (COLOMB); Northern Venezuela
(VENEZ); Arid northwestern Venezuela and extreme northeastern
Colombia (DRYVEN); Guyana (GUYANA); Trinidad (TRIN);
all R. gracilis (GRACIL); all R. alleni (ALLENI); all specimens of
the unnamed species ( MIRA) .

Although sexual dimorphism in size was detected within some
samples, f-tests did not show it to be consistently significant. On
the average, females are as much as 4% larger than males. However,
in R. minufiUa from Venezuela, males are slightly larger on the

average.

ACKNOWLEDGMENTS

I wish to thank the following persons and institutions for per-
mission to examine specimens in their care. Abbreviations as used
in lists of specimens examined are given in parentheses. Listed
alphabetically. Ticul Alvarez, Escuela Nacional de Ciencias Bio-
logicas, Mexico City (ENCB); Robert J. Baker, Texas Tech Uni-
versity (TTU); Alberto Cadena, Universidad de los Andes, Bogata
(LTAB); E. Lendell Cockrum, University of Arizona (UA); James
S. Findley, Museum of Southwestern Biology, University of New
Mexico (UNM); Charles O. Handley, Jr., National Museum of
Natural History (NMNH); Robert S. Ploffmann, Museum of Natural
History, Lhiiversity of Kansas (KU); Emmet T. Hooper, Museum of
Zoology, University of Michigan (UM); Karl F. Koopman, Ameri-
can Museum of Natural History (AMNH); Barbara Lawrence,
Museum of Comparative Zoology, Hai-vard University (MCZ);
George H. Lowery, Jr., Museum of Zoology, Louisiana State Univer-
sity (LSU); Helen Matuskowitz, Academy of Natural Sciences of
Philadelphia (ANSP); Donald R. Patten, Los Angeles County Mu-
seum of Natural History (LACM); James L. Patton, Museum of
Vertebrate Zoology, University of California, Berkeley (MVZ);
Randolph L. Peterson, Royal Ontario Museum (ROM); Jose
Ramirez-P., Instituto de Biologia, Universidad Nacional Autonoma
de Mexico (UNAM); David J. Schmidly, Texas Cooperati\e WM-
life Collection, Texas A&M University (TCWC); Luis de la Torre,
Field Museum of Natural History (FMNH).

SYSTEMATICS OF THE GENUS RHOGEESSA 7

J. E. Hill of the British Museum of Natural History (BMNH)
and C. O. Handley of the National Museum of Natural History are
due a debt of gratitude for making available to me notes and meas-
urements of those holotypes on deposit in the British Museum. I
thank Robert J. Baker, Brent L. Davis, and V. Rick McDaniel for
showing me unpublished results of their study of karyotypic varia-
tion in Rhogeessa. Robert S. Hoffmann kindly provided editorial
assistance. My study was financed, in part, by Penrose Fund Grant
6320 from the American Philosophical Society, by Biomedical Sci-
ences Support Grant RR-07037, and by computing fimds allocated
through the Division of Biological Sciences, College of Liberal Arts
and Sciences, Universitv of Kansas.

ANALYSIS OF GHARAGTERS

Goodwin (1958), in his revision of the genus Rhofi,eessa, stated
that the genus (exclusive of Baeodon) could be separated into three
species: "a big eared species with thin flying membranes [iiraciJis],
a large species with short ears and thick membranes [tiimida], and
a small species [pawtila]." He further stated that "The propor-
tionate length of the forearm and color of thc> pelage cannot be
relied on as determining factors except in a \'er\' general way.
Granial measurements can also be confusing unless accompanied
by comparative material." As might be anticipated from these intro-
ductory remarks, Goodwin did not state clearly how to distinguish
among parvula, tumida, and the \'ari()us subspecies into which he
divided them.

As suggested by Goodwin, skin and skull measurements are in-
adequate for distinguishing among species, except in the case of
R. aUeni, which is larger in most cranial variates than any other
species examined, and the new species, R. mira, which is smaller
than any specimens of R. parviihi from Michoacan or adjoining
states. However, in the multi\ariate analyses, notably the principal
component analysis, the canonical \'ariate analysis, and the dis-
criminant function analysis, the measurements taken were adequate
to discriminate many of the samples from each other (see table
4, and Figs. 4, 7, 8, 12). The discriminant function analysis indi-
cated that greatest length of skull and forearm were the best and
second best discriminators, respecti\'ely. Thus, I chose these two
variables to demonstrate geographic variation by means of uni-
variate analyses.

Color of fur. — \^ariation in relative positions of light and dark
bands seems to separate alleni and (gracilis from all other species.
Individual variation in contrast between bases and tips renders
pelage coloration of limited usefulness in identification of individual
specimens of the other four species, but dKFerences are evident
when series are compared. Molting individuals take on a grayish

8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

cast because dark gray hairs grow up throughout the old pelage
before it is shed. Thus, newly molted individuals are much darker
in color than individuals in old pelage. In the species which have
pale bases and darker tips, individual color variation is accounted
for primarily by differences in tip color, whereas color differences
among samples and species typically result from variation in basal
color.

Length of fur. — Although some individual variation in fur length
occurs, this character is sufficiently constant within a population to
separate certain species, and to illustrate geographic variation
among populations in a single species.

Distribution of fur on uropatagium. — This character evidences
so little individual variation that it is a fairly reliable means of
separating one of the species from the other five species.

Degree of toothwear. — Tooth wear was negligible in about 90
percent of specimens examined, except in the case of R. minutilla,
in which more than 35 percent of the specimens had moderate to
heavy toothwear.

Lingual cingulum of CI. — In all species except gracilis and mira
two (rarely one) cusps are present on the lingual cingulum of the
CI. These cusps are variable in development, from higher and
sharper than the example shown in figure lA, to lower and more
rounded. They tend to be obscured in individuals with heavy
toothwear. In the species lacking cusps, the cingulum is perfectly
smooth and straight, lacking even the slightest swelling (Fig. IB).

Bacula. — With the exception of R. alleni, bacula of the various
species are not sharply differentiated from each other. Although

B

Fic. 1. Upper liulit canine of two species of Rlio^crssa,
lingual \ie\v. A. R. jxtiviila, UA 10319, Jalisco; B. R. mira,
UNAM 8594, Michoacdn. x25.

SYSTEMATICS OF THE GENUS RHOGEESSA 9

bacular shape shows substantial geographic and indixidnal varia-
tion within species, it also seems to differ among species in areas
of sympatry and near sympatry (Fig. 2). Single bacula of parvida,
tumida, gracilis, and aUeni were illustrated and described by Brown,
et ah (1971), but these authors did not examine bacular variation.
Microstructure of hair. — Benedict (1957) examined hairs from

d

A
a//en/

B
gracilis

C

-parvuia

D

E
mira

tumida

L

J L

M N

—minutilia — '

K
tumida

Fig. 2. Top and lateral outline views of bacula of 14 specimens of
Rhogeessa. A. R. aUeni, KU 97307, Michoacan; B. R. gracilis, KU 92951,
Jalisco; C. R. parvuia, KU 97080, Sinaloa; D. R. parvuia, UNAM 8865, Guer-
rero; E. R. mira, UNAM 8953. Michoacan; F-L, all R. tumida: F. KU 55172,
Tamaulipas; G. KU 82923, Veracruz; H. LACM 18683, Chiapas; I. TCWC
24127, Nicaragua; ]. MVZ 113936, Colombia; K. KU 119073, Venezuela;
L. AMNH 66824, Ecuador; M. R. minutilia, NMNH 441783, Venezuela;
N. R. minutilia, NMNH 441805, Venezuela. X25.

10

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 3. Scanning electron micrographs of hairs of six species of Rho^ccssa.
A. R. mira, UNAM 8593, Michoacan; B. R. paivula, KU 107494, Jalisco;
C. R. minuiiUa, AMNH 130673, Venezuela; D. R. allcni, KU 61171, Oaxaca;
E. R. ftimida, KU 119073, Venezuela; F. R. gracilis, KU 97050, Jalisco. xl400.

SYSTEMATICS OF THE GENUS RHOGEESSA 11

most of the known genera of bats, including hairs from a specimen
of R. parvida. Unfortunately, she neither illustrated a hair from
Rhogeessa nor stated where the specimen was captured, and she
had only a light microscope at her disposal. Benedict (1957) and
Quay ( 1970 ) implied that hair structure was most useful as a taxo-
nomic tool at the generic and suprageneric levels. Although the
relationship (if any) of geographic, sexual, age, and individual
variation to hair structure cannot be evaluated here because of
small sample sizes, it is evident (Fig. 3) that there are differences
in hair structure among species of Rhogeessa.

The two similar lowland species of western Mexico, R. mira and
R. parvula, differ from all other species in that each scale is rotated
90° along its linear axis, in relation to adjoining scales; in the other
species scales are in alignment with each other. Based on hair
structure, two other groupings seem to be evident. In R. alleni and
R. gracilis the scales appear to be cone-shaped rings which are
bilaterally symmetrical, subtending a full 360° (the coronal scales
of Benedict). In R. tumida and K. minuiiUa the scales are alternate
and overlapping (imbricate scales of Benedict) with the visible
portion subtending an angle of 180° or perliaps slight!)' greater.

SYSTEMATIC ACCOUNTS

Rhogeessa H. Allen, 1866

Rhogeessa H. Allen, 1866:285 [Type species. — Rhogeessa parvula H. Allen,
1866, designated hy G. S. Miller, 1897 in accordance with page priority].

Baeodon G. S. Miller, 1906:85 [Type species. — Rhogeessa alleni Thomas, 1892,
by original designation]. \'alid as a subgenus.

Description.— As given by H. Allen (1866) and G. S. Miller
(1897) except as noted below. Skull drawings appear in Hall and
Kelson (1959; R. parvula) and G. S. Miller (1897; R. tumida); skull
photographs are included by Goodwin and Creenhall (1961; R.
tumida and R. minutilla). The i3 varies in size from minute (visable
only under magnification) to only slightly smaller than i2; il and i2
tricuspid, with outer cusp much lower than other two cusps; i3 bi-
cuspid or unicuspid; lingual cingulum of CI well developed, usually
with accessory cusps; parastyle, paracone, protocone, and first and
second commissures are only structures present on M3; width of
posterior half of m3 substantially less than width of anterior half;
maxillary cheek teeth converging anteriorly; basisphenoid pits ab-
sent; rostrum narrower than braincase; membranes relati\'ely thin;
baculum saddle-shaped at proximal end, with well-developed proxi-
mal lateral knobs, poorly-developed proximal median knob, and
long, usually narrow shaft, circular or elliptical in cross section
near distal end; length 0.50 to 0.80 mm; penis short, 3-4; baculum
extends to within 0.5 of distal end of penis.

12

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14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Comparisons. — Although Rhog^eessa shares its dental formula
with several other genera of bats, only Nycticeius has been sug-
gested to be a close relative. The only diagnostic differences noted
by any authors relate to the lower incisors, which are well spaced,
of uniform size, and have three cusps of equal size in Nycticeius. In
my comparison of N. humeralis, type species of its genus, with
RJwgeessa, the following additional differences were observed in
Nycticeius: mesostyle of M3 present, but low; third commissure of
M3 present; width of posterior half of m3 only slightly less than
width of anterior half; maxillary toothrows parallel; basisphenoid
pits present; rostiami nearly as wide as braincase; membranes rela-
tively thick; baculum with enlarged distal knob, relatively small
proximal lateral knobs, and poorly developed proximal median
knob (see Plate I in Hamilton, 1949); length about 2.25 to 2.6 in
four specimens; penis long, about 7-8; baculum extends from mid-
point of penis to a point about 1.5 short of distal end of penis. The
karyotype of N. humeralis differs in numerous ways from those of
R. tumida and R. parvula (Baker and Patton, 1967). Karl Koopman
(pers. com.) checked the list of characters listed above in seven
additional species of Nycticeius (schlieffeni, poUidus, greyi, hal-
stoni, rueppeUi, hirundo, and aJ])ofuscus) . Information on bacula
and karyotypes was not available. Most of the other characters
seemed to be consistent among the specimens of the species ex-
amined by Dr. Koopman, except penes were seen only for greyi,
balstoni, and hirundo; the mesostyle and third commissure of M3
and the relative widths of anterior and posterior halves of m3 in
rueppeUi were more like Rhogeessa than in other species of Nycti-
ceius; basisphenoid pits were rather weakly developed in some
species.

Subgenus Baeodon G. S. Miller, 1906

Baeodon G. S. Miller, 1906:85 [Type species. — Rhogeessa alleni Thomas, 1892,
by original designation].

Description. — As in account of genus, except as follows: i3
unicuspid, peglike, one sixth to one eighth cross-sectional area of i2;
postorbital width narrow relative to overall .skull size; .skull large
relative to body; first phalanx of third digit long relative to third
metacarpal.

Rhogeessa (Baeodon) alleni Thomas

Rhogeessa alleni Thomas, 1892:477 [Holotype.— BMNH 93.2..5.25 from Santa
Rosalia, near Autlan, Jalisco, Mexico; A. C. Buller, collector].

Baeodon alleni— G. S. Miller, 1906:85.

Holotype.— V>r\i\<,]\ Museum of Natural History 93.2.5.25, adult
female, presei-ved in alcohol, faded, skull removed. Measurements
in table 3.

SYSTEMATICS OF THE GENUS RHOGEESSA

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16

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Description. — A very large species of Rhogeessa, with tribanded
dorsal fur and minute i3. Distal one fourth of dorsal hairs close to
Dresden Brown, center one half buffy, and basal one fourth gray;
ventrally, tips close to Light Ochraceous-Buff, bases gray; fur 4-6;
dorsum of uropatagium almost bare; average ratio of third meta-
carpal to first phalanx of third digit 2.20:1; sagittal crest present in
8 of 9 skulls; sagittal and occipital crests form "helmet" above oc-
ciput; i3 average diameter 0.1, with little variation; cusps on lingual
cingulum of CI small, one or two in number; skull large in all
dimensions except postorbital width.

Bacula. — The two R. alleni bacula examined are 0.44, 0.50 in
length, 0.20, 0.22 in depth, and 0.44, 0.28 in width. In addition to
their short lengths, these two bacula are notable for their short
shaft and widely flaring proximal lateral knobs, with convex proxi-
mal margins (Fig. 2A). Although these bacula are quite distinct

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for gracilis (triangle), allciii (square), and mira (star) represent single pooled
samples of all available specimens of those species.

SYSTEMATICS OF THE GENUS RHOGEESSA 17

from those of any other North American species of Rhogeessa, a
few South American specimens of R. tumida possess bacula with
similar development of the lateral knobs; however, the bacula
have a longer shaft and greater length than those of alleni.

Comparison. — Rhogeessa (Baeodon) alleni is larger in cranial
measurements (except postorbital width) than any specimens of
the three species with which it occurs sympatrically in southwestern
Mexico (Table 2). Among the samples of specimens from south-
western Mexico compared in table 2, ^tests showed that mean
ratios of third metacarpal to first phalanx were significant between
parvula and alleni, and between gracilis and alleni, at P=:<.001,
with no overlap. Rhogeessa alleni is separated from both by the
characters given in the subgeneric description of Baeodon. It is
easily separated from gracilis by the shorter ears and relatively
narrow basal band of the dorsal fur; it can be easily distinguished
from parvula by the three-banded dorsal fur (two-banded in par-
vula). In the multivariate analyses presented graphical!)' here,
Figs. 4, 6, 7, <S, 12, 14), alleni is distinctly separated from the other
species of Rhogeessa, supporting assignment of that species to a
monotypic subgenus. No specimens were misidentified by the dis-
criminant function analysis (Table 4).

Distribution. — Mountains of western Mexico, from central
Oaxaca north to central Jalisco (Fig. 5). Altitudinal range from
125 m to 2000 m. W^getation at the seven known localities seems
to be either desert scrub or tliorn forest. Altliough R. alleni may be
more widespread than these few records indicate, it is probably
restricted to western Mexico northwest of the Isthmus of Tehuan-
tepec.

Reproduction. — An individual caught on 6 August was not
pregnant, and another female taken on 29 November was not lac-
tating. An adult male captured on 28 March had testes 3 mm long.
No juveniles were found.

Remarks. — In removing alleni from the genus Rhogeessa and
erecting the genus Baeodon for it. Miller (1906) only mentioned a
single diagnostic character, the minute i3. Relative size of teeth
and even number of teeth var\' within other vespertilionid genera.
In R. parvula size of i3 is \ariable, and in some specimens ap-
proaches the condition observed in R. alleni. In most other char-
acters, alleni is clearly similar to other species of Rhogeessa, al-
though it has by far the largest skull of any species in the genus.
For these reasons there seems to be no justification for assignment
of R. alleni to a genus separate from Rhogeessa.

It should also be mentioned that the nominal species Baeodon
meijeri Pine is not a Rhogeessa (see Pine, et al. 1971) and will
not be treated in this revision.

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

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SYSTEMATICS OF THE GENUS RHOGEESSA 19

Subgenus Rhogeessa II. Allen, 1866

( see citation under account of genus )

Description. — As in account of genus, except that cross-sectional
area of i3 always greater than one sixth cross-sectional area of
i2; postorbital width wide relative to o\'erall skull size; skull small
relative to body; first phalanx of third digit usually short relative
to third metacarpal.

Rhogeessa (Rhogeessa) gracilis G. S. Miller

Rhogeessa gracilis G. S. Miller, 1897:126 [Holotype.— NMNH 70694 from
Piaxtla, Puebla, Mexico, elevation 1100 m; E. W. Nelson and E. A. Gold-
man, collectors].

Holotype. — National Museum of Natural History 70694, adult
male, preserved in alcohol with skin peeled back from skull; penis
and baculum removed, cleared, and stained; baculum 0.72 in
length, 0.20 in depth, 0.44 in \\'idth. Comparted with KU 97050.
Other measurements in table 3.

Description. — A largo species of Rhop^eessa with very long cars
(average length LS, based on collectors" measurements), long, three-
banded dorsal fur, and relatively narrow skull. Distal one fourth
of dorsal hairs close to Light Ochraceous-Buff; center one fourth a
paler buff, and basal one half dark grayish brown; ventrally, tips
close to Pinkish Buff, bases dark grayish brown; fur 6-7; dorsum
of uropatagivmi with sparse fur extending almost to knees; mean
of ratios of third metacarpal to first phalanx of third digit 2.56 to 1;
sagittal crest present in 7 of 8 skulls, but poorly developed, as are
occipital crests; i3, 0.2 or greater in width, nearly as large as 12;
lingual cingulum of CI low, smooth, lacking cusps; skull moderate
in size, but relatively narrow; slope of forehead steep; braincase
inflated.

Bacida. — Measurements of four R. pricilis bacula: 0.72-0.80 in
length, 0.16-0.28 in deptli, and 0.40-0.54 in width. As viewed from
above ( Fig. 2B ) the triangular outline of the baculum of gracilis
differentiates it from the other three species with which it occurs
sympatricall)'. In lateral profile the relatively straight shaft con-
trasts with the normally curved shaft of the other species.

Comparisons. — The large ears and unique color banding pattern
allow gracilis to be separated easih' from all other species. Like-
wise, the skull differs from those of other Rhogeessa in the relatively
steeply sloping forehead, inflated braincase, and narrow lateral
dimensions. In superficial appearance, the skull could be mistaken
for that of a Myotis. In other species of the genus, the skulls are
similar to those of Eptesicus, with respect to their widths and the
slightly sloping foreheads, and their tendency to form a supra-
occipital "helmet."

20

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

The distance phenogram (Fig. 6) separates grar/Z/.s rather well
from the otlier samples of similarly-sized individuals of other species.
The discriminant function analysis correctly identified all specimens
( Table 4 ) , and the plot of the first and second principal components
(not figured) placed gracilis well outside the main cluster of sam-

r-

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120 Chiapas

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l)ased on multivariate analysis of 9 mensural characters. Computed from the
distance matrix. The cophenetic correlation is 0.860.

SYSTEMATICS OF THE GENUS RHOGEESSA 21

pies. Altliough other multivariate methods were of little value,
they suggested that gracilis is more closely related to the other
species of the subgenus Rhogeessa than it is to Baeodon.

Distribution. — Mountains of western Mexico, from Jalisco,
Oaxaca, and Puebla ( Fig. 5 ) . Altitudinal range from 600 m to 2000
m. The three specimens in the University of Kansas collection were
taken at three localities described as follows by the collector: "pine-
oak forest"; "oak forest"; "deciduous forest along stream, pine-oak
forest on hills." However, it is not safe to assume that gracilis is
restricted to this kind of vegetation, because one of the other speci-
mens was taken at 600 m, where the predominating vegetational
association in western Mexico is thorn forest and desert scrub.

Reproduction. — A female captmcd on 15 May contained two 17
mm embryos. Two subadults taken on 27 July were nearly adult
in size. No other information concerning reproduction in the species
is available.

Rhogeessa (Rhogeessa) parvula H. Allen

Rhogeessa parvula H. Allen, 1866:285 [Holotype.— ANSP 1832 from Tres
Marias Islands, Nayarit, Mexico; Col. Grayson, collector].

Rhogeessa tumida major Goodwin, 1958:4 [Holotype. — AMNH 175263 from
San Bartolo Yautepec, Oaxaca, Mexico, 800 m; Thomas MacDougall, col-
lector] .

Holotype. — Academy of Natural Sciences of Philadelphia 1832,
adult, sex indeterminant (male, according to Allen, 1866); skin
disintegrated; skull intact but damaged, i3 notably smaller than il
or i2; compared with KU 105565 and KU 107494 from Jalisco, and
KU 90754 from Sinaloa. Measurements in table 3.

Description. — A small to moderate-sized Rhogeessa with short
ears and relatively hairy uropatagium. Distal one third of dorsal
hairs. Hair Brown to \\^irm Buff; basal two thirds, which may or
may not contrast with tips, pale grayish to buff to pale yellow;
ventrall)', tips from Cartridge Buff to Light Ochraceous-Buff, bases
concolor to slightly contrasting with tips; fur 3-7; sparse to mod-
erately thick fur on dorsum of uropatagium usually extends halfway
from knees to feet; ratio of third metacarpal to first phalanx of third
digit averaging 2.66 to 1 in area of sympatry with alleni and gracilis;
sagittal crest absent in 115 of 131 skulls; i3 \'aries in size from nearly
equal to i2, to minute, but never as small as in alleni; lingual cingu-
lum of CI well-developed, bearing two cusps of variable size; in
some, cusps much higher than in example of figure lA; skull small
to moderate in size; forehead slope slight; postorbital width narrow
in relation to length of skull.

Bactda. — The average dimensions of 28 bacula are: length,
0.63 (0.48-0.76); depth', 0.13 (0.08-0.20); and width, 0.34 (0.22-
0.42). Bacula of parvula are characterized by a long, slender, curved

22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

-a

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fl
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Dinajvd -j; vpimn} -y vpiuint y Dpnuni -y

SYSTEMATICS OF THE GENUS RHOGEESSA 23

shaft, roughly elHptical in cross-section, and by short proximal
knobs, extending at a 45° angle to the sagittal plane of the baculum,
separated proximally by an indentation, and curving ventrally
around urethra (Figs. 2C and 2D); width approximately one half
length of baculum. They differ from most tumida bacula in being
more slender, from both the dorsal and lateral aspects. Juveniles
and some adults (sexually immature individuals?) with shorter
shaft and knobs.

Comparisom. — Rhogeessa (Rhogeessa) parvula are easily dis-
tinguished from alleni and gracilis, as outlined under the accounts
of those species. This species differs from R. mira in its greater
size and the presence of cusps on cingulum of CI. In the Isthmus
of Tehuantepec parvula and tumida are known from localities only
50 miles distant, and are probably sympatric. Here the two species
cannot be distinguished on the basis of size. (Table 2). Chiapan
specimens of tumida, as compared to Oaxacan specimens of parvula,
are differentiated by darker tips of dorsal fur, which contrast more
sharply with bases. The uropatagium is virtually bare, and the i3 is
nearly as large as the i2.

The two most consistent characters for separating parvula and
tumida, fur coverage on uropatagium and relative size of i3 and
i2, seem to be exaggerated in tlie Isthmus of Tehuantepec, where
sympatry may occur. No specimens from that area exhibit inter-
mcdiacy in these characters. For these reasons it seems advisable
to regard parvula and tumida as species.

The discriminant function analysis (Table 4) correctly identi-
fied 76 of 115 specimens; of tlie 39 incorrectly identified, all but 8
were identified as being R. tumida. The plot of the second and
fourth principal components (Fig. 4) reveals a close relationship
between parvula and the two samples of tumida from the Yucatan
Peninsula, but shows good separation between parvula and the
other species, including most samples of tumida. The canonical
scattergrams (Figs. 7 and 8) indicate a wide overlap with tumida.

Distribution and geographic variation. — Western Mexico, from
central Sonora south and east to the Isthmus of Tehuantepec ( Fig.
9). Altitudinal range from sea level to 1600 m; however, the highest
elevation recorded from Nayarit and states to the nortli is 600 m.
Vegetational associations from which R. parvula is known include
scrub desert, thorn forest, tropical deciduous forest, and pine-oak
forest. Broad-leafed and evergreen forests (mostly cloud forest)
of various compositions occur locally in portions of the range of
parvula, but I am unable to establish definitely that any specimens
were taken in those kinds of vegetation.

Variation in R. parvula is essentially clinal. Size measurements
indicate increasing size from nortli to soutli (Table 2), with topo-
types from the Tres Marias Islands close to specimens from Sonora.

24

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

The north-south cline and widely overlapping non-significant sub-
sets are well illustrated by the measurements of forearm (Fig. 10)
and greatest length of skull (Fig. 11). Distance coefficients suggest
that the cline is steeper south of Jalisco (Fig. 15), but there is no
logical point at which a line separating two subspecies might be
drawn. There is a tendency for specimens from more southerly
localities to have fur extending farther out the uropatagium, to be
brighter in color, to have Icmger fur, to have relatively smaller i3,
and to have a more well developed sagittal crest; all of these trends
appear to be clinal, with perhaps the most rapid changes occurring
in Michoacan (one known locality) and the western half of Guer-
rero (no known localities). Were it not for the intermediacy of
the series from Michoacan, it could easily be maintained that the
northern and southern populations represented two subspecies (or
two species, fide Goodwin, 195<S ) .

Reproduction and molt. — Pregnant females were recorded from
the months of February ( 27th ) , March, April, May, and early June.

3.6
26

^ 1.6

>

0.6

S-0.4

I -,.4

o

-0-24

o

^-3,4

CO

-44
-5.4

n 1 1 i 1 r

— /? mi'ra

— WC. MEXICO

— SW MEXICO

— NW MEXICO-"

— /?. o//eni

R.parvula

-I 1 1 1 1 1 1 1 1 1 1 1 1 r

-7.4 -5.4 -3.4 -1.4 0.6 2.6 4.6

First Canonical Variable

Fig. 7. Scatter diagram from first two canonical varial^les for three species
of Rhof^ecssa from western Mexico. The lines were formed l)y connecting the
outlying dots within each of the 5 pooled samples. The spaces enclosed by
the lines originally contained the follo\\ing number of dots, each representing
an iiidi\idual specimen: R. iniia, 13; WCMEX, 75; SWMEX, 28; NWMEX,
12; R. allctii, 7. Specimens of unknown identity may be plotted on canonical
scattergrams for purposes of identification. See Appendix 1.

SYSTEMATICS OF THE GENUS RHOGEESSA

25

Lactating females were taken during April (27th), May, June, and
July (3rd). Because few embryo measurements are available, there
is no indication of any definite growth trends or patterns. The only
juveniles (flying young) recorded were taken on 14 June, 18 July,
9 September, and 18 September. The presence of flying young and
embryos in GuerrcTO in June and February-March, respectively,
suggests that parturition occurs earlier in the southern portion of
the range of R. parvula. At least eight females from Sinaloa and
Sonora contained two embryos, but two bats from Guerrero and
one from Sonora contained only one embryo each. One female was
molting on 4 July, which would be anticipated, if she had just
weaned her young. The new pelage was dark gray, contrasting
sharply with the pale, old pelage. Two males from July and several
females from August and September exhibited very dark-colored
tips, suggestive of recent molt. Negative data showed that some
females collected on various dates during March, April, May, June,
July, and September were not pregnant. Weights for 30 non-
pregnant individuals varied from 3 to 8. No seasonal trends in
weight could be detected.

o

I

o
o

"c
o
c
o

o

o
o

CD

4.6

3.6

2.6
1.6

0.6
-0.4
-14
-24

— E MEXICO

— CENTAM t NICPAN + GRACILIS

— YUCATAN _

-5.4

T 1

-3.4

-| 1 r

-1.4

0.6
First Canonical Variable

2.6

Fig. 8. Scatter diagram from first two canonical \arial)les for tlircc i^ooled
samples of RJiogccssa luinida. The following numbers of dots are enclosed by
die lines: EMEX, 85; CENTAM -{- NICPAN + GRACIL, 90; Yucatan, 5.

26

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

20

10

cs-^-:

A minulilla
• tumida
^^ parvula

0 200

30

600

0 200 600

20

100

Fig. 9. Map showing localities from which were examined specimens of
one of three species of Rlwgecssa. The shaded area represents the distril^ution
of R. parviila, the unshaded area, R. tumida. The triangles representing iiiintt-
tilla are all in northwestern Venezuela and adjacent Colombia, plus the island
of Margarita.

Remarks. — Goodwin (1958) used size as the major criterion for
separating parvula and tumida. He lacked specimens from Jalisco,
Nayarit, and Sinaloa where a clinal change occurs; therefore, it is
not suiprising that lie identified the larger specimens from south-
western Mexico as tumida. Specimens of tumida from adjoining
Chiapas and Veracruz are much like Oaxacan parvula in size. Good-
win also assigned populations of small individuals from the Yucatan
Peninsula and parts of South America to parvula. In addition to
the nominate subspecies, he assigned four other subspecies to par-
vtda. One of these, minutilla, is here regarded as a species; the
other three are relegated to the synonymy of tumida.

Rhogeessa (Rhogeessa) mira, new species

Holottjpe. — Universidad Nacional Autonorna de Mexico 8594,
adult male, skin, skull, and baculum, from 20 km N El Infiernillo,
Michoacan, Mexico, elevation 125 m; collected by Ticul Alvarez,
29 November 1964. Gollcctor's measurements, overall length, 70;
tail, 33; foot, 6; ear, 12; length of testis, 9; external and cranial meas-
urements in Table 3.

Etymology. — The specific epitlict mira is feiuinine for the Latin
mirus, meaning wonderful, astonishing, or extraordinary. These

SYSTEMATICS OF THE GENUS RHOGEESSA

27

parvula-

Loca ity

Mean Su

bsets

~0axaca(9)

30.9

Guerrero(6)

30.5

MichoacanO)

29.9

CoimadS)

28.9

Jalisco (16)

28.9

Sonora(l9)

282

Sinaloa-S(8)

28.

Sinaoa-N(29)

279

_Nayarit(24)

27.7

Michoacan(l4)

26.0

mira

Fig. 10. SS-STP analysis showing geograpliic \anation in
forearm length in Wiogeessa parvula. The vertical bars con-
nect samples within each non-significant subset (at the 0.05
level). For example, the mean of Sinaloa-S is not significantly
different from the mean of Sonora, but is significantly difl:erent
from the mean of mira.

Locality

parvula-

Mean Subsets

~0axaca(9)

3.5

Guerrero( 4)

3.2

Michoacan(8)

3.0

Ja isco( 4)

2.7

Colima ( 5)

2.6

Sina oa-S(7)

2.3

Sinaoa-N(25)

2.2

Nayarit(24)

2.1

_Sonora( 3)

2.0

Michoacan( 3)

4

mira

Fig. 11. SS-STP analysis showing geographic variation in
greatest length of skull of Rhogeessa parvula.

were my initial reactions upon examining a specimen of R. mira
and comparing it with the mucli larger species taken at the same
locality, R. oUeni and R. parvula.

Description. — An extremely small Rhogeessa with smooth-edged
lingual cingulum of CI and external characteristics much as in
R. parvula. Distal one third of dorsal liairs Buckthorn Brown to
Bully Brown; basal two thirds more buflV than tips, not contrasting
strongly with tips; ventrally, overall color like that of bases of dorsal

28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

hairs, tips not contrasting; fur 4; sparse fur on dorsum of uropa-
tagium extends to or past knees; sagittal crest absent; i3 only slightly
smaller than i2; lingual cingulum of CI smooth, lacking the slightest
suggestion of cusps; skull very small; forehead slope moderate.

Baculum. — Measurements of the single baculum examined (Fig.
2E) are: length 0.66, depth 0.12, and width 0.34. It could not be
distinguished from bacula of some R. parvula.

Comparisons. — R. mira is easily distinguished from the other
three species in western Mexico on the basis of size. It resembles
gracilis in its smooth lingual cingulum of CI. Otherwise it could
not possibly be confused with gracilis or alleni. The lack of cingual
cusps, small overall size, and relatively large size of i3 separate it
from sympatric parviila. Nine specimens of parvula from the type
locality of mira are all redder and yellower than any specimens of
mira caught at the same time, or at any other time. Furthermore,
there is no size overlap among specimens of the two species caught
at the type locality. Certain specimens of parvula from Sonora and
the Tres Marias Islands are almost as small as mira, but tend to have
a less steeply sloping forehead, wider and longer rostrum, cingual
cusps on CI, smaller, more crowded i3, and less fur on uropatagium.
Small individuals of tiimicla and minutilla from Yucatan, Venezuela,
and Ecuador differ in color, skull outline, cingual cusps, nearly bare
uropatagium, and shape of baculum.

Generally, the multivariate analyses placed mira well away from
most samples of all other species. Based on these analyses alone,
mira would seem almost as distinct from other Rhogeessa as in R.
alleni. However, in external and cranial characters not included
in the multivariate analyses, mira is more similar to the others,
especially to parvula. In the univariate analysis mira, which had the
lowest means among all samples for most of the variates measured,
usually formed a nonsignificant subset with Sinaloa and Sonora,
with Sonora, or alone (as in Figs. 10 and 11).

Remarks. — All the specimens from 7 km N El Infiernillo were
caught in nets over a small cement water trough, in an area of
cactus-mesquite vegetation (Alvarez and Aviiia, 1965). Collected
at the same time were R. alleni, R. parvula, Balantiopten/x plicata,
Macrotus tvaterhouseii, and Micronycteris megalotis. The Rhogeessa
were taken on five nights, 25 and 27 March, and 26, 28, and 29
November, all in 1964. The capture of three species of Rhogeessa
during one night, in one net, is of interest, because each of the
three is in a non-overlapping size range. This suggests successful
partition of the feeding niche, possibly based on prey size. The
failure to achieve this kind of diversity within the genus Rhogeessa
outside of western Mexico could be accounted for by the presence
of additional Neotropical spc^cies of the genus Mi/otis in those areas,
although this suggestion is highly speculative at this time. It is also

SYSTEMATICS OF THE GENUS RHOGEESSA 29

noteworthy that R. niim is among the smallest of all speeies of bats,
although it exceeds in size specimens of the species Tylomjcteris
pachypus of Southeast Asia, and possibly specimens of certain
species of Pipistrellus, notably P. mimiis.

Rhogeessa (Rhogeessa) tumida H. Allen

Rhogeessa tumida H. Allen, 1866:286 [Holotype.— ANSP 1831 (skin), NMNH

37329 (cranium), and NMNH 84021 (lower jaw) from Mirador, Veracruz,

Mexico, ca. sea level; Dr. Sartorius, collector].
Rhogeessa io Thomas, 1903:382 [Holotype.— BMNH 94.9.25.1 from Valencia,

Carabobo, Venezuela; A. Mocquerys, collector].
Rhogeessa veUlla Thomas, 1903:383 [Holotype.— BMNH 99.8.1.5 from Puna,

Isla Puna, Gulf of Guyaquil, Ecuador, elev. 10 m; P. O. Simons, collector].
Rhogeessa homhijx Thomas, 1913:569 [Holotype.— BMNH 13.10.29.1 from

Condoto, Choco, Colombia, elev. 300 ft; Dr. H. G. F. Spurrell, collector].
Rhogeessa tumida riparia Goodwin, 1958:5 [Holotype. — AMNH 69968 from

Cumanacoa, Sucre, Venezuela, elev. 700 ft; G. H. H. Tate, collector].
Rhogeessa paixtda aeneus Goodwin, 1958:6 [Holotype. — AMNH 91234 from

Chichen-Itza, Yucatan, Me.xico, elev. ca. 10 m; R. T. Hatt, collector].

Holotype. — "Museum of Smithsonian Institution" 8195, male, is
the number listed by H. Allen (1866) as the holotype of tumida.
Part of the confusion regarding the location, status, and numbers
of the various parts of the holotype was discussed by Hall (1952).
The skin, in alcohol and appearing to be a female, is in the Academy
of Natural Sciences of Philadelphia, no. 1831. Two labels are with
the specimen; one reads "1831 Harr. Allen Rhogeessa tumida Mira-
dor, Mex. TYPE H. Allen." The other bears only the number 8195.
The cranium, NMNH 37329, is now in a vial bearing that number.
The accompanying lower jaw, numbered NMNH 84021, seems to
match the cranium perfectly. In another \ial, labelled NMNH
84021, is a cranium bearing that number and a lower jaw numbered
NMNH 37329. These also seem to match perfectly, which is sig-
nificant, because the two skulls in question differ in size. The vial
holding NMNH 84021 is also labelled "formerlv 7842" which would
make it the paratype of R. parvula. It appears that the lower jaws
were switched before the skulls were numbered. In any case
NMNH 37329 (cranium) is obviously of the species now occurring
in eastern Mexico, and should be regarded as the holotype. The
holot\pes of R. tumida and R. t. riparia were compared with KU
29886 from \^eracruz, TCW'C 19756 from Nicaragua, UNM 27545
from Oaxaca, and AMNH 91234 from Y\icatan (holotype of R. p.
aeneus), among others. Handley (in litt.) compared the holotype
of R. io with NMNH 374017, R. tumida from Urama, Venezuela, and
J. E. Hill (in litt.) compared the holotypes of R. io and R. velilla
with each other. Handley also compared the holotype of R. velilla
with NMNH 312113, R. tumida from Ft. Kobbe, Panama. Measure-
ments are in table 3.

30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Description. — A small to iiiodcrate-sized Rlw^eessa with short
ears and nearly bare uropatagium; otherwise highly variable. Distal
one third of dorsal hairs Fuscous-Black to Pinkish Cinnamon, ex-
cept in Venezuela and Guyana, where darkest specimens Brussels
Brown, and palest specimens somewhat darker than those from other
parts of the species' range; bases usually buffy gray to buffy yellow,
and may or may not contrast sharply with tips; ventrally, tips Buffy
Brown to Light Ochraceous-Buff, concolor to somewhat paler at
bases; fur 3-4 (rarely 5); sparse hairs on dorsum of uropatagium
rarely extend as far as knees; sagittal crest present in approximately
half of all specimens examined, but present in only one fourth of
specimens from Colombia, Venezuela, Trinidad, and Guyana; i3
usually smaller than i2, but may equal i2; even when i3 equals i2,
i3 has less well-developed cusps; lingual cingulum of CI with two
small cusps, sometimes only a suggestion of cusps present; skull
small to moderate in size; forehead slope slight; "helmet" often
present in North American specimens, rarely in South America.

Bacida. — Twenty-nine bacula of North American specimens have
dimensions as follows: length, 0.65 (0.50-0.80); depth, 0.19 (0.12-
0.28); and width, 0.52 (0.36-0.84). In fourteen specimens from
South America, means are: length, 0.66 (0.48-0.80); depth, 0.19
(0.14-0.24); width, 0.60 (0.44-0.74). The only apparent difference
between the two groups is in width, and this is not statistically
significant.

In shape, both individual and geographic variation are apparent
(Fig. 2F-L). Generally, as compared to bacula of pawula, the
bacula of tumida possess wider shafts, wider spreading (laterally),
and wider (distal-proximal) proximal knobs; their width exceeds
one half their length. In Central America the bacula of tumida
tend to be Y-shaped, due to deeper indentation between ends of
proximal knobs. The distal free portion of shaft is shorter than that
portion from which knobs extend. In some, additional widening
of shaft results in a roughly triangular shape. In Colombia, Y-
shaped, triangular, and intermediate specimens appear; whereas
in Venezuela, Guyana, and Trinidad, the Y-shape is rare, and in
some the proximal knobs extend laterally at an angle of 90° to
the shaft.

Comparisons. — In the Isthmus of Tehuantcpec R. tumida differs
from R. pawula as outlined under the account of that species. In
general, the bare uropatagium and relatively large i3 of tumida
distinguish all tumida from all parvida. Other characters tend to
be geographically variable and thus will not separate consistently
the two species. In northwestern Venezuela and adjacent Colombia,
where tumida apparently occurs sympatrically with minutilla, tu-
mida is larger on the average, and has proportionately greater post-
orbital width (Table 2). Rho<i,eessa tumida is always darker in
color (both fur and membranes). It has a greater difference in

SYSTEMATICS OF THE GENUS RHOGEESSA 31

length between forearm and third metaearpal, and the tibia is
shorter. Toothwear is rarely heavy. In most of the extensive range
of tumida there is no other species of Rhogeessa present.

The discriminant function analysis correctly identified 194 of 212
specimens as tumida (Table 4). Two were misidentified as minii-
tilla, one as gracilis, and 15 as parvida. In the pooled sample of 75
from Central America, a region where a bewildering amount and
kind of geographic variation occurs, only 15 were correctly identified
as being from Central America. The value of the other multivariate
analyses is discussed under pawida. It is noteworthy that the canoni-
cal scattergram (Fig. 12) seems to separate tumida in South Amer-
ica surprisingly well from minutilla.

Distribution and geographic variation. — Tamaulipas, south in
Caribbean lowlands, including the Yucatan Peninsula, to Pacific
versant in Chiapas, throughout Central America, and South America,
locally south to southern Brazil, Bolivia, and Ecuador (Fig. 9).
Sea level to 1500 m in North America, with the higlior elevations
from Chiapas through Costa Rica; sea level to 1200 m in South
America. A large majority of all specimens were collected from
below 500 m. Rhogeessa tumida seems to be known from virtually
every major vegetational association in tropical North America and
northern South America, within the limits imposed by elevational
restrictions. It is difficult to generalize in terms of geographic
variation in size, since clines are poorly developed and skin and
skull measurements do not indicate the same kinds of trends. Mexi-
can specimens (excluding Yucatan) tend to be the largest, fol-
lowed by specimens from Central America (excluding Caribbean
coast) and then specimens from South America (including Yucatan,
but excluding Pacific coast of C()loml:):a) .

The SS-STP analyses for fort^arm and greatest length of skull
(Fig. 13) illustrate geographic clianges in size. Note that specimens
from Costa Rica (interior), from Nicaragua (Caribbean lowlands),
and from eastern Mexico have the largest forearms, with the inter-
mediate sized bats being from the Pacific versant of Central America
( including Chiapas ) and southern Campcchc. Most of the smallest
specimens are from South America, but those from the northern
Yucatan Peninsula are the most diminutive of all. Based on greatest
length of skull, the same group of samples (with the addition of
Chiapas) is largest in size, but they are arranged in a different
sequence. The middle-sized group is similar except for sequence
and the loss of the El Salvador-Nicaragua (Pacific versant) sample
which drops to a position among the South American samples. The
Colombian sample also drops significantly in the standings. How-
ever, two specimens from the Pacific coast of Colombia are as large
as specimens from Nicaragua (Caribbean) and are not included
in the sample from Colombia.

The distance coefficients (Fig. 14) differ substantially in value

32

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

ill various parts of tlie range and support my coutrntion tluit varia-
tion is essentially non-clinal. The largest values in the eonterminous
portion of the range are those between southern Campeehe and
Veracruz, and between Campeehe and Pacific coastal Chiapas. Val-
ues between Campeehe and Guatemala, El Salvador, and Honduras
are much lower, however. Few specimens are available from the
Yucatan Peninsula, but it appears that a South-North cline of de-
creasing size is operative, resulting in specimens from Yucatan and
Quintana Roo which are more Lke South American specimens in
size than like those from adjacent areas. From Honduras south and
east through western Panama, it appears that clines from small to
large and pale to dark operate along a Pacific-Caribbean axis. Two
specimens from the Pacific coast of Colombia are also large and
dark. It seems probable that the large, dark population of Carib-
bean Central America extends across Panama in the far east to join
the large, dark bats of Colombia, but ncj specimens are available

4.6 -
3.6 -

J3

2.6

O

V

5

1.6

D
O

0.6

C

o

-04

U

O

•o

-14

c

o

-24

iO

-34

-44

I .1

COLOMBtVENEZ+GUYANAtTRINIDAD
minutilla

-3.4 -1.4 0.6 2.6 4.6

First Canonical Variable

6.6

Fig. 12. Scatter diagram from first two canonical variables for Rhogeessa
tnmida (4 pooled samples) and R. minutilla in Sonth America. The following
numbers of dots are enclosed by the lines: COLOMB -}- \^NEZ + GUYANA
+ TRINIDAD, 49; mimitiUa, 62.

SYSTEMATICS OF THE GENUS RHOGEESSA

33

to support that hypothesis. If true, it will pose some rather inter-
esting systematic problems.

Farther south along the Pacific Coast of South America, two
specimens known from Puna Island, Ecuador, are both very small
in size, and are similar to specimens from the Caribbean coast of
Colombia and Venezuela. Two specimens from Brazil are very
much like specimens from Chiapas and Guatemala, in size and
color. I was not able to examine the single specimen from Bolivia
in the Field Museum of Natural History. South of Venezuela and
central Colombia, R. tumida is represented by so few specimens
that it is not possible to speculate on geographic xariation or even
on the limits of geographic distribution. Because of the absence
of specimens among large collections of bats from some parts of
Brazil, Peru, Bolivia, Paraguay, and Uruguay, I believe that R.
tumida is truly a rare species in much of South America, and may
be only locally distributed.

Color is both individually variable, based on age of pelage, and
geographically variable. Eastern Mexican specimens tend to be
closer to the red and yellow end of the spectrum, with little contrast

Subsets Locality Mean Subsets

Chiapas(7) 13.5

Veracruz-N(8) 13.5

Tamaulipas2(l7) 13.4

Tamaulipasl(32) 134

Nicaragua(Carib)(5)l33

Veracruz-C(7) 13.2

Veracruz-S(l2) 13.1

SanLuisPotosi(l9) 13.1

Costa Rica (13) 13.1

Honduras (12) 13.0

Cannpeche(7) 12.8

GuatennalaO) 12.8

Panama (17) 12.7

Venez.-NE(7) 12.7

Trinidad (5) 12.7

Venez-lnterior(4) 12.6 1

Guyana (11) 12.6

ElSalvador(15) 126

Venez.-Coastl(5) 12.5

Venez.- Coast 2(6) 12.5

Colombia (8) 12.4

Yucatan (6) 124

Locality Mean

Tamaulipas2(37) 31.6
Tamaulipas I (15) 31.5
Veracruz-N(8) 31.2
San Luis Potosi(l8) 31.2
Costa Rica (5) 30.2
Veracruz-C(8) 30.1
Nicaragua(CaribX6)30.0
Veracruz-S(12) 29.9
Honduras (1 2) 297
Panama (17) 29.6

Trinidad (6) 29.5

Chiapas(9) 294

Guatemala(9) 29.2
ElSalvador(l6) 290
Campeche(8) 29.0
Colombia (9) 28.7

Venez.-Coast2(6) 28.7
Guyana (13) 28.7

Venez.- lnterior(5) 28.2
Venez.-NE(7) 28.0
Venez.- Coast I (8) 27.7
Yucatan (6) 27.1

Fig. 13. SS-STP analy.si.s .showing geographic \ ariation in foR-ann
and in greatest lengtli of skull (right) of Rhogeessa tumida.

(left;

34

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

SYSTEMATICS OF THE GENUS RHOGEESSA 35

between bases and tips. Specimens from the Yucatan Peninsula,
Chiapas, Caribbean coast of Nicaragua and Panama, and Pacific
coast of Colombia tend to have bufBer bases and darker brown or
black tips, which contrast strongly with bases. Specimens from the
Pacific coast of Central America tend to resemble eastern Mexican
specimens in color. In South America, most of the color combina-
tions known from North America occur, but many redder and yel-
lower specimens also occur, with differing combinations of base
and tip color. Throughout the range, males tend to be darker than
females; however, this observation may represent an artifact, be-
cause many series of specimens have been collected during early
summer, when males have probably recently molted and females
have not.

Based on the data obtained in this studv, I have been unable to
divide the wide-ranging R. tumida into sympatric species, allopatric
species, or into subspecies, although one or more of these actions
ultimately may be necessary. The preliminary karyotypic studies of
R. J. Baker (pers. com.) indicate that animals having at least four
different karyotypes exist within the geographic range of R. tumida.
Two different karyotypes arc represented among animals from
southern Chiapas, and individuals from the Caribbean lowlands of
Nicaragua, and from Trinidad, differ karyotypically from specimens
collected in Pacific coastal Central America. At this point it is not
possible to evaluate these karyotypic trends in terms of the syste-
matics of R. tumida.

Reproduction ami molt. — Relatively few^ preparators of Rho-
g^eessa specimens have recorded data on reproductive condition.
More information is available on tumida than the other species, even
though much of it is negatix'c. Pregnant females were taken in May
in Tabasco. Nine bats caught in Guatemala, El SaK^ador, Honduras,
and Nicaragua were pregnant in the period from 14 February
through 28 April. Gravid females were taken in Costa Rica on 9
March, and in Panama on 25 February. A pregnant individual from
Venezuela was captured on 23 March, and another, possibly preg-
nant, on 12 October. Lactating females were collected on various
dates from 24 May to 20 June in Tamaulipas and 30 April to 12 June
in Veracruz. Others were taken on 12 June in Oaxaca, May in
Tabasco, 18 July in Nicaragua, and 29 March and 7 May in Costa
Rica. Bats recorded as not pregnant, or as not lactating, were caught
in most months of the year, but few of these were from the spring
or early summer months.

Based on these data, it appears that gestation, parturition, and
lactation extend from mid-February through mid-July. Obviously,
there is variation in dates for reproductive e^'ents among females.
To what extent this is geographically correlated cannot be deter-
mined, however. Of ten specimens for which the number of em-
bryos was recorded, three had but one embryo, and the remainder

36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

two. Juveniles were taken on 23 dates scattered during the period
21 May through 27 August. One Venezuelan juvenile was caught
on 4 October. Because this individual was captured at a locality
within 5°N of the equator, it seems possible that a somewhat dif-
ferent breeding cycle prevails at that latitude.

Males with enlarged testes (6-7) were taken during September,
November, December, and January. Bats from other months had
testes ranging in size from 2 to 5, suggesting that maximum produc-
tion of spermatozoa may occur during the autumn and early winter,
as in Temperate Zone vespertilionid bats. Five males were molting
in late May and June, and three females in July, suggesting co-
ordination with reproductive cycles.

Remarks. — Included in the synonymy are five names applied to
subspecies of either tumida or parvula by Goodwin (1958). Rho-
g,eessa io Thomas and R. f. riparia Goodwin are names applied to
Venezuelan specimens. There is little geographic variation in this
area, and, although the holotype of riparia is a larger individual
than the holotype of io, it would be pointless to apply these names
to subspecies. There is no sharp break in Panama or Colombia
which would tend to validate the recognition of a distinct South
American subspecies.

Rhogeessa velilla Thomas, based on a specimen from Ecuador,
may be applicable to a subspecies, but the specimens which might
show a pattern of geographic variation deserving of nomenclatural
recognition have not yet been collected. The holotype of velilla
does not differ in size from the holotype of io, and J. E. Hill (in
litt.) noted that they were almost identical. The holotype of R.
bomhyx, from the Pacific coastal lowlands of Colombia, is typical
of the large individuals which seem to occur along the Caribbean
versant of Central America (from Nicaragua southward) and the
west coast of Colombia. At present I choose not to recognize
homhijx as a species or subspecies because of the apparent clinal
transition from small pale individuals on the Pacific coast of Central
America to large, dark individuals on the Caribbean coast. The
substantial gaps in eastern Panama and Colombia suggest that
subsequent collecting may reveal a more accurate rendition of
variation in this region.

Rhogeessa parvula aeneus Goodwin is a name applied to small
individuals from Yucatan and northern Quintana Roo. According
to my notes "This skull [holotype of tumida] is almost identical to
AMNH 91234, holotype of flenef/.s . . . (although latter is smaller)."
The size of specimens from various localities in Campeche, southern
Quintana Roo, and Belice suggests a clinal change. I prefer not to
give nomenclatural recognition to \ari()us segminits of a clinc",
especially when the dividing line would be of necessity arbitrary.

SYSTEMATICS OF THE GENUS RHOGEESSA 37

Rhogeessa (Rhogeessa) minutilla G. S. Miller

Rhogeessa minutilla G. S. Miller, 1897:139 [Holohpe.— NMNH 63216 from

Margarita Island, Venezuela; Lt. Wirt Robinson, collector].
Rhogeessa parvula ininutilla — Goodwin 1958:7.

Holotyps. — National Museum of Natural Histoiy 63216, adult
male, skin and skull. This specimen was compared with 98 other
specimens of minutilla from Venezuela and Margarita in the NMNH,
as well as with 51 specimens of tumida from Venezuela and Trini-
dad. Measurements of holotype appear in table 3.

Description. — Essentially as in description of Venezuelan tumida.
Major differences will be noted under "Comparisons." Distal one
third of dorsal hairs Cinnamon Buff to near Sudan Brown; basal
two-thirds contrasting with tips. Light Buff in most specimens;
ventrally, Warm Buff to Light Ochraceous-Buff to Cinnamon Buff;
fur 4-5; sagittal crest present in 26 of 76 skulls.

Bacula. — Mean dimensions of eleven bacula from specimens of
minutilla (all from northwestern Venezuela) are: length. 0.50
(0.40-0.60); depth, 0.17 (0.12-0.22); width, 0.50 (0.40-0.62). As
compared to bacula of South America tumida, the bacula of minu-
tilla are shorter and narrower; lateral knobs extend at a higher angle
from shaft, and approach a right angle in some specimens (Figs.
2M and 2N). Some minutilla bacula r(\s(>mble closely those of some
tumida.

Comparisons. — Despite their great similaritx', there arc some
consistent differences between tumida and )ninutilla. The two spe-
cies are compared in tables 2 and 5.

Univariate analyses give poor separation of tumida and minutilla
in South America, but the multi\ariate analyses allow a somewhat
different perspective. The plot of the second and fourth principal
components ( Fig. 4 ) and of the first and second canonical variables
(Fig. 12) show remarkably good separation betvveen these two
species. The discriminant function table (Table 4) indicates that
42 of 62 specimens of minutilla were correctly identified. Ten speci-
mens of the 12 misidentified as tumida were identified as tumida

Table 5. A comparison of major characters which distinguish Rhogeessa
tumida and R. rninutiUa in northwestern Venezuela.

R. tumida

R. minutilla

Fur and membranes darker

"helmet" rarely present

POW greater

tibia shorter

heavy toothwear rare

tragus shorter

third metacarpal at least

1 mm shorter than forearm

fur and membranes paler
"helmet" usually present
POW less
tibia longer

heavy toothwear common
tragus longer

third metacarpal about 0.5 mm
shorter than forearm

38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

from Yucatan, and eight as parvula. Only two were misidentified
as tiimida from South America.

Distribution. — Arid lowhmds of Margarita Ishind, northwestern
Venezuehi, and adjoining portion of the Guajira Peninsuhi of Co-
lombia (Fig. 9) Rhogeessa minutilla occurs from sea level to 500 m.
Virtually all of this region is vegetated by desert scrub and thorn
forest. Although there appear to be minor differences in size and
color among the samples examined, there are no obvious trends
that might suggest geographic variation.

Reproduction and molt. — No pregnant females have been re-
corded; however, numerous specimens collected in June and July
were recorded as being not pregnant. Juveniles were taken from
late June through mid-July. Five males from June and July had
small testes, measuring 2-3. Molting females were taken on 30 June,
17 July, and 18 July; a single molting male was recorded from 16
July.

Remarks. — The type locality (Margarita Island) is approxi-
mately 500 km east of the main range of minutilla; its separation
by an area where only tumida seems to occur is a strangely anomo-
lous situation. Yet the three specimens from Margarita can be easily
matched by bats from northwestern Venezuela. It seems probable
that during a more arid (interglacial?) period the range of minu-
tilla was continuous along the northern coast of South America, and
reached at least one of the offshore islands. On the other hand, the
lowered sea levels characteristic of periods of glaciation may have
facilitated a crossing from the mainland to Margarita, or vice versa.
Arid habitat in the areas occupied by minutilla today is separated
by areas of more mesic vegetation. No specimens of minutilla seem
to have been taken in areas where tumida is common, but a few
specimens of tum,ida have come from within or very near the main
range of minutilla.

DISCUSSION

The multivariate statistical analyses ( utilizing only mensural
characters) were undertaken with the intent of illustrating the
degree to which dots representing sample means or individual speci-
mens would cluster in multidimensional space. I hoped these
analyses would give some clue to evolutionary (and thus systematic)
relationships, as well as demonstrate geographic variation.

The results of the multivariate analyses ( Figs. 4, 6, 7, 8, 12 and
14) were helpful, but hardly conclusive; in some cases results were
confusing or even contradictory. In general, they seem to indicate
that R. alleni is phenetically distant from the other species of
Rhogeessa, thus supporting my subgeneric separation of that species.
Although Rhogeessa gracilis and R. mira are closer plienetically to
the other species than is alleni, the degree of phenetic separation

SYSTEMATICS OF THE GENUS RHOGEESSA 39

varies among the difFcrent analyses. Rhogeessa s^racilis seems to be
closer to the others than R. mira. However, when one examines
characters not included in the statistical analyses, the reverse seems
to be true; i.e., mira is closely related to parvula, and gracilis seems
very distinct from all other species.

Rhogeessa parvula, R. tumida, and R. minutiUa, on the other
hand, are shown to be closely related by both the multivariate sta-
tistics and the other characters discussed in the species accounts.
Thus it appears that there are four closely related species, which
probably evolved by isolation from a common ancestor, plus two
peripheral species (alleni and gracilis) which were separated from
the common ancestor of the other four at a much earlier point
in time. It is also possible that the genus is polyphyletic and that
alleni, and even more likely, gracilis are similar to Rhogeessa only
by convergence. When gracilis is karyotyped, some light may be
shed on this question.

Because four species occur sympatrically in western Mexico, it
seems safe to assume that Rhogeessa evolved there from a primitive
vespertilionine bat, which perhaps also gave rise to Eptesiciis and
Pipistrellus. Th(\se latter two genera, however, are almost cosmo-
politan, and encompass many species. If we therefore* postulate a
much longer evolutionary history for them, then Rhogeessa is either
much younger as a genus, or has been blockc^d in its efforts to ex-
pand its distributional range and diversify into additional species
by such factors as competitive exclusion by other insectivorous spe-
cies, or by other evolutionary and zoogeographic problems.

Periodic climatic changes before and during the Pleistocene
probably led to periods of isolation between lowland populations
on the western and eastern coasts of Mexico, making possible the
occurrence of separate but similar species (tumida and parvula)
on the coasts. The eastern population spread southward throughout
Central America and northern South America, finding climates much
like those in eastern Mexico. In order for speciation to have oc-
curred in northwestern Venezuela, producing R. mimitilla, ances-
tral R. tumida must have reached South America before or during
the Pleistocene, so that the climatic changes necessary to isolate
a population there could have taken place. Presently, R. tumida
may be in the process of invading central and southern South
America, where it seems to be rare and local in distribution.

In western Mexico the lowland population must have been
broken into two segments at one point in time, to allow speciation
of R. mira from R. parvula. How and when this might have oc-
curred is not clear. Because alleni and gracilis normally are found
at higher elevations than parvula or mira, it can be speculated that
geographic isolation of different mountain masses and associated
contrasts in climate and vegetation led to their speciation at an

40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

early date, and tliat these same factors have maintained the present
restricted distributional ranges of these two species.

Figures 4, 5, and 14 tell us something about multidimensional
geographic variation. Because the dots are not in a straight line,
figure 4 demonstrates that the rate of clinal change differs among
variables, and figure 14 shows that the rate of clinal change differs
among samples in different portions of the species' geographic
ranges. Figure 4 shows increasing size from upper right to lower
left, and figure 5 groups samples of similar size.

KEY TO THE SPECIES OF RHOGEESSA

1. Greatest length of skull more than 14.5; i3 unicuspid, 1/6
to 1/8 cross-sectional area of i2; ratio of third metacarpal to

first phalanx of third digit averaging 2.20 to 1

Subgenus Baeoclon; R. olleni

Greatest length of skull less than 14.5; i3 often bicuspid,
having cross-sectional area greater than 1/6 that of i2; aver-
age ratio of third metacarpal to first phalanx of third digit
more than 2.20 to 1 Subgenus Rhogeessa, 2

2. Ears averaging 18; dorsal fur obviously 3-banded, darker at
bases R. gracilis

Ears averaging 15 or less; dorsal fur 2-banded, paler at
bases 3

3. Greatest length of skull averaging 11.4; lingual cingulum

of CI smooth, lacking cusps R. mira

Greatest length of skull averaging 11.9 to 13.5; lingual cin-
gulum of CI not smooth, usually with cusps 4

4. Uropatagium sparsely to heavily furred from base to point
halfway from knees to foot; i3 usually much smaller than

i2 R. pawiila

Uropatagium furred only at base; i3 usually only slightly
smaller than i2 5

5. Color pale; helmet usuall)- present; POW averaging 2.9 or
3.0; third metacarpal averaging 0.5 mm shorter than fore-
arm R. minutiUa

Color dark; helmet usually absent; POW averaging 3.1 or
3.2; third metacarpal at least 1 mm shorter than forearm .,
R. ftim id a

SUMMARY

More than 650 specimens of bats of the Neotropical genera
Rhogeessa and Baeodon were examined to determine the number
of genera and species that should be recognized, as well as to evalu-

SYSTEMATICS OF THE GENUS RHOGEESSA 41

ate geographic xariation within each species. In addition to the
usual morphological characters, large numbers of bacula were meas-
ured and compared, and hairs were examined by scanning electron
microscopy. The preliminary results of a study of karyotypic varia-
tion in RJwgeessa by R. J. Baker and associates were taken into
account. Both multivariate and unixariate statistical methods were
utilized.

One species, Rho<^eessa alleni, is assigned to the subgenus Baeo-
dun, and is a large-skulled form restricted to western Mexico. The
remaining five species are assigned to the subgenus Rho<ieessa.
Rhogeessa parvuhi, restricted to western Mexico, is characterized
by a hairy uropatagium. Rhogeessa gracilis, also restricted to west-
ern Mexico, possesses much larger ears than do the other species.
A new species, R. mira, is presently known only from two localities
in Michoacan, and is substantially smaller than the other three
species inhabiting western Mexico. Rhogeessa tumida, an extremely
variable (and perhaps composite) species, ranges from Tamaulipas
to Brazil and Bolivia. Rhogeessa minutilkh a pale desert species, is
restricted to Margarita Island, northwestern Venezuela, and the
adjoining Guajira Peninsula of Colombia.

RESUMEN

Con el fin de determinar el numero de generos y especies que
deben ser reconocidos y tambien analizar la xariacion dentro de
cada especie, se examinaron mas de 650 especimenes de murcielagos
neotropicales do los generos Rhogeessa y Baeodon. Ademas de los
caracteres morfologicos usualmente empleados, sc midieron y com-
pararon los baculos, y se examinaron los pelos por medio de un
microscopio electronico de "scanning." Fueron tomados ademas
en consideracion, el estudio hecho por R. J. Baker y companeros,
sobre la variacion en los kariotipos de Rhogeessa; y finalmente para
el analisis estadistico se uso los metodos de univarianza y multi-
varianza.

Una de las especies, Rhogeessa alleni, una forma de craneo
grande, restringida a el Occidente de Mexico, es asignada al sub-
genero Baeodon. Las otras cinco especies son asignadas a el sub-
genero Rhogeessa. Rhogees.sa parvida, restringida a Mexico occi-
dental, esta caracterizada por tener un uropatagio cubierto de pelo.
Rhogeessa gracd's, tambien restringida a Mexico occidental, tiene
orejas que son mas grandes que las de las otras especies. Una
especie nueva, R. mira, conocida solamente de dos localidades de
Michoacan, es la mas pequena de las quatro especies en Mexico
occidental. Rhogeessa tumida, una especie extremadamente vari-
able y posiblemente un "composite species" (compuesta de mas
de una especie), se c>\tiende desde Tamaulipas hasta Brasil y
Bolivia. Rhogeessa minutilla, una especie de color palido, encon-

42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

trada en areas deserticas, es restringida a la Lsla Margarita, el
Noroeste de Venezuela, y la peninsula de la Guajira de Colombia
y Venezuela.

SPECIMENS EXAMINED

For each species, countries are listed from north to south, and
from west to east. States, departments and provinces are given in
the same sequence following each country, and individual localities
are listed in the same manner following each state, department or
province. Localities which have not been precisely located on a
map are placed in (quotation marks. Abljreviations for museums
given with specimen numbers are those listed in the Acknowledg-
ments section of this paper. Localities from which specimens were
examined are represented on the distribution maps only insofar
as space permits. Many map spots cover several (as many as five)
localities.

Rhogeessa allcni

MEXICO: Jalisco: Piedra Gorda, ca. 8 km NW Soyatlan del Oro, 1600
m, UA 10292. Michoacdn: 20 km N El Infiernillo, UNAM 8597; 7 km N El
Infiernillo, ca. 125 m, KU 97307. Puehla: 10 mi W Acatlan, 6000 ft, TCWC
8480. Oaxaca: 2 mi NNW Tamazulapan [=Tamazulapam], 1990 m, KU
61170-1; Cuicatlan, 590 m, KU 29439; 2 mi N, 6 mi W Nejapa, KU 68773.

Rhogeessa gracilis

MEXICO: Jalisco: 5 mi NE Hiieiuquilla, 6200 ft, KU 108976; 10 mi
SE Talpa de Allende, 5350 ft, KU 97050; 17 km SE Talpa de Allende, 5200
ft, KU 92951. Piiebla: Piaxda, about 1100 m, NMNH 70691, 70694. Oaxaca:
Valero Taujano, 2 mi W Tomallim [=Tomellin?], MVZ 78315-6; Ceiro San
Felipe, San Felipe del Agiia, about 1700 m, AMNH 186901; "Istbmus of
Tehauntepec," NMNH 38294.

Rhogeessa parvula

MEXICO: Sonora: Estero Tastiota, sea level, MVZ 85290; 28 mi E
Mazatan, 500 m, UNM 18587; W Side Alamos, KU 24853-4; near Alamos,
UA 2717; Casa Las Delicias, Alamos, UA 2969; La Aduana, LACM 1.3247;
8 mi (by road) S Rio Alamos, Alamos, about 200 m, UA 16956, 16981-3;
11.3 mi SSE Alamos, Rio Cuchijaga, UA 14270; "Rancho Guirocoba," LACM
9700. Sinaloa: 1 mi S El Cajon, 1800 ft, KU 100401; 16 km NNE Choix.
1700 ft, KU 90752, 90754; 1 mi S, 6 mi E El Carrizo, KU 105563; 7 mi
ESE Sanalona, 600 ft, KU 100402; La Cruz, 30 ft, KU 90755, 90757; Elota,
sea level, UA 6358, LACM 19011; Vz mi E Piaxtla, sea level, KU 61163-8; km
marker 1289 on Mex. hwy. 15, 9 km W Rio Piaxtla, sea level, UA 15991-3,
17189-90; 14 mi N Mazatlan, sea level, UA 13477, 13978, LSU 10431-3;
Punta de Cauca, 11 mi N, 2.5 mi E Mazatlan, LACM 12498-500; Mazatlan,
LACM 16531; Chupaderos, on Rio Panuco, about 26 mi NE Villa Union,
800 ft, UA 9725, LACM 19080-1; 2 mi E Palmito, KU 97078-81; 5 mi NE
Concordia on Mex. hwy. 40, 100 m, UA 11059; 12 km N Villa Union, 400
ft, KU 95878; Escuinapa, 100 ft, TCWC 14471; 5 mi WSW Plomosas, 800
ft, KU 97082, 97084. Durango: Santa Ana, 12 km E Cosala, Sinaloa, 1300
ft, KU 90759. Nayarit: Huajicori, Rio del Bajar, 120 m, UA 9724, LACM
19079; 4 km S Playa Novilleros, sea level, UA 11037; a])out 40 mi E Acaponeta,

SYSTEMATICS OF THE GENUS RHOGEESSA 43

La Cuchara, LACM 13821; Ties Marias Islands, San Juanito Island, AMNH
180543-4; Tres Marias Islands, Maria Madre Island, AMNH 180522, NMNH
92413; "Tres Marias Islands," NMNH 84021, ANSP 1832; 5 mi NE San
Bias, }i mi N Singaita, sea level, UA 18552-3; 8 mi E San Bias, about 200
m, UA 8278-9; LACM 12503-5, 12507, 13244-6, 13436-41; % mi E San
Bias, 10 ft, KU 39723-5; 4 mi S, 5 mi E San Bias, UNM 16858. Jalisco:

9 mi N Guadalajara, 4000 ft, KU 107493-4; 2 mi S La Cuesta, 1500 ft, KU
111613-5; Piedia Gorda, about 8 km NW Soyatlan del Oro, about 1600 m,
UA 10294; El Salitre, about 4 km N Soyatlan del Oro, about 1600 m, UA
10590, 10602; Rio de Aguacate, about 4 km E Soyatlan del Oro, about 1300
m, UA 10307; about 4 km S Soyatlan del Oro, about 1500 m, UA 10319-20;
Cuitzamala, 25 ft, KU 105564; 2 mi N Tenacatita, 25 ft, KU 105565-6; 10
mi NNE Pihuamo, 3500 ft, KU 108974-5. Colima: Rancho Tavernillas,
about 35 km NW Pueblo Juarez, 350 m, UA 8838, LACM 19082; Tabamillas,
6 km N Agua Zarca, UNAM 6321-3; Puelilo Juarez, 330 m, UNAM 5643;
2 km N Tlapeixtes, near Manzanillo, sea le\el, UA 10676; 1 km N Tlapeixtes,
near Manzanillo, sea level, UA 10686; Colima, about 500 m, NMNH 52065-6,
52102; "4 mi S Cerro de Ortega," UA 3299; 2 mi E Cuyutlan, LACM 11271-7;
5 mi SE Armeria, LACM 11729-30. Michoacan: 20 km N El Infiemillo,
UNAM 8592; 7 km N El InBernillo, ENCB 990-7. Guerrero: "Guerrero,"
NMNH 187713; 4.3 km N Teloloapan, 1480 m, UNAM 8864-7; Ojo de Agua,
3.4 km N Mexicapan, 1480 m, UNAM 9984-5, 10684-5; El Ojo de Agua de
Chapa [7 km SSE Teloloapan], UNAM 1990-2; 8 km SW Teloloapan, 1300
m, UNAM 11869; Agua del Obispo, 3300 ft, TCWC 6351; Xaltianguis, about
500 m, NMNH 269315-6. Morclos: Rio Oaxtepec, 1 km S Oaxtepec, 890
m, UNAM 9706. Oaxaca: Santo Domingo, NMNH 73269; San Carlos Yau-
tepec, San Bartolo Yautepec, 800 m, AMNH 167467, 175263; 9 mi NW
Tehauntepec, near sea level, UNM 27545; Tehauntepec, sea level, AMNH
178744; 8 km NW Salina Cruz, ENCB 3487-8; 20 mi W Tapanatepec, sea
level, UA 13440-2; Rio Ostuta, 4 mi W Zanatepec, sea level, AMNH 186406.

Rhogeessa mira

MEXICO: Michoacan: 20 km N El Infiernillo, UNAM 8593-6; 7 km
N El Ineernillo, ENCB 998-1007.

Rlwgeessa tumida

MfiXICO: Tamaulipas: Santa Maria, 870 m, AMNH 148196; 4 mi N
La Pesca, sea level, KU 55192; 3 mi N La Pesca, .sea level, KU 55191,
55193, 55208; 2 mi N La Pesca, sea level, KU 55198-208; 1 mi N La Pesca,
KU 55194-7; 2 mi S. 10 mi W Piedra, Sierra de Tamaulipas, 1200 ft, KU
55152-63; 55165-89; 3 mi S, 16 mi W Piedra, Sierra de Tamaulipas, 1400
ft, KU 55190; 30 mi N El Mante, Rio Cielito, TCWC 25679. San Luis Potosi:

10 mi WSW Ebano, LSU 4047; 19 km SW Ebano, LSU 4942-53; 3 km N
Taninul, 650 ft, LSU 4936-41. Veracruz: 25 mi W Tampico, KU 82922;
El Hi go, 6 km NNW Ejido El Chote, about 75 m, UNAM 11063; 12.5 mi
N Tihuatlan, 300 ft, KU 88427-34; Boca del Rio, 10 ft, KU 29886; Rio Blanco,
20 km W Piedras Negras, 400 ft, KU 19231; 24 mi S Veracniz, near sea
level, AMNH 203917; San Andres Tuxtla, 360 m, UNAM 9485; 9 mi ENE
Catemaco, ca. 200 m, UNAM 7751-2; 24 mi S Santiago Tuxtla, Los Tuxtlas,
TCWC 9488; Mirador, sea level, NMNH 37329 (skull) + ANSP 1831 (skin
of NMNH 37329); Achotal, FMNH 14149; 2.3 km W, 3 km SSW, 2.5 km SW,
and 4.1 km S Tenochtitlan, 50 m, UM 116286-91, 116293-98. Oaxaca: 3 km
W Estacion Vicente, Municipio de Acatlan, 60 m, UNAM 11492-3. Tabasco:
Rancho El Tumbo, 4 km E F.F.C.C. El Zapote, Macuspana, UNAM 1717,
1959-60. Chiapas: Rancho San Fernando, 42 km W Cintalapa, 500 m, UA
15711; 32 mi SW Cintalapa, Rancho San Miguel, TTU 11274-5; 2 mi SW

44 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Las Cruces, about 600 m, KU 61169; Finca San Salvador, 15 km SE San
Clemente, 1000 ni, KU 102619; 11.9 mi SE Ties Picos, Puente El Mosquito,
TTU 11354-5; Pijijiapan, 10 m, UM 96523-4; 5 km SE Pijijiapan, 100 ft,
TCWC 14466-9; Prusia, 1000 m, UM 88196-8; Huehuetan, 35 m, NMNH
78600; 7 mi ENE Tapachula, about 300 m, KU 68772; 3.8 mi SW Tapacbula,
TTU 11133; 14 km SW Tapachula, 50 m, KU 120535. Yucatan: 10 mi W
Progresso, LACM 18167; Piste, 10 m, KU 92000; Chichen-Itza, AMNH
91234, UNAM 1808, MCZ 10807, 32868, UM 79938. Campeche: 5 km S
Champoton, 10 m, KU 92007-9; La Tuxpena, NMNH 170858; Isla del Carmen,
1 km S Puerto Real, 3 m, KU 92010; Balchacaj [=Balchacah?], Lagima de
Terminos, FMNH 47394; 65 km S, 128 km E Escarcega, KU 93536. Quiniaua
Roo: 4 km WSW Puerto Juarez, 5 m, KU 92001; Pueblo Nuevo X-Can, 10
m, KU 92002-3; Isla Cozumel, 4 km N San Miguel, KU 92004-5; 4 km NNE
Felipe Carrillo Puerto, 30 m, KU 92006.

GUATEMALA: "Moca Guatalon," MCZ 28144. El Peten: La Libertad,
170 m, AMNH 144697. San Marcos: Finca [El] Porvenir, FMNH 50063.
Quetzaltenango: Finca Los Pirineos, near Santa Maria de Jesus, ca. 1500 m,
FMNH 50062. Suchitepeqiiez: Moca, about 1000 m, FMNH 41649, 41828-30,
41856-7. Santa Rosa: Finca Santa Isabel, about 200 m, FMNH 74012; Asti-
llero, 25 ft, KU 64992-3.

BELICE: Belize: Turneffe Island, Calabash Cay, LSU 7148.

HONDURAS: "Patuca River," NMNH 36062. Cortez: 23 mi bv road
N San Pedro Sula, ca. sea level, TTU 13297. Yoro: Portillo Grande, 4100
ft, MCZ 33869. Copdn: Copan, 660 m, TCWC 19752. Santa Barbara: 12
km N Santa Barbara, TTU 13295-6; 7 mi N Santa Barbara, 120 m, TCWC
19753-5. Olancho: 10.3 mi by road SSW Dulci Nombre de Culmi, TTU
13298, 13300-1. Comayapna: 3 km W Comayagua, 580 m, TCWC 21106-7.
Distrito Central: La Flor Archaga, ca. 800 m, MCZ 28976; Comayaguela,
1000 m, MCZ 28975. La Paz: El Pedrero, 3000 ft, AMNH 126901-4. Fran-
cisco Morazdn: 2 mi S El Zamorana, ca. 1200 m, TCWC 11017; Escue'a
Agricola Panamericana, MCZ 45401. El Paraiso: Chichicaste, 480 m, TCWC
22108.

EL SALVADOR: Chalatenango: San lose del Sacare, 3600 ft, M\^
13099S; 20 km W Chalatenango, 250 m, TCWC 19751. Cuscatldn: Colima,
MVZ 1.30990-1. Usulatan: Puerto del Triunfo, sea level, MW 130994-5. San
Miguel: Rio San Miguel, 13° 25' N, 225 ft, MVZ 130991-3.

NICARAGUA: Matagalpa: 6 km N Tuma, 550 m, TCWC 24126-7;
Uluce [=Uluse?], AMNH 29863; 1 km NE Esquipulas, 420 m, KU 115148.
Chinandega: 6.5 km N, 1 km E Cosiguina, 10 m, KU 115145-7; Hda. Bella-
\ista, 720 m, Volcan Casita, KU 106284; Chinandega, about 20 m, AMNH
28996. Boaco: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, KU 111233-4.
Carazo: 3 km N, 4 km W Diriamba, 600 m, KU 11123.5-6. Zelai/a: 10 km
W Rama, 40 m, TCWC 19756; 4.5 km NW Rama, TTU 13313; 3 km NW
Rama, TTU 13317-8. Rivas: 6.9 mi E San Juan del Sur, TTU 13320.

COSTA RICA: "Pacific Coast," AMNH 4966. "Finca San Miguel,"
FMNH 67305-6. Alajuela: San Torte, between San Carlos and Altamira,
ROM 60464. Giianacaste: Samara [=Samari?], LACM 26669. Puntarenas:
Boca del Barranca, LACM 23767, 25258. San Jo.se: Rio Corrogres, about 2
km NW Santa Ana, 820 m, LSU 12989-93; Finca Lornessa, aliout 2 km NW
Santa Ana, 850 m, LSU 14750-7.

PANAMA: "Panama," MCZ 33525. Bocas del Toro: 7 km SSW Chan-
guinola, sea level, NMNH 315776; Almirante, sea level, NMNH 315774-5;
Isla Bastimentos, sea level, NMNH 335419. Chiriqui: Bugaba, alxiut 200
m, MCZ 10446; 8 m\ SE Progresso, sea le\el, NMNH 363107. Code: 3 mi
W Churubr, 300 ft, NMNH 331910; Santa Clara, sea level, NMNH 296265.
Los Santos: Guanico Arriba, about 200 m, NMNH 323605. Canal 7Ame:
Ft. Sherman, sea level, NMNH 396406; Summit Gardens, about 100 m.

SYSTEMATICS OF THE GENUS RHOGEESSA 45

NMNH 304930; San Pablo, NMNH 171212-3, 223330; Ciimndu, NMNH
309219; Ft. Kobbe, sea level, NMNH 309220; Ft. Kobbe Beach, sea level,
NMNH 312108-13, 313777-82; Ancon, MCZ 28100; Tapia, AMNH 140472.
Darien: La Palma de Darien, sea level, MVZ 113936.

COLOMBIA: Magdalene: "San Alberto," LACM 16781; Rio Guaimaral,
Valledupar, 168 m, NMNH 281310; Colonia Agricola Caracolicito, Santa
Malta, about 200 m, NMNH 281921. Bolivar: Catiral Upper Rio San Jorge,
FMNH 69526. Norte de Santander: 10 mi N Cueta [=Cucuta], FMNH
18725-6. Tolima: Honda, Las NLirgaritas, ROM 49045, 49048; Guamo, about
300 m, UAB 426. Cundinamarca: Mesitas de Colegio, 1200 ni, AMNH
207922, LACM 19064-5. Valle: Rio Raposo, about 100 m, NMNH 334744.
Htiila: 16 km NE Mllavieja, 1600 ft, M\'Z 113936.

VENEZUELA: Zulia: 30 mi E Maracaibo, sea level, NMNH 260191;
Perija, Rio Cogollo, sea level, FMNH 21984; 48 km WNW Encontrados, 54
m, NMNH 441790; 38 km WNW Encontrados, 37 m, NMNH 441791. Trujillo:
23 km NW Valera (nr. Agua Santa), 90 m, NMNH 372488. Merida: Sta.
Elena, Rio Guachi, FMNH 21985. Yaracmj: 8 km N, 18 km W San Felipe,
near Minas de Aroa, 400 m, NMNH 441773-5; 19 km NW Urama, km 40,
100 m, N\L\H 372487, 372489, 374016-8. Aragua: nr. Rancho Grande,
sea level, NNLNH COH7876-7. Miranda: 5 km S, 5 km E Caracas, Encan-
tados, 570 m, NMNH 441776; 1 km E Rio Chico, sea level, NMNH 387738;
5 km E Rio Chico, nr. Puerto Tuy, sea level, NMNH 387736-7. Sucre:
Takal, 11 km SSW Cumana, al^out 200 m, KU 119074; 2.5 km SW Cumana,
5 m, KU 119072; Cuchivano, 700 ft, ANLXH 69968; 4 km S, 25 km E Caru-
pano (nr. Manacal), 170 m, NMNH 409487. Giidrico: Calabozo, Est. Biol.,
NMNH UCV4871. Monaga.s: Jusepin, about 100 m, KU 119073; 54 km SE
Maturin, Mata de Bejuco, 18 m, NMNH 441792-4; 60 km SE Maturin, LACM
14355. Apitre: 60 km NE Pto. Paez, Hato Cariben, La \'illa, 76 m, NMNH
374019-20; 8 km NW Pto. Paez, Cerro de los Murcielagos, 76 m, NMNH
374021. Amazonas: 65 km SSW Puerto Ayacucho, Morocoy, 161 m, NMNH
409488.

TRINIDAD: "Trinidad," NMNH 141885. St. George: Maracas Valley,
TTU 5409, 5457; Maracas, 500 ft, AMNH 183163; Maracas Valley, W^aterfalls
Rd., about 1100 ft, AMXH 176379, ROM 314.38-9, 31453; Port of Spain,
sea level, FMNH 51157-8. S^ Patrick: San Rafael Rancho Estate, Santa
Maria [=St. Mary's?], TTU 5257; Siparia, aliout .sea le\el, AMNH 179965-6,
182924, 183861.

GUYANA: Rupununi: Dadanawa House, ROM 32348; Cotanrili. Isl., 5
mi above Dadanawa, left Inink Rupunimi Ri\er, ROM 59709-10; nr. Shea
v. Kuitaro, ROM 43093; Tacatu R. area, 40 mi SW Dadanawa, ROM 58843;
(remaining localities not precisely located, but near four prc\iously listed
localities)' Ikoui Tau, ROM 40764, 40766-7; Kuitaro River, ROM ' .32783,
46211; Raa Wau, ROM 40798; Tamtoon, ROM 36842; Weri More, ROM 44527.

ECUADOR: Puna: Puna Island, San Ramon, sea level, AMNH 66824.

BRASIL (not mapped): Maranhao: Alto Parnahyha? [Alto Parnaiba,
9° 06' S, 45° 57' W?], FMNH 26465. Mato Grosso: 264 km (bv road) N
Xavatina, Serra do Roncador, 12° 51' S, 51° 46' W, 1750 ft, NMNH 393759.

Rhogeessa minutilla

COLOMBIA: Guajira: 119 km N, 32 km W Maracaibo, Venezuela, 15 m,
NMNH 441781-2.

\T:NEZUELA: Zulia: 114 km N, 32 km W Maracaibo, 12 m, NMNH
441777; 114 km N, 28 km W Maracailx), 15 m, NMNH 441778-80, 441783-9;
Rio Aurare, sea level, FMNH 18711; Empeladi Savanna, FMNH 18742-5.
Falcon: Capatarida, 55 m, NMNH 441795-9, 441801, 441819-36; 6 km SSW
Capatarida, Santa Rosa, NMNH 441771-2, 441800, 441802-13, 441815-18.

46 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Lara: Rio Tocuyo, 500 m, AMNH 130671-2, 130674-6, 130711-4, 131192;
10 km N El Tocuyo, Casen'o Boro, 528 m, NMNH 443183, 443186, 455992,
455994, 455997-9, 456000-1, 456009-15, 456020-24, 456028-9, 456031-2,
456035-7, 456039. Margarita Island: "Margarita Island," NMNH 63216,
113497; Nueva Esparta, 2 km N, 30 km W Porlamar (nr. Teatas de Maria
Guevara), 10 m, NMNH 405828.

APPENDIX 1

To plot unknown specimens on any of the three canonical scatter-
grams (Figs. 7, 8, and 12), first make the nine standard measure-
ments outlined in the section on materials and methods, and substi-
tute into the following formulae:

Xaxis ( First canonical variable ) = (-0.46) ( FA-29.04) - 0.16 (3MC-28.28)
+ 0.47(GLS-12.90) - 0.94(DB-4.66) - 2.09(POW-3.14) +
0.05(MW-6.78) - 1.44(M2M2-5.20) - 0.21 (ClCl-3.66) -
1.64(MAX-4.48).

Yaxis (Second canonical variable) = 0.08 ( FA-29.04) + 0.42 (3MC-28.28)
+ 0.58(GLS-12.90) - 0.11(DB-4.66) + 5.59(POW-3.14) -
1.47(MW-6.78) + 0.95(M2M2-5.20) -0.42 (ClCl-3.66) -
5.12(MAX-4.48).

The two coordinates thereby obtained can then be plotted on the
appropriate scattergram in the usual manner.

LITERATURE CITED

Allen, H.

1866. Notes on the Vespertilionidae of tropical America. Proc. Acad. Nat.
Sci. Philadelphia, 18:279-288.
Alvarez, T. and C. E. Avixa

1965. Baedon [sic] alleni, Rhogcessa tumida major and R. p. parvula
newly reported for Michoacan, with notes on the qualitatixe dif-
ferentiation of the two rhogeessas. Southwest. Nat., 10:75-76.
Anderson, S.

1960. The baculimi in microtine rodents. Univ. Kansas Publ. Mns. Nat.
Hist., 12:181-216.
Baker, R. J. and J. L. Patton

1967. Karyotypes and karyotypic \ariation of North American vesper-
tilionid bats. J. Mamm., 48:270-286.
Benedict, F. A.

1957. Hair structure as a generic character in bats. Univ. California
Publ. Zool., 59:285-548.

Blackith, R. E. and R. A. Reyment

1971. Multivariate morphometries. Academic Press, London, vii-|-412 pp.
Brown, R. E., H. H. Gexoways and J. K. Jones, Jr.

1971. Bacula of .some Neotropical bats. Mammalia, 35:456-464.
DoH-soN, G. E.

1878. Catalogue of the Chiroptera in the . . . British Museum. London,
xlii-f-567 pp.
Goodwin, G. G.

1958. Bats of the genus Rhogcessa. Amer. Mus. Novitates, 1923:1-17.

SYSTEMATICS OF THE GENUS RIIOGEESSA 47

Goodwin, G. G. and A. M. Gheenhall

1961. A review of the bats of Trinidad and Tobago. Bull. Amer. Mus.
Nat. Hist., 122:187-302.
Hall, E. R.

1952. Taxonomic notes on Mexican bats of the genus Rhogeessa. Univ.
Kansas Publ. Mus. Nat. Hist., 5:227-232.
Hall, E. R. and K. R. Kelsox

1959. The mammals of Nortli America. Ronald Press, New York, l:xxx+
546+79 pp.
Hamilton-, W. J., Jr.

1949. The bacula of some North American vespertilionid bats. J. Mamm.,
30:97-102.
Handley, C. O., Jr.

1959. A revision of American bats of the genera Euderma and Plecotus.
Proc. U. S. Nat. Mus., 110:95-246.
JoxEs, J. K., Jr., J. D. Smith and R. W. Turner

1971. Noteworthy records of bats from Nicaragua, with a checklist of the
chiropteran fauna of the country. Occas. Pap., Mus. Nat. Hist.
Univ. Kansas, 2:1-35.
KoopMAx, K. F. and E. L. Cockrum

1967. Bats, pp. 109-150, in Recent mammals of the World, a synopsis
of families (S. Anderson and J. K. Jones, Jr., eds. ), Ronald Press,
New York, \'iii+453 pp.
LaVal, R. K.

1973. A revision of the Neotropical bats of the genus Myotis. Bull. Los
Angeles Co. Mus. Nat. Hist., 15:1-54.
Miller, G. S.

1897. Revision of the North American bats of the family Vespertilionidae.

N. Amer. Fauna, 13:1-135.
1897. Description of a new bat from Margarita Island, Venezuela. Proc.

Biol. Soc. Washington, 11:139.
1906. Twelve new genera of bats. IhicL, 19:83-86.
Pine, R. H., D. G. Garter and R. K. LaVal

1971. Status of Banerus Van Gelder and its relationships to other nyc-
tophiine bats. J. Mamm., .52:663-669.

Quay, W. B.

1970. Integument and derivatives. Pp. 1-56, in Biology of bats (W. A.
Wimsatt, ed.). Academic Press, New York, 2:\iii + 477 pp.
Ridgway, R.

1912. Color standards and color nomenclature. Pri\ately published,
Washington, D. C.

Simpsox. G. G.

1945. The principles of classification and a classification of mammals.
Bull. Amer. Mus. Nat. Hist., 85:1-350.
Smith, J. D.

1972. Systematics of the chiropteran family Mormoopidae. Misc. Publ.,
Mus. Nat. Hist. Univ. Kansas, 56:1-132.

Tate, G. H. H.

1942. Results of the Archbold Expeditions. No. 47. Review of the
vespertilionine bats, with special attention to genera and species
of the Archbold Collections. Bull. Amer. Mus. Nat. Hist., 80:
221-297.
Thomas, O.

1892. Description of a new Mexican bat. Ann. Mag. Nat. Hist., (6) 10:

477-478.
1903. Two South American forms of Rhogeessa. Ihicl, (7) 11:382-383.

1913. New mammals from South America. Ihid., (8) 12:567-574.

University of Kansas Publications
MUSEUM OF NATURAL HISTORY

The University of Kansas Publications, Museum of Natural
History, beginning with volume 1 in 1946, was discontinued
with volume 20 in 1971. Shorter research papers formerly pub-
lished in the above series are now published as Occasional
Papers, Museum of Natural History. The Miscellaneous Publica-
tions, Museum of Natural History, began with number 1 in 1946.
Longer research papers are published in that series. Monographs
of the Museum of Natural History were initiated in 1970. All
manuscripts are subject to critical review by intra- and extra-
mural specialists, final acceptance is at the discretion of the
publications committee.

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may obtain the Occasional Papers and Miscellaneous Publica-
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Editor: Linda Truer
Managing Editor: William E. Duellman

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