iS
HARVARD UNIVERSITY
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Library of the
Museum of
Comparative Zoology
(US ISSN 0027-4100)
The Systematics of Neotropical
Orb-weaving Spiders in the
Genus Metepeira (Araneae: Araneidae)
WILLIAM H. PIEL
(
MCZ
LIBRARY
JUL 3 2001
HARVARD
UNIVERSITY
HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS, U.S.A.
VOLUME 157, NUMBER 1
8 JUNE 2001
(US ISSN 0027-4100)
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THE SYSTEMATICS OF NEOTROPICAL ORB-WEAVING SPIDERS IN
THE GENUS METEPEIRA (ARANEAE: ARANEIDAE)
WILLIAM H. PIEU
CONTENTS
Abstract _ ___. 1
Introduction 2
Acknowledgments 2
Materials and Methods 3
Collections Examined 3
Locality Data Storage and Manipulation 4
Examination and Illustration 4
Metepeira F. O. P.-Cambridge 5
Key to Female Metepeira 12
Key to Male Metepeira 17
Metepeira foxi Group 19
1. Metepeira datona ChamberHn and Ivie 20
2. Metepeira desenderi Baert 21
3. Metepeira grandiosa grandiosa
Chamberlin and Ivie 23
4. Metepeira grandiosa alpina
Chamberlin and Ivie 24
Metepeira vigilax Group 26
5. Metepeira cajahamba New Species .... 26
6. Metepeira glornerabilis (Keyserling) ... 28
7. Metepeira vigilax (Keyserling) 30
8. Metepeira rectangula (Nicolet) 32
Metepeira labyrinthea Group 33
9. Metepeira spinipes F. O. P.-Cambridge .. 34
10. Metepeira lacandon New Species 37
Metepeira nigriventris Group 38
11. Metepeira nigriventris (Taczanowsld) 38
12. Metepeira tarapaca New Species 40
13. Metepeira calamuchita New Species .. 42
14. Metepeira galatheae (Thorell) 43
15. Metepeira karkii (Tullgren) 46
Metepeira conipsa Group 47
16. Metepeira compsa (Chamberlin) 48
17. Metepeira roraima New Species 53
18. Metepeira gressa (Keyserling) 54
Metepeira incrassata Group 56
19. Metepeira maija New Species 56
20. Metepeira inca New Species 58
' Museum of Comparative Zoology, Harvard Uni-
versity, Cambridge, Massachusetts 02138. Current
address: Institute of Evolutionaiy and Ecological Sci-
ences, Leiden University, 2311 GP Leiden, The
Netherlands; piel@rulsfb.leidenuniv.nl.
21. Metepeira gosoga Chamberlin and Ivie
59
22. Metepeira ohnec New Species 60
23. Metepeira comanche Levi 62
24. Metepeira pimungan New Species 62
25. Metepeira triangidaris (Franganillo) .. 63
26. Metepeira arizonica Chamberlin and
Ivie 66
27. Metepeira atascadero New Species .... 67
28. Metepeira incrassata F. O. P.-
Cambridge 68
Metepeira ventura Group 71
29. Metepeira ventura Chamberlin and
Ivie 71
30. Metepeira revillagigedo New Species 73
31. Metepeira celestun New Species 74
32. Metepeira uncata F. O. P.-Cambridge ... 76
33. Metepeira crassipes Chamberlin and
Ivie 77
34. Metepeira chilapae Chamberlin and
Ivie 78
Metepeira ininima Group 80
35. Metepeira petatlan New Species 80
36. Metepeira minima Gertsch 82
37. Metepeira pacifica New Species 84
38. Metepeira jamaicensis Archer 86
Literature Cited 88
Index 91
Abstract. Of the 39 species and three subspecies
of the orb-weaver genus Metepeira in the Americas,
36 species and two subspecies are known to occur
outside of the U.S. and Canada. Yet, despite their
conspicuous webs, diurnal foraging, and relatively
common presence, the taxonomy of Metepeira is
poorly understood, probably because the genitalia are
small and difficult to distinguish. In fact, many names
for species south of the U.S. were, at some time, in-
correctly synonymized with the name Metepeira la-
byrinthea. In this paper, 14 new species are named
(Metepeira atascadero, M. cajahamba, M. calamuchi-
ta, M. celestun, M. inca, M. lacandon, M. maya, M.
ohnec, M. pacifica, M. petatlan, M. pimungan, M. re-
villagigedo, M. roraima, M. tarapaca); 11 new junior
Bull. Mus. Comp. ZooL, 157(1): 1-92, June, 2001 1
2 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
synonyms are reported (M. acostai, M. bani, M. dom-
inicana, M. grinnelli, M. latigyna, M. perezi, M. san-
ta, M. salei, M. seditiosa, M. vaurieorum, M. virgi-
nensis); five cases of erroneously synonymized names
are reversed; 22 species and two subspecies are re-
described (M. arizonica, M. triangidaris, M. chilapae,
M. Comanche, M. compsa, M. crassipes, M. datona,
M. desenderi, M. galatheae, M. glo7nerabilis, M. go-
soga, M. grandiosa alpina, M. grandiosa grandiosa,
M. gressa, M. incrassata, M. jamaicensis, M. karkii,
M. minima, M. nigriventris, M. rectangula, M. spi-
nipes, M. uncata, M. ventura, M. vigilax); and a key
to all Metepeira species is presented. In addition, sev-
eral ecological and life history observations are re-
ported for various species.
INTRODUCTION
The absence of a comprehensive revi-
sion of Neotropical Metepeira has left the
taxonomy of this group in shambles. Over
the years, a fair number of species have
been named, particularly by A. F. Archer,
R. V. Chamberlin, and W. Ivie. However,
these efforts have been sporadic and, for
the most part, scant. For example, the de-
scription of Metepeira dominicana (Ar-
cher, 1965) provides little information oth-
er than "form typical of Metepeira in all
respects," a few measurements, and two
unrecognizable figures. Even when species
are properly described they have far less
taxonomic value when published alone, in
the absence of a full comparative revision.
The poor understanding of Metepeira
taxonomy has persisted despite great eco-
logical and behavioral interest in this ge-
nus. Indeed, many species are obligate or
facultative social species and offer excel-
lent models for investigating genetic and
environmental factors that influence colo-
ny formation (e.g., Uetz and Cangialosi,
1986; Uetz et al, 1987). The monumental
work carried out over many years by G. W.
Uetz has made great strides in our under-
standing of gregarious social behavior in
spiders and in risk-sensitive foraging the-
ory in general (e.g., Uetz, 1996). Still, in
the absence of solid taxonomic literature,
behavioral ecologists have been forced to
apply informal names to their study ani-
mals (e.g., Metepeira "atascadero" in Uetz
[1989] or Metepeira "Species A" in Viera
[1989]), but this practice can lead to trou-
ble. In one case, the behavior of several
different species was initially studied un-
der the false assumption that they all be-
longed to the same species (e.g., Uetz et
al., 1982). Clearly, a strong taxonomic
foundation is important for further biolog-
ical work.
Ultimately, the relatively small, indis-
tinct genitalia and the relatively homoge-
neous abdominal patterns are to blame for
the weakness in our knowledge of Mete-
peira taxonomy. Many of these species are
undoubtedly hard to distinguish, and this
fact has surely intimidated arachnologists
from taking on the painful task of revising
the group. In the absence of good distin-
guishing characteristics, the catalogs of
Bonnet (1957) and Roewer (1942) synon-
ymized the names of many Neotropical
species with the name Metepeira lahyrin-
thea. Levi's (1977) revision of Nearctic
species observes that M. labyrinthea is ac-
tually limited to the eastern United States.
One task in this revision consists of reas-
serting the names of species that were im-
properly synonymized and clarifying the
diagnostic characters that are needed to
identify them.
ACKNOWLEDGMENTS i
This paper is part of my Ph.D. thesis for
the Department of Organismic and Evo-
lutionary Biology, Harvard University. I ain
indebted to many people for their help,
assistance, and encouragement in this pro-
ject. I am especially thankful for the ded-
ication and support of my advisors, Her-
bert W. Levi and Edward O. Wilson. I am
grateful that my colleagues in the Depart-
ment of Invertebrate Zoology provided
such a pleasant place to work: Edward
Cutler, Ardis Johnston, Laura Leibensper-
ger, Damhnait McHugh, Diana Sherry,
Van Wallach, and Dee Woessner, among
others.
Field collecting and new specimen ac-
quisitions were made possible with the
help of Gita Bodner, Fundacion Capacitar,
Tim Coonan (CINP), Fred Coyle, Dawn
Metepeira • Piel
Fitzpatrick, Germania Jacome, Antdnia
Monteiro, Tila Perez, George Putnam,
Linda Rayor, Grace Smith (NAWF), and
George Uetz. I am particularly indebted to
George Uetz for his assistance and corre-
spondence.
I am thankful for the comments by
those who read this paper — especially to
the members on my thesis committee: H.
W. Levi, N. E. Pierce, and E. O. Wilson.
I am also indebted to Kathy Horton for
her help in formatting and preparing the
manuscript and to the Colles Fund for de-
fraying the costs of publication. Curators
at various institutions who lent me speci-
mens are listed in the Materials and Meth-
ods section. I cannot overstress the value
of museum collections and expert curators,
without which research in taxonomy would
not be possible. Museum collections are
the most important tools available for un-
derstanding biodiversity.
MATERIALS AND METHODS
Collections Examined. The taxonomic
revision was carried out on specimens bor-
rowed from the following collections. The
abbreviations correspond to those listed
with each record after eveiy species de-
scription. I am grateful to the museums,
curators, and staff that graciously loaned
the material.
ADC A. Dean, Texas A&M University,
College Station, Texas, United
States
AMNH American Museum of Natural
History, New York, United
States; N. Platnick, L. Sorkin
BMNH Natural History Museum, Lon-
don, England; P. Hillyard
CAS California Academy of Sciences,
San Francisco, California, Unit-
ed States; C. Griswold
CV Carlos Valderrama A.; Bogota,
Colombia
FSCA Florida State Collection of Ar-
thropods, Gainesville, Florida,
United States; G. B. Edwards
IRSNB Institut Royal des Sciences Na-
turelles de Belgique, Brussels,
Belgium; L. Baert
JAK J. A. Kochalka, Ciudad Univer-
sitaria, Paraguay
JEC J. Carico, Lynchburg, Virginia,
United States
JMM J. Maes, Leon, Nicaragua
MACN Museo Argentino de Ciencias
Naturales, Buenos Aires, Argen-
tina; E. A. Maury, C. L. Scioscia
MCN Museu de Ciencias Naturais,
Fundagao Zoobotanica do Rio
Grande do Sul, Porto Alegre,
Rio Grande do Sul, Brazil; E. H.
Buckup, M. A. L. Marques
MCZ Museum of Coniparative Zool-
ogy, Harvard University, Cam-
bridge, Massachusetts, United
States; H. W. Levi
MECN Museo Ecuatoriano de Ciencias
Naturales, Quito, Ecuador; Ger-
mania Estevez Jacome
MEG M. E. Galiano, Buenos Aires,
Argentina
MLJC Maria Luisa Jimenez, Centro de
Investigaciones Bioldgicas del
Noroeste, La Paz, Mexico
MLP Museo de Universidad Nacional,
La Plata, Argentina; R. F. Arro-
zpide, C. Sutton
MNRJ Museu Nacional, Rio de Janeiro,
Brazil; A. Timotheo da Costa
MNSD Museo Nacional de Historia
Natural, Santo Domingo, Re-
publica Dominicana; Felix Del
Monte
MUSM Museo de Historia Natural,
Universidad Nacional Mayor de
San Marcos, Lima, Peru; D. Silva
MZSP Museu de Zoologia, Universida-
de de Sao Paulo, Sao Paulo, SP,
Brazil; P. Vanzolini, J. L. Leme
MZUF Museo Zoologico de "La Spe-
cola" Universita di Firenze,
Florence, Italy; S. Whitman
NRMS Naturhistoriska Riksmuseet,
Stockholm, Sweden; T. Krones-
tedt
PAN Polska Akademia Nauk, Warsza-
4 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
wa, Poland; J. Proszynski, A. Slo-
jewska, W. B. Jedryczkowski
REL R. E. Leech, Edmonton, Alber-
ta, Canada
SMF Forschungsinstitut Sencken-
berg, Frankfurt am Main, Ger-
many; M. Grasshoff
SR Susan Riechert, Knoxville, Ten-
nessee, United States
USNM National Museum of Natural
History, Smithsonian Institution,
Washington, D.C., United
States; J. Coddington, S. F.
Larcher
ZMB Zoologisches Museum der
Humboldt Universitat, Berlin,
Germany; M. Moritz
ZMUC Zoologisk Museum, Gopenha-
gen, Denmark; H. Enghoff, N.
Scharff
ZSM Zoologische Staatssammlung,
Munich, Germany
Locality Data Storage and Manipula-
tion. Locality data from each collection vial
were entered into a database designed us-
ing Glaris FileMaker Pro®. Geographic
coordinates were added to locality data
that lacked them using maps, USBGN gaz-
etteers, and on-line databases (http://164.
214.2.59/gns/html/ and http://mapping.
usgs.gov/www/gnis/). Occasionally locality
information was illegible or unknown or
one of several homonymous sites. In such
cases a reasonable, educated guess was
inade and a "[?]" designation was append-
ed to the locality. In some cases the itin-
erary of a collector was reconstructed from
other known records, and the ambiguous
locality was assigned a coordinate halfway
between the previous and following known
collection sites. The locality database
worked in concert with the mapping pro-
gram Atlas Pro® to generate thematic
maps on the fly. These maps helped in the
process of delimiting species and discov-
ering cryptic species.
Elevation (in meters) was estimated for
each locality that lacked this information.
In some cases, elevation was estimated us-
ing contour maps, such as DMAAG ONG
aeronautical maps; in most cases, elevation
was estimated using NOAA data with an
on-line database server (http://phylogeny.
harvard.edu/~piel/find.htiril).
The enhanced locality database was
used to reveal ecological and life history
traits. Seasonality of species was expressed
by plotting a circular histogram showing
the relative amount of collecting activity
per 5-day interval (Figs. 300-337). While
locality dates alone cannot control for the
seasonal activity of human collectors, these
data at least provide an estimate of spider
seasonal abundance, if only approximate.
Some syiupatric species show incongruous
seasonal abundance, which is at least some
evidence that seasonality of spider collec-
tors does not unduly overshadow the sea-
sonality of the spiders themselves.
Examination and Illustration. Speci-
mens were examined under 80% ethanol
in a dish with light and dark sand grains
for specimen support. Digital photographs
of preserved specimens were taken
through a Nikon SMZ-10 photomicro-
scope using a Panasonic WV-CL320 GGD
video camera, chosen for its high sensitiv-
ity to light. Video images were captured
using a Quicklmage®24 digitizer and ed-
ited on a Quadra 700 Macintosh® com-
puter. The computer allows relatively in-
expensive pictures to be printed rapidly on
a 1,200 dpi Xante® Accel-a- Writer 8200
laser printer. Digital pictures were used to
help sort out individuals to species, to cre-
ate publishable pictures of gross dorsal
and ventral markings, and to aid in the il-
lustration of genitalia. As an aid in illustra-
tion, the digital pictures functioned as a
camera lucida because they assured accu-
racy when drawing the proportions of gen-
ital parts and scle rites. Usually a digital
picture was laid over carbon paper and an
outline of the genitalia was transferred to
coquille board underneath. The illustra-
tion continued on the coquille board using
a Staedtler OmniGhrom® pencil and a
drafting pen with India ink and then was
scanned at 600 dpi on a LaGie Silverscan-
Metepeira • Piel
ner II®. The resulting digital image was
edited in Adobe Photoshop® and reduced
in size to 1,200 dpi. The edited figures
were finally arranged on plates using Can-
vas®.
External genital structures were manip-
ulated with pins to reveal hidden parts.
The terminal division on the male palp is
hinged, so it had to be pried open to see
the embolus and embolic apophyses prop-
erly. In females, mating plugs had to be
removed from epigynal openings using
pins. Sometimes the entire epigynum was
partly cut from the body so as to see it
from a posterior view.
Internal genital structures were studied
by clearing them in clove oil and examin-
ing them using an Olympus BH-2 com-
pound microscope. Sketches were made
directly on the computer in Canvas® by
aiming the camera lucida at the computer
monitor. While internal genital structures
helped in the process of delimiting spe-
cies, they did not prove to be as useful as
external genital structures in describing
species; thus, these working sketches are
not figured herein.
Measurements of the spiders were tak-
en using a Leitz stereo dissecting micro-
scope with a calibrated reticule. Sizes of
leg articles, eyes, and carapace, were per-
formed on one specimen of each sex, for
each species. The respective localities of
the candidate specimens were indicated in
the descriptions. This study placed little
reliance on spider leg measurements be-
cause they are not usually very useful in
spider taxonomy, and because Metepeira
species are notorious for their variability in
size (Levi, 1977; Piel, 1996).
All eye sizes were reported as a ratio of
the posterior median eye diameters to the
diameter of every other eye type. For ex-
ample, in the case of "ratio of eye diame-
ters: posterior medians and anterior me-
dians 2.0, anterior laterals 0.5, posterior
laterals 1.0," the reader should interpret
the anterior medians to be half the size of
the posterior medians, and the anterior lat-
erals to be twice the size of the posterior
medians. Eye separations were expressed
in terms of their own diameters, or in
terms of the anterior lateral eyes when be-
tween eyes of different types. Oval eyes
were measured as an average of the lon-
gest and shortest lengths.
In parallel with the last revision of Me-
tepeira (Levi, 1977), leg measurements
were made on each article distal to the tro-
chanter for the first leg and on the com-
bined lengths of the patellae and tibiae for
all remaining legs. Variation in total body
size was provided as an average, minimum,
and maximum of the total lengths from a
number of mature specimens, usually cho-
sen from a wide geographic spread.
Metepeira F. O. P. -Cambridge
Metepeira F. O. P.-Cambridge, 1903: 457. Type spe-
cies by original designation M. spinipes F. O. P.-
Cambridge 1903. The name is feminine.
Diagnostic Abstract. Web combines bar-
rier or scaffolding structure surrounding a
classic araneid orb with a retreat suspend-
ed in air (Fig. 1). Like a raccoon with its
facial colors reversed, the eye region is
lighter than any other part of the carapace
(Fig. 2). The venter has a wide median
white line set on a black background that,
with only some exception, extends anteri-
orly on the sternum (Fig. 3). With one ex-
ception, the total lengths of distal leg ar-
ticles (metatarsus and tarsus) exceed that
of the middle articles (patella and tibia).
The median apophysis has two distinctive
flagella (F in Fig. 5) and, in some species,
an easily recognizable keel (K in Fig. 5).
The dorsal abdominal markings (the foli-
um) look like an inverted fleur-de-lis, al-
lowing easy recognition of the genus in the
field (Fig. 2).
Description and Diagnosis. For field
ecologists, the most obvious and distinctive
feature of Metepeira is the combination of
orb and barrier web (Fig. 1). The barrier
\^eh forms scaffolding around an almost
vertical orb and supports the spiders re-
treat, which is thus suspended away from
any substrate.
In contrast to most araneids, the cara-
6 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
pace of Metepeira is lightest in the eye re-
gion. However, this distinctive feature
varies within the genus: in the case of M.
rectangula (Nicolet, 1849), the hghter re-
gion takes up ahiiost half the carapace
(Fig. 65); in the case of Metepeira F. O.
P.-Cambridge, 1903, the lighter region is
usually limited to the anterior edges of the
carapace (Fig. 2). White, downy hairs of-
ten cover the carapace but are especially
white and conspicuous on the lighter parts
of the carapace outside the eye region. In
some species, such as M. spinipes, these
hairs make the carapace look gray or sil-
very when the spider is alive, but dark
brown when the spider is in ethanol.
The eyes of Metepeira are not particu-
larly unusual. Eye separations relative to
eye diameters increase with spider size:
larger spiders tend to have relatively great-
er eye separations. In either sex, the pos-
terior median eyes are between 1.1 and
1.7 times the size of the anterior medians,
and the separation between posterior me-
dian eyes is between 0.4 and 0.7 of that
between anterior median eyes. The sepa-
ration between the anterior median eyes
and the anterior lateral eyes is between 1
and 3.7 times the size of anterior median
eyes in males and between one and five
times the size in females. The diameter of
the anterior median eyes exceeds the
height of the clypeus.
The shape of the female abdomen rang-
es from wider than long and rhomboid (M.
datona. Fig. 12) to roundish (e.g., M. de-
senderi. Fig. 20; M. rectangula. Fig. 65),
to longer than wide and oval (e.g., M. inca.
Fig. 169). The dorsal folium has a recog-
nizable white fleur-de-lis pattern, usually
on a dark background, its edges shaped by
a wavy, zig-zag white outline (Fig. 2). The
dorsum of live spiders is often more red-
dish — a pigment that rapidly dissolves in
alcohol.
Somewhat less common among other
araneids is the median white line on the
venter of the abdomen (Fig. 3), which is
present (though shortened) even in the
most darkly pignriented species. However,
unique among araneid genera is the com-
bination of median white line on the ven-
ter and median white line on a black or
brown sternuin. Some Metepeira species
lack a complete white line on the sternum,
but even those, such as M. datona, that
usually have an entirely black sternum
nonetheless sho\v hints of white markings
in some specimens. Characteristics found
in the carapace, abdomen, and sternum of
Metepeira are also found in Araneus
koepckeorurn Levi, but this last species
lacks the white line on the venter.
With the exception of M. datona, and in
some cases, M. desenderi, all Metepeira
species have a combined metatarsus and
tarsus that is longer than the combined pa-
tella and tibia. This feature is unusual
among araneids and is not found in Kaira
O. P. -Cambridge or other likely relatives
to Metepeira (Levi, 1977; Piel and Nutt,
1997).
In most species the leg articles are
ringed, usually with brownish black on the
distal and dorsal surfaces of each article,
except for the patellae and tarsi which are
usually entirely dark. In mainly tropical
and high-altitude species, the coxae are
mostly black (e.g.. Fig. 75), but in desert/
mesquite species they appear yellowish
white (e.g.. Fig. 28). I
Unlike many other araneids — and per-
haps because of the small male size — the
coxa on leg I of male Metepeira lacks the
hook and corresponding groove typically
found on femur II. In addition, males lack
a tooth on the lateral side of the endite,
and they lack a basal tooth on the palpal
femur. The phylogenetic analysis of
Scharff and Coddington (1997) incorrectly
codes Metepeira as having a tooth on the
endite. However, had the authors coded
this character as absent, they would have
decreased the length of their preferred
tree because the nearest relatives hypoth-
esized for Metepeira {Kaira, Zijgiella, and
Singa) also lack this tooth.
Macrosetae usually concentrate on arti-
cles that contact other spiders during mat-
ing or grappling. In contrast to most gen-
Metepeira • Piel 7
Barrier Web
White Dorsal Light Eye Region
Fleur-de-Lis / ^^ Median White
Pattern x AW* ^Ma Line on Sternum
Orb Web
Median White
Line on Black
Venter
vigilax nigriventris incrassata minima
foxi labyrinthea compsa vcntura
Figure 1. Web of immature Metepeira grandiosa alpina from Chihuahua, Mexico.
Figure 2. Dorsum of adult female Metepeira crassipes.
Figure 3. Venter of adult female Metepeira tarapaca new species.
Figure 4. Hypothetical phylogenetic relationships among Metepeira species groups. Shaded branches indicate species groups
that live in South America; open branches indicate species groups that live in North America, Central America, and the Caribbean.
Abbreviations: DEA, distal embolic apophysis; K, keel of median apophysis; TA, terminal apophysis; (+), character state gain;
(-), character state loss.
Figures 5, 6. Male palpus. 5, mesal view, Metepeira compsa. 6, ventral view of distal embolic division, Metepeira labyrintliea
(Hentz).
Abbreviations: BEA, basal embolic apophysis; C, conductor; DEA, distal embolic apophysis; E, embolus; F, flagellum on median
apophysis; K, keel of median apophysis; MA, median apophysis; TA, terminal apophysis; TO, terminal division.
Figures 7-13. Metepeira datona Chamberlin and Ivie (sp. 1; 17°53'N, 76°19'W). 7, male palpus, mesal. 8, epigynum, posterior.
9, epigynum, ventral. 10, male, dorsal. 11, male, ventral. 12, female, dorsal. 13, female, ventral.
Scale bar: dorsum and venter figures 1 .0 mm.
8 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
era related to Araneus, Metepeira has con-
centrated macrosetae on femur I instead
of tibia II (Scharff and Coddington, 1997).
Female Metepeira have between two and
five macrosetae on die anterior side of the
femur, and between zero and seven on the
anteroventral side. Males typically have
inore setae than their conspecific females:
four to nine on the anterior side and two
to nine on the anteroventral side. Variation
in the number of macrosetae appears to
correlate with body size. In most species,
the inale palpal tibia and patella each have
two strong macrosetae (Levi, 1977, fig. 8).
Compared with other araneid genera,
Metepeira have rather small and similar
genitalia, which on the one hand makes
the genus easy to recognize, but on the
other hand makes species tough to iden-
tify. The small epigynum is fleshy, variable
in shape, and weakly sclerotized. Unlike
Araneus, Metepeiras scape never has a
pocket but always ends with a pointed tip
(e.g.. Fig. 31). The cleared epigynum —
and in many cases the uncleared epigyn-
um — reveals a pair of sclerotized spherical
structures where the embolus is inserted,
as well as ducts to pass semen to the larger,
spherical seminal receptacles. In some
species, these spherical structures are wide
apart (e.g.. Figs. 16, 17), in others they are
tubular (e.g.. Figs. 39, 40), but in many,
they are closer together (e.g.. Figs. 93, 94).
Frequently the deeper, large seminal re-
ceptacles can be seen through uncleared
tissue (e.g.. Figs. 201, 295).
The male palp is more distinctive. In
particular, the median apophysis (MA in
Fig. 5), while not always a good character
for separating closely related species, is ex-
cellent when it comes to identifying the
genus. Two flagellae (F in Fig. 5) grace-
fully curve off the base of the median
apophysis, and in some species, a toothed
or smooth keel (K in Fig. 5) extends in the
opposite direction. This design is also seen
in Kaira, Aculepeira, and Amazonepeira,
but none of these have flagellae that ap-
pear so integral to the base structure.
The tenninal division on the Metepeira
palp is very similar in almost all species.
When this structure is pulled up, a basal
embolic apophysis (also known as an em-
bolar lamella) can be seen in the shape of
a club or spatula (E in Figs. 5, 6). Soiue-
times a distal embolic apophysis can be
seen if it is not hidden from view by an
overhanging terminal apophysis. When the
terminal apophysis is large and sclero-
tized — which is die case in all but the Me-
tepeira foxi species group — it has a rec-
ognizable toothed notch, like the mouth
on a wrench (Fig. 6). Virgin males have a
cap on the embolus that remains in the
epigynum after mating and presumably
serves as a barrier to subsequent mating
(Levi, 1977). The shape of the embolus
cap varies from tiny (e.g.. Fig. 178) to
short but wide (e.g., Fig. 199) to large and
winged (Fig. 46). Finally, the terminal di-
vision lacks a stipes — a sclerite between
the radix and the embolus that is frequent-
ly found in other genera related to Ara-
neus (Scharff and Coddington, 1997).
Natural History. All Metepeira species
build a unique web that combines an orb
with a barrier web (Levi, 1977; Lopez,
1993). As with Cijrtophora Simon or Me-
cynogea Simon (Levi, 1997), the retreat of
Metepeira hangs in the air, away from sub-
strate, and is suspended by a scaffolding
structure created by the barrier web (Fig.
1). The spider detects vibrations in the
web and gains quick access to the hub us-
ing a signal line that runs from the retreat
to the center of the orb (Fig. 1). Tan col-
ored egg sacs are strung together, usually
above the retreat, and the most recently
laid eggs are nearest to the spider. In some
species the egg sacs and retreat are deco-
rated with insect parts (e.g., M. spinipes);
in other species they are carefully wrapped
by leaves and woven together (e.g., M. da-
tona). Unlike the webs oi Cijrtophora and
Mecynogea, the orb web of Metepeira is
oriented vertically, and the number of radii
and sticky spirals are more typical of other
araneines.
In some species, such as M. pimungan
(personal observation) and, to a lesser de-
Metepeira • Piel
gree, M. incrassata (G. Uetz, personal
communication), juveniles and adults
without eggs will live on webs lacking a
suspended retreat. Instead, the spider sits
on a white disk-shaped stabilimentum in
the center of the hub. Of 110 M. pimun-
gan specimens observed on San Miguel Is-
land, about 40% occupied webs of this
type. In two cases the disk stabilimenta
were partly separated from the hub by
barrier web lines and were further bent
over to form a partly covered protective
retreat for the spider. This observation
makes it possible to imagine that the disk
stabilimentum seen in M. pimiingan re-
sults from the fusion of the suspended re-
treat with the hub.
When food supplies are plentiful, spi-
ders of all kinds show an increased toler-
ance for one another and an increased ten-
dency to aggregate (e.g., Gillespie, 1987;
Rypstra, 1986). The suspended retreats
and barrier webs of Metepeira, Cyrtopho-
ra, and Mecynogea may actually further fa-
cilitate in the formation of aggregations by
easing dependency on substrate availabili-
ty and by providing a common support sys-
tem (Burgess and Witt, 1976; Uetz, 1986).
In any case, colony formation is known to
occur in all three genera (e.g., Rypstra,
1979), but especially in Metepeira. Small
colonial aggregations of two to 10 individ-
uals occur in M. datona (Spiller and
Schoener, 1989), M. minima (personal ob-
servation), M. glomerabilis (R. Baptista,
personal communication), and M. atascad-
ero new species (e.g., Uetz and Hodge,
1990). Medium-size colonies of 10 to 30
individuals occur in M. pimungan (person-
al observation), M. gressa (Viera and Cos-
ta, 1988), M. nigriventris (L. Rayor, per-
sonal coinmunication), M. tarapaca (V.
Roth, locality label), and M. spinipes (e.g.,
Uetz, 1988a). Large colonies, sometimes
in the thousands of individuals, commonly
occur in M. incrassata (e.g., Uetz and
Hodge, 1990). Near rivers and in other
lush habitats, M. tarapaca colonies can
reach 200 individuals (M. Roy, personal
communication). These cases of social be-
havior, broadly spread across seven differ-
ent species groups, may mean that aggre-
gation is a frequently lost and relatively old
trait, or it may mean that species are prone
to converge and evolve the same behavior
independently.
Either way, much research has focused
on elucidating the selective forces behind
colonial behavior in Metepeira. In partic-
ular, Uetz (1988a,b, 1996) has provided
strong support for the hypothesis that Me-
tepeira forage using a risk-sensitive strat-
egy. He suggests that spiders in abundant
habitats seek to minimize individual vari-
ance in prey capture by aggregating in col-
onies, whereas spiders in poor habitats
seek to maximize variance by living soli-
tarily — perhaps in a risky attempt to find
areas of local prey maxima. The diversity
of social tendencies among species is
therefore commensurate of the diversity of
ecological habitats that they inhabit.
Indeed, Metepeira species thrive in a
wide array of habitats, though often they
are quite harsh. These include wet, mon-
tane cloud forests in Mexico and Panama
(M. incrassata, M. olmec); tropical and wet
agricultural areas (M. uncata, M. vigilax,
M. glomerabilis, M. roraima); high-eleva-
tion pine forests (M. lacandon, M. nigri-
ventris, M. grandiosa alpina); Canadian
bogs (M. grandiosa palustris); deciduous
forests in the eastern U.S. (M. labyrin-
thea); Caribbean coastal shrubbery (M. da-
tona, M. minima, M. triangularis, M. ja-
maicensis, M. maya, M. celestun); Mexican
mesquite grasslands (M. atascadero, M.
chilapae); Patagonian dunes and scrub (M.
galatheae) and pampas grass (M. karkii);
diy Californian buckwheat and sage (M.
crassipes, M. ventura, M.foxi, M. grandio-
sa grandiosa); and arid and semiarid de-
serts (M. arizonica, M. inca, M. ventura,
M. crassipes). Although some species (e.g.,
M. galatheae, M. spinipes, M. cornpsa) cov-
er vast geographic areas and live in many
different habitats, many species are more
biogeographically restricted. In fact, sev-
eral species follow narrow ecological zones
that decrease in elevation with distance
10 Bulletin Museum of Comparative Zoology, Vol. 157, No.
from the equator (e.g., M. rectangula, M.
vigilax, M. cajahamha. Fig. 36; M. arizon-
ica. Fig. 213).
Close cohabitation with different inter-
and intrageneric species is not uncommon.
Colonies of M. incrassata are known to
contact webs of Nephila clavipes Linnaeus
(Hodge and Uetz, 1996) and Mecijnogea
ocosingo and Gasteracantha cancriformis
(personal observation). Often M. crassipes,
M. Ventura, M. foxi, and M. grandiosa
grandiosa are collected together (Levi,
1977), as are M. minima and M. celesiun
(personal observation). Species that have
been collected from identical localities,
though not necessarily at the same time,
include: M. chilapae and M. spinipes; M.
chilapae and M. atascadero; M. karkii and
M. galatheae; M. calamuchita new species,
M. gressa, and M. galatheae; M. rectan-
gula, M. calamuchita, and M. galatheae;
M. compsa and M. gressa; M. vigilax and
M. compsa; M. glomerahilis and M. vigilax;
M. compsa and M. glotnerabilis; M. comp-
sa and M. nigriventris; M. compsa and M.
mcfl; M. datona and M. jamaicensis; and
M. datona and M. triangularis.
Despite the wide biogeographic ranges
of M. compsa (Puerto Rico and south to
Argentina, Map 8) and M. datona (His-
paniola and north to Florida, Map 1), they
nonetheless come geographically close to
one another but do not overlap. It is hard
to imagine that the hurricanes that fre-
quently pass through the Caribbean, as
well as the homogeneous island environ-
ments, would not gradually cause these
two species distributions to overlap. Per-
haps these abrupt, disjunct distributions
are a rare example of competitive exclu-
sion in Metepeira, which in other species
is not thought to be an important factor
(Wise, 1983).
Sphecid wasps are predators on Mete-
peira. Locality labels indicated that M. pa-
cifica has been found in the nests of Try-
pargilum nitidum, T. tenoctitlan, and T.
hensoni. Jimenez and Tejas (1994) report
that M. crassipes is the most frequent prey
item in the nests of Trypargilum triden-
tatum. Colonial spiders, such as M. incras-
sata, are especially vulnerable to wasps,
other spiders, sarcophagid flies (e.g., Ar-
achnidomyia Undue, A. rayorae), and
hummingbirds (Hieber and Uetz, 1990;
Lopez, 1989; Rayor and Uetz, 1990).
Species Groups. Nearctic Metepeira
were divided into two species groups: the
M. labyrinthea group and the M. foxi
group, based on the pattern on the ster-
num and the shape of the median apoph-
ysis (Levi, 1977). Baert (1987) questioned
the taxonomic usefulness of the M. foxi
species group (M. foxi, M. grandiosa, M.
datona) because he found that M. desen-
deri has both a keel on the median apoph-
ysis and a white sternal line (Figs. 15,
21) — a combination that is incoinpatible
by Levi's scheme. Nonetheless, the geni-
talia of M. desenderi closely ally this spe-
cies with the M. foxi group, so I am re-
defining the M. foxi group based on purely
genitalic characters. This is likely to be a
basal, paraphyletic group (Fig. 4) (Piel and
Nutt, 1997).
Seven additional species groups are dis-
tal to the M. foxi. These remaining species
are united by sharing a large terminal
apophysis that is sclerotized and usually
studded with teeth or denticles. The M.
vigilax group (M. vigilax, M. cajahamha,
M. glomerahilis, M. rectangida) are united
by large emboli with long scooplike basal
embolic apophyses (Fig. 60). Unlike the
remaining species, the terminal apophysis
in this group — albeit large — does not ac-
tually overhang or hide the embolus. In
addition to an overhanging terminal
apophysis, the remaining taxa are also
united by a distal embolic apophysis that
either protrudes (Fig. 76), curves off (Fig.
185), or is secondarily lost (Fig. 264). The
M. labyrinthea group (M. labyrinthea, M.
lacandon, M. spinipes) share a toothless,
smooth keel on the median apophysis
(Figs. 67, 69).
The M. nigriventris group and the M.
compsa group together share a median
apophysis with teeth on the face of the
keel (Figs. 92, 149). The M. incrassata
Metepeira • Piel
11
Map 1
• glomerabilis
+ cajabamba
Map 3
Map 2
grandiosa alpina
grandiosa grandiosa'
4»^
• vigilax
+ rectangula
Map 4
Maps 1, 2. Metepeira fox; species group. 1, M. datona, M. desenderi. 2, M. grandiosa grandiosa. M. grandiosa alpina.
Maps 3, 4. Metepeira vigilax species group. 3, M. glomerabilis. M. cajabamba. 4, M. vigilax, M. rectangula.
group, the M. ventura group, and the M.
minima group all lack a keel on the me-
dian apophysis (Figs. 164, 222, 293). How-
ever, both the M. compsa group and the
M. incrassata group have epigyna with
similar oval or round sclerotized rims
(Figs. 151, 166), so it is likely that these
are paraphyletic and consist of species
leading up to a major North American
(without a keel) and South American (with
a keel) phylogenetic split (Fig. 4).
The South American branch includes
the M. compsa group (M. compsa, M. ro-
raima, M. gressa) and, more distally, the
M. nigriventris group (M. nigriventris, M.
tarapaca, M. calamuchita, M. galatheae,
M. karkii). This latter group is united by a
distinctive and derived scape, which pro-
jects out and down, creating a noticeable
arch and overhang (Fig. 86).
The remaining species all lack a keel on
the median apophysis, and with one ex-
ception {M. inca), they live exclusively in
North America (from the Caribbean and
Panama to Nevada). The M. incrassata
group (M. gosoga, M. maya, M. inca, M.
Comanche, M. olmec, M. atascadero, M. ar-
izonica, M. incrassata, M. triangularis, M.
pimungan) are very likely paraphyletic.
The epigynum on each species seems au-
tapomorphic and difficult to unite with any
others. Some species (M. gosoga, M. maya,
M. inca) have a pointed or projecting distal
embolic apophysis (Fig. 171). Others have
12 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
a distal embolic apophysis that curves off
sharply but does not project forward (Fig.
185). Finally, others have a distal embolic
apophysis that curves off gently, almost to
the point of hiding the existence of an
apophysis (Fig. 206).
The M. minima group (M. jamaicensis,
M. minima, M. pacifica, M. petatlan) and
the M. Ventura group (M. uncata, M. Ven-
tura, M. celestun, M. chilapae, M. revilla-
gigedo new species, M. crassipes) are unit-
ed by derived characters: both have thin
flagellae arising from a thin base on the
median apophysis (Fig. 264), and both
share the secondary loss of the distal em-
bolic apophysis. The M. minima group is
clearly monophyletic; its species all have
their flagellae further set off from the me-
dian apophysis on a separate, narrow stalk
(Fig. 286).
Key to Female Metepeira
1 Epigynal openings strongly sclerotized,
round but tilted so that they appear
oval from a ventral view (Figs. 40, 48,
55) 2
— Epigynal openings weakly sclerotized
(e.g.. Fig. 208), and if round, they are
not tilted and do not appear oval from
a ventral view (Fig. 131) 4
2(1) Epigynal openings shaped like the en-
trance to a snail shell (Fig. 55); His-
paniola, Brazil, Argentina (Map 4)
( 7 ) vigilax
— Epigynal openings tubular (Figs. 40, 48)
3
3(2) Epigynal openings strongly tilted (Fig.
40); Peru (Map 3) (5) cajabamba
— Epigynal openings weakly tilted (Fig. 48);
Colombia to Brazil (Map 3)
(6) glomerabilis
4(1) Epigynal openings large and gaping, cre-
ating large atria inside (Figs. 61, 62);
Chile and Argentina (Map 4)
( 8 ) rectangula
— Epigynal openings not gaping and not
creating large atria (e.g.. Fig. 69) 5
5(4) Weak posterior lobes on the epigynum
create a single, wide epigynal depres-
sion (Figs. 9, 17, 24, 31) or epigynum
with crescent-shaped sclerotized open-
ings on either side of a thin scape
(Levi, 1977, fig. 87). Dark, sclerotized
spheres below epigynal openings are
greatly separated (Figs. 8, 16) 6
— Stronger posterior lobes on the epigynum
create separate, smaller epigynal de-
pressions that are not crescent-shaped,
or if crescent-shaped, the scape is thick
and puffy (e.g.. Figs. 86, 102, 143, 166,
187, 208). Dark, sclerotized spheres
below epigynal openings are closer to-
gether (e.g.. Figs. 101. 142) 11
6(5) Sternum with longitudinal white line
(Fig. 21); Galapagos Islands (Map 1)
(2) desenderi
- Sternum entirely black or brown (Figs.
13, 28, 35) 7
7(6) Abdomen wider than long (Fig. 12); Flor-
ida to Hispaniola (Map 1) (1) datona
- Abdomen longer dian wide (Figs. 27, 34)
8
8(7) Coxae as black as sternum (Levi, 1977,
fig. 98); Canada-U.S. border (Levi,
1977, map 2)
(Levi, 1977: 212) grandiosa palustris
— Coxae yellow or orange and lighter than
sternum (Figs. 28, 35) 9
9(8) Epigynum with crescent-shaped sclero-
tized openings on eitlier side of die
scape (Levi, 1977, fig. 87); western
U.S. and Canada (Levi, 1977, map 2)
(Levi, 1977: 210) foxi
— Epigynum with wide, transverse depres-
sion (Figs. 23, 24, 30, 31) 10
10(9) Scape wide and stubby (Fig. 24); Baja
California north to Canada (Map 2)
(3) grandiosa grandiosa
- Scape triangular (Fig. 31); in mountains
from north-central Mexico to Canada
(Map 2) (4) grandiosa alpina
11(5) Scape diickness equal to greater than
width of epigynal depressions (e.g..
Figs. 86, 119, 131, 143, 180, 201, 266) '
12
— Scape narrower than epigynal depres-
sions, or epigynal depressions in the
shape of longitudinal slits (e.g.. Figs.
238, 252, 280) 35
12(11) Base of scape originates anteriorly and
projects ventrally before cui'ving pos-
teriorly. This projection creates an
overhang and a noticeable gap between
the scape and the genital openings
(e.g.. Figs. 85, 86, 101, 102, 119, 143)
13
— Scape does not create a noticeable gap or
overhang (e.g.. Figs. 77, 78, 130, 131,
165, 166, 265) 18
13(12) Rim of epigynal depressions slightly
sclerotized and oval-shaped (Fig. 143);
northern Brazil, French Guiana, and
Colombia (Map 8) (17) roraima
- Epigynal depressions without distinct rim
(Fig. 123) or not sclerotized (e.g.. Fig.
102) 14
14(13) Sternum black, coxae mostly black (Fig.
Metepeira • Piel
13
A calamuchita
karkii
mgnventris f ( ^
Map 5
▲ galatheae
• tarapaca
Map 7
• spinipes
■ lacandon
Map 6
+ compsa
• roraima
■ gressa
Map 8
Maps 5, 7. Metepeira nighventris species group. M. calamuchita, M. I<arl<ii, M. nigriventris, M. galattieae, M. tarapaca.
Map 6. Metepeira labyrintfiea species group. M. spinipes, M. lacandon.
Map 8. Metepeira compsa species group. M. compsa, M. roraima, M. gressa.
91), and carapace without lighter me-
dian mark (Fig. 90); high altitudes in
Bolivia and Peru near Lake Titicaca
(Map 5) (11) nigriventri.s
Sternum with median white line (Fig.
99), or if sternum is black, then either
the carapace has a median lighter ar-
row-shaped mark (Fig. 113) or the cox-
ae are mostly yellow (Figs. 114, 128)
15
15(14) Stenium brown to black with parallel
lines on either side of median white
line on venter: the parallel lines are
tliicker anteriorly than posteriorly (Fig.
128). Lower lip on epigynum thick and
bulbous (Fig. 123); southern Argentina
and southern Chile (Map 5) .... (15) karkii
— Sternum with median white line, or if
sternum is entirely brown to black,
then parallel lines on either side of
14 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
white line on venter are either absent
(Fig. 114) or equally thick anteriorly as
posteriorly (Fig. 99). Epigynum with-
out thickened lower lip (Figs. 94, 102,
118, 119, 120) 16
16(15) Dark, sclerotized spheres in epigynal
openings small (Figs. 119, 120). Ante-
rior lip of epigynum rounded off,
sometimes with openings shifted pos-
teriorly and wiinkled portion of the
scapes hood shifted more to the epi-
gynum proper (Fig. 118); Chile and
Argentina (Map 7) (14) galatheae
— Dark, sclerotized spheres in epigynal
openings large, and anterior lip of epi-
gynum not rounded off. Openings al-
ways located midway down the epigyn-
um (Figs. 94, 102) .._._ 17
17(16) General shape of epigynum is triangular
with posterior width greater than an-
terior width. Scooped-out depressions
project more posteriorly than laterally
(Fig. 102); north-central Argentina
(Map 5) (13) calamiichita
— General shape of epigynum square.
Scooped-out depressions project more
laterally than posteriorly (Fig. 94);
northern Chile and southern Peru
(Map 7) (12) tarapaca
18(12) Epigynal openings almond-shaped, not
noticeably sclerotized, and at a 40° to
60° angle from axis of spider (Figs. 69,
78) 19
— Epigynal openings not almond-shaped
(e.g.. Figs. 180, 187, 201), or if al-
mond-shaped, slightly sclerotized and
at an angle from spider's axis of 80° to
100° (Figs. 151, 159, 166) 22
19(18) Almond-shaped openings created by de-
pression where scape arises from epi-
gynum (Fig. 173); California, Arizona,
northwestern Mexico (Map 9)
(21) gosoga
— Almond-shaped openings created by
membranes inside depressions and not
associated with scape (Figs. 69, 78) ..... 20
20(19) Distinct C-shaped depression created
where die scape arises from the epi-
gynum (Fig. 78). Black marks inside al-
mond-shaped openings not cross-eyed
in appearance (Fig. 78). Black ster-
num, usually with white spot in center
(Fig. 83); mountainous regions of Chia-
pas, Mexico (Map 6) (10) lacandon
— Indistinct depression created where the
scape arises from the epigynum (Fig.
69). Black marks inside almond-shaped
openings cross-eyed in appearance
(Fig. 69). Sternum black, with or with-
out a median white line. If with only a
portion of a median white line present.
usually only at the posterior end of the
sternum (Fig. 75). Never with only one
white spot in center 21
21(20) Because of interspecific variability and
polymorphism, females of the follow-
ing two species are almost impossible
to separate reliably without molecular
sequence data. Small ribosomal sub-
unit (12S) mtDNA sequence data has
tlie following diagnostic markers. Base
14261: ACGGT; base 14285: ATTTT;
base 14361: ACTAC; base 14394:
CTTAT; base 14412: ATTA. (Base
numbers refer to homologous sites in
tlie mitochondrion of Drosophila yak-
uba, as reported by Clary and Wolsten-
holme [1985].) One quarter of scape
extends below lower lips of epigynum
(Levi, 1977, fig. 14); New England to
Florida and west to eastern Texas
(Levi, 1977, map 1, but not including
points appearing in Mexico)
(Levi, 1977: 196) labyrinthea
— For 12S mtDNA sequence data, the fol-
lowing sequences are diagnostic. Base
14261: AGGAT; base 14285: ATCTT;
base 14361: ACCAC; base 14394:
CTAAT; base 14412: TTTA. One third
of scape extends below lower lips of
epigynum (Levi, 1977, fig. 21; Fig. 69);
Mexico City north to California (Map
6) (9) .spinipes
22(18) Epigynal openings small, round, and
sclerotized (Fig. 131). Openings some-
times hidden by wide scape (Fig. 134);
Puerto Rico to Argentina and Chile
(Map 8) (16) compsa
— Epigynal openings not small, round, and
sclerotized 23
23(22) Rim of epigynal openings sclerotized and
in an oval or teardrop shape (Figs. 151,
159, 166, 187) 24
— Rim of epigynal openings not sclerotized
in an oval or teardrop shape (Figs. 180,
194, 201, 208) 27
24(23) Epigynal openings oval, small, and partly
hidden by scape (Fig. 151); northern
Argentina, Uruguay, and southern Bra-
zil (Map 8) (18) gressa
— Epigynal openings teardrop-shaped (Fig.
187) or large and oval but not hidden
by scape (Figs. 159, 166) _. 25
25(24) Epigynal openings teardrop-shaped (Fig.
187); northeastern Mexico to Texas
(Levi, 1977, map 1; Map 11)
(23) Comanche
— Epigynal openings oval-shaped (Figs.
159, 166) 26
26(25) Lower lip of epigynum pointed (Fig.
166), abdomen white (Fig. 169); north-
ern tip of Peru (Map 11) (20) inca
Metepeira • Piel
15
+ arizonica -x
• gosoga
♦ incrassata
■ pimungan
Map 9
pacifica
petatlan
Map 10
+ Comanche
X inca
♦ olmec
Map 11
• uncata
♦ crassipes
■ revillagigedo
Map 12
Map 13
■ Ventura
▲ celestun
• chilapae
jamaicensis
minima b; s
jcmaya
+ atascadero
• triangularis
Map 14
^;:^5y-.^
Map 15
Maps 9, 11, 14. Metepeira incrassata species group. 9, M. arizonica, M. gosoga, M. incrassata, M. pimungan. ^^, M. comanche,
M. inca, M. olmec. 14, M. maya, M. atascadero, M. triangularis.
Maps 10, 15. Metepeira minima species group. 10, M. pacifica, M. petatlan. 15, M. minima, M. jamaicensis.
Maps 12, 13. Metepeira ventura species group. 12, M. uncata, M. crassipes, M. revillagigedo. 13, M. ventura, M. celestun, M.
chilapae.
— Lower lip of epigynum thickened but not
pointed (Fig. 159), abdomen dark (Fig.
162); southern Mexico and Belize to
Costa Rica (Map 14) (19) maya
27(23) Rim of epigynal openings sclerotized and
shaped hke a pair of sunglasses (Fig.
201); eastern Cuba and Hispaniola
(Map 14) (25) triangularis
- Rim of epigynal openings not sclerotized
or, if sclerotized, nol in shape of sun-
glasses (e.g.. Figs. 208, 224) 28
28(27) Posterior epigynal lobes converge behind
the scape so that epigynal depressions
appear closed off froin ventral view
(Figs. 180, 208, 216, 224) 29
- Posterior epigynal lobes end before they
disappear behind the scape so that epi-
gynal depressions appear open poste-
16 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
riorly from ventral view (Figs. 194,
245, 259, 266) 32
29(28) Scape relatively long and thin, epigynal
depressions large and round, large
black spheres take up almost all the
space in the depressions (Fig. 180);
southern Veracruz to Panama (Map 11)
(22) olmec
— Scape relatively short, fat, and fleshy.
Epigynal depressions not perfectly
round; black spheres do not take up
most of the space in the depression
(Figs. 208, 216, 224) 30
30(29) Epigynum puffy, scape so thick that de-
pressions on either side of scape ap-
pear crescent-shaped (Fig. 208); south-
western U.S. to central Mexico (Map
9) (26) arizonica
— Epigynum not puffy, scape not so fat that
depressions become crescent-shaped
(Figs. 216, 224) 31
31(30) Very social. Epigynal depressions large,
disk-shaped, with shiny-smooth scler-
otized inner surfaces and thin posterior
lips (Fig. 224). Sternum mostly black,
sometimes with anterior white marks.
Venter without U-shaped mark circum-
scribing median white line posteriorly.
Coxae mostly dark brown (Fig. 228);
mainly in southern Veracruz (Map 9)
(28) incrassata
— Mostly solitary. Epigynal depressions
small, oval, with reduced shiny-smooth
sclerotized surfaces and thick posterior
lips (Fig. 216). Sternum with median
white line or only white mark at pos-
terior end. Venter with U-shaped mark
circumscribing median white line pos-
teriorly. Coxae mostly yellow (Fig.
221); central Mexican plateau (Map
14) (27) atascadero
32(28) Small square-shaped epigynal depres-
sions on either side of scape (Figs. 259,
266) 33
— Large, more rounded epigyniil depres-
sions (Fig. 194) or with straight to S-
shaped edges mostly covered by scape
(Fig. 245) 34
33(32) Dark spheres inside epigynal depressions
appear slightly walleyed. Scape rela-
tively narrower at base: about the
width of the depressions (Fig. 266);
southeast and south-central Mexico
(Map 13) (34) chilapae
— Dark spheres inside epigynal depressions
appear slightly cross-eyed. Scape rela-
tively wider at base: about twice the
width of the depressions (Fig. 259);
northwestern Mexico and California
(Map 12) (33) crassipes
34(32) Large, rounded epigynal depressions
with large black spheres inside (Fig.
194). Sternum black with posterior
dewdrop-shaped mark. Fair of short
parallel lines on either side of ventral
median white line (Fig. 198). Carapace
with large anterior white region (Fig.
197); San Nicolas Island off southern
California (Map 9) (24) pimungan
— Epigynal depressions with straight to
slightly S -shaped edges, mostly hidden
by a wide triangular scape (Fig. 245).
Sternum with wide median white line.
No parallel white lines on either side
of ventral median white line (Fig. 249).
Carapace with small anterior white re-
gion (Fig. 248); Yucatan Peninsula
(Map 13) (31) celestun
35(11) Epigynal depressions wider than long
(Figs. 231, 252) 36
— Epigynal depressions longer than wide
(e.g.. Figs. 238, 273, 280, 288) 37
36(35) Black comma shapes inside epigynal de-
pressions (Fig. 231). Sternum with me-
dian white line (Fig. 235). Dorsum
lightly pigmented (Fig. 234); north-
western Mexico and coastal California
(Map 13) (29) Ventura
— Black S-shaped marks inside epigynal de-
pressions (Fig. 252). Sternum black
with dewdrop mark at posterior end
(Fig. 256). Dorsum darkly pigmented
(Fig. 255); Guatemala and Costa Rica
(Map 12) ___ (32) uncata
37(35) Epigynal depressions indistinct anterior-
ly. Black comma-shaped marks inside
depressions and covered by translucent
membranes (Fig. 238); Isla Socorro of
the Archipielago de Revillagigedo
(Map 12) (30) revillogigedo
— Epigynal depressions distinct anteriorly.
Black spheres shifted laterally and lo-
cated outside the depressions (e.g..
Figs. 273, 280, 295) 38
38(37) Epigynal depressions slit-shaped and
usually narrower than scape (Figs. 280,
281); northwestern Mexico and Yuca-
tan Peninsula (Map 15) (36) minima
— Epigynal depressions oval and wider than
scape (Figs. 273, 288) 39
39(38) Dark spheres larger than epigynal open-
ings (Fig. 273); west coastal Mexico
(Map 10) (35) petatlan
— Dark spheres smaller than epigynal open-
ings (Figs. 288, 295) 40
40(39) V-shaped ridge under die scape. Dark
spheres located behind the junction
where the ridge meets the lateral edge
of the epigynal depressions (Fig. 288);
Honduras to Costa Rica (Map 10)
(37) pacifica
— Straight ridge under the scape. Dark
Metepeira • Piel
17
spheres located laterally and outside of
the junction where the ridge meets the
lateral edge of the epigynal depressions
(Fig. 295); Cayman Islands, Jamaica,
and Haiti (Map 15) .— (38) jamaicensis
Key to Male Metepeira
1 Terminal apophysis tliin, small, fleshy,
without teeth or sclerotized parts (Figs.
7, 15, 22, 29) 2
- Terminal apophysis enlarged, meaty, with
teeth or sclerotized parts (e.g.. Figs.
38, 60, 84, 199) 7
2(1) Terminal apophysis narrow; embolus
curled clockwise like the tip on a cork-
screw (Figs. 7, 15) 3
- Terminal apophysis wide; embolus tilted
up and L-shaped (Figs. 22, 29) 4
3(2) Curled embolus with a ridge on the up-
per surface and raised on a pedicel;
basal embolic apophysis enormous
(Fig. 15). Sternum witli wide median
white line (Fig. 21); Galapagos Islands
(Map 1) (2) desenderi
- Curled embolus smooth on its upper sur-
face and not raised on a pedicel; basal
embolic apophysis hardly noticeable
(Fig. 7). Sternum entirely black (Fig.
11); Florida to Hispaniola (Map 1) - --
( 1 ) datona
4(2) L-shaped embolus at an acute (< 90°) an-
gle (Levi, 1977, figs. 91-93); western
U.S. and Canada (Levi, 1977, map 2)
(Levi, 1977: 210) /oxi
- L-shaped embolus at angle of 90° or
greater (Figs. 22, 29) 5
5(4) Median apophysis widi rounded projec-
tion on dorsal side; jagged posterior
edge of keel (Figs. 22, 29) 6
- Median apophysis without projection —
flat on dorsal side; rounded posterior
edge of keel (Levi, 1977, fig. 105);
along the Canadian and U.S. border,
north to Nova Scotia and British Co-
lumbia, south to Maine and North Da-
kota (Levi, 1977, map 2)
(Levi, 1977: 212) grandiosa palustris
6(5) Lower, transverse part of L-shaped em-
bolus longer than vertical part (Fig.
29); mountains from north-central
Mexico to Canada (Map 2)
(4) grandiosa alpina
- Lower, transverse part of L-shape em-
bolus shorter than or equal to vertical
part (Fig. 22); Baja California north to
Canada (Map 2) .. (3) grandiosa grandiosa
7(1) Terminal apophysis does not entirely
overhang the embolus. Embolus long
and robust, wdth a long and thin gap
created between the embolus and the
basal ennbolic apophysis (Figs. 38, 46,
53, 60) 8
- Terminal apophysis often overhangs the
embolus, covering it from view. Em-
bolus not long and robust, without
long, thin gap between embolus and
basal embolic apophysis (e.g.. Figs. 67,
79, 92, 121, 141, 185) 11
8(7) Longer flagellum as thick as shorter fla-
gellum. Keel short, slim, and feather-
shaped. Embolus as wide as base of
median apophysis (Fig. 53); Hispanio-
la, Brazil, Argentina (Map 4) .... (7) vigilax
- Longer flagellum thicker than shorter fla-
gellum. Keel absent (Fig. 38) or wide
and arrowhead-shaped. Embolus thin-
ner than base of median apophysis
(Figs. 38, 46, 60) 9
9(8) Keel absent or greatly reduced (Fig. 38);
Pei-u (Map 3) (5) cajabamba
- Keel present (Figs. 46, 60) 10
10(9) Large embolus as long as basal embolic
apophysis (Fig. 60). Normal embolic
cap; Chile and Argentina (Map 4)
(8) rectangula
- Small embolus shorter than basal embol-
ic apophysis, often seen with winged
embolic cap (Fig. 46); Colombia to
Brazil (Map 3) (6) glomerabilis
11(7) Median apophysis with keel (e.g., Figs.
92, 141)
- Median apophysis without keel (e.g..
Figs. 157, 178, 286); all North Ameri-
can or Caribbean species except for M.
inca
12(11) Keel without teeth; smooth (e.g.. Figs.
67, 76); North America 13
- Keel with teeth on face; rough (e.g.. Figs.
84, 92, 100); South America 15
13(12) Distal embolic apophysis sleek, pointed,
and feather-shaped when viewed from
underside of terminal division (Fig.
70); Mexico City north to California
(Map 6) (9) spinipes
- Distal embolic apophysis spoon-shaped
(Fig. 6) or widened with bump (Fig.
79) 14
14(13) Distal embolic apophysis spoon-shaped
(Fig. 6). Sternum reddish brown with
median white line. New England to
Florida and west to eastern Texas
(Levi, 1977, map 1, excluding points in
Mexico) (Levi, 1977: 196) labijrinthea
- Distal embolic apophysis widened with
bump (Fig. 79). Sternum black with or
without faint white mark in center
(Fig. 81); mountainous regions of Chia-
pas, Mexico (Map 6) (10) lacandon
15(12) Distal embolic apophysis a simple exten-
sion that projects forward, parallel to
the embolus. Keel usuallv rounded
12
22
18 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
(Fig. 129); Puerto Rico to Argentina
and Chile (Map 8) (16) compsa
— Distal embolic apophysis raised up or
projected away from the embolus. Keel
usually pointed or jagged (Figs. 84, 92,
100, 121, 141, 149) 16
16(15) Distal embolic apophysis points up and
away from the embolus. Keel usually
jagged (Figs. 141, 149) 21
— Distal embolic apophysis wide and raised
up on boss. Keel usually pointed (Figs.
84, 92, 100) 17
17(16) Dewlap extension under embolus (Fig.
100) curves under, narrowing the gap
between the embolus and the basal
embolic apophysis by one-half (Fig.
103); north-central Argentina (Map 5)
(13) calarnuchita
No such dewlap (Figs. 84, 92, 110, 121).
Gap between embolus and basal em-
bolic apophysis narrow by less than
one-half the widest distance (Figs. 87,
95, 117, 124) 18
18(17) Outside edge of distal embolic apophysis
gently rounded when viewed from un-
derside of terminal division (Fig. 124);
southern Argentina and southern Chile
(Map 5) (15) karkii
— Outside edge of distal embolic apophysis
with distinct bump when viewed from
underside of terminal division (Figs.
87, 95, 117) 19
19(18) Outside edge of distal embolic apophysis
with rounded bump that is sclerotized
and has a distinct line rising up from
embolus (Fig. 117); Chile and Argen-
tina (Map 7) (14) galatheae
— Outside edge of distal embolic apophysis
with pointed bump, less sclerotized
than embolus proper and without a dis-
tinct line rising up from the embolus
(Figs. 87, 95) 20
20(19) Thick neck joining embolus and basal
embolic apophysis. Bump on distal em-
bolic apophysis peeks out from under
the terminal apophysis (Figs. 92, 95).
Sternum usually with white spot or me-
dian white line (Fig. 97); northern
Chile and southern Peru (Map 7)
(12) tarapaca
— Thin neck joining embolus and basal em-
bolic apophysis. Bump on distal em-
bolic apophysis does not peek out from
under the terminal apophysis (Figs. 84,
87). Sternum always black, general col-
oration dark (Figs. 88, 89); high alti-
tudes in Bolivia and Peru near Lake
Titicaca (Map 5) (11) nigriventris
21(16) Distal embolic apophysis thinner tlian
embolus near die junction of the two.
Embolus tip is curved gendy (Fig.
149); northern Argentina, Uruguay,
and southern Brazil (Map 8) ... (18) gressa
— Distal embolic apophysis the same size as
embolus near the junction of the two.
Embolus tip is curved abruptly (Fig.
141); northern Brazil, French Guiana,
and Colombia (Map 8) (17) roraima
22(11) Distal embolic apophysis a projecting
bump (Figs. 157, 164, 171) 23
— Distal embolic apophysis curved off
(Figs. 178, 185, 199, 214), rounded
(Figs. 192, 206), or absent (Figs. 264,
278, 293) 25
2.3(22) Distal embolic apophysis raised up from
the embolus and pointed (Fig. 171);
California, Arizona, northwestern Mex-
ico (Map 9) (21) gosoga
— Distal embolic apophysis not raised up,
but projecting forward and rounded off
(Figs. 157, 164) 24
24(23) Main flagellum on median apophysis thin
and initially as thick as base. Embolus
and embolic cap shortened (Fig. 164);
northern tip of Peru (Map 11) ... (20) inca
— Main flagellum on median apophysis
thick, but initially thinner than base.
Embolus and embolic cap elongated
(Fig. 157); southern Mexico and Belize
to Costa Rica (Map 14) (19) mnija
25(22) Embolus shaped like the nib on a foun-
tain pen, with bump near tip on op-
posite side of distal embolic apophysis
(Figs. 214, 217); central Mexican pla-
teau (Map 14) (27) atascadero
— Embolus witliout such bump near tip
(e.g.. Fig. 222) 26
26(25) Distal embolic apophysis abruptly ends
in sharp curve and flagellae not set off
on a narrow stalk (Figs. 178, 185, 199,
222) 27
— Distal embolic apophysis gently cui-ved
off (Figs. 229, 236, 243), rounded
(Figs. 192, 206), or absent (e.g.. Fig.
293) 30
27(26) Flagellae veiy thin and equal in length.
Embolus long and arching, with wide
but short embolus cap. Overhanging
terminal apophysis covers only a distal
portion of the sclerotized part of the
embolus (Fig. 199); eastern Cuba and
Hispaniola (Map 14) (25) triangularis
— Flagellae normal in thickness and usually
of different lengths. Terminal apophy-
sis centered above the entire sclero-
tized portion of the embolus (Figs.
178, 185, 222) 28
28(27) Not known to be very social. Height of
embolus plus distal embolic apophysis
just before the latter curves off sharply
is equal to or greater than length of
embolus tip distal to this point (Figs.
Metepeira • Piel
19
178, 185). Sternum with median white
markings (Figs. 182, 189) 29
- Highly social species. Height of embolus
plus distal embolic apophysis just be-
fore die latter curves off shai^ply is less
than the length of the embolus tip dis-
tal to this point (Fig. 222). Sternum
mostly black; sometimes with anterior
white marks. Coxae mostly dark brown
(Fig. 226); mainly in southern Vera-
cruz, Mexico (Map 9) (28) incrassata
29(28) Darker, sclerotized portion of the embo-
lus does not extend over the hump of
the distal embolic apophysis. Base of
embolus narrower than widest part of
the first flagellum (Fig. 178); southern
Veracruz, Mexico to Panama (Map 11)
(22) olmec
- Darker, sclerotized portion of the embo-
lus extends over the hump of the distal
embolic apophysis. Width of embolus
base the same or greater than widest
part of the first flagellum (Fig. 185);
northeastern Mexico to Texas (Levi,
1977, map 1; Map 11) (23) comanche
30(26) Distal emboUc apophysis rounded off to
form convex curve (Figs. 192, 206) 31
- Distal embolic apophysis gently falls off
to form concave shape (Figs. 229, 236,
243) or absent (e.g.. Fig. 293) 32
31(30) Embolus S-shaped (Fig. 192). Sternum
black with posterior white mark. Ven-
ter without white anchor shape inark
(Fig. 196); San Nicolas Island off
southern California (Map 9)
(24) pimiingan
- Embolus convex on upper surface,
straight on lower surface (Fig. 206).
Sternum black with median white line.
Venter with faint white anchor-shaped
mark posterior to median white line
(Fig. 210); southwestern U.S. to cen-
tral Mexico (Map 9) (26) arizonica
32(30) Flagellae on median apophysis set off on
separate, narrow, stalk (e.g.. Figs. 278,
286) 38
- Flagellae on median apophysis not set off
on a separate, narrow, stalk (e.g.. Figs.
229, 264) 33
33(32) Larger flagellum twice as wide as smaller
flagellum (Fig. 236); Isla Socorro of the
Archipielago de Revillagigedo (Map
12) (30) revillagigedo
- Larger flagellum less than twice as thick
as smaller flagellum (e.g.. Figs. 229,
243) 34
34(33) Flagellae on median apophysis relatively
thin (Figs. 250, 264) 35
- Flagellae on median apophysis thicker
(Figs. 229, 243, 257) 36
35(34) Embolus with sharp bend near the tip
(Fig. 264). Sternum with median white
line (Fig. 268); southeast and south-
central Mexico (Map 13) (34) chilapae
— Embolus with gentle bend further from
the tip (Fig. 250). Sternum mosdy
black with small white mark at poste-
rior end of sternum (Fig. 254); Gua-
temala and Costa Rica (Map 12)
(32) uncata
36(34) Embolus with sharp bend near the tip
(Fig. 229); northwestern Mexico and
coastal California (Map 13) ... (29) ventura
— Embolus with gentle bend further from
the tip (Figs. 243, 257) 37
37(36) Sclerotized portion of embolus about as
long as the longer flagellum (Fig. 243);
Yucatan Peninsula (Map 13)
(31) celestun
— Sclerotized portion of embolus shorter
than the longer flagellum (Fig. 257);
northwestern Mexico and California
(Map 12) (33) crassipes
38(32) Embolus thick and tapering to a point
(Figs. 271, 278) 39
— Embolus thin and needlelike (Figs. 286,
293) 40
39(38) Longer flagellum greater than half the
length of the cymbium (Fig. 271); west
coastal Mexico (Map 10) (35) petatlan
— Longer flagellum less than half the length
of the cymbium (Fig. 278); northwest-
ern Mexico and Yucatan Peninsula
(Map 15) (36) minima
40(38) Flageflae set off on long, thin stalk (Fig.
293); Cayman Islands, Jamaica, and
Haiti (Map 15) (38) jamaicensis
— Flagellae set off on short, diicker stalk
(Fig. 286); Honduras to Costa Rica
(Map 10) (37) pacifica
Metepeira foxi Group
The M. foxi species group {sensu Levi,
1977) and M. desenderi share veiy similar
genitaha, especially among males. Conse-
quently, I am expanding the M. foxi group
to include M. desenderi, but with the ex-
clusion of the black sternum as a diagnos-
tic character. The M. foxi group {sensu
lato) includes males with an embolus that
lacks a distal apophysis. The embolus curls
almost 180° clockwise around a reduced or
fleshy terminal apophysis (Figs. 7, 15, 22,
29). In other species groups the embolus
is straighter and does not curl (e.g.. Fig.
60), and the terminal apophysis is large,
overhanging the embolus, and often scler-
otized with teeth (e.g.. Fig. 121). Epigynal
20
Bulletin Museum of Comparative Zoologtj, Vol. 157, No. 1
features uniting females in the M. foxi spe-
cies group are harder to discern. Generally
speaking, the epigynum has a stubby,
shorter, often triangular scape and weaker
posterior lobes that permit a wider view
into the epigynal openings (compare Figs.
9, 17, 24, 31 with Figs. 143, 252).
1 . Metepeira datona
Chamberlin and Ivie
Figures 7-13, 312; Map 1
Metepeira datona Chamberlin and Ivie, 1942: 68, fig.
196, ? . Female holotype from Daytona Beach,
Florida, USA, in the AMNH, examined. Levi,
1977: 208-210, figs. 78-86, S, 9. Brignofi, 1983:
275,
Metepeira inenna Bryant, 1945: 378. Female holo-
type from Cap Haitian, Haiti in the MCZ, exam-
ined. First synonymized by Levi, 1977: 208—210.
Description. Female from Morant
Point, Mammee Bay, Jamaica. Carapace
light around eyes with short lateral poste-
rior extensions, sometimes with thin lon-
gitudinal white line (Fig. 12). Legs ringed.
Femur I with row of two macrosetae on
anterior side; three light setae on anter-
oventral side. Anterior half of dorsal foli-
um white, posterior half black, margined
by white; two halves separated by trans-
verse white line. Abdomen widest in cen-
ter (Fig. 12). Venter with a longitudinal
white line. Pair of white spots on either
side of spiracle (Fig. 13). Sternum usually
entirely black, though soinetimes with thin
anterior and posterior marks, suggestive of
median white line (Fig. 13). Ratio of eye
diameters: posterior medians and anterior
medians 1.0, anterior laterals 1.3, posterior
laterals 1.1. Anterior median eyes separat-
ed by 1.4 diameters, posterior median eyes
by 0.9, anterior median eyes separated
from anterior laterals by 1.6 diameters of
anterior lateral eyes, lateral eyes separated
by 0.3 their diameters. Total length 4.2
mm. Carapace 1.7 mm long, 1.4 wide.
First femur 2 mm, patella and tibia 2.4,
metatarsus 1.5, tarsus 0.8. Second patella
and tibia 1.9 inm, third 1.1, fourth 1.6.
Male from Morant Point, Mammee Bay,
Jamaica. Carapace yellowish brown, light
around eyes. Median white mark anterior
to thoracic furrow (Fig. 10). Legs same
color as carapace, lightly ringed. Femur I
with row of three macrosetae on anterior
side, two on anteroventral side. Center of
dorsum with transverse white line; poste-
rior half darker than anterior half; thin me-
dian black line; margin of folium white
(Fig. 10). Venter, sternum as in female
(Fig. 11). Ratio of eye diameters: posterior
medians and anterior medians 1.0, anterior
laterals 1.3, posterior laterals 1.2. Anterior
median eyes separated by 1.8 diameters,
posterior median eyes by 1.1, anterior me-
dian eyes separated from anterior laterals
by 1.5 diameters of anterior lateral eyes,
lateral eyes separated by 0.5 their diame-
ters. Total length 3 mm. Carapace 1.4 mm
long, 1.1 wide. First femur 2.7 mm, patella
and tibia 2.7, metatarsus 1.9, tarsus 0.8.
Second patella and tibia 2.1 mm, third 1.0,
fourth 1.3.
Diagnosis. The dorsal folium and ab-
dominal shape is distinctive in M. datona.
Typically, the females abdomen is widest
in the middle, forming a rhomboid in
shape (Fig. 12), and both sexes have a
transverse white line dividing the dorsal
folium, with wide black markings on the
posterior half (Figs. 10, 12). A ventral view
of the epigynum in M. datona reveals a
ridge under the scape that almost forms a
straight line and which cui'ves up at the
ends (Fig. 9) instead of a V-shape (Figs.
17, 24, 31). The openings to the epigynum
consist of small slits that flank a wide de-
pression (Fig. 9), compared to larger
openings that are relatively closer together
(Figs. 17, 24, 31). The smooth embolus on
M. datona is somewhat more compressed
as it curls clockwise like the tip on a cork-
screw, in contrast to the slightly ridged
embolus supported on a stalk (Fig. 15) or
the strongly ridged and tilted embolus
(Figs. 22, 29). Like M. desenderi (Fig. 15)
but unlike M. grandiosa grandiosa (Fig.
22) and M. grandiosa alpina (Fig. 29), tlie
posteroventral edge of the keel on the me-
dian apophysis is rounded, as opposed to
pointed (Fig. 7). Finally, M. datona has the
Metepeira • Piel
21
thinnest and fleshiest terminal apophysis
in the genus (Fig. 7).
Variation. Average body length of 15 fe-
males examined 3.8 mm, range 2.9 to 4.8
mm. Average body length of seven males
examined 2.6 mm, range 1.8 to 3 mm.
Natural History. Mature adults have
been collected throughout the year (Fig.
312). According to locality labels, M. da-
tona are found near the beach on cactus,
palm, low scrub, mangrove, and bamboo.
Females are known to wrap their egg sacs
in dead leaves.
Distribution. At sea level in Florida, Ba-
hamas, British West Indies, and the Do-
minican Republic (Levi, 1977, map 2; Map
1).
Records Examined. BAHAMAS Abaco Cays: Aba-
co, 26°29'N, 77°5'W, 2.xii.l964 (W. B. Peck, MCZ);
Allons Cay, 26°59'N, 77°40'W, 9.V.1953 (E. B. Hay-
den, AMNH); Hopetown, Elbow Cay, 26°33'N,
76°57'W, 5.V.1953 (E. Hayden, AMNH); New Plym-
outh, Green Turtle Cay, 26°50'N, 77°23'W, 7.V.1953
(G. Rabb, AMNH); Whale Cay 26°43'N, 77°14'W,
12.i.l964 (W. B. Peck, CAS). Acklins Id.: Atwood's
Harbor, 22°13'N, 74°18'W, 15.ix.l958 (A. W. Scott Jr.,
MCZ); Salina Point, 22°13'N, 74°18'W, 15.viii.l958
(R. Robertson & A. W. Scott Jr, MCZ). Andros: Fresh
Creek, 24°26'N, 77°57'W, 23.iv.1953 (L. Giovannoh,
AMNH); Mangrove Cay, 24°15'N, 77°39'W,
26. iv. 1953 (E. Hayden, AMNH); Nicolls Town,
25°8'N, 78°0'W, 14.iii.l967 (A. M. Nadler, AMNH).
Berry Islands: Frazier's Hog Cay, 25°24'N, 77°50'W,
29.iv.1953 (E. Hayden, AMNH); Little Harbor Cay,
25°34'N, 77°43'W, l.vl953 (Hayden & Giovannoh,
AMNH). Crooked Island: North shore of Cripple
Hill, 22°49'N, 74°16'W, 15.ix.l958 (A. W. Scott Jr,
MCZ); NW end, Gordon (= Gun) Bluff, 22°50'N,
74°20'W, 15.viii.l958 (R. Robinson & A. W. Scott Jr,
MCZ). Crooked Island Group: Long Cay 22°37'N,
74°20'W, 7.iii.l953 (Hayden & Giovannoh, AMNH).
East Plana Cay: 22°37'N, 73°.33'W, 4.iii.l953 (E.
Hayden, AMNH). Exurnas: Musha Cay 23°50'N,
76°15'W, 29.xii.1985 (A. Boutard, MCZ); Warderick
Wells Cay 24°22'N, 76°36'W, 9.i.l953 (L. Giovannoh,
AMNH), ll.i.l953 (Hayden & Giovannoh, AMNH).
Grand Bahamas Island: Dundee Bay, 26°30'N,
79°15'W [?], 25.xii.1965 (L. Pinter, MCZ); near Fre
port, pine-palmetto, 26°34'N, 78°27'W [?],
25.vii.1965 (L. Pinter, MCZ). Long Island: Clarence
Town, 23°6'N, 74°59'W, 10.iii.l953 (L. Giovannoh,
AMNH); Deadman's Cay, 23°14'N, 75°14'W,
ll.iii.l953 (E. Hayden, AMNH). New Providence Is-
land: 7 mi W of Nassau, 25°5'N, 77°28'W, 4.i.l953
(Hayden & Giovannoli, AMNH). North Bimini:
25°44'N, 79°15'W, 25.i.l950 (C. M. Bogert, AMNH),
6.vi.l950 (M. Gazier & F. Rindge, AMNH), 15.V.1951
(W. J. Gertsch & M. Gazier, AMNH), 15.vi.l951 (M.
Gazier, P & C. Vaurie, AMNH), 15.vii.l951 (C. & P
Vaurie, AMNH), 13.xii.l952 (A. M. Nadler, AMNH),
28.xi. 19.59 (A. M. Nadler, AMNH). Rmn Cay: near
Port Nelson, 2.3°38'N, 74°50'W, 16.ih.l953 (Hayden
& Giovannoli, AMNH). South Bimini: 25°42'N,
79°17'W, 12.vi.l950 (M. Gazier & F Rindge,
AMNH), 22.vi.1950 (M. Gazier & F. Rindge,
AMNH), 15.V.1951 (W. J. Gertsch & M. Gazier,
AMNH), 15.vii.l951 (C. & P Vaurie, AMNH),
4.vLii.l951 (C. & P Vaurie, AMNH), 4.xii.l952 (A. M.
Nadler, AMNH), 25.ih.1953 (A. M. Nadler, AMNH);
Gun Cay 25°35'N, 79°20'W, 15.vi.l951 (AMNH).
BRITISH WEST INDIES Caicos Islands: Long Cay,
21°28'N, 71°33'W 10.ii.l953 (E. Hayden, AMNH);
South Caicos, from webs in upper beach zone,
21°31'N, 71°30'W, 3.iv.l973 (D. W Buden, MCZ);
West Caicos, 21°39'N, 72°28'W, 4.U.1953 (Hayden &
Giovannoh, AMNH), 5.ii.l953 (Hayden, Rabb, &
Giovannoli, AMNH). Grand Cayman Island:
19°20'N, 81°10'W, 15.ii.l960 (R. A. Levidn, MCZ).
CUBA Oriente: Banes, 20°58'N, 75°43'W, 2.viii.l955
(A. F. Archer, AMNH); Ensenada Nispero, Giudamar,
19°58'N, 75°52'W, 9.xi.l945 (P Alayo, AMNH); Jur-
agua, 19°56'N, 75°40'W, l.x.1955 (P Alayo, AMNH);
Santa Fe [?], 20°22'N, 75°53'W (A. F Archer,
AMNH). DOMINICAN REPUBLIC Barahona: Pla-
ya Los Patos, 17°58'N, 71°11'W, 31.viii.l976 (J. A.
Ottenwalder, MNSD). La Altagracia: Punta Cana,
Isla Saona, 18°8'N, 68°34'W (Felix E. Del Monte &
K. Guerrero, MNSD). HAITI Dept. du Nord: Cap-
Haitien, 19°46'N, 72°13'W, 15.iii.l934 (E. Bryant,
Utawana Exp., MCZ). JAMAICA Portland: between
Boston and Blue Hole, 18°6'N, 76°37'W, 29.vii.1955
(A. F Archer, AMNH). Saint Andrews: Hope Gar-
dens, Gordontown, 18°2'N, 76°45'W, 27.vii.1955 (A.
F Archer, AMNH). Surrey: Morant Point, Mammee
Bay, 17°53'N, 76°19'W, 14.X.1957 (A. M. Chickering,
MCZ); Palisadoes, 17°56'N. 76°46'W, ll.xi.l967 (A.
M. Chickering, MCZ); Palisadoes Area, 17°56'N,
76°46'W, l.xi.l957 (A. M. Chickering, MCZ). U.S.
VIRGIN ISLANDS Saint Thomas: Morant Point,
17°55'N, 76°10'W, 25.vii.1985 (G. B. Edwards,
FSCA).
2. Metepeira desenderi Baert
Figures 14-21, 307; Map 1
Epeira labyrinthea:— Banks, 1902: 60. Banks, 1924:
97. Misidentification.
Metepeira sp.: — Roth and Craig, 1970: 116.
Metepeira desenderi Baert, 1^87: 145, figs. 16-21, S ,
9 . Male holotype from Isla Pinzon, Galapagos, Ec-
uador, in the IRSNB. Platnick, 1989: 341.
Note. Holotype not examined because the figures
in Baert (1987) are clear and because this is the
only Metepeira species found on the Galapagos.
Description. Female from east slope of
Isla Santa Cruz, Galapagos Islands, Ecua-
22 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
dor. Carapace yellowish brown, white
around eyes, lighter median line (Fig. 20).
Legs yellowish white, slightly darker an-
nulations on distal ends of articles. Femur
I with row of four macrosetae on anterior
side; two to four on anteroventral side.
Dorsal fleur-de-lis pattern broken into four
white patches with anterior pair often larg-
er than in most species. Posterior pair
straighter than usual, forming a cross in
center of folium (Fig. 20). Venter olive-
brown with median white line surrounded
by white U-shaped marking (Fig. 21). Pair
of white spots on either side of spiracle.
Sternum brownish black with wide, white
line widening anteriorly (Fig. 21). Ratio of
eye diameters: posterior medians and an-
terior medians 1.0, anterior laterals 1.3,
posterior laterals 1.3. Anterior median
eyes separated by 1.5 diameters, posterior
median eyes by 0.9, anterior median eyes
separated from anterior laterals by 2.3 di-
ameters of anterior lateral eyes, lateral eyes
separated by 0.2 their diameters. Total
lengdi 5.5 mm. Carapace 2.3 mm long, 1.7
wide. First femur 2.7 mm, patella and tibia
3, metatarsus 2.2, tarsus 0.8. Second pateUa
and tibia 2.4 mm, third 1.4, fourth 2.
Male from same locality as female. Car-
apace yellowish brown with wide white
median mark (Fig. 18). Legs ringed like
female. Femur I with row of four macro-
setae on anterior side; four to seven on an-
teroventral side. Dorsum and venter as in
female, though median white line on ster-
num sometimes broken (Figs. 18, 19). Ra-
tio of eye diameters: posterior medians
and anterior medians 0.9, anterior laterals
1.2, posterior laterals 1.1. Anterior median
eyes separated by 2.2 diameters, posterior
median eyes by 1.4, anterior median eyes
separated from anterior laterals by 2.3 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.5 their diameters. To-
tal length 3.2 mm. Carapace 1.6 mm long,
1.3 wide. First femur 2.6 nam, patella and
tibia 2.6, metatarsus 2.3, tarsus 0.8. Sec-
ond patella and tibia 2.2 mm, third 1.0,
fourth 1.6.
Diagnosis. Within the M. foxi species
40 80 120 160
Collection day (since Jan 1 )
200
Figure 14. The length of mature M. desendeh collected on
specific days of the year over a period of 84 years. There is a
trend from smaller spiders early in the season to large spiders
later in the season.
Scale of abscissa: -80 = October 12 of the previous year; 40
= February 9; 160 = June 9.
Symbols: Females (o), stippled regression line (t^ = 40%);
males (•), solid regression line (i^ = 42%).
group, the female abdomen shape of M.
desenderi is closest to M. datona, though
not quite as rhomboid (compare Fig. 20
with Figs. 12, 27, 34). The scape on M.
desenderi is narrower and less triangular
than other M. foxi species, and the de-
pression under the scape forms a distinct
U-shaped smile that is not seen in the oth-
ers (compare Fig. 17 with Figs. 9, 24, 31).
Like M. datona (Fig. 7) but unUke M.
grandiosa grandiosa (Fig. 22) or M. gran-
diosa alpina (Fig. 29), the posteroventral
edge of the keel on the median apophysis
is rounded as opposed to pointed (Fig. 15).
Similar to M. datona (Fig. 7), the embolus
of M. desenderi is curved like a slightly
flattened corkscrew. Unlike M. datona, the
embolus seems to be supported by a ped-
icel off a large basal embolic apophysis
(Fig. 15). In contrast, a pedicel is not ev-
ident in M. datona, and the basal embolic
apophysis is barely visible (Fig. 7).
Variation. Average body length of 45 fe-
males exaiTiined 5.8 mm, range 3.8 to 8.5
mm. Average body length of 13 males ex-
amined 4.9 mm, range 2.9 to 6.5 mm.
Natural History. Notes on the collection
labels of Y. Lubin and R. Silberglied in-
dicate that M. desenderi is active at night
Metepeira • Piel
23
and that the retreat is composed of Opun-
tia bark and croton leaves. According to
Baert (1987), M. desenderi is found in
large numbers in arid ecological zones on
all islands. Although some mature speci-
mens have been collected in November
and August, most specimens are found be-
tween Januaiy and June (Fig. 307), cor-
responding to the "warm and wet" season
in the Galapagos (van der Werff, 1978).
The size of mature M. desenderi speci-
mens appears to correlate with the collec-
tion date: female spiders taken in August
are, on average, 80% larger than those tak-
en in November (Fig. 14). This trend may
indicate that M. desenderi can vary the
number of instars before maturity. Alter-
natively, the intermolt growth may be
greater for spiders with their antepenulti-
mate or penultimate instars occurring dur-
ing the warmer and wetter season.
Distribution. Endemic to the Galapagos
Islands (Map 1).
Records Examined. ECUADOR Galapagos: Albe-
marle, Tagus Cove, 0°16'S 91°22'W, 23.1.1899
(AMNH), 8.ii.l899 (AMNH), 21.iii.l899 (AMNH),
23.iii.1899 (AMNH); Archipielago de Galapagos,
0°0'N, 90°30'W (Williams Exped., 1923, MCZ); Ba-
hia Conway, Indefatigable Island, 0°33'S, 90°32'W,
17.iii.l935 (Exline-Peck, CAS); Barrington Island,
0°49'S, 90°4'W, l.viii.l929 (H. H. Cleaves, CAS);
Campion, nr. Floreana (Santa Maria), 1°15'S,
90°27'W, l.vi.l981 (Y. Lubin, MCZ); Charles Island,
1°17'S, 90°26'W, 10.V1899 (AMNH); Indefatigable
Island, 0°38'S, 90°23'W, 27.iv.1899 (AMNH),
18.vi.l929 (Pinchot South Sea Exp, USNM); Isla
Abingdon, 0°35'N, 90°44'W, 25.vi.1899 (AMNH); Isla
Albemarle, 0°30'S, 91°4'W, 13. i. 1899 (AMNH),
20.ii.l899 (AMNH), 20.iii.l899 (AMNH); Isla Bin-
dloe, 0°19'N, 90°29'W, 20.vi.l899 (AMNH); Isla
Hood, 1°23'S, 89°39'W, 18.V.1899 (AMNH),
26.V1899 (AMNH); Isla Pinta, S Coast, 0°35'N,
90°44'W, 25.V1964 (D. Q. Cavagnaro, CAS); Isla Pla-
za, 0°35'S, 90°9'W, 7.iii.l970 (R. Silberglied, MCZ),
26.xi.1973 (Y. Lubin, MCZ); Isla Santa Cruz, Acade-
my Bay, 0°44'30"S, 90°17'30"W, 13.iii.l970 (R. Sil-
berglied, MCZ); Isla Santa Cruz, E slope, 0°38'S,
90°23'W, 16.ivl964 (D. Q. Cavagnaro, CAS); Isla
Santa Cruz, Estacion Cientifica Charles Darwin,
0°44'S, 90°18'W, 24.i.l964 (D. Q. Cavagnaro & R. O.
Schuster, CAS), 12.ii.l964 (Cavagnero & Schuster,
CAS), 3.xi.l973 (Y. Lubin, MCZ), 24.xi.1973 (Y Lu-
bin, MCZ), 25.xi.1973 (Y. Lubin, MCZ); Isla Santa
Fe, S coast, 0°50'S, 90°4'W, 30.i.l983 (Y. Lubin,
MCZ); James, 0°16'S, 90°42'W, 22.ivl899 (AMNH);
Narborough Island, 0°25'S, 91°30'W, 28.iii.1899
(AMNH); Sombrero Chino, Rocas Bainbridge, SE of
Santiago, 0°21'S, 90°34'W, 31.iii.l970 (R. Silberglied,
MCZ); Tower Island, Darwin's Bay [?], 0°17'N,
89°59'W (AMNH); Tower Islands, 0°20'N, 89°58'W,
7.ivl925 (N.Y. Zoological Society, AMNH); W coast
of Albemarle Island, 0°11'N, 91°21'W, 9.iii.l935
(AMNH).
3. Metepeira grandiosa grandiosa
Chamberrm & Ivie
Figures 22-28, 321; Map 2
Metepeira grandiosa Chamberlin and Ivie, 1941: 17,
figs. 24—26, 9 . Female holotype from Ben Lo-
mond, California, USA in the AMNH, examined.
Metepeira palomara Chamberlin and Ivie, 1942: 72,
figs. 200-204, 9, 6. Female holotype and para-
types from Mt. Palomar, California, in the AMNH,
examined. First synonymized by Levi, 1977: 214.
Metepeira grandiosa grandiosa: — Levi, 1977: 214,
figs. 112-116, ?, (J.
Note. Levi (1977) opted to collapse M. paliistris,
M. grandiosa, M. alpina, M. dakota, and M. palo-
mara into three subspecies of M. grandiosa, with
the caveat that more data may show the three sub-
species to be distinct species. D. Buckle (personal
communication) claims that his own recent obser-
vations suggest that M. grandiosa alpina and M.
grandiosa palustHs are separate species. However,
since the bulk of M. grandiosa specimens are out-
side of the geographic range of this revision, and
in the absence of molecular data, I will follow
Levi's (1977) recommendation and leave these as
separate subspecies.
Description. Female from Los Angeles,
California, USA. Carapace light around
eyes with inedian white line extending to
thoracic furrow (Fig. 27). Legs ringed. Fe-
mur I with row of three to four macrosetae
on anterior; one on anteroventral. Dorsum
with usual folium, though lighter and more
speckled than in most species (Fig. 27).
Venter with a wide longitudinal white line.
Pair of white spots on either side of spi-
racle (Fig. 28). Sternum entirely black
(Fig. 28). Ratio of eye diameters: posterior
medians and anterior medians 1.0, anterior
laterals 1.2, posterior laterals 1. Anterior
median eyes separated by 1.9 diameters,
posterior median eyes by 1.3, anterior me-
dian eyes separated from anterior laterals
by 3 diameters of anterior lateral eyes, lat-
eral eyes separated by 0.4 their diameters.
Total length 6.5 mm. Carapace 3.2 mm
24 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
long, 2.3 wide. First femur 3.5 mm, patella
and tibia 3.7, metatarsus 3.2, tarsus 1.1.
Second patella and tibia 3.2 mm, third 1.9,
fourth 2.8.
Male from Parque Nacional Sierra San
Pedro Martir, Baja California Norte, Mex-
ico. Carapace as in female. Legs lightly
ringed. Macrosetae on femur I variable —
usually row of four macrosetae on anterior
side, five on anteroventral side. Dorsum as
in female (Fig. 25). Venter and sternum as
in female (Fig. 26). Ratio of eye diameters:
posterior medians and anterior medians
1.1, anterior laterals 1.5, posterior laterals
1.5. Anterior median eyes separated by 1.7
diameters, posterior median eyes by 0.8,
anterior median eyes separated from an-
terior laterals by 2.1 diameters of anterior
lateral eyes, lateral eyes separated by 0.3
their diameters. Total length 3.6 mm. Car-
apace 1.9 mm long, 1.5 wide. First femur
2.9 mm, patella and tibia 2.8, metatarsus
2.5, tarsus 0.9. Second patella and tibia 2.3
mm, third 1.3, fourth 1.9.
Diagnosis. The epigynal scape of M.
grandiosa grandiosa is shorter and stub-
bier than in other members of the M. foxi
species group (compare Fig. 24 with Figs.
9, 17, 31). Unlike M. datona and M. de-
senderi (Figs. 7, 15), the corkscrew em-
bolus is tilted up with a heavy ridge (Fig.
22), but unlike M. grandiosa alpina (Fig.
29), it is more graceful and not as sharply
bent. The posteroventral edge of the keel
on the median apophysis in M. grandiosa
grandiosa is not as pointed (Fig. 22) as in
M. grandiosa alpina (Fig. 29) but not
curved as in M. datona and M. desenderi
(Figs. 7, 15).
Variation. Body length of females varies
from 5.4 to 8.5 mm; males from 3.5 to 5.1
mm (Levi, 1977).
Natural History. Mature specimens
have been collected from March to Sep-
tember (Fig. 321; Levi, 1977) in yellow
pine forests and on Eriogonum fascictda-
tum bushes. Elevations range from sea lev-
el to 2,000 m.
Distribution. Coastal mountainous re-
gions from British Columbia to Baja Cal-
ifornia Norte (Levi, 1977, map 2; Map 2).
Records Examined. MEXICO Baja California Nor-
te: Parque Nacional Sierra San Pedro Martir,
30°45'N, 115°13'W, l.vii.l977 (C. E. Griswold, CAS).
USA Arizona: Sycamore Canyon [?] Santa Cruz Co,
31°28'N, 110°42'W, 9.ix.l978 (G. F. Knowlton,
MCZ). California: Los Angeles, 34°3'N, 118°15'W
(Davidson, MCZ); Winchester, Double Butte,
33°42'N, 117°5'W, 20.iv,1974 (W. Icenogle, MCZ).
4. Metepeira grandiosa alpina
Chamberlin and Ivie
Figures 29-35, 325; Map 2
Metepeira dakota Chamberlin and Ivie, 1942: 73, figs.
205—207, ? , c? . Male holotype and female paratype
from Noonan, North Dakota, USA, in the AMNH,
examined. Name synonymized by Levi, 1977: 212—
214.
Metepeira alpina Chamberlin and Ivie, 1942: 74. Fe-
male holotype and female paratypes from Fish
Lake, Utah, USA, in tlie AMNH, examined.
Metepeira grandiosa alpina: — Levi, 1977: 212-214,
figs. 99, 100, 106-111, 9,6. BrignoH, 1983: 276.
ISlote. As first revisor, Levi (1977) chose to use
the name M. alpina because its type specimen was
collected closer to the center of the subspecies dis-
tribution.
Description. Female from Charcas, San
Luis Potosi, Mexico. Carapace light
around eyes with lateral posterior exten-
sions and median white line extending td
thoracic furrow (Fig. 34). Legs lightly
ringed. Femur I with row of four macro-
setae on anterior side; one on anteroven-
tral side. Dorsum with usual folium,
though lighter than in most species (Fig.
34). Venter with a wide longitudinal white
line. Pair of white spots on either side of
spiracle (Fig. 35). Sternum entirely black
(Fig. 35). Ratio of eye diameters: posterior
medians and anterior medians 1.0, anterior
laterals 1.1, posterior laterals 1.1. Anterior
median eyes separated by 1.7 diameters,
posterior median eyes by 0.7, anterior me-
dian eyes separated from anterior laterals
by 2.8 diameters of anterior lateral eyes,
lateral eyes separated by 1.1 their diame-
ters. Total length 6.7 mm. Carapace 3 mm
long, 2.4 wide. First femur 3.5 mm, patella
and tibia 3.8, metatarsus 3.4, tarsus 1.2.
Metepeira • Piel 25
desenderi
(2)
grandiosa grandiosa
(3)
m^ grandiosa alpina
V (4)
Figures 15-21. Metepeira desenderi Baert (sp. 2; 0°38'S, 90°23'W). 15, male palpus, mesal. 16, epigynum, posterior. 17,
epigynum, ventral. 18, male, dorsal. 19, male, ventral. 20, female, dorsal. 21, female, ventral.
Figures 22-28. Metepeira grandiosa grandiosa Chamberlin and Ivie (sp. 3 [22, 25, 26] 30°45'N, 115°13'W; [23, 24, 27, 28]
34°3'N, 118°15'W). 22, male palpus, mesal. 23, epigynum, posterior. 24, epigynum, ventral. 25, male, dorsal. 26, male, ventral.
27, female, dorsal. 28, female, ventral.
Figures 29-35. Metepeira grandiosa alpina Chamberlin and Ivie (sp. 4 [29,32,33] 25°56'N, 105°22'W; [30,31,34,35] 23°8'N,
101°7'W). 29, male palpus, mesal. 30, epigynum, posterior. 31, epigynum, ventral. 32, male, dorsal. 33, male, ventral. 34, female,
dorsal. 35, female, ventral.
Scale bars: dorsum and venter figures 1 .0 mm.
26 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
10000
1000
■I 100 ;
° vigilax
X rectangula
• cajabamaba
JQ I I I I ''I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I
-45 -35 -25 -15 -5 5 15 25
Latitude (Degrees North)
Figure 36. The elevation of collection localities for M. vigilax,
M. rectangula, and M. cajabamba at their corresponding lati-
tudes. Species-specific altitudes appear to decrease with dis-
tance from the equator. Elevations estimated from NOAA da-
tabase of 5- by 5-minute geographic tiles. Regression line of
M. vigilax does not include data points north of the equator.
Symbols: M. vigilax (o), M. rectangula [X], M. cajabamba [•].
Second patella and tibia 3.3 mm, third 1.9,
fourth 2.9.
Male from Santa Maria del Oro, Duran-
go, Mexico. Carapace yellowish brown,
light around eyes. Median white triangular
mark anterior to thoracic furrow (Fig. 32).
Legs same color as carapace. Macrosetae
on femur I variable — usually row of four
macrosetae on anterior side, five to six on
anteroventral side. Dorsum, venter, and
sternum as in female (Figs. 32, 33). Ratio
of eye diameters: posterior medians and
anterior medians 1.0, anterior laterals 1.4,
posterior laterals 1.2. Anterior median
eyes separated by 1.5 diameters, posterior
median eyes by 0.8, anterior median eyes
separated from anterior laterals by 2.3 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.3 their diameters. To-
tal length 5.2 mm. Carapace 2.5 mm long,
2 wide. First femur 4.5 mm, patella and
tibia 4.6, metatarsus 4.5, tarsus 1.3. Sec-
ond patella and tibia 3.6 mm, third 1.8,
fourth 2.8.
Diagnosis. The epigynal scape of M.
grandiosa alpina has the widest base of all
members of the M. foxi species group, and
the lobes surrounding the central depres-
sion are soiuewhat fatter (compare Fig. 31
witli Figs. 9, 17, 24,). Unlike M. datona
and M. desenderi (Figs. 7, 15), the cork-
screw embolus is tilted up with a heavy
ridge (Fig. 29), but unlike M. grondiosa
grandiosa (Fig. 22), it is thicker and with
a sharper L-shape bend instead of a more
cui'ved L-shape. The posteroventral edge
of the keel on the median apophysis in M.
grandiosa alpina is pointier (Fig. 29) than
in M. grandiosa grandiosa (Fig. 22) and
not curved as in M. datona and M. desen-
deri (Figs. 7, 15).
Variation. Body length of females varies
from 4.0 to 6.8 mm; males from 3.1 to 5.3
mm (Levi, 1977).
Natural History. According to Levi
(1977), mature specimens have been col-
lected from June to August in meadows of
bunchgrass, browsed aspen, oak, juniper,
and sagebrush (Fig. 325). Elevations are at
around 2,000 m.
Distribution. North American Rockies
from southern Alberta and Saskatchewan
to Central Mexico (Levi, 1977, map 2;
Map 2).
Records Examined. MEXICO Durango: Santa Ma-
ria del Oro, 25°56'N, 105°22'W, 28.vii.1947 (W. J.
Gertsch, AMNH). San Luis Potosi: Charcas, moun-
tain side, 23°8'N, 101°7'W, 7.vii.l934 (MCZ). USA.
Colorado: Cimarron, 38°27'N, 107°33'W, 21.vii.l959
(H. W. & L. Levi, MCZ); Hayden Creek, Sangre de
Cristo Mtns., 38°25'N, 105°35'W [?], Il.vii.l961 (H.
W. & L. Levi, MCZ). Utah: SE shore of Bear Lakei
41°59'N, 111°20'W, 3.vii.l978 (G. F. Knovi^lton,
MCZ).
Metepeira vigilax Group
Spiders in the M. vigilax group (Mete-
peira cajabamba, Metepeira glome rabilis,
Metepeira vigilax, Metepeira rectangula)
are characterized by large, sclerotized epi-
gynal openings, not unlike a snails shell
(Figs. 40, 48, 55, 62), and long, robust,
emboli with large scooplike basal embolic
apophyses (Figs. 38, 46, 53, 60).
5. Metepeira cajabamba new species
Figures 36, 38-45, 315; Map 3
Holotype. Male from Cajabamba, Cajamarca, Peru,
25. Lx. 1955, W Weyi-auch, in CAS. The specific
name is a noun in apposition after the localit)'.
Description. Female paratype from Ca-
jabamba, Cajamarca, Pei-u. Carapace dark
Metepeira • Piel 27
brown, light around eyes with lateral pos-
terior extensions (Fig. 43). Proximal two-
thirds of femora white, remainder dark
browii. Remaining articles lightly annulat-
ed. Femur I with row of three macrosetae
on anterior side; none on anteroventral
side. Dorsal folium dark; white fleur-de-lis
pattern reduced with thin branches (Fig.
43). Venter of abdomen black with wide
white median line. Pair of small white
spots on either side of colulus (Fig. 44).
Sometimes the posterior end of median
line ends in anchor shape (Fig. 45). Ster-
num black, sometimes with (Fig. 44) or
without (Fig. 45) median white line. Ratio
of eye diameters: posterior medians and
anterior medians 1.1, anterior laterals 1.4,
posterior laterals 1.2. Anterior median
eyes separated by 1.6 diameters, posterior
median eyes by 0.9, anterior median eyes
separated from anterior laterals by 3 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.2 their diameters. To-
tal length 6.4 mm. Carapace 3 mm long,
2.3 wide. First femur 3.3 mm, patella and
tibia 3.5, metatarsus 2.9, tarsus 0.9. Sec-
ond patella and tibia 2.9 mm, third 1.7,
fourth 2.7.
Male holotype. Carapace dark brown,
white around eyes with lateral posterior
extensions and median white mark (Fig.
41). Proximal third of femora white, re-
mainder dark brown. Remaining articles
lightly annulated. Femur I with row of
three macrosetae on anterior side; one on
anteroventral side. Dorsal folium dark;
white fleur-de-lis pattern indistinct with
thin and broken branches (Fig. 41). Venter
of abdomen black with median white
mark. Pair of small white spots on either
side of spiracle and colulus (Fig. 42). Ster-
num black, often with median white line
(Fig. 42). Ratio of eye diameters: posterior
medians and anterior inedians 1.0, anterior
laterals 1.3, posterior laterals 1. Anterior
median eyes separated by 1.5 diameters,
posterior median eyes by 0.9, anterior me-
dian eyes separated from anterior laterals
by 2.3 diameters of anterior lateral eyes,
lateral eyes separated by 0.3 their diame-
ters. Total length 4 mm. Carapace 1.9 mm
long, 1.5 wide. First femur 3 mm, patella
and tibia 3, metatarsus 2.7, tarsus 0.9. Sec-
ond patella and tibia 2.4 mm, third 1.2,
fourth 1.8.
Diagnosis. Female M. cajahamba and
M. glomerabilis differ from the other spe-
cies in the M. vigilax group (M. vigilax and
M. rectangula) by the smaller and more
tubelike openings to the epigynum (com-
pare Figs. 40, 48 with Figs. 55, 62). Fe-
male M. cajabamha differs froin M. glom-
erabilis by having epigynal openings that
are more oval (Fig. 40) than round (Fig.
48) when viewed ventrally, having the epi-
gynal openings farther apart (compare Fig.
40 with Fig. 48), and by having the scler-
otized tubelike openings more anteriorly
directed (Fig. 40) than parallel to the epi-
gynal groove (Fig. 48). Male M. cajabarnba
and M. glomerabilis differ from other spe-
cies in the M. vigilax group by the smaller,
thinner, and more graceful emboli (com-
pare Figs. 38, 46 with Figs. 53, 60). Me-
tepeira cajahamba differs from M. glom-
erabilis by lacking a keel on the median
apophysis, an only slightly fatter embolus,
and having a normal embolus cap, in con-
trast to a winged embolus cap (Fig. 46). A
larger portion of the prosoma is dark in M.
cajahamba (Fig. 43) as compared to M.
glom,erabilis (Fig. 51).
Variation. Average body length of nine
females examined 6.2 mm, range 5.4 to 7.5
mm. Average body length of four males
examined 4 mm, range 3.7 to 4.9 mm.
Natural History. With the exception of
two specimens, mature M. cajabarnba
specimens have been collected in May
through October (Fig. 315). Altitudes of
collection localities appear to correlate
steeply with latitude (Fig. 36). Median el-
evation, about 500 m, with a range from
near sea level to 3,500 m.
Distribution. Ecuador and Peru (Map
3).
Records Examined. PERU Ancash: Callejon de
Huaylas, 9°10'S, 77°45'W, 15.viii.l988 (V. and B.
Roth, CAS). Cajamnrca: Cajabarnba, 12.427, 7°37'S,
78°3'W, 25.ix.1955 (W. Weyrauch, CAS). La Libet-tad:
28 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
Cerro Campana, La Cumbre, 8°1'S, 79°5'W,
10.x. 1966 (A. F. Archer, AMNH). Liiiia: 20 km E An-
con, 11°47'S, 76°59'W, l.xi.l953 (W. Weyrauch,
CAS); 23 mi N Pativilca, 10°42'S, 77°47'W, 15.1.1955
(E. I. Schlinger and E. S. Ross, CAS); 5 km NW
Chilca, 12°29'S, 76°48'W, 12.ix.l954 (E. I. Schlinger
and E. S. Ross, CAS); between moudis of Rio Chillon
and Ancon [?], 11°54'S, 77°7'W, 5.viii.l953 (M.
Koepcke and Koepcke, MUSM); Cajacay [?], Rio
Fortaleza, 10°40'S, 77°52'W, 6.iii.l956 (W. Weyrauch,
CAS); Canta, Rio Chillon, 11°28'12"S, 76°37'23"W,
12.V.1951 (W. Weyrauch, CAS); Cerro Caracoles,
12°23'S, 76°45'W 15.b£.1951 (W. Weyi-auch, CAS);
Lima, 12°3'S, 77°3'W (K. Jelski and Stolzman, PAN);
Lomas de Iguanil [?] (Huaral), 11°29'51"S,
77°12'12"W, 14.vi.l986 (D. Silva, MUSM).
6. Metepeira glomerabilis (Keyserling)
Figures 37, 46-52, 314; Map 3
Epeira glomerabilis Keyserling, 1892: 154, fig. 113,
9 , <S . Male and female syntypes from Taquara, Rio
Grande do Sul and Serra Vermelha, Rio de Janeiro,
Brazil, in BMNH, examined. Male here designated
lectotype.
Araneus glomerabilis: — Petrunkevitch, 1911: 294.
Roewer, 1942: 843.
Aranea .santa Chamberlin, 1916: 254, pi. 19, fig. 10,
9 . Female holotype from Santa Ana, 3,000 m, Cuz-
co, Peru, in MCZ, examined. NEW SYNONYMY.
Metepeira santa: — Chamberlin and Ivie, 1942: 67.
Platnick, 1993: 449.
Metepeira glomerabilis: — Chamberlin and Ivie, 1942:
68, fig. 181.
Metazygia gloiTierabilis: — Levi, 1991: 179. Platnick,
1993: 448. Erroneous transfer.
Description. Female from Punapi, Tun-
gurahua, Ecuador. Carapace brown; white
arrowhead mark between eye region and
thoracic furrow (Fig. 51). Legs dusky
brown, hghter on ventral surfaces of fem-
ora I and II; lighter on dorsal surfaces of
patellae, tibiae, and metatarsi I and II. Fe-
mur I with row of two to three macrosetae
on anterior side; none on anteroventral
side. Dorsal folium dark, speckled white,
and white fleur-de-lis pattern with thin
branches (Fig. 51). Venter black with
short, wide, white median line ending in a
T-shape; pair of small white spots on either
side of spiracle (Fig. 52). Sternum black
with median white line widening anteriorly
(Fig. 52). Ratio of eye diameters: posterior
medians and anterior medians 1.0, anterior
laterals 1.3, posterior laterals 1.2. Anterior
median eyes separated by 1.3 diameters,
posterior median eyes by 0.9, anterior me-
dian eyes separated from anterior laterals
by 1.7 diameters of anterior lateral eyes,
lateral eyes separated by 0.2 their diame-
ters. Total length 5 mm. Carapace 2.4 mm
long, 1.7 wide. First femur 2.2 mm, patella
and tibia 2.3, metatarsus 1.9, tarsus 0.9.
Second patella and tibia 1.9 mm, third 1.2,
fourth 1.8.
Male from Puiiapi, Tungurahua, Ecua-
dor. Carapace as in female, except for a
greater separation between white colora-
tion around eyes and arrowhead mark
(Fig. 49). Legs light tan, dark distally on
femora. Femur I with row of three macro-
setae on anterior side; four on anteroven-
tral side. Dorsal folium and venter as in
female (Figs. 49, 50). Sternum black with
partly broken median white line widening
anteriorly (Fig. 50). Ratio of eye diame-
ters: posterior medians and anterior me-
dians 1.1, anterior laterals 1.3, posterior
laterals 1.2. Anterior median eyes separat-
ed by 1.8 diameters, posterior median eyes
by 1.2, anterior median eyes separated
from anterior laterals by 2 diameters of an-
terior lateral eyes, lateral eyes separated by
0.3 their diameters. Total length 3.5 mm.
Carapace 1.7 mm long, 1.4 wide. First fe-
mur 2.5 mm, patella and tibia 2.4, meta-
tarsus 2.3, tarsus 0.9. Second patella and
tibia 1.9 mm, third 1.0, fourth 1.7.
Diagnosis. Female M. ^omerabilis and
M. cajabamba differ from the other spe-
cies in the M. vigilax group (M. vigilax and
M. rectangula) by the smaller and more
tubelike openings to the epigynum (com-
pare Figs. 40, 48 with Figs. 55, 62). Me-
tepeira glomerabilis differs from M. caja-
bamba by having epigynal openings that
are more round (Fig. 48) than oval (Fig.
40) when viewed ventrally, having the epi-
gynal openings closer together (compare
Fig. 48 with Fig. 40), and by having the
sclerotized tubelike openings more parallel
to the epigynal groove (Fig. 48) than an-
teriorly directed (Fig. 40). Male M. glom-
erabilis and M. cajabamba differ from oth-
er species in the M. vigilax group by the
smaller, thinner, and more graceful emboli
Metepeiba • Piel
29
(compare Figs. 38, 46 with Figs. 53, 60).
Metepeira glomerabilis differs from M. ca-
jabaniba by having a keel on the median
apophysis, an only slightly slimmer embo-
lus, and a larger, winged embolus cap
(compare Fig. 46 with Fig. 38). A larger
portion of the prosoma is white in M.
glo77ierabilis (Fig. 51), compared to M. ca-
jabaniba (Fig. 43). In addition, the mar-
gins of the folium, particularly in the male,
are whiter than in other species in the
group (compare Fig. 49 and Keyserling
[1892, fig. 113b] with Figs. 41, 56, 63).
Variation. Average body length of 28 fe-
males examined 5.1 mm, range 3.8 to 7.3
mm. Average body length of 10 males ex-
amined 3.1 mm, range 2.4 to 4.3 mm. The
base of the embolus varies from relatively
thin (Fig. 46) to somewhat thicker, as in
M. cajabamba (Fig. 38).
Natural History. At first it would appear
that this species is not seasonal — mature
specimens have been collected throughout
the year (Fig. 315). However on closer in-
spection, there seems to be a seasonal shift
with elevation: mature spiders are found
at low altitudes (0—500 m) between March
and October, at medium altitudes (500—
1,500 m) between August and March, and
at high altitudes (1,500-4,000 m) between
December and June (Fig. 37). In coastal
regions, R. Baptista (personal communi-
cation) reports that this species forms
small aggregations of two to 10 spiders.
Distribution. Colombia, Ecuador, Peru,
Bolivia, Paraguay, and southern Brazil
(Map 3). Elevations range froin sea level
to 4,000 m.
Records Examined. BOLIVIA Chuquisaca: Mon-
teagudo, 19°49'S, 63°59'W, 24.xii.1984 (L. E. Pena,
AMNH). BRAZIL EspiHto Santo: Fazenda Santa
Maria [?], Apiaca, 21°4'S, 41°25'W, 22. ix. 1985 (R. L.
C. Baptista, MZSP). Mato Grosso: Chavantina,
14°40'S, 52°21'W, 15.vi.l947 (J. C. Carvalho, MNRJ);
Fazenda Cei-vo, Tres Lagoas, 20°48'S, 51°43'W,
18. ix. 1964 (Exp. Depto. ZooL, MZSP); Utiariti,
13°2'S, 58°17'W, 15.vlii.l961 (Lenks, MZSP). Minas
Gerais: Lavras, 21°14'S, 45°0'W, 20.x. 1978 (W. Don
Fronk, MCZ); Pedra Azul, 16°1'S, 41°16'W,
15.xii.l970 (F. M. Oliveira, AMNH); Peti Forest Res.
[?] Santa Barbara, on bushes in cerrado, 19°56'S,
43°24'W, 28.viii.1986 (R. L. C. Baptista, MZSP). Pa-
Peru, Ecuador, Colombia
Brazil, Paraguay, Bolivia
50 100 150 200 250 300 350 400 450 500 550
Collection Day (since Jan 1)
Figure 37. The elevation of collection localities for mature spi-
ders of M. glomerabilis on specific days of the year between
1939 and 1990. There appears to be a shift in seasonal mat-
uration times that corresponds better with elevation than with
latitude. Elevations estimated from NOAA database of 5- by
5-minute geographic tiles.
Scale of abscissa: 1 00 = April 9; 300 = October 27; 500 =
May 15 of the following year.
Symbols: Peru, Ecuador, Colombia [■]; Brazil, Paraguay, Bo-
liva [♦].
rand: Praia do Leste, Paranagua, 2.5°46'S, 48°31'W,
4.V.1967 (P. Biasi, MZSP). Rio de Janeiro: Guaratiba,
22°58'S, 42°48'W, 28.viii.1976 (J. A. R Dutra, MZSP);
Ilha de Santana, Macae, 22°25'S, 41°44'W, 18.x. 1986
(R. L. C. Baptista, MZSP); Itaipu, Niteroi, 22°56'S,
43°5'W, 20.ivl985 (R. L. C. Baptista, MZSP). Rio
Grande do Norte: Fazenda Canaan [?], Macaiba,
5°51'S, 35°21'W, 15.ix.l951 (M. Alvorenga, MZSP).
Rondonia: Vila Rondonia, 10°52'S, 61°57'W, 9.ii.l961
(Pereira & Machado, MZSP). Santa Catarina: Nova
Teutonia, 27°3'S, 52°24'W, 12.vl949 (SMF). Sao
Paulo: Barueri, 23°31'S, 46°53'W, 13.iii.l966 (K. Len-
ko, MZSP); Campos do Jordao, 22°44'S, 45°35'W,
15.iii.l945 (Wygod, MZSP); Institute Oceanografico,
Ubatuba, 2.3°26'S, 45°4'W, 1.5.vl967 (P Montouchat,
MZSP); km 1 Rod, Rio Santos [?], Ubatuba, 23°26'S,
45°4'W, 12.X.1985 (R. L. C. Baptista, MZSP),
13.X.1985 (R. L. C. Baptista, MZSP); km 3 Rod, Rio
Santos [?], Ubatuba, 23°26'S, 45°4'W, 6.i.l985 (R. L.
C. Baptista, MZSP). COLOMBIA Cundinamarca:
Sabana de Bogota, 4°43'N, 74°10'W, 10.xii.l990 (C.
Valderrama, CV). ECUADOR Guaijas: Guayaquil,
2°10'S, 79°.54'W, 18.iii.l942 (H. E. F. & D. E. F,
CAS), 22.iii.1942 (Landis, CAS). Pichlncha: 1.5 mi N
Quito, 0°0'N, 78°30'W, 23.ii.1955 (E. I. Sehlinger &
E. S. Ross, CAS). Tungurahua: Baiios, 1°24'S,
78°25'W, 15.ivI939 (W. C. Macintyre, MCZ); Puiiapi,
1°22'S, 78°28'W, 19.vi.l943 (D. L. F. & H. E. F,
CAS). PARAGUAY Alto Parana: Taguararaya [?],
25°30'S, .54°50'W (AMNH). Caazapa: Villa Pastoreo,
25°53'S, 55°45'W (D. Wees, MCZ). PERU Cajamar-
ca: Cajamarca, 7°10'S, 78°31'W, 15.ii.l942 (W. Wey-
rauch, AMNH). Lanihaijeque: 10 km S Chiclayo,
7°59'S, 77°17'W, 19.iii.l951 (E. S. Ross & Michel-
bacher, CAS). Piura: Cerro Prieto, La Brea, 4°41'S,
81°6'W (CAS); Higueron (Las Lomas) [?], 4°19'S,
80°26'W, 29.vii.194I (D. L. F & H. E. F, CAS).
VENEZUELA Sucre: 1 km S Villa Frontado, Rd. to
30 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
Caripe, 10°27'N, 63°37'W, 12.ii.l984 (J. Coddington,
USNM).
7. Metepeira vigilax (Keyserling)
Figures 36, 53-59, 327; Map 4
Epeira vigilax Keyserling, 1893: 211, fig. 156, 6.
Male holotype from Taquara, Rio Grande do Sul,
Brazil, in BMNH, examined.
Araneus vigilax: — Petrunkeviteh, 1911: 324. Roewer,
1942: 856.
Metepeira dominicana Archer, 1965: 132, figs. 12, 18,
$ . Female holotype from west of Bani, Dominican
Republic, in AMNH. Holotype lost. Brignoli, 1983:
275. NEW SYNONYMY.
Metepeira vigilax: — Levi, 1991: 180. Platnick, 1993:
449.
Note. Although the type for M. dominicana is
lost, the name has been identified by using Archer's
(1965) description and illustration.
Description. Female from Trujillo, west
of Bani, Dominican Republic. Carapace
light around eyes with lateral posterior ex-
tensions (Fig. 58). Legs dark, light rings on
proximal ends of leg articles. Femur I with
ro\v of four macrosetae on anterior side;
three on anteroventral side. Dorsal folium
darker than in most species; fleur-de-lis
usually reduced to two white spots (Fig.
58). Venter brownish gray with lighter
margins. Wide, short, median white line
with pair of white spots on either side of
spiracle (Fig. 59). Sternum brownish black
with wide, white line widening anteriorly
(Fig. 59). Ratio of eye diameters: posterior
inedians and anterior medians 1.2, anterior
laterals 1.2, posterior laterals 1.1. Anterior
median eyes separated by 1.8 diameters,
posterior inedian eyes by 1.1, anterior me-
dian eyes separated from anterior laterals
by 3.5 diameters of anterior lateral eyes,
lateral eyes separated by 0.1 their diame-
ters. Total length 9.2 mm. Carapace 3.9
mm long, 3.2 wide. First femur 4.1 mm,
patella and tibia 4, metatarsus 3.3, tarsus
1.2. Second patella and tibia 3.4 mm, third
2.3, fourth 3.4.
Male froin same locality as female.
Black carapace with white around eyes and
extending posteriorly; white wedge mark
in center (Fig. 56). Legs ringed like fe-
male. Femur I with row of three luacro-
setae on anterior side; three on anterov-
entral side. Dorsum and venter as in fe-
male (Figs. 56, 57). Median white line may
be limited to posterior end of sternum
(Fig. 57). Ratio of eye diameters: posterior
medians and anterior inedians 0.9, anterior
laterals 1.1, posterior laterals 1. Anterior
median eyes separated by 1.6 diameters,
posterior median eyes by 1.3, anterior me-
dian eyes separated from anterior laterals
by 2 diameters of anterior lateral eyes, lat-
eral eyes separated by 0.2 their diameters.
Total length 3.9 mm. Carapace 2 mm long,
1.5 wide. First femur 2.6 mm, patella and
tibia 2.5, metatarsus 2.4, tarsus 0.9. Sec-
ond patella and tibia 2.1 mm, third 1.2,
fourth 1.7.
Diagnosis. Female M. vigilax differ
from those of other species in the M. vi-
gilax species group by the shape of the
epigynal openings: from a ventral view the
openings are oval and angled inward pos-
teriorly (Fig. 55); from a posterior view,
the edges of the openings are more par-
allel to the body (Fig. 61) as opposed to
more pei^pendicular to the body (Figs. 39,
61). Male M. vigilax differ from other spe-
cies because the embolus is larger and
more robust (compare Fig. 53 with Figs.
38, 46, 60); the two flagella on the median
apophysis are of more similar width (com-
pare Fig. 53 with Fig. 38), and the keel on
the median apophysis is slim and feather-
shaped (Fig. 38), in contrast to arrow-
shaped (Figs. 46, 60) or absent (Fig. 38).
The dorsal folium differs from other Me-
tepeira species by having a wide black me-
dian stripe at the posterior end of the ab-
domen (Fig. 58). In Brazilian and Bolivian
specimens this stripe often extends all the
way to the black anterior shoulders of the
dorsum, forming a wide T-shape mark.
Variation. Specimens from Argentina
tend to be more lightly pigmented than
those from more northern localities. White
markings on the eye region of Brazilian
and Bolivian specimens surround only the
lateral eyes, in contrast to those on His-
paniolan specimens, which cover the en-
tire eye region.
Natural History. Mature adults have
Metepeira • Piel 31
cajabamba
(5)
Figures 38^5. Metepeira cajabamba new species (sp. 5 [38-44] 7°37'S, 78°3'W; [45] 9°10'S, 77°45'W). 38, male palpus,
mesal. 39, epigynum, posterior. 40, epigynum, ventral. 41 , male, dorsal. 42, male, ventral. 43, female, dorsal. 44, female, ventral.
45, female, ventral.
Figures 46-52. Metepeira glomerabilis (Keyserling) (sp. 6; 1°22'S, 78°28'W). 46, male palpus, mesal. 47, epigynum, posterior.
48, epigynum, ventral. 49, male, dorsal. 50, male, ventral. 51, female, dorsal. 52, female, ventral.
Figures 53-59. Metepeira vigilax (Keyserling) (sp. 7 [53-57] 18°27'N, 72°17'W; [58,59] 18°17'N, 70°22'W). 53, male palpus,
mesal. 54, epigynum, posterior. 55, epigynum, ventral. 56, male, dorsal. 57, male, ventral. 58, female, dorsal. 59, female, ventral.
Scale bars: dorsum and venter figures 1 .0 mm.
32 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
been collected throughout the year except
Januaiy, February, and March (Fig. 327).
Spiders have been found among electric
wires four meters above ground. Locality
elevations range from near sea level to
1,400 m and follow an ecological zone that
decreases in elevation with distance from
the equator (Fig. 36). Over equivalent lat-
itudes, M. vigilax lives at less than one-
tenth of the elevation of M. rectangula
(Fig. 36).
Distribution. Hispaniola, Bolivia, Brazil,
and coastal Argentina (Map 4). The dis-
junct distribution between Hispaniolan
and South American populations may be
due to human-assisted migration.
Records Examined. ARGENTINA Buenos Aires:
Celsya-Pereyra [?], 34°50'S, 58°6'W (MACN); Ze-
laya, 34°21'S, 58°52'W (MACN). BOLIVIA La Paz:
Apolo, 14°43'S, 68°31'W, 10.viii.l989 (L. E. Pena,
AMNH). Santa Cruz: Estacion Robore, above creek,
18°20'S, 59°45'W, 27.ix.1955 (Azambuya, CAS).
BRAZIL Espirito Santo: Fazenda Santa Maria [?],
Apiaca, 21°4'S, 41°25'W, 14.V.1988 (R. L. C. Baptista,
MZSP). Rio Grande do Sul: Sao Leopoldo, 29°46'S,
5r9'W, 14.vi.l964 (Celia Valle, MZSP). DOMINI-
CAN REPUBLIC Aziia: El Pueriio, Majagual and
Peralta, I8°34'N, 70°47'W, 10.xi.l979 (E. Marcano,
MNSD). Prov. Tnijillo Vdldez: W Bani, 18°17'N,
70°22'W, 8.viii.l958 (A. F. Archer & E. de Boyrie
Moya, AMNH). HAITI Departement de L' Quest:
Kenscoff, 18°27'N, 72°17'W, 15.xii.l929 (J. C. Myers,
AMNH), 17.iv.l935 (AMNH).
8. Metepeira rectangula (Nicolet)
Figures 36, 60-66, 306; Map 4
Epeira rectangula Nicolet, 1849: 500, female holo-
type from Valdivia, Chile, in MNHN.
Metepeira labyrinthea: — Petrunkevitch, 1911: 298.
Roewer, 1942: 868. Bonnet, 1957: 2821. Erroneous
synonymy.
Metepeira rectangulata: — Chamberlin and Ivie, 1942:
71. Unjustified emendation.
Note. The name was identified using drawings of
the holotype (H. W. Levi, personal illustrations).
Description. Female from Angol, Mal-
leco, Chile. Carapace reddish brown with
long white setae behind lateral eyes. An-
terior third of carapace white, median
white line reaching thoracic furrow (Fig.
65). Proximal halves of femora white, re-
mainder black with distal white marks on
dorsal surfaces. Patellae inostly black, re-
maining articles white with black marks at
base of setae. Femur I with row of three
to five macrosetae on anterior side; three
to four on anteroventral side. Anterior
margin of dorsal abdomen black; dorsal fo-
lium yellowish with brown speckles, white
fleur-de-lis pattern with wide branches
(Fig. 65). Venter black with wide white
median line, flanked by pair of thin white
lines; pair of white spots on either side of
spiracle connected by white line (Fig. 66).
Sternum black with posterior white mark
(Fig. 66). Batio of eye diameters: posterior
medians and anterior medians 1.0, anterior
laterals 1.1, posterior laterals 1.1. Anterior
median eyes separated by 1.7 diameters,
posterior median eyes by 1.0, anterior me-
dian eyes separated from anterior laterals
by 3.3 diameters of anterior lateral eyes,
lateral eyes separated by 0.1 their diame-
ters. Total length 8.4 mm. Carapace 3.7
mm long, 2.9 wide. First femur 4.2 mm,
patella and tibia 4.1, metatarsus 3.7, tarsus
1.2. Second patella and tibia 3.4 mm, third
2.1, fourth 3.1.
Male from Angol, Malleco, Chile. Car-
apace reddish brown, anterior third white,
median white line extending to thoracic
furrow (Fig. 63). Proximal halves of fem-
ora, white, distal halves black. Patellae
black, remaining articles white widi black
spots at base of setae. Feinur I with row
of six to eight macrosetae on anterior side;
seven to 11 on anteroventral side. Dorsal
abdomen white, marbled, and speckled
brown (Fig. 63). Venter dark brown with
wide white median mark, flanked by pair
of thin white lines; pair of white spots on
either side of spiracle connected by white
line (Fig. 64). Sternum dark brown with
partly broken median white line (Fig. 64).
Batio of eye diameters: posterior medians
and anterior medians 1.0, anterior laterals
1.2, posterior laterals 1.1. Anterior median
eyes separated by 1.7 diameters, posterior
median eyes by 1.1, anterior median eyes
separated from anterior laterals by 2.9 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.2 their diameters. To-
tal length 6.1 mm. Carapace 3.2 mm long.
Metepeira • Piel
2.5 wide. First femur 5.2 mm, patella and
tibia 5, metatarsus 4.9, tarsus 1.2. Second
patella and tibia 4.1 mm, third 2, fourth
3.1.
Diagnosis. Female M. rectangula differ
from the other species in the M. vigilax
group (M. vigilax, M. glomerahilis, M. ca-
jahamba) by the shape of the epigynal
openings: from a ventral view they are
larger, wider, and more gaping than in the
other species (compare Fig. 62 with Figs.
40, 48, 55). Males differ from other spe-
cies in the group by having an embolus
that is relatively larger than that of M.
glomerahilis and M. cajabaniba, yet small-
er than that of M. vigilax (compare Fig. 60
with Figs. 38, 46, 53). Among all members
of the M. vigilax group, the keel on the
median apophysis of M. rectangula is the
largest and most robust, and the dorsal fo-
lium has the lightest coloration (Fig. 65).
Va nation. Average body length of 11 fe-
males examined 8.6 mm, range 5.8 to 10
mm. Average body length of six males ex-
amined 6 mm, range 4.1 to 7 mm. Speci-
mens from two localities in western Ar-
gentina resemble M. vigilax, and may be
hybrids.
Natural History. This species appears to
follow a narrow ecological zone that de-
creases in elevation with increasing south-
ern latitude (Fig. 36). Median elevation,
about 500 m. Mature specimens have
been collected January through April (Fig.
306). Specimens from localities south of
the 36th parallel tend to be found in Jan-
uaiy and Februaiy, whereas those north of
the 36th parallel tend to be found in
March and April.
Distribution. Chilean Andes between
31° and 38° south (Map 4).
Records Examined. ARGENTINA Cordoba: Cala-
muchita, 32°4'S, 64°33'W, 15.iii.l954 (J. M. Viana,
MACN). Mendoza: Mendoza, 32°53'S, 68°49'W,
30.iii.l965 (H. W. Levi, MCZ). CHILE Bio-Bio: 4 km
E road to Pinto, 36°42'S, 71°53'W, 4.i.l976 (B. Mo-
reno, AMNH); Road to Pemuco, Ci-uce del Carmen,
36°56'S, 72°4'W, 10.i.l976 (G. Moreno, AMNH). Co-
qnimbo: lUapel: Salamanca: Fundo Tahuinco,
31°44'S, 71°5'W, 30.iv.l946 (R. Doneso, AMNH).
Malleco: Angol, 37°48'S, 72°43'W (D. S. Bullock,
CAS), 10.iii.l945 (E. A. Chapin, USNM). Maule: 10
km S Curico, 35°4'S, 71°14'W, 15.iii.l968 (L. E.
Peiia, MCZ); Cordillera de Parral [?], 36°9'S,
71°50'W, 25.ii.1956 (L. E. Pena, IRSNB); Linares,
35°51'S, 71°36'W (L. E. Peiia, IRSNB); Miraflores,
Pedag. [?]. 35°55'S, 71°39'W (Toro, AMNH).
O'Higgins: Fundo Millahue, Cunaco, 34°36'S,
71°16'W, 30.iv.l961 (AMNH). Region Metropolitana:
Melipilla, 33°42'S, 71°13'W (L. E. Pena, IRSNB).
Valparaiso: Casablanca, 33°19'S, 71°25'W, 15.ii.l955
(Edwdn Reed, AMNH).
Metepeira labyrinthea Group
Levi (1977) described the M. labyrin-
thea group veiy broadly — it included spe-
cies with a longitudinal white line down
the sternum and a short keel on the me-
dian apophysis. Here, this species group is
much narrowed to include only three
North American species: Metepeira laby-
rinthea, Metepeira lacandon, and Metepei-
ra spinipes. Males of these three species
are unique among Metepeira by having a
toothless, smooth keel on the median
apophysis. In addition, their distal embolic
apophysis rises away (anteriorly) from the
embolus proper and projects forward (ven-
trally) until it is almost even with the em-
bolus tip (Figs. 67, 76). In contrast, other
Metepeira species with distal embolic
apophyses have the embolus tip extend far
beyond the projection of the apophysis
(e.g., Fig. 171). The female epigynum has
a characteristic shape. The scape is thick
and fleshy and the epigynal openings have
membranes that make them look distinctly
(Fig. 69) or indistinctly (Fig. 78) almond-
shaped. The epigyna of the M. labyrinthea
species group are easily confused with that
of the closely related species, M. gosoga
(Fig. 173). Although most differences be-
tween M. labyinthea group and M. go,soga
are only obvious in the epigynum, it
should be noted that the dark marks inside
the epigynal openings of the former ap-
pear to look cross-eyed, but this cannot be
said for the latter. In this work, only the
species collected at localities south of the
U.S. /Mexico boarder are treated.
34 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
9. Metepeira spinipes
F. O. P. -Cambridge
Figures 67-75, 335; Map 6
Metepeira spinipes F. O. P.-Cambridge, 1903: 459,
figs. 9, 10, (J, ?. Male holotype from Mexico City,
Mexico, in BMNH, examined. Roewer, 1942: 868.
Epeira labyrinthea grinnelli Coolidge, 1910: 281, 9.
Holotype from Palo Alto, California, lost.
Araneus spinipes: — Petrunkevitch, 1911: 317.
Aranea labyrinthea grinnelli: — Moles, 1921: 42.
Metepeira douglasi Chamberlin and Ivie, 1941: 18,
figs. 21-23, 9 . Female holotype from Santa Ana,
California, in AMNH, examined. Chamberlin and
Ivie, 1942: 66, figs. 169-170. First synonymized
with M. labyrinthea grinnelli by Levi, 1977: 198.
Metepeira labyrinthea grinnelli: — Roewer, 1942: 868.
Metepeira labyrinthea: — Bonnet, 1957: 2822. Erro-
neous synonyiTiy.
Metepeira grinnelli: — Levi, 1977: 198, figs. 21-27, S ,
? . NEW SYNONYMY.
Description. Female from Huitzilac,
Morelos, Mexico. Brown carapace with an-
terior portion darker reddish brown, white
behind lateral eyes (Fig. 74). Legs yellow-
ish, femora reddish brown distally, otlier
articles dark brown distally. Femur I with
row of four to five macrosetae on anterior
side; two to seven on anteroventral side.
Anterior shoulders of abdomen black.
Dorsal folium with usual Metepeira pat-
tern, though largest branches of white
fleur-de-lis shape usually widened into
large spots (Fig. 74). Venter of abdomen
black with wide white median line (Fig.
75). Pair of small white spots on either side
of spiracle. Sternum black with posterior
white mark that in some cases extends an-
teriorly to the labium (Figs. 73, 75). Ratio
of eye diameters: posterior medians and
anterior medians 1.1, anterior laterals 1.0,
posterior laterals 0.9. Anterior median
eyes separated by 1.9 diameters, posterior
median eyes by 1.3, anterior median eyes
separated from anterior laterals by 2.9 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.2 their diameters. To-
tal length 9.2 mm. Carapace 3.7 mm long,
2.9 wide. First femur 4.4 mm, patella and
tibia 4.7, metatarsus 4.6, tarsus 1.3. Sec-
ond patella and tibia 4.1 mm, third 2.4,
fourth 3.6.
Male from Huitzilac, Morelos, Mexico.
Carapace, dorsum, venter, sternum as in
female (Figs. 71, 72). Distal portions of leg
articles reddish black, elsewhere yellowish.
Femur I with row of four to six macrosetae
on anterior side; five to nine on anterov-
entral side. Ratio of eye diameters: poste-
rior medians and anterior medians 1.1, an-
terior laterals 1.2, posterior laterals 1.1.
Anterior median eyes separated by 1.8 di-
ameters, posterior median eyes by 1.1, an-
terior median eyes separated from anterior
laterals by 3.2 diameters of anterior lateral
eyes, lateral eyes separated by 0.3 their di-
ameters. Total length 7.5 mm. Carapace
3.8 mm long, 2.8 wide. First femur 6.2
mm, patella and tibia 6.3, metatarsus 7.1,
tarsus 1.8. Second patella and tibia 5.3
mm, third 2.5, fourth 4.
Diagnosis. Inside each epigynal depres-
sion of M. spinipes and M. lacandon is a
membrane that forms a slanted, oval-
shaped opening (Figs. 69, 78). Within each
oval-shaped opening is a dark mark, which
in M. spinipes takes up a small part of that
opening, resulting in a cross-eyed appear-
ance (Fig. 69). Also, in M. spinipes (Fig.
69), the edges of the epigynal depressions
are less distinct than in M. lacandon (Fig.
78). The distal embolic apophysis of M.
spinipes is arrow-shaped (Fig. 79), in con-
trast to a wider shovel-shape in M. lacanA
don (Fig. 79).
Variation. Average body length of 54 fe-
males examined 9.1 mm, range 5.5 to 12.4
mm. Average body length of 55 males ex-
amined 6.6 mm, range 3 to 10.8 mm.
Natural History. Mature specimens
have been collected between August and
early November (Fig. 335). Elevations
range from near sea level in California to
2,600 m in central Mexico. Variation in
sexual dimoi-phism appears to correlate
with habitat and social structure (Piel,
1996). Webs are found in dry regions
among mesquite, Opuntia, Agave (ma-
guey), cultivated Yucca, and Cactus. Spi-
ders live in medium to small social colo-
nies, which vary in size in accordance with
local habitat quality (Uetz, 1988a,b). This
behavioral and ecological relationship is
Metepeira • Piel 35
4 M
82
lacandon
(10)
Figures 60-66. Metepeira rectangula (Nicolet) (sp. 8; 37°48'S, 72°43'W). 60, male palpus, mesal. 61, epigynum, posterior. 62,
epigynum, ventral. 63, male, dorsal. 64, male, ventral. 65, female, dorsal. 66, female, ventral.
Figures 67-75. Metepeira spinipes F. O. P. -Cambridge (sp. 9 [67-72,74,75] 19'=0'29"N, 99°15'50"W; [73] 39°18'N, 123°48'W).
67, male palpus, mesal. 68, epigynum, posterior. 69, epigynum, ventral. 70, male embolic division, ventral. 71, male, dorsal. 72,
male, ventral. 73, female, ventral. 74, female, dorsal. 75, female, ventral.
Figures 76-83. Metepeira lacandon new species (sp. 10; 16°45'N, 92°38'W). 76, male palpus, mesal. 77, epigynum, posterior.
78, epigynum, ventral. 79, male embolic division, ventral. 80, male, dorsal. 81, male, ventral. 82, female, dorsal. 83, female,
ventral.
Scale bars: dorsum and venter figures 1.0 mm.
36 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
thought to occur as a result of the spiders
pursuing a risk-sensitive foraging strategy
(Uetz, 1996).
Distribution. Oregon to central Mexico
(Levi, 1977, map 1; Map 6).
Records Examined. MEXICO Agiiascalientes: Hwy
45, 5.3 mi N Aguascalientes, 21°57'N, 102°17'W,
7.ix.l967 (R. E. Leech, REL). Baja California Sun
Sierra Laguna, 17 air mi ENE Todos Santos,
23°34'N, 110°0'W, 15.xii.l979 (C. E. Griswold, CAS).
Chihuahua: 22.4 mi S Miiaaca, 28°24'N, 107°26'W,
23.viii.1950 (R. Smith, AMNH). Durango: 10 mi E
El Salto, 23°12'N, 105°52'W, 8.viii.l947 (W. J.
Gertsch, AMNH); 11 km W Suehil, 23°35'N,
104°5'W, 5.ix.l984 (W. J. Pulawsld, CAS); 6 mi NE
El Salto, 23°19'N, 105°50'W, ll.viii.l947 (W. J.
Gertsch, AMNH); Las Puentes [?], 26°49'N,
106°2'W, 23.vii.1947 (W. J. Gertsch, AMNH); Oti-
napa, 24°11'N, 105°2'W, 12.viii.l947 (W. J. Gertsch,
AMNH); Palos Colorados, 24°2'N, 104°54'W,
5.viii.l947 (W. J. Gertsch, AMNH); Providencia.
26°44'N, 105°56'W, 24.viii.1947 (A. M. Davis,
AMNH); SW Durango, 23°59'17"N, 104°45'47'W,
22.X.1994 (W. H. Piel, MCZ). Guanajuato: 30 mi SE
Leon, 6 mi SE Silao, 20°52'N, 101°21'W, 6.ix.l964
(Jean & Wilton Ivie, AMNH); 6.3 mi NW Leon,
21°13'N, 101°43'W, 6.ix.l967 (R. E. Leech, REL);
between Moroleon & Cuitzeo, 20°5'36"N,
101°9'28'W, 20.X.1994 (W. H. Piel, MCZ); near San
Miguel de Allende, 20°55'N, 100°45'W, 16.lx.1976
(C. E. Grisv^^old, CAS); S San Miguel de Allende,
20°46'36"N, 100°47'28"W, 20.x. 1994 (W. H. Piel,
MCZ); San Miguel de Allende. Road SW town,
20°52'N, 100°56'W 25.x. 1982 (George Uetz, MCZ).
Hidalgo: 18 mi E Huichapan, off Hwy 45, 20°23'N,
99°22'W, 25.viii.1984 (W D. Sissom, C. Myers & L.
Bom, MCZ); 4 mi N Tizayuca, 19°54'N, 98°59'W
20..xi.l946 (E. S. Ross, CAS); 41 km N Zimapan,
20°54'N, 99°13'W, 10.viii.l991 (W H. Piel & G. S.
Bodner, MCZ); Apulco, 20°19'N, 98°20'W, 6.X.1947
(H. Wagner, AMNH); Ozumbilla, 20°9'N, 101°16'W,
2.X.1957 (R. Dreisbach, MCZ); Pachuca, 20°7'N,
98°44'W, 30.viii.l957 (R. Dreisbach, MCZ); Tenango
de Doria, 20°19'N, 98°13'W, 5.x. 1947 (H. Wagner,
AMNH). Jalisco: 12 mi S Mazamitla, 19°47'N,
103°8'W, 5.xii.l948 (H. B. Leech, CAS); Charco
Ondo, 30 km W Ojuelos, 21°47'N, 101°53'W
25. ix. 1945 (H. Wagner, AMNH). Mexico: Ixtapan de
la Sal, 18°50'N, 99°41'W, 24.viu.1946 (H. Wagner,
AMNH); Nevado de Toluca, 19°18'N, 99°44'W,
8.iv.l979 (George Uetz, MCZ); San Juan Teotihu-
acan, 19°41'N, 98°52'W, 4..xi.l939 (C. M. Bogert &
H. E. Vokes, AMNH); Tenancingo, 18°58'N,
99°36'W, 6.LX.1946 (H. Wagner, AMNH), l.x.1946
(H. Wagner, AMNH), 15.x. 1946 (H. Wagner,
AMNH); Tenango del Valle, 19°7'N, '99°33'W,
25.viii.1946 (H. Wagner, AMNH), 27.viii.1946 (H.
Wagner, AMNH); Teotihuacan, 19°41'N, 98°52'W
31.viii.l959 (A. F. Ai-cher, AMNH); Tepotzotlan,
19°43'N, 99°13'W, 26.X.1982 (George Uetz, MCZ),
21. ii. 1983 (George Uetz, MCZ), 5.x. 1983 (George
Uetz, MCZ); Toluca, 19°18'N, 99°44'W, 10.viii.l978
(George Uetz, MCZ), l.viii.l986 (George Uetz,
MCZ); Toluca, at bottom of mountain near Parque
Cierra Morelos, 19°18'N, 99°44'W, 10.viii.l978
(George Uetz, MCZ); Toluca, E of town on Paseo
ToUocan [?], 19°18'N, 99°42'W, 23.X.1982 (George
Uetz, MCZ). Mexico D. F.: 19°25'N, 99°10'W,
12.X.1940 (H. Wagner, AMNH), 28.xii.1940 (R. H.
Crandall, AMNH), 15.ix.l943 (H. & D. Frizzell,
AMNH); Contreras, 19°18'N, 99°17'W, 4.xii.l944 (H.
Wagner, AMNH), 15.ix.l965 (N. L. H. Krauss,
AMNH); Delegacion Tlalpan, Colonia Santa Ursula
Xitla, 19°16'0"N, 99°10'25'W, 12.x. 1994 (W H. Piel,
MCZ); Desierto de los Leones, 19°22'N, 99°16'W,
15.ix.l941 (H. Wagner, AMNH); El Xitle, 18°61'N,
99°17'W [?], 12.viii.l942 (C. Tellez, AMNH); Haci-
enda Cordoba, 19°26'N, 99°10'W [?], 29.X.1944 (H.
Wagner, AMNH); Ouieros, 18°62'N, 99°17'W [?],
5.vii.l943 (M. Cardenas, AMNH); Mexico City,
19°25'N, 99°10'W, l.xi.l941 (C. Velo, AMNH),
25.Lx.1957 (R. Dreisbach, MCZ); Mixcoac, 19°23'N,
99°11'W (AMNH), 13.X.1940 (A. F. Archer, AMNH);
Mixenac, 19°25'N, 99°10'W 13.X.1940 (H. Wagner,
AMNH); Pedregales, 18°60'N, 99°17'W [?],
15.viii.l909 (AMNH); Petregal [?], 18°60'N,
99°17'W l.xii.l943 (AMNH); Rancho Cordoba,
19°27'N, 99°10'W, 29.X.1944 (H. Wagner, AMNH);
Tlaplan, 19°17'N, 99°10'W, 7.viii.l991 (W. H. Piel &
G. S. Bodner, MCZ). Michoacan: 25 mi W La Barca
nr Lago de Chapala, 20°17'N, 102°34'W, ll.ix.l976
(C. E. Griswold & Jackson, CAS); between Patzcuaro
& Uruapan, 19°29'19"N, 101°48'20"W, 19.X.1994 (W
H. Piel, MCZ); Hills N of Patzcuaro, 19°45'N,
101°36'W, 24.viii.1959 (A. F. Archer, AMNH); Hwy
110, 4 mi W. Jiquilpan, 19°59'N, 102°47'W,
2.viii.l967 (R. E. Leech, REL); Hwy 15, 9.5 mi W
Morelia, 19°42'N, 101°16'W; 18.viii.l967 (R. E.
Leech, REL); Lake Chapala, NW of Cojumatlan,
20°10'N, 102°53'W, 7.ix.l966 (Jean & Wilton Ivie,
AMNH); Monte de Zacapu, 19°47'N, 101°50'W,
24.viii.1959 (A. F. Archer, AMNH). Morelos: Cuer-
navaca, 18°55'N, 99°15'W (AMNH), 15.ix.l941 (H.
Wagner, AMNH), 18.xi.l946 (M. G. Bradt, AMNH);
Cueniavaca/Tepotzotlan, interchange between 1-95 &
115, 18°55'N, 99°13'W, 7.viii.l978 (MCZ); Huitzilac,
19°2'2"N, 99°16'13'W^, 13.X.1994 (W. H. Piel, MCZ);
North of Cuernavaca, 18°58'11"N, 99°14'37"W,
11.x. 1994 (W. H. Piel, MCZ); S of Huitzilac,
19°0'29"N, 99°15'50"\'V, 16.x. 1994 (W H. Piel, MCZ).
Puehla: 6 mi E Rio Frio, 19°20'N, 98°35'W,
22.viii.1964 (Jean & Wilton Ivie, AMNH); Puebla,
19°3'N, 98°12'W, 21.x. 1982 (George Uetz, MCZ).
San Luis Potosi: 3 km W Pilares, 21°55'34"N,
100°48'6'W, 21.X.1994 (W. H. Piel, MCZ); Cuidaddel
Maiz, 22°24'N, 99°36'W, 25.viii.1954 (R. Dreisbach,
MCZ). Sonora: 46 mi S Agua Prieta on Highway 10,
31°0'N, 109°16'W, 15.viii.l959 (B. A. Branson,
AMNH); Hermosillo, 29°4'N, 110°55'W, 20.lx.1952
(B. Malldn & V. E. Thatcher, AMNH); Sierra Man-
Metepeira • Piel
37
zanal, 30°50'N, 110°10'W, 14.ix.l976 (Roth &
Schroepfer, MCZ). Tlaxcala: Huamantla, 19°19'N,
97°56'W [?], 15.vii.l981 (C. Gold, CAS). Veracruz:
15 mi W. Banderilla, 19°39'N, 97°8'W, 31.X.1973 (S.
C. Williams & C. L. Mullinex, CAS); 15 mi West of
Jalapa, 19°32'N, 97°9'W, 23.vi.1946 (A. M. & L. I.
Davis, AMNH). Zacatecas: 13 mi N. Sombrerete,
23°44'32"N, 103°47'10"W 22.X.1994 (W H. Piel,
MCZ); Canutillo, 24°47'N, 101°31'W, 14.viii.l947 (W.
J. Gertsch, AMNH); S. Zacatecas, 22°45'7"N,
102°29'37"W, 22.X.1994 (W. H. Piel, MCZ). USA. Ar-
izona: Southwestern Research Station, Chiricaliua
Mtns., 31°35'N, 109°14'W, 20.viii.l976 (V. Roth,
MCZ). California: 26 mi W. Santa Rosa on Hwy 116,
38°31'N, 123°4'W, 19.ix.l976 (M. E. Thompson,
MCZ); Mendocino, 39°18'N, 123°48'W, 18.viii.l959
(W J. Gertsch, MCZ); Monterey, 36°36'N, 121°54'W,
l.ix.l949 (A. F. Archer, MCZ); Pacific Grove,
36°37'N, 121°56'W (R. V. Chamberlin, MCZ); Palo
Alto, 37°27'N, 122°9'W (Doane, MCZ); Salt marsh
on N shore of San Pablo Bay, Vallejo, 38°8'N,
122°27'W (D. Spiller, MCZ).
10. Metepeira lacandon new species
Figures 76-83, 332; Map 6
Holotijpe. Male from San Cristobal, Chiapas, Mexico.
The specific name is a noun in apposition after the
Indian people who live in Chiapas. Holotype de-
posited in the AMNH.
Description. Female paratype from San
Cristobal, Chiapas, Mexico. Reddish cara-
pace, slightly darker in anterior half, ligh-
ter behind lateral eyes (Fig. 82). Leg ar-
ticles yellowish, gradually turning reddish
brown distally. Femur I with row of four
or five macrosetae on anterior side; two to
four setae on anteroventral side. Anterior
shoulders of abdomen black. Branches of
white fleur-de-lis shape in dorsal folium
thinner than in most species (Fig. 82).
Venter of abdomen black with wide white
median line that extends about half the
distance between epigynal groove and
spinnerets (Fig. 83). Pair of very small
white spots on either side of spiracle. Ster-
num black, often with central w^hite spot
(Fig. 83). Ratio of eye diameters: posterior
medians and anterior medians 1.0, anterior
laterals 1.3, posterior laterals 1.2. Anterior
median eyes separated by 1.7 diameters,
posterior median eyes by 1.0, anterior me-
dian eyes separated from anterior laterals
by 4.5 diameters of anterior lateral eyes,
lateral eyes separated by 0.1 their diame-
ters. Total length 11.5 mm. Carapace 4.6
mm long, 3.6 wide. First femur 5.3 mm,
patella and tibia 5.3, metatarsus 4.9, tarsus
1.6. Second patella and tibia 4.4 mm, third
2.7, fourth 4.
Male holotype. Carapace, dorsum, ven-
ter, sternum as in female (Figs. 80, 81).
Distal halves of femora, tibia reddish
brown, elsewhere yellowish. Patellae,
metatarsi reddish. Femur I with row of
four or five macrosetae on anterior side;
six or seven on anteroventral side. Ratio of
eye diameters: posterior medians and an-
terior medians 1.1, anterior laterals 1.5,
posterior laterals 1.3. Anterior median
eyes separated by 1.7 diameters, posterior
median eyes by 0.9, anterior median eyes
separated from anterior laterals by 3.3 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.3 their diameters. To-
tal length 6.8 mm. Carapace 3.5 mm long,
2.7 wide. First feinur 5.3 mm, patella and
tibia 3.9, metatarsus 5.6, tarsus 1.7. Sec-
ond patella and tibia 4.4 mm, third 2.3,
fourth 3.4.
Diagnosis. Inside each epigynal depres-
sion of M. lacandon and M. spinipes is a
membrane that forms a slanted, oval-
shaped opening (Figs. 69, 78). Within each
oval-shaped opening is a dark mark, which
in M. lacandon takes up only a large part
of that opening, resulting in a less cross-
eyed appearance (Fig. 78). Also, in M. la-
candon (Fig. 78), the edges of the epigynal
depressions are luore distinct than in M.
spinipes (Fig. 69). The distal embolic
apophysis of M. lacandon is more shovel-
shaped (Fig. 79) than the thinner, arrow-
shaped one in M. .spinipes (Fig. 70).
Variation. Average body length of three
females examined 8.5 mm, range 7.5 to 10
mm. Average body length of four males
examined 5.8 mm, range 4.2 to 6.8 mm.
Natural History. Mature specirnens
were collected between July and Septem-
ber (Fig. 332) from oak— pine woodland.
Elevations range from 1,700 to 2,300 m.
Distribution. Mountainous regions of
Chiapas, Mexico (Map 6).
38 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
Records Examined. MEXICO Chiapas: 12 km NW
Comitan, 16°23'N, 92°15'W, 30.\dii.l976 (E. S. Ross,
CAS); 4 mi SE San Cristobal, 16°42'N, 92°36'W,
23.viii.1966 (Jean & Wilton Ivie, AMNH); 5 km W
San Cristobal de Las Casas on HWY 190, 16°44'N,
92°41'W, 27.vii.1983 (W. Maddison & R. S. Anderson,
MCZ); 5 mi W San Cristobal, 16°45'N, 92°41'W,
24.viii.1966 (Jean & Wilton Ivie, AMNH); San Cris-
tobal, 16°45'N, 92°38'W, 13.ix.l947 (H. Wagner,
AMNH); San Cristobal de las Casas, 16°45'N,
92°38'W, 22.vii.1947 (C. J. & M. Goodnight,
AMNH); Tenejapa, 16°49'N, 92°31'W, 22.vii.1950 (C.
J. & M. Goodnight, AMNH).
Metepeira nigriventris Group
There are five species in the Metepeira
nigriventris group: Metepeira nigriventris,
Metepeira tarapaca, Metepeira calamuchi-
ta, Metepeira galatheae, and Metepeira
karkii. These closely related species are of-
ten hard to distinguish because their gen-
italia are similar, yet highly variable within
a species. This species group is easily rec-
ognized by the distinctive shape of the
scape and similarities in palp moiphology.
Typically the base of the scape originates
anteriorly and projects ventrally before
curving posteriorly. This projection creates
an overhang and a noticeable gap between
the scape and the genital openings (e.g.,
Figs. 85, 86, 101, 102). The embolus is
thick and has a large, prominent, distal
apophysis that hides under the terminal
apophysis (Figs. 84, 92, 100, 110, 121).
1 1 . Metepeira nigriventris (Taczanowski)
Figures, 84-91, 310; Map 5
Epeira nigriventris Taczanowski, 1878: 151, fig. 6, ?.
Female lectotype from Lake Junin, Peru, in PAN,
type lost. Keyserling, 1893: 217, fig. 161, 9,3.
Araneus nigriventris: — Chamberlin, 1916: 248. Bon-
net, 1955: 550.
Metepeira nigriventris: — Chamberlin and Ivie, 1942:
74, figs. 211-214, 9,3. Platnick, 1993: 449.
Note. Although the type is lost, the type locality
and Taczanowski s descriptions are sufficient to rec-
ognize the species.
Description. Female from 12 km west of
Tarma, Junin, Peru. Carapace dark brown,
light around eyes with lateral posterior ex-
tensions (Fig. 90). Proximal halves of leg
articles yellow, distal halves black. Femur
I with row of four macrosetae on anterior
side; five on anteroventral side. Dorsum
darker and white fleur-de-lis pattern small-
er than in most species (Fig. 90). Venter
mostly black with reduced, short, thin,
white median line (Fig. 91). SternuiTi en-
tirely black. Ratio of eye diameters: pos-
terior medians and anterior medians 1.0,
anterior laterals 1.4, posterior laterals 1.4.
Anterior median eyes separated by 1.8 di-
ameters, posterior median eyes by 1.2, an-
terior median eyes separated from anterior
laterals by 5 diameters of anterior lateral
eyes, lateral eyes separated by 0.4 their di-
ameters. Total length 11 mm. Carapace 5.1
mm long, 4.2 wide. First femur 5.5 mm,
patella and tibia 5.8, metatarsus 5.5, tarsus
1.9. Second patella and tibia 5.3 mm, third
3.3, fourth 4.5.
Male from same locality as female. An-
terior margin of chelicerae with large,
swollen tooth and several denticles. Cara-
pace reddish brown with lighter eye re-
gion, lateral posterior extensions, and long,
thin median white line (Fig. 88). Femur 1
with row of about five macrosetae on an-
terior side; nine on anteroventral side.
Coloration of legs, dorsum, venter, and
sternum as in female (Figs. 88, 89). Ratio
of eye diameters: posterior medians and
anterior medians 1.1, anterior laterals 1.3,
posterior laterals 1.3. Anterior median
eyes separated by 1.8 diameters, posterior
median eyes by 1.2, anterior median eyes
separated from anterior laterals by 3.7 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.4 their diameters. To-
tal length 9 mm. Carapace 4 mm long, 3.2
wide. First femur 5.8 mm, patella and tibia
6.2, metatarsus 6.5, tarsus 1.7. Second pa-
tella and tibia 4.5 mm, third 2.8, fourth 4.
Diagnosis. Metepeira nigriventris is eas-
ily distinguished from other species in the
M. nigriventris group by its dark pigmen-
tation. As its name implies, the sternum is
black and the white ventral mark on the
abdomen is reduced to a much shorter and
thinner line (Figs. 88-91). While the ster-
num of M. karkii is similarly dark, the dor-
sal and ventral markings on the abdomen
are much lighter (compare Figs. 88-91
Metepeira • Piel 39
nigriventris
(11)
i9& 9^^^ 99
« tarapaca
(12)
A. iBi
H 106 107 _
calamuchita
105 - (13)
Figures 84-91. Metepeira nigriventris (Taczanowski) (sp. 11; ir25'S, 75°48'W). 84, male palpus, mesal. 85, epigynum, pos-
terior. 86, epigynum, ventral. 87, male embolic division, ventral. 88, male, dorsal. 89, male, ventral. 90, female, dorsal. 91,
female, ventral. , „„ .
Figures 92-99 Metepeira tarapaca new species (sp. 12; 21°39'S, 69°33'W). 92, male palpus, mesal. 93, epigynum, postenor.
94, epigynum, ventral. 95, male embolic division, ventral. 96, male, dorsal. 97, male, ventral. 98, female, dorsal. 99, female,
ventral. , ,.,„.,•
Figures 100-107. Metepeira calamuchita new species (sp, 13; 32°4'S, 64°33'W). 100, male palpus, mesal. 101, epigynurri,
posterior. 102, epigynum, ventral. 103, male embolic division, ventral. 104, male, dorsal. 105, male, ventral. 106, female, dorsal.
107, female, ventral.
Scale bars: dorsum and venter figures 1.0 mm.
40 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
with Figs. 125-128). The epigynum of M.
nigriventris has a wider membrane over
the openings that is visible on either side
of a very wide scape (Fig. 86). In contrast,
this feature is hidden behind a thinner
scape in its Hkely sister species, M. tara-
paca (Fig. 94). The overall shape of tlie
epigynum (Fig. 86) and the pair of small
notches in the posterior lobes (Fig. 85)
also make this species distinctive. The
male palp of M. nigriventris has an em-
bolus that is relatively sliinmer and more
graceful than those of other species in the
M. nigriventris species group (compare
Fig. 84 with Figs. 92, 100, 109-110, 121).
The shape of the embolus and its distal
apophysis differs from other species (com-
pare Fig. 87 with Figs. 95, 103, 112, 124).
Variation. Average body length of 25 fe-
males examined 9.5 mm, range 7.2 to 11.5
mm. Average body length of 16 males ex-
amined 7.4 mm, range 5.3 to 9.5 mm.
Natural History. Spiders are commonly
found around Lake Titicaca living in me-
dium and large colonies among power
lines, Bolivian pines. Cactus, Bacharis,
rock outcroppings, and tall grasses (L.
Rayor, personal communication, and vari-
ous locality labels). Mature speciinens
have been collected throughout the year
except September and October (Fig. 310).
Median elevation, 3,900 m.
Distribution. High altitude regions of
southern Peru and western Bolivia (Map
5).
Records Examined. ARGENTINA Jujuy: Puma-
huasi, 22°17'S, 65°41'W, 8.xi.l970 (L. E. Pena,
MCZ). BOLIVIA La Paz: 45 mi S La Paz, 17°9'S,
67°36'W, 25.ii.1951 (E. S. Ross & Michelbacher,
CAS); 70 mi S La Paz, 17°30'S, 67°36'W, 25.ii.1951
(E. S. Ross & Michelbacher, CAS); La Paz, Avenida
Sport Club, 16°30'S, 68°9'W, 4.i.l959 (A. M. Nadler,
AMNH); La Paz, in garden of house, 16°30'S,
68°9'W, 15.iv.l959 (R. Walsh, AMNH); Lake Titicaca,
Copacabana, Yampupata, & Isla del Sol, 16°10'S,
69°5'W, 17.V.1995 (L. Rayor, MCZ); near La Paz,
16°30'S, 68°9'W, 24.V.1958 (R. Walsh, AMNH); S end
of Lake Titicaca, 100 km NW La Paz, 16°10'S,
69°5'W, 5.vii.l958 (R. Walsh, AMNH); Tialiuanaco,
Puma Puerto Ruins, 16°33'S, 68°42'W, l.ii.l973 (Ann
Moreton, MCZ). Oruro: 6 km N Challapata, 18°51'S,
66°47'W, 23.ii.1951 (E. S. Ross & Michelbacher,
CAS); Gorge Uhuschlucht, near Oruro, 17°59'S,
67°9'W, 7.ii.l954 (Forster & Schindler, ZSM). Potosi:
Villazon, 22°6'S, 65°36'W, 30.xii.l984 (L. E. Peiia,
AMNH). PERU Apurimac: Chincheros, 13°30'48"S,
73°42'47"W, 12.xii.l980 (C. Gold, CAS); Puna near
Abancay, 13°38'2"S, 72°52'52"W, 15.xii.l947 (W. Wey-
rauch, CAS). Ayacucho: Puquio, 14°42'S, 74°8'W,
15. iv. 1950 (F. Blancas, MUSM); San Antonio (Pu-
quio), 14°47'S, 74°7'W, l.xi.l985 (D. Silva, MUSM).
Cusco: Cheqquerec, 13°23'S, 72°8'W, 2.ix.l993 (J.
Ochoa Camara, MCZ); Cusco, 13°31'6"S,
71°58'41"W, 8.viii.l965 (P & B. Wygodzinsky,
AMNH). junin: 8 mi W Tarma, 11°25'S, 75°48'W,
6.i.l955 (E. I. Schlinger & E. S. Ross, CAS); Cochas
Bajo, 11 km W Tarma, 11°25'21"S, 75°46'11"W,
27iii.l988 (J. Palmer & D. Smith, MCZ); Cochas
Bajo, 11 km W Tarma, rock ledge in agricultural val-
ley, 11°25'21"S, 75°46'11"W, 29.iii.1988 (J. Palmer,
MCZ); Huancayo, 12°4'S, 75°14'W, 15.vi.l947 (W.
Weyrauch, AMNH); Oroya, 11°32'S, 75°54'W,
12.iv.l914 (M. P. Anderson, AMNH). Lima: Bosque
de Zarate, 11°53'S, 76°27'W, 18.i.l981 (J. Francke,
MUSM). Puno: 10 mi N Ayaviri, 14°45'S, 70°35'W,
l.iii.l951 (E. S. Ross & Michelbacher, CAS); Cama-
cane, 15°55'S, 69°50'W, 20.xi.l955 (L. E. Peiia,
IRSNB); Isla Taquih, Lago Titicaca, 15°46'S,
69°41'W, 23.xii.1980 (C. Gold, CAS); Juh (col. Chu-
cuito), 16°13'S, 69°27'W, 7.xi.l952 (F. Blancas,
MUSM); near Chucuito, Lago Titicaca, 15°50'S,
69°48'W, 10.iii.l953 (M. Koepcke, MUSM); Puna,
Lake Titicaca, 15°50'S, 70°2'W, 15.vi.l947 (W Wey-
rauch, AMNH); Puno, 15°50'S, 70°2'W (Soukup,
AMNH); Yunguyo, downtown plaza, 16°15'S, 69°5'W,
31.i.l973 (Ann Moreton, MCZ).
12. Metepeira tarapaca new species
Figures 92-99, 305; Map 7 |
Holotype. Male from Quillagua, Antofagasta, Chile,
4.ii.l965, L. E. Pena, in MCZ. The specific name
is a noun in apposition after a Chilean province
where it is abundant.
Description. Female paratype from
Quillagua, Antofagasta, Chile. Light red-
dish brown carapace, white in center and
around eyes with lateral posterior exten-
sions (Fig. 98). Legs yellowish white,
ringed brown at distal ends of articles. Fe-
mur I with row of four macrosetae on an-
terior side; three on anteroventral side.
Dorsal folium white with black speckles
(Fig. 98). Venter of abdomen black with
wide, white median line, sometimes
flanked by thinner white lines that togeth-
er form a U-shape posteriorly (Fig. 98).
Sternum black with median white line,
sometimes broken (Fig. 99). Ratio of eye
Metepeira • Piel 41
4500j
400O-
350O-
3000--
250a-
20oa-
1500- -
1000--
500- -
.-
cP o °
20 40 60 80 100
Percent of median white line on sternum
Figure 108. Elevation of collection localities of mature female
M. tarapaca with differing amounts of whiite on the sternum.
Spiders with a median white line covering 100% of the sternum
length are found at a wide altitude range. Spiders with a me-
dian white mark covering only a short length of the sternum
are only found at high elevations. Elevations estimated from
NOAA database of 5- by 5-minute geographic tiles.
diameters: posterior medians and anterior
medians 1.0, anterior laterals 1.3, posterior
laterals 1.2. Anterior median eyes separat-
ed by 1.4 diameters, posterior median eyes
by 1.0, anterior median eyes separated
from anterior laterals by 3.4 diameters of
anterior lateral eyes, lateral eyes separated
by 0.1 their diameters. Total length 7.8
mm. Carapace 3.2 mm long, 2.8 wide.
First femur 4.3 mm, patella and tibia 4.4,
metatarsus 4, tarsus 1.2. Second patella
and tibia 3.8 mm, third 2.1, fourth 3.2.
Male holotype. Light reddish brown
carapace, lighter around eyes and white
mark in center (Fig. 96). Legs yellowish
white, gradually growing darker toward
distal ends of articles. Femur I with row
of four macrosetae on anterior side; five on
anteroventral side. Dorsal folium, venter,
and sternum as in female (Figs. 96, 97).
Ratio of eye diameters: posterior medians
and anterior medians 1.0, anterior laterals
1.2, posterior laterals 1.1. Anterior median
eyes separated by 1.5 diameters, posterior
median eyes by 1.0, anterior median eyes
separated from anterior laterals by 2.5 di-
ameters of anterior lf Comparative Zoology, Vol. 157, No. 1
-40 -35
Latitude (Degrees North)
Figure 109. Days on which mature M. galatheae were col-
lected between 1937 and 1989 with latitude of the collection
locality. Seasonality appears to be more restricted in southern
regions than in northern regions.
Scale of abscissa: -150 = August 3; 1 = January 1; 150 =
May 30.
scape with a corresponding variation in the
position of the epigynal openings, it is
nonetheless possible to distinguish the fe-
males of M. galatheae from those of other
species in the M. nigriventris group.
Whether ventrally or posteriorly posi-
tioned, the epigynal openings and the
darkened shadows of the sclerotized re-
ceptacles beneath them are relatively
smaller than those of M. calamuchita, M.
tarapaca, and M. nigriventris (compare
Figs. 118-120 with Figs. 86, 94, 102). Also,
M. galatheae lacks the swollen posterior
lobes present in M. karkii (compare Figs.
118-120 with Fig. 123). The shape of the
embolus of M. galatheae varies significant-
ly (compare Fig. 109 with Fig. 110). How-
ever, unlike other species, the embolus of
M. galatheae has a distinct, round and
swollen protrusion (Fig. 117).
Variation. Average body length of 80 fe-
males examined 7.9 mm, range 4.8 to 12.5
mm. Average body length of 16 males ex-
amined 5.9 mm, range 3.5 to 8.1 mm. Epi-
gyna vary considerably. Many, similar to
the holotype, open ventrally and resemble
a posteriorly widened version of M. tara-
paca (Figs. 119, 120); cleared epigyna
show relatively straight ducts connecting
the epigynal openings with the seminal re-
ceptacles. These have a short distance be-
tween the openings and the hood of the
scape. Others, usually in southern Chile,
have posterior openings and look surpris-
ingly different (Fig. 118); cleared epigyna
show S -shaped ducts connecting the open-
ings with the seminal receptacles. These
have an extended wrinkled area between
the openings and the hood of the scape.
However, several females [e.g., CHILE
Bio-Bio: Chilian, 8.xi.l976 (G. Moreno,
AMNH); Las Lajuelas, ll.i.l976 (G. Mo-
reno, AMNH)] have epigyna that appear
to be intermediate between the two forms.
Furthermore, no somatic features were
found to be sufficiently different, and little
corresponding variation was found among
sympatric males. It is possible that further
collecting efforts will discover correspond-
ing males, and future molecular studies
may show that speciation has, in fact, oc-
curred. But in the meantime, I am opting
to treat both varieties as belonging to the
same species.
Natural History. Although mature spec-
imens have been collected throughout the
year (Fig. 304), the seasonality of this spe-
cies appears to depend on its latitude. At
the 45th southern parallel, spiders are usu-
ally found in late November and Decem-
ber; at the 40th parallel, spiders occur be-
tween October and Febrtiary; and at the
30th parallel, they are collected year round
(Fig. 108). Median elevation, 550 m. Spi-
ders are found on Patagonian scrub,
dunes, and wire fences.
Distribution. Chile and Argentina (Map
7).
Records Examined. ARGENTINA Buenos Aires:
Felipe Sola, 38°1'S, 62°50'W, 15.1.1944 (Prosen,
MLP); Patagones, 40°48'S, 62°59'W, 15.ii.l937 (J. M.
Viana, MACN); Sierra de la Ventana, 38°9'S,
61°48'W, 15.iii.l939 (J. C. Gario, MACN). Catamar-
ca: Mutquin, 28°19'S, 66°10'W, 15.1.1963 (O. de Fer-
rariis, AMNH). Chubut: 15 km S Epuyen, 42°22'S,
71°21'W, 15.1.1986 (P A. Goloboff, N. I. Platnlck, &
R. T. Schuh, AMNH); 19.5 km E Shaman, 44°27'S,
70°30'W, 19.xi.l966 (E. I. Schllnger & M. E. Irwin,
CAS); 3 km N Puerto Lobos" 41°59'S, 65°6'W,
14.xii.l966 (E. I. Schllnger & M. E. Irwin, CAS); 35
km E Esquel, 42°54'S, 70°53'W, 18.xl.l966 (E. I.
Schhnger & M. E. Irwin, CAS); El Hoyo [?], 42°4'S,
71°30'W (A. Kovacs, AMNH), 10.1.1962 (Andor Ko-
vacs, AMNH); Epuyen, 42°15'S, 71°23'W, 18.xl.l962
(Andor Kovacs, AMNH); Leleque, 42°28'S, 71°6'W,
12.11.1965 (Andor Kovacs, AMNH); Los Manantlales,
Metepeira • Piel
45
A A
Figures 110-120. Metepeira galatheae {ThoreW) (sp. 14 [110-114,116,117,120] 46°33'S, 71°57'W; [115,119] 29°50'S, 70°2'W;
[118] 33°30'S, 71°25'W). 110, male palpus, mesal. Ill, male, dorsal. 112, male, ventral. 113, female, dorsal. 114, female,
ventral. 115, epigynum, posterior. 116, epigynum, posterior. 117, male embolic division, ventral. 118-120, epigynum, ventral.
Figures 121-128. Metepeira /car/f/V (Tullgren) (sp. 15; 51°38'S, 69°13'W). 121, male palpus, mesal. 122, epigynum, posterior.
123, epigynum, ventral. 124, male, dorsal. 125, male, ventral. 126, female, dorsal. 127, female, ventral.
Scale bars: dorsum and venter figures 1.0 mm.
N of Comodoro-Rivadavia, 45°28'S, 69°29'W,
19.xi.l985 (L. E. Peiia, AMNH); N of Camarones,
Cantera, Namuncura, 44°46'S, 65°42'W, 17..>d.l9S5
(L. E. Pena, AMNH); Rio Turbio, 42°13'S, 71°41'W
(Andor Kovacs, AMNH), 12.1.1962 (Andor Kovacs,
AMNH). Cordoba: 12 mi W Sampacho, 33°23'S,
64°43'W, 7.ii.l951 (E. S. Ross & Michelbacher, CAS);
Arguello, 31°21'S, 64°15'W, 15.xii.l943 (J. A. De Car-
lo, MACN); Calamuchita, 32°4'S, 64°33'W,
15.xii.l940 (J. M. Viana, MACN); Sampacho, 33°23'S,
64°43'W, 7.ii.l951 (E. S. Ross & Michelbacher, CAS).
Mendoza: Between Beazley and San Rafael, 34°10'S,
67°29'W, 4.iii.l983 (L. E. Pefia, AMNH); Mendoza,
32°53'S, 68°49'W, 30.iii.l965 (H. W. Levi, MCZ); Us-
pallata, 32°35'S, 69°20'W, 7.iii.l983 (L. E. Pefia,
AMNH). Neuquen: Catan Lil, Charaliuilla, 39°45'S,
46 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
70°37'W, 15.ii.l971 (O. de Ferrariis, AMNH); Cuba
del Leon [?], 39°9'S, 70°53'W, 15.1.1975 (Mauiy,
MACN); Lago Alumine, 38°55'S, 71°9'W, 15.1.1976
(O. de Ferrariis, AMNH); Zapala, 38°54'S, 70°4'W,
15.1.1958 (J. R. Navas, MACN). Rio Negro: Cerro
Alto [?], 41°8'S, 70°40'W (MACN); Co. Leones,
source of Rio Limay, 40°33'S, 70°26'W, 28.11.1959 (J.
R. Navas, MACN); El Bolson, 41°58'S, 71°31'W,
1.11.1961 (A. Kovacs, AMNH), 17.X.1961 (Andor Ko-
vacs, AMNH); El Bolson, 41°58'S, 71°35'W, 2.11.1965
(Andor Kovacs, AMNH); Ceneral Roca, 39°3'S,
67°32'W, 15.lx.1964 (Bachmann, MEG). Salta: Mauiy
[?], 24°40'S, 65°45'W, 15.1.1975 (MACN). San Juan:
10 km N Matagusanos, 31°10'S, 68°38'W, 13.1.1983
(L. E. Peiia, AMNH). San Luis: INTA Experimental
Station, E of Villa Mercedes, 33°40'S, 65°27'W,
8.xil.l967 (C. R. Ward, CAS). Santa Cruz: 2.4 km S
Fltz Roy, 47°2'S, 67°15'W, 12.xll.l966 (E. I. Schllnger
& M. E. Irwin, CAS). Tucumdn: San Miguel de Tu-
cuman, IML gardens, 26°49'S, 65°13'W, 19.xil.l979
(L. A. Stange, FSCA). BRAZIL Mato Grosso: Campo
Grande, 20°27'S, 54°37'W, 7.11.1952 (M. Alvarenga,
MZSP). CHILE Aconcagua: W end tunnel, 85 km S
Illapel, 32°49'S, 71°7'W, 29.xl.1950 (E. S. Ross &
Mlchelbacher, CAS). Alsen: 8 km W Chile Chlco,
46°33'S, 71°57'W, 22.xi.1966 (E. I. Schllnger & M.
E. Ii^wln, CAS); Chile Chlco, near lake, 46°33'S,
71°43'W, 21.xl.1966 (E. I. Schllnger & M. E. Invln,
CAS). Antofagasta: 6 km N Muelle de Pledra, N Tal-
tal, 25°21'S, 70°30'W, 4.11.1942 (Junius Bird, AMNH);
Caleta Hueso Parado, Taltal, 25°22'S, 70°28'W,
1.11.1941 (Junius Bird, AMNH); Coblja, 22°33'S,
70°16'W (ZMUC); Quebrada Paposo, 25°2'S,
70°27'W, 3.11.1989 (L. Stange, FSCA). Araucania: Pe-
mehue [?], 38°3'S, 71°43'W (L. E. Pena, IRSNB);
Villarrlca, 36°16'S, 72°13'W, 25.xl.1963 (G. F Ed-
munds, AMNH). Atacama: 50-60 km S Copiapo,
27°51'S, 70°20'W, 24.vlll.1966 (E. I. Schllnger & M.
E. Irwin, CAS); Copiapo, 27°22'S, 70°20'W (Cartis,
MNRJ); Rio Copiapo, by the sea, 27°19'S, 70°56'W,
13.vl.1968 (L. E. Pena, MCZ). Bio-Bio: 4 km E road
to Pinto, 36°42'S, 71°53'W, 4.1.1976 (B. Moreno,
AMNH); Chilian, 36°36'S, 72°7'W, 2.1.1976 (G. Mo-
reno, AMNH), 21.11.1978 (G. Moren, MCZ); Chilian,
In cemetery, 36°36'S, 72°7'W, 8.xl.l976 (G. Moreno,
AMNH); Cuesta de QuUmo, Chilian, 36°38'S,
72°12'W, 13.xl.1976 (G. Moreno, AMNH); El Aban-
Ico, 37°20'S, 71°31'W, 30.xll.l950 (E. S. Ross &
Mlchelbacher, CAS); Las Lajuelas, 36°39'S, 72°8'W,
11.1.1976 (G. Moreno, AMNH). Concepclon,
36°50'S, 73°3'W (L. E. Pena, IRSNB); Concepclon:
Salta de Rio Laja, 37°13'S, 72°23'W, 30.1.1951 (E. S.
Ross & Mlchelbacher, CAS); Nuble: 50 km E San
Carlos, 36°25'S, 71°6'W, 26.xll.1950 (E. S. Ross &
Mlchelbacher, CAS); Nuble: Cordillera de Chilian
[?], 36°51'S, 7r24'W, 1.11.1947 (L. E. Peiia, IRSNB);
Rio Andallen, 36°44'S, 73°1'W, 25.111.1979 (S. Gu-
tierrez, MCZ). Coquinibo: 20 ml E La Serena,
29°54'S, 70°56'W, 3.vll.l950 (E. S. Ross & Mlchel-
bacher, CAS); 5 ml N Ovalle, 30°31'S, 71°12'W,
l.xil.l950 (E. S. Ross & Mlchelbacher, CAS); Banos
del Toro, 29°50'S, 70°2'W, 15.11.1947 (L. E. Pena,
IRSNB); Cerro Tahnay, 30°50'29"S, 71°37'14"W,
29.xl.1961 (A. F. Archer, AMNH); Cuesta las Cardas,
Ovalle Rd., 30°17'S, 71°16'W, 13.xl.l961 (R. Wagen-
knecht, AMNH); Hacienda Illapel, 31°36'S, 71°7'W,
3.X1.1954 (L. E. Pena, IRSNB), 19..X.1966 (E. I.
SchUnger, M. E. Irwin, & L. E. Pena, CAS); Illapel:
Salamanca: Fundo Quelen, 31°52'S, 70°52'W,
30.lv.1961 (A. F. Archer, AMNH); La Serena,
29°54'28"S, 7ri5'15"W, 15.11.1947 (L. E. Pena,
IRSNB); Loma de Penuelas, 6 km S La Serena,
29°57'S, 71°18'W, 28.xl.1961 (A. F. Archer, AMNH);
Qullacan, 16 km E La Serena, 29°54'S, 71°5'W,
2.x. 1961 (R. Wagenknecht, AMNH). Los Lagos: Purr-
anque, 40°55'S, 73°10'W, 15.11.1955 (Edwin Reed,
AMNH); Rio Bueno, 40°19'S, 72°58'W (L. E. Pena,
IRSNB); Valdlvla: Neltume, 39°48'S, 71°57'W,
23.xi.1988 (V. & B. Roth, CAS). Malleco: Angol,
37°48'S, 72°43'W, 29.1.1951 (E. S. Ross & Mlchel-
bacher, CAS). Maule: Linares, 35°51'S, 71°36'W (L.
E. Pena, IRSNB), 15.1.1947 (L. E. Pena, IRSNB);
Mlraflores, Pedag. [?], 35°55'S, 71°39'W (Toro,
AMNH). O'Higgins: Cheplca, 34°44'S, 71°17'W,
15..X11.1947 (L. E. Peiia, IRSNB). Region Metropoli-
tana: 34 km W Sandago, 33°30'S, 71°25'W,
19.xll.1950 (E. S. Ross & Mlchelbacher, CAS); Ba-
tuco, nr. Santiago, 33°13'S, 70°47'W (Gull. Mann,
AMNH); Lampa, 33°17'S, 70°54'W, l.v.1979 (L. E.
Peiia, AMNH). Santiago: El Golf [?], 33°30'S,
71°25'W, 9.iv.l961 (A. F Archer & J. Aros, AMNH);
Santiago, 33°30'S, 71°25'W (L. E. Peiia, IRSNB),
1.11.1973 (W. C. Sedgvdck, MCZ). Talca: 22 ml N Tal-
ca, 35°7'S, 71°40'W, 22.xll.1950 (CAS). Valparaiso:
Concon, in cow farm, 32°55'S, 71°31'W, 4.111.1962 (H.
Morales, AMNH); La Cruz, 32°53'S, 71°16'W,
18.1.1973 (W. C. Sedgwick, MCZ); Llay-Llay, 32°51'S,
70°58'W, 20.1.1973 (W. C. Sedgwick, MCZ); Los Mai-
tenes [?], 32°59'S, 71°15'W, 14.X.1954 (L. E. Peiiai
IRSNB); Quintay, 33°11'S, 71°42'W, 19.11.1967 (E. I.
Schhnger, CAS); Valparaiso, 33°2'S, 71°38'W (Edwin
Reed, AMNH).
15. Metepeira /car/c/7 (Tullgren)
Figures 121-128, 303; Map 5
Araneus karkii Tullgren, 1901: 219, 259. Female lio-
lotype from Kark, Chile in the SMNH, examined.
Metepeira labyrinthea: — Roewer, 1942: 868. Bonnet,
1957: 2821. Erroneous synonymy.
Description. Female from Rio Gallegos,
Santa Cruz Province, Argentina. Carapace
reddish brown with white setae, light
around eyes with lateral posterior exten-
sions (Fig. 127). Legs light yellow, articles
annulated distally. Femur I with row of
three to four macrosetae on anterior side;
one to four on anteroventral side. Dorsum
covered with denser, longer, black and
Metepeira • Piel
47
white setae than in most species. Folium
mostly white with brown speckles (Fig.
127). Venter brownish gray with wide
white median line; pair of large white spots
on either side of spiracle. A pair of thin
white lines, parallel to and on either side
of median line, sometimes connect to a
pair of thinner transverse white lines: one
just posterior to the epigynal groove, one
just anterior to the spinnerets (Fig. 128).
Sternum dark reddish brown (Fig. 128).
Ratio of eye diameters: posterior inedians
and anterior medians 1.2, anterior laterals
1.4, posterior laterals 1.5. Anterior median
eyes separated by 1.8 diameters, posterior
inedian eyes by 1.0, anterior median eyes
separated from anterior laterals by 4.2 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.3 their diameters. To-
tal length 9.5 inm. Carapace 4 mm long,
3.4 wide. First femur 4 mm, patella and
tibia 4.4, metatarsus 3.7, tarsus 1.2. Sec-
ond patella and tibia 3.9 mm, third 2.5,
fourth 3.8.
Male from Rio Gallegos, Santa Cruz
Province, Argentina. Carapace reddish
brown with lighter eye region, lateral pos-
terior extensions, and median arrowhead
inark (Fig. 125). Legs light yellow, articles
distally annulated reddish brown. Femur I
with row of three to four macrosetae on
anterior side; five to six on anteroventral
side. Dorsal folium mostly white with
brown speckles (Fig. 125). Venter and
sternuiTi as in female (Fig. 126). Ratio of
eye diameters: posterior medians and an-
terior medians 1.0, anterior laterals 1.3,
posterior laterals 1.2. Anterior inedian
eyes separated by 1.7 diameters, posterior
inedian eyes by 1.2, anterior median eyes
separated from anterior laterals by 2.8 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.4 their diameters. To-
tal length 5.7 mm. Carapace 2.8 mm long,
2 wide. First femur 4.3 mm, patella and
tibia 4.3, metatarsus 4.1, tarsus 1.2. Sec-
ond patella and tibia 3.5 mm, third 1.9,
fourth 2.9.
Diagnosis. Females are easily separated
from other species in the M. nigriventris
group by the thick posterior epigynal lobes
(compare Fig. 123 with Fig. 119). The dis-
tal embolic apophysis does not protrude
out from under the terminal apophysis
(Fig. 124) as it does in M. galatheae, M.
tarapaca, and M. nigriventris (Figs. 87, 95,
119). The embolus of M. karkii differs
from M. calaniuchita by lacking the in-
wardly curved "dewlap" under the embo-
lus (compare Fig. 121 with Fig. 103).
Variation. Average body length of 13 fe-
males examined 6.8 mm, range 5 to 8.2
mm. Average body length of five males ex-
amined 4.4 mm, range 2 to 5.3 mm. Dor-
sal folia vaiy from white with little contrast
and indistinct fleur-de-lis to darker with
more contrast and distinct fleur-de-lis.
Natural History. This species appears to
be strongly seasonal: mature specimens
have been collected between November
and March (Fig. 303). Median elevation,
300 m. Spiders are found in pampas (tree-
less grassland).
Distribution. Lower altitudes in south-
ern Argentina and Chile (Map 5).
Records Examined. ARGENTINA Chubut: Puerto
Piramides, Peninsula Valdes, 42°34'S, 64°17'W,
12.xi.l988 (V. & B. Rodi, CAS). Neuqtien: Laguna
Blanca, 39°3'S, 70°23'W, 15.iii.l959 (J. Nara,
MACN); Zapala, 38°54'S, 70°4'W, 15.1.1958 (J. R. Na-
vas, MACN); Zapala, Laguna Blanca, 38°54'S,
70°4'W, 15.1.1959 (J. R. Navas, MACN). Rto Negro:
Cerro Alto [?], 41°8'S, 70°40'W (MACN); Coronel
Juan Jose Comez, 39°2'S, 67°39'W, 15..\i.l945 (Ibarra
Grasso, MLP); Ne-Luan, 41°25'S, 68°45'W (MACN).
Santa Cruz: Laguna Calafate, Precordllfera [?],
50°55'S, 70°9'W, 22.1.1967 (P San Martin, MCZ); Rio
Gallegos, 51°38'S, 69°13'W, 20.1.1967 (P San Martin,
MCZ). BOLIVIA Santa Cniz: Patagonia: Estancia
Monte, cerca Rio Coyby [?], 50°14'S, 68°55'W (B.
Brown, AMNH). CHILE Magallanes: 4 km W La-
guna Amarga, 50°59'S, 72°49'W, 8.xil.l966 (E. I.
Schlinger & M. E. Irwin, CAS); Kark, 51°17'30"S,
72°32'30"W, 13.111.1899 (E. Nordenskiold, NRMS).
Metepeira compsa Group
The three species in the M. co77ipsa
group include Metepeira compsa, Metepei-
ra roraima, and Metepeira gressa. Along
with the Metepeira nigriventris group, this
group has a median apophysis with teeth
on the face of the keel. Unlike the Mete-
48 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
peira nigriventris group, this group has
smaller, slimmer distal embolic apophyses
that do not arch up over the embolus tip
(compare Fig. 149 with Fig. 84). The
openings to the epigynum have distinct,
sclerotized edges and are clearly visible ei-
ther as circles (Figs. 131, 134) or eye-
shaped ovals (Figs. 143, 151).
1 6. Metepeira compsa (Chamberlin)
Figures 129-140, 311; Map 8
Aranea compsa Chamberlin, 1916: 252, fig. 6, ? . Fe-
male holotype from Ollantaitambo, Cusco, Peru, in
the MCZ, examined. Bonnet, 1955: 462.
Araneus lahijrintheus: — Petnmkevitch, 1926: 27. Er-
roneous synonymy.
Metepeira virginensis Chamberlin and Ivie, 1942: 70,
figs. 188-190, 9,8. Female holotype from St.
Thomas, U.S. Virgin Islands, in AMNH, examined.
NEW SYNONYMY.
Metepeira latigijna Chamberlin and Ivie, 1942: 70,
figs. 191, 192, 5 . Female holotype from Porto Ale-
gre [Baliia], Brazil, in AMNH, examined. NEW
SYNONYMY.
Metepeira compsa: — Chamberlin and Ivie, 1942: 71,
figs. 193-195, $, c?.
Metepeira labyrinthea: — Biyant, 1942: 346.
Metepeira perezi Archer, 1965: 132, figs. 14, 16, 3,
9 . Male holotype from subexperiment station, Is-
abela, Puerto Rico, in AMNH, examined. Brignoli,
1983: 276. NEW SYNONYMY.
Metepeira vaurieonim Archer, 1965: 133, figs. 15, 19,
S, ?. Male holotype from Usine de Robert, Mar-
tinique, in AMNH, lost. BrignoU, 1983: 276. Lo-
pez, 1993: 10, 11, figs. 1-4, 11, 6. NEW SYN-
ONYMY.
Note. Although the type for M. vaurieonim is
lost, all examined records from Martinique (includ-
ing ones from "Usine de Robert") belong to M.
compsa.
Description. Female from Savonet, Cu-
rasao, Netherlands Antilles. Carapace light
around eyes with lateral posterior exten-
sions (Figs. 135, 137). Only tibia IV
ringed. Femur I with row of four macro-
setae on anterior side; two or three fine
setae on anteroventral side. Dorsum of ab-
domen with usual Metepeira folium,
though whiter than usual in some speci-
mens (Figs. 135, 137). Venter with wide
white median line flanked by two thin
(Fig. 136) or wide (Fig. 138) white lines;
pair of white spots on either side of spi-
racle. Sternum has wide median white line
widening anteriorly, sometimes broken
(Figs. 136, 138). Ratio of eye diameters:
posterior medians and anterior medians
1.0, anterior laterals 1.3, posterior laterals
1.2. Anterior median eyes separated by 1.3
diameters, posterior median eyes by 0.7,
anterior median eyes separated from an-
terior laterals by 1.8 diameters of anterior
lateral eyes, lateral eyes separated by 0.2
their diameters. Total length 3.9 mm. Car-
apace 1.8 mm long, 1.4 wide. First femur
1.9 mm, patella and tibia 1.9, metatarsus
1.5, tarsus 0.7. Second patella and tibia 1.7
inm, third 0.9, fourth 1.5.
Male from Savonet, Curasao, Nether-
lands Antilles. Carapace light around eyes
with lateral posterior extensions. Slightly
lighter median triangular mark anterior to
thoracic furrow (Fig. 139). Legs lightly
ringed. Femur I with row of four macro-
setae on anterior side, four on anteroven-
tral side. Dorsum, venter, and sternum as
in female (Figs. 139, 140). Ratio of eye
diameters: posterior medians and anterior
medians 1.0, anterior laterals 1.3, posterior
laterals 1.2. Anterior median eyes separat-
ed by 1.3 diameters, posterior median eyes
by 0.6, anterior median eyes separated
from anterior laterals by 1.4 diameters of
anterior lateral eyes, lateral eyes separated
by 0.3 their diameters. Total length 3.6
mm. Carapace 1.8 min long, 1.4 wide.
First femur 2.7 mm, patella and tibia 2.8,
metatarsus 2.4, tarsus 0.9. Second patella
and tibia 2.3 mm, third 1.2, fourth 1.8.
Diagnosis. Unlike other species, the
openings to the epigynum of M. compsa
are small and almost perfectly round and
sclerotized around the riin (Fig. 131). In
Peruvian and Argentinean populations the
scape can be extremely wide, often entire-
ly covering a ventral view of the openings
(Fig. 134). The distal embolic apophysis
on the male palp extends straight from its
base and parallel to the embolus tip (Figs.
129, 132), in contrast to those of other spe-
cies in the M. compsa species group that
lift away from the embolus tip (Figs. 141,
149).
Variation. Average body length of 22 fe-
Metepeira • Piel 49
roraima
(17)
Figures 129-140. Metepeira compsa (Chamberlin) (sp. 16 [129-131,135,136,139,140] 12°20'N, 69°7'W; [132] 4°30'S, 81°8'W;
[133,134] 4°51'S, 80°46'W; [137,138] 17°5'N, 61°42'W). 129, male palpus, mesal. 130, epigynum, posterior. 131, epigynum,
ventral. 132, male palpus, mesal. 133, epigynum, posterior. 134, epigynum, ventral. 135, female, dorsal. 136, female, ventral.
137, female, dorsal. 138, female, ventral. 139, male, dorsal. 140, male, ventral.
Figures 141-148. Metepeira roraima new species (sp. 17 [141-147] 3°22'N, 60°19'W; [148]3°21'N, 76°33'W). 141, male palpus,
mesal. 142, epigynum, posterior. 143, epigynum, ventral. 144, male, dorsal. 145, male, ventral. 146, female, dorsal. 147, female,
ventral. 148, female, dorsal.
Scale bars: dorsum and venter figures 1 .0 mm.
males examined 5 mm, range 3.5 to 7.3
mm. Average body length of 18 males ex-
amined 3.6 mm, range 2.4 to 4.7 mm.
Enormous variation in the shape of the
scape can be seen in this species: popula-
tions in the Caribbean and northeastern
South America have a narrow scape (Fig.
131), whereas populations in Argentina
and Peru have a thick scape (Fig. 134).
Somewhat less consistently parallel differ-
ences can be seen in the shape of the me-
dian apophysis: flagella are centered in Ca-
ribbean and northeastern South America
(Fig. 129) but shifted to the left in Argen-
tina and Peru (Fig. 132). Further subtle
differences appear in the shape of the em-
50 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
bolus tip and embolus arm (compare Fig.
129 with Fig. 132). These male and female
genital differences are not sufficiently con-
sistent to provide solid evidence for spe-
ciation; thus, these populations will be
treated as one species.
Natural History. Mature speciinens
have been collected throughout the year
(Fig. 311). Altitudes range from just above
sea level (for Caribbean and eastern South
American localities) to 4,000 m (for Pe-
ruvian localities). Spiders have been found
in everything from second growth mixed
exotics (e.g., mango, citrus, banana) and
mangroves to grasses and semidesert chap-
arral.
Distribution. Puerto Rico to northern
Argentina, but absent from the Amazon
(Map 8).
Records Examined. ARGENTINA Buenos Aires:
Punta Lara, 34°49'S, 57°59'W, 15.ii.l941 (F. Moneos,
MACN). Chaco: Basail, 27°52'S, 59°18'W, 18.iv.l942
(MACN). Corrientes: Paso de la Patria, 27°19'S,
58°35'W, 15.1.1966 (M. E. Galiano, MEG). Entre
Rws: Parana, 31°44'S, 60°32'W (Rosenzwaig, MLP);
Salto Grande, 31°13'S, 57°56'W, 15.iii.l964 (M. E.
Galiano, MEG). Misiones: Posadas, 27°23'S, 55°53'W,
15.ix.l963 (M. E. Galiano, MEG). Neuquen: Piedra
del Aguila, 40°3'S, 70°5'W, 18.vii.l978 (Mision Cien-
tifica Danesa, ZMUC). Santa Fe: Tostado, 29°14'S,
61°46'W (A. Giai, MACN). BOLIVIA La Paz: Apolo,
14°43'S, 68°31'W, 10.viii.l989 (L. E. Pefia, AMNH).
BRAZIL Bahia: Arquipelago dos Abrolhos, 17°40'S,
38°50'W, 28.xii.1887 (U.S. E.G., Voy. of Albatross,
USNM); Parque Ondina, Salvador, 12°59'S, 38°31'W,
25.vii.1962 (A. F. Archer, AMNH); Porto Alegre,
18°5'S, 39°34'W (AMNH). Minas Gerais: Pedra /lzuI,
16°1'S, 41°16'W, 15.xii.l970 (F. M. Oliveira, AMNH).
Periiambuco: Pernambuco [?], 8°3'S, 34°54'W,
12.iii.l927 (SMF), 8.iii.l955 (SMF). Rio de Janeiro:
Lagomar [?], Macae, near sea, 22°23'S, 41°47'W,
17.vii.l986 (R. L. C. Baptista, MZSP). Rio Grande do
Sal: Montenegro, 29°42'S, 51°28'W, 3.xi.l977 (E. H.
Buckup, MGN); Rambo [?], 30°4'0"S, 51°11'W
(MNRJ); Sao Leopoldo, 29°46'S, 51°9'W, 14.X.1965
(Gelia Valle, MZSP). Sao Paulo: Rubiao, Jr. [?], Bo-
tucatu, 22°52'S, 48°26'W, 25.iv.1988 (R. L. G. Bap-
tista, MZSP); Sao Paulo, Guamja, 24°0'S, 46°16'W,
24.vii.1983 (R. Sievers, AMNH). BRITISH WEST
INDIES Anegada: 18°45'N, 64°20'W, 12.xi.l966
(Harry Beatty, AMNH); center, nr. salt pond,
18°45'N, 64°20'W, 4.vi.l966 (Island Project Staff,
Univ of Puerto Rico, AMNH); West end, 18°45'N,
64°22'W 4.vi.l966 (Island Project Staff, Univ. of
Puerto Rico, AMNH); Anegada Settlement: 18°45'N,
64°20'W, 5.vi.l966 (Island Project Staff, Univ. of
Puerto Rico, AMNH). Antigua: Goolidge Airport,
17°6'N, 61°51'W, 15.xi.l967 (N. L. H. Krauss,
AMNH); Devils Bridge, 17°5'N, 61°42'W, 30.vi.l963
(Rick & E. N. Kjellesvig-Waering, AMNH); Jolly
Beach, 17°4'N, 61°53'W 20.ix.l963 (E. N. Kjellesvig-
Waering, AMNH); Lignum Vitae Bay: Jolly Beach [?],
17°4'N, 61°53'W, 19.vi.l968 (E. N. Kjellesvig-Waer-
ing, AMNH); Redonda Island, from webs spun be-
tween boulders on beach, 16°55'N, 62°19'W,
10.ivl9.56 (J. F G. Glarke, USNM); Reeds Point, nr.
Jolly Beach; 17°4'N, 61°53'W, 2.vii.l963 (E. N. Kjel-
lesvig-Waering, AMNH); Saint John's, 17°6'N,
61°51'W, 15.viii.l967 (N. L. H. Krauss, AMNH),
15.xii.l967 (N. L. H. Krauss, AMNH). Dead Mans
Ghest [?]: 18°22'N, 64°.34'W, 26.vl966 (Island Pro-
ject Staff, Univ. of Puerto Rico, AMNH). East Seal
Dog: 18°30'N, 64°25'W 7.vi.l966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). George Dog Is-
land: 18°30'N, 64°27'W, 7.vi.l966 (Island Project
Staff, Univ. of Puerto Rico, AMNH); 4 mi W Virgin
Gorda Isl, high on diy cliffs, 18°30'N, 64°27'W,
25.iii.1979 (K. Johnson, AMNH). Ginger Island:
18°24'N, 64°28'W, 25.V.1966 (Island Project Staff,
Univ of Puerto Rico, AMNH). Great Dog: 18°29'N,
64°27'W, 7.vi.l966 (Island Project Staff, Univ. of
Puerto Rico, AMNH). Green Cay, near Tortola:
18°27'N, 64°42'30'W, 14.viii.l965 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Island O. near
Anegada Setdement: 18°45'N, 64°20'W, 5.vi.l966 (Is-
land'Project Staff, Univ. of Puerto Rico, AMNH). Is-
land R. near Anegada Settlement: 18°45'N, 64°20'W,
4.vi.l966 (Island Project Staff, Univ. of Puerto Rico,
AMNH). ]ost Van Dyke: 18°28'N, 64°45'W,
30.viii.l965 (H. Heatwole, R. Levins & F Mac-
Kenzie, AMNH). Large mangrove patch nr. Settle-
ment Anegado: 18°45'N, 64°20'W [?], 5.vi.l966 (Is-
land Project Staff, Univ. of Puerto Rico, AMNH). Lit-\
tie Camanoe: 18°28'N, 64°33'W, 25.vl966 (Island
Project Staff, Univ. of Puerto Rico, AMNH). Little
Josi Van Dyke: 18°27'N, 64°43'W, 27.vii.1965 (Island
Project Staff, Univ of Puerto Rico, AMNH). Little
Tobago: 18°26'N, 64°51'W 4.iv.l966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Montserrat:
Gage's Soufriere [?], 16°43'N, 62°10'W 28.vii.1972
(N. L. H. Krauss, AMNH); Plymouth, 16°42'N,
62°13'W, 15.xi.l967 (N. L. H. Krauss, AMNH). Neck-
er Island: 18°33'N, 64°21'W, 6.vi.l966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). nr. Anegada Set-
tlement: Byer's Bache [?], 18°45'N, 64°20'W,
5.vi.l966 (Island Project Staff, Univ. of Puerto Rico,
AMNH). Peter Island: 18°22'N, 64°35'W, 6.vii.l965
(Island Project Staff, Univ of Puerto Rico, AMNH),
12.V.1966 (Percy Chubb, AMNH). Prickly Pear Is-
land: 17°10'N, 61°48'W, 6.vi.l966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Saint Christo-
pher Island: Basseterre, 17°18'N, 62°43'W, 6.ii.l968
(B. Malkin, AMNH). Salt Island: 18°23'N, 64°31'W,
24.V.1966 (Island Project Staff, Univ. of Puerto Rico,
AMNH). Sandy Key: near Tortola, 18°13'N, 63°7'W,
31.viii.l965 (H. & A. Heatwole, AMNH). Tobago Is-
land: 18°27'N, 64°48'W, 2.iv.l966 (Island Project
Metepeira • Piel
51
Staff, Univ. of Puerto Rico, AMNH). Tortola:
18°27'N, 64°36'VV, 8.vii.l958 (A. F. Archer, C. Hel-
sley, & M. Sanderson, AMNH); Beef Island, 18°27'N,
64°31'W, 21.vii.l965 (H. Heatwole, R. Levins & F.
MacKenzie, AMNH); Greater Camanoe Isl, 18°29'N,
64°32'W, l.vii.l965 (Island Project Staff, Univ. of
Puerto Rico, AMNH); Little Thatch Island, 18°23'N,
64°42'W, 16.viii.l965 (Island Project Staff, Univ. of
Puerto Rico, AMNH); Long Bay Estate, 18°24'N,
64°41'W, 24.vii.1965 (Island Project Staff, Univ. of
Puerto Rico, AMNH); Marina Key [?], 18°27'N,
64°36'W, 4.vii.l965 (H. Heatwole & R. Levins,
AMNH); Prospect Reef, S Roadtown, 18°25'N,
64°36'W, 23.iii.1979 (K. Johnson, AMNH), 31.iii.l979
(K. Johnson, AMNH); Road to town, 18°27'N,
64°36'W, 15.vii.l972 (N. L. H. Krauss, AMNPI);
Sandy Spit [?], 18°27'N, 64°36'W, 14.viii.l965 (Island
Project Staff, Univ. of Puerto Rico, AMNH). Toi-tola
Island: 18°27'N, 64°36'W, 15.vii.l965 (Island Project
Staff, Univ of Puerto Rico, AMNH). Virgin Gorda:
18°30'N, 64°24'W, 15..xi.l966 (Hany Beatty, AMNH);
Baths & Devils Bay, 18°26'N, 64°27'W, 25.vi.1966
(Island Project Staff, Univ. of Puerto Rico, AMNH);
Cooper Mine Trail 18°26'N, 64°26'W, 25.vi.1966 (Is-
land Project Staff, Univ. of Puerto Rico, AMNH); Sa-
vana Bay & Pond Bay, 18°28'N, 64°25'W, 27.vi.1966
(Island Project Staff, Univ. of Puerto Rico, AMNH);
Trelhs Bay (W end of island), nr. harbor, 18°27'N,
64°26'W, 27.iii.1979 (K. Johnson, AMNH); Virgin
Gorda Mountain, 18°30'N, 64°24'W, 26.vi.1966 (Is-
land Project Staff, Univ. of Puerto Rico. AMNH).
Virgin Islands Area: Island Q [?], 18°45'N, 64°20'W,
5.vi.l966 (Island Project Staff, Univ. of Puerto Rico,
AMNH). West Seal 'Dog Island: 18°29'N, 64°28'W,
7.vi.l966 (Island Project Staff, Univ. of Puerto Rico,
AMNH). CHILE Tarapaca: Arica, 18°29'S, 70°20'W,
28.i.l973 (W. C. Sedgwick, MCZ); Azapa Arica,
18°31'S, 70°11'W, 9..xi.l955 (L. E. Peiia, IRSNB);
Lluta, 18°24'S, 70°19'W, 12.xi.l955 (L. E. Pena,
IRSNB); Rio Lluta, 18°24'S, 70°19'W, I2.xi.l955 (L.
E. Pena, IRSNB); Sobraya, 18°32'S, 70°9'W,
10.xi.l955 (L. E. Peila, IRSNB); Taltape, Camarones
Valley, 18°59'S, 69°47'W, 29.i.l973 (MCZ). COLOM-
BIA Magdalena: Cabaila "Villa Culebra," 10 km E
Station Marta, 11°12'N, 74°7'W, 15.X.1985 (H.-G.
Miiller, SMF); Casajera [?], 11°0'N, 74°15'W,
l.ii.l974 (J. A. Kochalka, JAK); Cienaga, 11°1'N,
74°15'W, 30.i.l974 (J. A. Kochalka, JAK); Gaira,
11°11'N, 74°13'W, 15.xii.l975 (W. Eberhard, MCZ).
DOMINICAN REPUBLIC Barahona: Patos,
10°38'N, 61°52'W, 24.ix.1944 (R. H. Montgomery,
AMNH). FRENCH WEST INDIES Guadeloupe:
Deshaies, 16°18'N, 61°48'W, 28.vi.1960 (C. & P Vau-
rie, AMNH); Domaine Duclos [?], 16°16'N, 61°31'W,
25.vi.1960 (C. & P Vaurie, AMNH), 15.vii.l960 (C.
& P. Vaurie, AMNH); Marie-Galante, in citrus,
15°56'N, 61°16'W, 15.iii.l977 (W. H. Whitcomb,
FSCA); Pointe-a-Pitre, Ilet a Boissard, 16°14'N,
61°34'W, 26.vi.1960 (C. & R Vaurie, AMNH); Terre-
de-Haut, Les Saintes, 15°58'N, 61°35'W, 2.vii.l960
(C. & P. Vaurie, AMNH). Martinique: Ansemitian [?],
Trois-Ilets, 14°33'N, 61°2'W, 10.vi.l960 (C. & P Vau-
rie, AMNH); Diamant, 14°29'N, 61°2'W, 17.vi.l960
(C. & P Vaurie, AMNH), 18.vi.l966 (C. & P Vaurie,
AMNH); Fort de France, 14°36'N, 61°5'W,
15.xii.l950 (N. L. H. Krauss, MCZ); Pointe Ferret,
La Caravelle, 14°45'N, 60°54'W, 19.vi.l960 (C. & R
Vaurie, AMNH); Sainte-Anne, 14°26'N, 60°53'W,
20.vi.l966 (C. & P Vaurie, AMNH); Usine de Robert
[?], 14°41'N, 60°57'W, 16.vi.l960 (C. & R Vaurie,
AMNH). GRENADA nr Saint Georges: 12°3'N,
61°45'W, 3.vi.l950 (Leo Isaacs, AMNH). NETH-
ERLAND ANTILLES Bonaire: Red Pond [?],
12°12'N, 68°15'W, 3.i.l968 (B. Malkin, AMNH). Cu-
ragao: 12°11'N, 68°58'W, 13. i. 1968 (B. Malkin,
AMNH); Fuik (Oostpunt), muddauber nests, 12°4'N,
68°49'W, 20.xii.l962 (H. W. Levi & B. de Jong,
MCZ); Groot Sint Joris, plantation, 12°14'N, 69°3'W,
22.xii.1962 (H. W. Levi & B. de Jong, MCZ); Groote
Berg, 12°11'N, 69°0'W, 19.xii.l962 (H. W. & L. Levi,
MCZ); Hato, 12°11'N, 68°58'W, 28.xii.1967 (B. Mal-
kin, AMNH), 7.i.l968 (B. Malkin, AMNH); Piscadera
Baai, 12°8'N, 68°59'W, 18.xii.l962 (H. W. Levi,
MCZ), 20.xii.l962 (H. W. Levi, MCZ); Savonet;
shady ravine, 12°20'N, 69°7'W, 28.xii.1962 (H. W.
Levi, MCZ); SE of airport, 12°10'N, 68°54'W,
20.xii.l962 (H. W. Levi & B. de Jong, MCZ); Siberie,
12°14'N, 69°3'W, 25.xii.1962 (H. W. Levi, MCZ); Sint
Jan, 12°15'N, 69°6'W, 25.xii.1962 (H. W. Levi & B.
de Jong, MCZ); Willemstad, Jewish Cemetary,
12°7'N, 68°57'W, 24.xii.I962 (H. W. Levi, MCZ). Sint
Eustatius: Oranjestad, 17°29'N, 62°59'W, 18.i.l968
(B. Malkin, AMNH). Sint Maarten: nr. Juliana Air-
port [?], 18°4'N, 63°4'W, 24.ii.1965 (H. Heatwole &
F. MacKenzie, AMNH). PARAGUAY Alto Parana:
Taguarazaya [?], 25°30'S, 54°50'W (AMNH). PERU
Ancash: Huaraz, 9°32'S, 77°32'W, 6.xii.l980 (C. Gold,
CAS). Apuriniac: 35 mi E Abancay, 13°38'S, 72°22'W,
5.iii.l951 (E. S. Ross & Michelbacher, CAS); 40 mi
E Abancay, 13°38'S, 72°20'W, 4.iii.l951 (E. S. Ross
& Michelbacher, CAS); Abancay, 13°38'2"S,
72°52'53"W, 6.iii.l951 (E. S. Ross &' Michelbacher,
CAS). Cajamarca: Jaen, 5°42'34"S, 78°48'32"W,
17.V.1967 (A. F. Archer, AMNH). Cusco: 40 mi W
Cusco, 13°32'S, 72°33'W, 5.iii.l951 (E. S. Ross &
Michelbacher, CAS); Hacienda Urco, near Galea,
13°22'S, 71°54'W, 19. ix. 1939 (Karl P Schmidt,
AMNH); Huacerpay [?], 13°37'S, 72°13'W, 10.ix.l993
(J. Ochoa Camara, MCZ); Ollantaitambo, 13°15'17"S,
72°15'48"W, 15.vii.l911 (Yale Pemvian Expedition,
AMNH), 15.xii.l980 (C. Gold, CAS); Pisac,
13°25'21"S, 71°50'48'W, 13.-xii.l980 (C. Gold, CAS);
Urubamba, 13°18'28"S, 72°6'55"W, 15.i.l965 (Carras-
co, MCZ), 18.ii.l965 (H. W. Levi, MCZ), 6.viii.l987
(D. Silva, MUSM). Huanuco: Huanuco, 9°55'S,
76°14'W, 6.X.1946 (J. C. Palhster, AMNH). lea: km
367 between lea and Nazca, sandy semidesert,
14°24'S, 75°23'W, 22.i.l952 (W. Weyrauch, CAS);
Nazca, 14°50'S, 74°57'W, 15. iv. 1951 (P. Aguilar,
GAS). Piura: Gerro Prieto, La Brea, 4°41'S, 81°6'W
(CAS); Mallares, 4°51'S, 80°46'W, 8.vi.l941 (H. E. F.
& D. E. F, CAS), 13.vii.l941 (H. E. F. & D. E. F,
52 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
CAS); N of Negritos, Farinas Valley, 4°41'S, 81°18'W,
9.X.1938 (D. L. & H. E. Frizzell, CAS); nr. Negritos,
4°42'S, 81°18'W, 5.m.l939 (H. E. F, CAS); nr. Se-
chura, 5°33'S, 80°51'W, 4.xi.l941 (D. L. F, CAS); Ne-
gritos, 4°42'S, 81°18'W, 15.xi.l934 (H. E. F, CAS),
15.iii.l941 (H. E. F, CAS); Farinas, 4°30'S, 81°8'W,
7.V.1939 (D. L. & H. E. Frizzell, CAS), 31.V.1939 (D.
L. & H. E. Frizzell, CAS), 15.X.1939 (D. L. & H. E.
Frizzell, CAS); Farinas Valley, 4°30'S, 81°8'W,
6.iii.l939 (D. L. & H. E. Frizzell, CAS), 8.iv.l939 (D.
L. & H. E. Frizzell, CAS); Fortachuelo, 4°59'S,
79°54'W, 29.iv.1939 (M. H. & H. E. Frizzell, CAS);
Quebrada Mogollon, 4°32'S, 81°4'W, 30.iv.l939 (D.
L. & H. E. Frizzell, CAS), ll.vi.l939 (D. L. & H. E.
Frizzell, CAS); Rio Chira Valley, N of Amotape,
4°53'S, 81°1'W, 13,xi.l938 (D. L. & H. E. Frizzell,
CAS); S of Sechura, 5°33'S, 80°51'W, 25.X.1941 (D.
L. F, CAS); San Lorenzo [?], Zona Alta Este Herrera,
5°4'S, 79°47'W, 12.iv.l969 (F Aguilar, MCZ); Sullana,
4°53'S, 80°41'W, 8.X.1939 (D. L. & H. E. Frizzell,
CAS), 5.X.1941 (D. L. & H. E. Frizzell, CAS). Tum-
bes: 34 km E, 25 km N Funta Farinas, 4°18'S,
80°51'W, l.i.l939 (D. L. & H. E. Frizzell, CAS).
SAINT LUCIA Castries: 14°1'N, 61°0'W, 28.vii.1963
(E. N. Kjellesvig-Waering, AMNH). TRINIDAD &
TORAGO Tohago Island: Ruccoo Ray, 11°10'N,
60°48'W, 15.viii.l965 (E. N. Kjellesvig-Waering,
AMNH); Guayaguayare Foint, 10°8'N, 61°2'W,
14.ix.l963 (E. N. Kjellesvig-Waering, AMNH); Pi-
geon Foint, 11°10'N, 60°50'W, 18.viii.l937 (E. D.,
MCZ); Salybia Ray, by shore, 10°42'N, 61°2'W,
15.ii.l972 (J. A. L. Cooke, MCZ); Toco, 10°50'N,
60°57'W, 19. iv. 1964 (Erik N. Kjellesvig-Waering,
AMNH). Trinidad: Gasparee, 10°46'N, 61°19'W,
3.xi.l944 (R. Montgomery, AMNH). URUGUAY Co-
lonia: Fmita Gorda [?], 34°28'S, 57°51'W, 25.ii.1968
(R. Capocasale & L. Rruno, CAS). USA Puerto Rico:
Algodones Key, 18°12'N, 65°41'W, 15.x. 1964 (H.
Heatwole, R. Levins & F MacKenzie, AMNH); Ral-
neario [?] Guajataca, 18°21'N, 66°55'W, 4.vii.l958 (A.
F Archer, AMNH); Railos de Coamo, 17°59'N,
67°3'W, 2.iv.l990 (H. W & L. Levi, MCZ), 3.iv.l990
(H. W & L. Levi, MCZ); Rarranquitas, 18°11'N,
66°18'W, 28.xii.1977 (J. Coddington, USNM); below
Quebradillas along old RR track, 18°28'N, 66°56'W,
30.iii.l989 (H. W & L. Levi, MCZ); Cabeza de Ferro
Island, 18°15'N, 65°35'W, 16.i.l965 (H. Heatv^^ole &
F. MacKenzie, AMNH); Cayo Don Luis, 17°57'N,
66°58'W 12.i.l966 (Island Froject Staff, Univ. of
Fuerto Rico, AMNH); Cayo Norte, off Culebra,
18°20'N, 65°15'W, 14.ivl965 (H. Heatwole & F.
MacKenzie, AMNH); Cayo Pinerito, 18°15'N,
65°36'W, 24.ix.1964 (H. Heatwole & F. MacKenzie,
AMNH); Channel at Culebra, Isla del Diablo,
18°23'N, 65°40'W, 12.viii.l965 (Island Froject Staff,
Univ. of Fuerto Rico, AMNH); Cuevas de los Alfaros,
Rarrio Mora, 18°29'N, 67°1'W, 20.vii.l958 (A. F. Ar-
cher & Rolle, AMNH); Culebra Island, 18°19'N,
65°17'W, 19.vii.l965 (F. MacKenzie, AMNH); Cule-
bra, near Dewey, 18°18'N, 65°18'W, 10.viii.l965 (Is-
land Froject Staff, Univ. of Fuerto Rico, AMNH); Cu-
lebrita Island, 18°19'N, 65°14'W, 15.ivl965 (H. Hea-
twole & F MacKenzie, AMNH), ll.viii.l965 (Island
Froject Staff, Univ. of Fuerto Rico, AMNH); Dese-
cheo Is, 18°23'N, 67°29'W, 19.ii.l914 (AMNH); De-
secheo Island, 18°23'N, 67°29'W, 28.V.1965 (H. Hea-
twole, R. Levins & F MacKenzie, AMNH); Faro de
Cabo Rojo, 18°5'N, 67°9'W, 13.iii.l961 (F. Rolle,
AMNH); Frank Key, Isl. #13 nr. La Farguera area,
17°58'N, 67°3'W, 14.i.l966 (Island Froject Staff,
Univ. of Fuerto Rico, MCZ); Heatwole Island, off
Culebrita, 18°19'N, 65°13'W, 14.iv.l965 (H. Heatwo-
le & F. MacKenzie, AMNH); Hormiqueros, 18°9'N,
67°8'W, 16.ii.l962 (Aida Velez, AMNH), ll.iii.l962
(Aida Velez, AMNH), 19.iii.l962 (Aida Velez,
AMNH); Isabela, subexperiment station, 18°30'N,
67°1'W, 17.viii.l957 (A. F. Archer, AMNH); Jayuya,
coffee plantation, 18°13'N, 66°37'W, 23.iii.1986 (H.
W. & L. Levi, MCZ); Juana Diaz, 18°3'N, 66°31'W,
7.xi.l971 (J. E. Carico, JEC); Levins Rock [?],
18°12'N, 65°41'W, 15.X.1964 (H. Heatwole, R. Levins
& F. MacKenzie, AMNH); Loma Tinaja [?], S of La-
guna Cartagena, 17°59'N, 67°6'W 5.vii.l958 (A. F.
Archer & M. Sanderson, AMNH); Luquillo, near
beach, 18°23'N, 65°44'W, 24.i.l932 (A. S. Mills,
AMNH), 24.xii.1985 (V. & R. Roth, CAS), 27.iii.1988
(H. W & L. Levi, MCZ); Mayagiiez, 18°12'N,
67°9'W, 15.vii.l958 (A. F. Archer, MCZ); Mayagiiez,
5 km N university campus, 18°12'N, 67°9'W, 5.i.l964
(MCZ); Mayagiiez: Las Mesas, 18°11'N, 67°6'W,
20.xi.l960 (F. Rolle, AMNH); Mayagiiez, university
farm N university campus, 18°12'N, 67°9'W, 7.ii.l964
(MCZ); McKenzie Key, Key #3 [?], 18°12'N,
65°41'W, 29.X.1964 (H. Heatwole, R. Levins & F.
MacKenzie, AMNH); Mona Island, Serralles, 18°5'N,
67°54'W [?], 7.ivl944 (Reatty, MCZ); Muertos Is-
land, beating bushes, 17°54'N, 66°32'W 28.vl959
(Jordan & Martorell, AMNH); Muertos Island, mud)
nests of Sceliphron caementarium, 17°54'N, 66°32'W,
27.V.1959 (Medina & Martorell, AMNH); N slope Ti-
naja, nr. Cartagena Lagoon, beating and sweeping,
18°23'N, 67°10'W, 5.vii.l958 (M. W Sanderson,
AMNH); Palominitos, 18°20'N, 65°34'W, 16.vi.l965
(Island Froject Staff, Univ. of Fuerto Rico, AMNH);
Falomino Island, 18°21'N, 65°34'W, 7.xi.l964 (H.
Heatwole & F. MacKenzie, AMNH); Farguera,
17°59'N, 67°3'W, 25.iii.1990 (H. W & L. Levi, MCZ);
Fatillas, 18°0'N, 66°1'W, 3.iv.l931 (Mills & Leonard,
AMNH); Fico Atalaya, Rte. 2, nr Aiiasco, 18°18'N,
67°11'W, 3.vii.l958 (M. W. Sanderson, AMNH); Fi-
fieros Island, 18°15'N, 65°35'W, 24.ix.1964 (H. Hea-
twole & F. MacKenzie, AMNH); Flaya de Humacao,
18°10'N, 65°45'W, 23.V.1964 (MCZ); Quebradillas,
around hotel, 18°29'N, 66°56'W, 30.iii.l989 (H. W. &
L. Levi, MCZ); Ratones Island, 17°56'N, 66°17'W,
19.xi.l964 (H. Heatwole & R. Levins, AMNH); Rom-
ero III [?], 17°57'N, 67°0'W, 13.i.l966 (Island Froject
Staff, Univ. of Fuerto Rico, AMNH); Rubianes,
Caimito Rajo [?], Cord. Jaicoa, 18°26'N, 67°8'W,
19.vii.l958 (A. F Archer & Rolle, AMNH); Santurce,
18°27'N, 66°4'W, 28.iii.1931 (A. S. Mills, AMNH);
South of Corozo, Cabo Rojo salt flats, edge of salt
Metepeira • Piel
53
ponds, 17°56'N, 67°11'W, 23.iii.1990 (H. W. & L.
Levi, MCZ); Valle de Lajas, 18°1'N, 67°8'W, 3.vi.l958
(A. F. Archer, AMNH); xerie hills N of Guanica,
18°0'N, 66°55'W, 28.viii.1957 (A. F. Archer, AMNH);
Zancudo Island (Isleta Marina), 18°20'N, 65°37'W,
2..xi.l964 (H. Heatwole & F. MacKenzie, AMNH).
U.S. VIRGIN ISLANDS Big Cockroach: 18°24'N,
65°4'W, 7.vi.l966 (Island Project Staff, Univ. of
Puerto Rico, AMNH). Congo Cay: 18°22'N, 64°48'W,
12.xi.l966 (Island Project Staff, Univ. of Puerto Rico,
AMNH). Grass Cay: 18°22'N, 64°50'W, 12.xi.l966
(Univ. of Puerto Rico, AMNH). Great Saint James
Island: 18°19'N, 64°50'W, 13.xi.l966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Hans Lollik Is-
land: 18°24'N, 64°55'W, 6.iv.l966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Little Cock-
roaches: 18°25'N, 65°3'W, 7.vi.l966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Little Hans Lol-
lick: 18°25'N, 64°54'W, 5.iv.l966 (Island Project Staff,
Univ. of Puerto Rico, AMNH). Little Saint James Is-
land: 18°18'N, 64°50'W, 13.xi.l966 (Island Project
Staff, Univ. of Puerto Rico, AMNH). Mingo Cay:
18°22'N, 64°49'W, 12.xi.l966 (Island Project Staff,
Univ. of Puerto Rico, AMNH). Saint John: 18°20'N,
64°45'W, 9.iii.l925 (F. E. Lutz, AMNH), ll.vii.l958
(A. F Archer & M. Sanderson, AMNH), 15.xii.l967
(N. L. H. Krauss, AMNH); Cruz Ray, 18°20'N,
64°48'W, 27.ii.1964 (A. M. Chickering, MCZ); nr.
Cinnamon & Hart Bays on W half of island, 18°20'N,
64°46'W, 2.vlii.l976 (D. E. & D. N, Rosen, AMNH).
Saint Thomas: 18°24'N, 64°55'W, 24. ii. 1925
(AMNH), 24.vii.1925 (A. Petrunkevitch, CAS),
24.xi.1925 (AMNH); Crov^^n Mountain, 18°21'N,
64°58'W, 30.viii.l957 (A. F. Archer, AMNH),
7.vii.l958 (A. F. Archer, AMNH); Denmark Hill [?],
18°24'N, 66°55'W, l.ix.l957 (A. F Archer, AMNH);
Flagstok Hill, Stumpy Bay, 18°22'N, 65°0'W,
9.vii.l958 (A. F. Archer, AMNH); Harmans, Char-
lotte Amalia, 18°21'N, 64°56'W, 2.ix.l957 (A. F. Ar-
cher & family, AMNH); Hassell Island, 18°20'N,
64°56'W, l.ix.l957 (A. F Archer, AMNH), 15.ii.l964
(A. M. Chickering, MCZ), 10.viii.l966 (M. L. Pres-
sick, AMNH). St. Croix: Christiansted, 17°45'N,
64°42'W, 3.vi.l911 (AMNH), 15.i.l955 (A. M. Nad-
ler, AMNH); East End, 17°45'N, 64°40'W, 15.xii.l965
(Island Project Staff, Univ. of Puerto Rico, AMNH);
Green Key, 17°46'N, 64°40'W, 17.iv.l964 (H. Hea-
twole, MCZ); Mount Eagle, dryish forest, 17°46'N,
64°49'W, 15.xii.l965 (Island Project Staff, Univ. of
Puerto Rico, AMNH); Protestant Cay, 17°45'N,
64°42'W, 18.iv.l964 (H. Heatwole, MCZ); St. Croix,
17°45'N, 64°54'W, l.ix.l966 (Chickering, MCZ),
6.ix.l966 (Chickering, MCZ). VENEZUELA Depen-
dencias Federales: Patos, 10°38'N, 61°52'W,
23.ix.1944 (R. H. Montgomery, AMNH). Districto
Federal: Punta Tanaguarena, in coastal bldg. & gar-
den, 10°37'N, 66°48'W, 26.xii.1970 (W. B. Peck,
CAS).
1 7. Metepeira roraima new species
Figures 141-148, 316; Map 8
Holotype. Female from Rio Suioimu, Roraima, Brazil,
X.1966, M. Abrorenga, in MZSP The specific name
is a noun in apposition after the locality.
Description. Female holotype. Brown
carapace; lighter around eyes (Fig. 146).
Coxae, femora, tibiae, and patellae tan,
lighter ventrally Metatarsi, tarsi white. Fe-
mur I with row of three or four macrose-
tae on anterior side; none on anteroventral
side. Dorsal folium with typical Metepeira
fleur-de-lis pattern (Fig. 146). Venter of
abdomen brown with wide white median
line, flanked by thinner white lines that to-
gether form a T-shape posteriorly (Fig.
147). Pair of small white spots on either
side of T-shape mark. Sternum brown with
median white line (Fig. 147). Ratio of eye
diameters: posterior medians and anterior
medians 1.1, anterior laterals 1.5, posterior
laterals 1.3. Anterior median eyes separat-
ed by 1.2 diameters, posterior median eyes
by 0.6, anterior median eyes separated
from anterior laterals by 1.9 diameters of
anterior lateral eyes, lateral eyes separated
by 0.1 their diameters. Total length 4 mm.
Carapace 1.8 mm long, 1.4 wide. First fe-
mur 1.9 mm, patella and tibia 2.1, meta-
tarsus 1.8, tarsus 0.7. Second patella and
tibia 1.7 mm, third 1.0, fourth 1.6.
Male paratype from Rio Surumu, Ro-
raima, Brazil. Carapace brown; lighter
around eyes with median white mark (Fig.
144). Legs brown, white at base of femora.
Femur I with row of four to five macro-
setae on anterior side; four to five on an-
teroventral side. Dorsal folium, venter, and
sternum as in female (Figs. 144, 145). Ra-
tio of eye diameters: posterior medians
and anterior medians 1.1, anterior laterals
1.6, posterior laterals 1.4. Anterior median
eyes separated by 1.4 diameters, posterior
median eyes by 0.6, anterior median eyes
separated from anterior laterals by 1.6 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.2 their diameters. To-
tal length 2.9 mm. Carapace 1.3 mm long,
1.1 wide. First femur 1.9 mm, patella and
54 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
tibia 2, metatarsus 1.6, tarsus 0.6. Second
patella and tibia 1.5 mm, third 0.8, fourth
1.3.
Diagnosis. The epigynum of M. roraima
differs from that of M. compsa by having
oval openings (Fig. 143) instead of round
openings (Figs. 131, 134); it differs from
M. gressa by being more translucent and
by having a narrower scape (compare Fig.
143 with Fig. 151). Unlike M. gres.sa, the
embolus tip of M. roraima is more bent
and the distal apophysis is as wide as the
embolus tip (compare Fig. 141 with Fig.
149). Unlike M. compsa, the distal embolic
apophysis projects away from the embolus
tip (Fig. 141) instead of extending straight
from the base and parallel to the embolus
tip (Figs. 129, 132).
Variation. Average body length of eight
females examined 5 mm, range 4 to 7 mm.
Average body length of five males exam-
ined 3.1 mm, range 2.8 to 4 mm. Colom-
bian specimens are much darker than
those from northern Brazil and southern
Guyana, suggesting the possibility that
these populations represent separate cryp-
tic species. In fact, the carapace on Col-
ombian specimens is often jet black with
the white marks on the dorsal folium
much reduced (Fig. 148). However, there
is little corresponding genitalic difference,
especially among males; consequently,
these populations are deemed to be con-
specific.
Natural History. Mature M. roraima
specimens from eastern South America
have been collected in November and De-
cember; specimens from western South
America have been collected between
March and July (Fig. 316). Median eleva-
tion, 1,000 m.
Distribution. Western Colombia, north-
ern Brazil, and southern Guyana (Map 8).
Records Examined. BRAZIL Roraima: Rio Suru-
mu, Jerrit Rio Branco [?], 3°22'N, 60°19'W, x.1966
(M. Abrorenga, MZSP). COLOMBIA. Valle del Cau-
ca: Atuncela, 3°46'N, 76°42'W, 19.vii.l970 (W. Eber-
hard, MCZ); Cali, around house, 3°27'N, 76°31'W
(W. Eberhard, MCZ); Palmira, 3°32'N, 76°17'W,
l.iii.l964 (Ballo, CAS); Rio Panee, below Buenos Ai-
res, 3°55'N, 76°8'W, 5.iv.l970 (W. Eberhard, MCZ);
Rio Pance, near Cali., 3°21'N, 76°33'W, 8.V.1970 (W.
Eberhard, MCZ), 8.vi.l970 (W. Eberhard, MCZ),
15.vi.l970 (W. Eberhard, MCZ), 23.vi.1970 (W.
Eberhard, MCZ), 25.vi,1973 (W. Eberhard, MCZ).
GUYANA Upper Takiitu: Isherton, on lat. 2, 10 mi E
Rupununi River, 2°19'N, 59°22'W, 15.xi.l937 (W. G.
Hassler, AMNH).
18. Metepeira gressa (Keyserling)
Figures 149-156, 300; Map 8
Epeira gressa Keyserling, 1892: 166, fig. 123, 9 . Five
female syntypes from Taquara, Rio Grande do Sul,
Brazil, in BMNH, examined. One specimen des-
ignated lectotype.
Epeira seditiosa Keyserling, 1893: 212, fig. 157, S.
Male holotype from Rio Grande do Sul, Brazil, in
BMNH, examined. NEW SYNONYMY.
Araneus gressus: — Petrunkevitch, 1911: 314. Roewer,
1942: 844. Bonnet, 1955: 511.
Araneus seditiosus: — Petrunkevitch, 1911: 314.
Roewer, 1942: 852. Bonnet (1955: 592) erroneously
suggests that Petrunkevitch (1911: 314) synony-
mized Araneus scitulus (Blackwall, 1863) with Ar-
aneus seditiosus.
Eustala seditiosa: — Mello-Leitao, 1943: 179. Erro-
neous transfer.
Metazygia gressa: — Mello-Leitao, 1943: 187. Erro-
neous transfer.
Metepeira gressa: — Levi, 1991: 179. Platnick, 1993:
449.
Metepeira seditiosa: — Levi, 1991: 180. Platnick, 1993:
449.
Note. Examination of voucher specimens of Vi-
era suggest that in the behavioral studies of Viera,
(1986, 1989) and Viera and Costa (1988), tlie name!
"Metepeira sp. A" is, in fact, M. gressa.
Description. Female from Punta del Es-
pinillo, Montevideo, Uruguay. Chelicerae
brown, lighter on distal inside margins.
Carapace brown; yellowish white across
and just behind posterior eye row; median
white line reaching thoracic furrow, some-
times thickened into arrow shape (Fig.
155). Proximal halves of femora, white; re-
mainder black. Patellae black ventrally,
white dorsally. Distal halves of tibiae,
black; remainder white with black annu-
lation. Femur I with row of four macro-
setae on anterior side; none on antero-
ventral side. Dorsal folium white fleur-de-
lis pattern on black, margined with wavy
white lines (Fig. 155). Venter black with
short, wide, white median line; pair of
small white spots on either side of spiracle
Metepeira • Piel 55
(Fig. 156). Sternum black with median
white hne, often broken (Fig. 156). Ratio
of eye diameters: posterior medians and
anterior medians 1.2, anterior laterals 1.3,
posterior laterals 1.3. Anterior median
eyes separated by 1.8 diameters, posterior
median eyes by 1.0, anterior median eyes
separated from anterior laterals by 2 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.2 their diameters. To-
tal length 3.3 mm. Carapace 1.8 mm long,
1.4 wide. First femur 1.9 mm, patella and
tibia 2, metatarsus 1.6, tarsus 0.7. Second
patella and tibia 1.8 mm, third 1.1, fourth
1.6.
Male from Punta del Espinillo, Monte-
video, Uruguay. Coloration of chelicerae
and carapace as in female except median
line brighter, thickened into arrow shape
(Fig. 153). Leg coloration as in female. Fe-
mur I with row of four macrosetae on an-
terior side; five to six on anteroventral side.
Dorsal folium, venter, and sternum as in
female (Figs. 153, 154). Ratio of eye di-
ameters: posterior medians and anterior
medians 1.0, anterior laterals 1.5, posterior
laterals 1.4. Anterior median eyes separat-
ed by 1.3 diameters, posterior median eyes
by 0.8, anterior median eyes separated
from anterior laterals by 1.9 diameters of
anterior lateral eyes, lateral eyes separated
by 0.3 their diameters. Total length 3 mm.
Carapace 1.8 mm long, 1.3 wide. First fe-
mur 2.5 mm, patella and tibia 2.3, meta-
tarsus 1.9, tarsus 0.9. Second patella and
tibia 1.8 mm, third 1.1, fourth 1.6.
Diagnosis. The epigynum of M. gressa
differs from that of M. compso by having
oval openings (Fig. 151) instead of round
openings (Figs. 131, 134); it differs from
M. roraima by being less translucent and
by having a wider scape (compare Fig. 151
with Fig. 143). Unlike M. roraima, the em-
bolus tip in M. gressa is not as bent, and
the distal apophysis is thinner than the
point on the embolus where it is attached
(compare Fig. 149 with Fig. 141). Unlike
M. compsa, the distal embolic apophysis
projects away from the embolus tip (Fig.
149) instead of extending straight from the
base and parallel to the embolus tip (Figs.
129, 132).
Variation. Average body length of 14 fe-
males examined 4.8 mm, range 3.3 to 6.8
mm. Average body length of five males ex-
amined 3.4 mm, range 2.8 to 4.3 mm.
Natural History. In Uruguay, M. gressa
live in small colonies of up to five individ-
uals surrounding the inflorescences of
Eryngium sp. (Viera and Costa, 1988). The
number of sticky spirals (14 below the
hub, 22 above the hub) and radii (c. 40)
are the same irrespective of age; however
the length and width of the web differ be-
tween juveniles (c. 6 and 5 cm, respective-
ly) and adults (c. 9 and 7 cm, respectively)
(Viera, 1992). Mature specimens are most
often collected between September and
March at low elevation (Fig. 300).
Distribution. Northern Argentina, Par-
aguay, and Uruguay (Map 8).
Records Examined. ARGENTINA Buenos Aires:
Arrecifes, 34°3'S, 60°7'W, 17.1.1939 (Biraben, MLP);
Boulogne, 34°30'S, 58°34'W, 15.x. 1938 (Prosen,
MLP); Colon [?], 33°53'29"S, 61°6'35"W, 20.Lx.l944
(Torres, MLP); Moreno, 34°39'S, 58°48'W, 15.ii.l966
(Rossi and Maury, MACN); San Isidro, 34°27'S,
58°30'W, 15.xii.l937 (Peregra, MACN); San Mi-
guel—San Fernando, 34°29'S, 58°39'W, 15.vii.l940
(F. Morrios, MACN); Sierra de la Ventana, 38°9'S,
61°48'W, 15.iii.l939 (S. H. Bavio, MACN), 15.xi.l954
(Fritz, MACN), 31.X.1969 (Carlos Grisolia, MCZ),
15.vii.l972 (Amarrilla, MACN); Tigre, 34°25'S,
58°34'W (J. M. Viana, MACN). Entre Rios: Concep-
cion del Uruguay, 32°29'0"S, 58°13'42"W, 4.i.l941
(Prosen, MLP); Salto Grande, 31°13'S, 57°56'W,
15.iii.l964 (M. E. Galiano, MEG). Santa Fe: Floren-
cia Varelo [?], 28°2'S, 59°15'W, 15.xii.l939 (F. Morris,
MACN). Tucumdn: 30 km S Concepcion, 27°36'S,
65°35'W, 16.i.l983 (L. E. Pena, AMNH). PARA-
GUAY Itapiia: San Luis, 27°6'S, 56°36'W, 15.X.1908
(AMNH). URUGUAY Canelones: Montevideo: De-
tras del Cerro, Camino de las tropas [?], 34°45'S,
56°10'W, 6.ii.l963 (R. Capocasale and L. Bmno,
CAS). Colonia: Punta Gorda^?], 34°28'S, 57°51'W,
25. ii. 1968 (R. Capocasale and L. Bruno, CAS),
26.ii.1968 (R. Capocasale and L. Bmno, CAS). Mal-
donado: Colonia, Cerro de las Animas. Small stones
in native forest, 34°46'S, 55°19'W, 30.x. 1967 (P San
Martin, MCZ). Montevideo: Punta del Espinillo,
34°50'S, 56°26'W, 12.X.1983 (Carmen Viera, MCZ),
15.xii.l983 (Carmen Viera, CV). Treinta y Tres: Que-
brada de los Cuervos, 33°10'S, 54°27'W, 27.X.1990
(Lopez, Perez, Viera, MCZ).
56 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
Metepeira incrassata Group
Species in the M. incrassata group all
lack a keel on the median apophysis. Males
have a distal embolic apophysis that either
projects toward the embolus tip (e.g.. Fig.
157), forms a shai-p comer but does not
actually project forward (e.g.. Fig. 185), or
forms a smoother, rounded, and gradual
comer (e.g.. Figs. 192, 206). The M. in-
crassata species group includes Metepeira
maya, Metepeira inca, Metepeira gosoga,
Metepeira ohnec, Metepeira comanche,
Metepeira pimungan, Metepeira triangu-
laris, Metepeira arizonica, Metepeira atas-
cadero, and Metepeira incrassata.
19. Metepeira maya new species
Figures 157-163, 317; IViap 14
Holotype. Male from North Bay, Twin Cays, Stann
Creek District, Belize, 14.iii.l986, P. Sierwald, in
USNM. The specific name is a noun in apposition
after the Indian people of southern Mexico and
Central America.
Description. Female paratype from
Twin Cays, Stann Creek District, Belize.
Carapace brown with large white eye re-
gion, white lateral posterior extensions,
and short median posterior extension (Fig.
162). Legs white, ringed black on distal
ends of articles. Femur I with row of four
macrosetae on anterior side; none on an-
te roventral side. Dorsum of abdomen with
usual Metepeira folium, though frequently
with deep red pigmentation on anterior
half (Fig. 162). Venter wide, white median
line with pair of large white spots on either
side of spiracle (Fig. 163). Sternum with
wide median white line widening anteri-
orly with constriction in center (Fig. 163).
Ratio of eye diameters: posterior medians
and anterior medians 1.0, anterior laterals
1.3, posterior laterals 1.3. Anterior median
eyes separated by 1.2 diameters, posterior
median eyes by 0.8, anterior median eyes
separated from anterior laterals by 2 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.2 their diameters. To-
tal length 3.9 mm. Carapace 1.8 mm long,
1.4 wide. First femur 2.2 mm, patella and
tibia 2.1, metatarsus 1.6, tarsus 0.8. Sec-
ond patella and tibia 1.8 mm, third 1.0,
fourth 1.6.
Male holotype. Carapace dark with light
region around eyes and light triangular
mark anterior to thoracic furrow (Fig.
160). Legs darker than in female. Femur
I with row of four macrosetae on anterior
side; four on anteroventral side. Dorsum,
venter, sternum as in female except me-
dian white line on sternum usually broken
(Figs. 160, 161). Ratio of eye diameters:
posterior medians and anterior medians
1.0, anterior laterals 1.4, posterior laterals
1.2. Anterior median eyes separated by 1.2
diameters, posterior median eyes by 0.9,
anterior median eyes separated from an-
terior laterals by 1.7 diameters of anterior
lateral eyes, lateral eyes separated by 0.3
their diameters. Total length 3 mm. Car-
apace 1.5 mm long, 1.2 wide. First feiTiur
2.2 mm, patella and tibia 2.3, metatarsus
1.8, tarsus 0.9. Second patella and tibia 1.8
mm, third 1.0, fourth 1.4.
Diagnosis. Unlike others in the M. in-
crassata species group, M. maya and M.
inca both have distal embolic apophyses
that project foiAvard, forming rounded
bumps (Figs. 157, 164). The male palps of
these two species differ by the thickness
of the flagellae and base on the median)
apophysis: thicker in M. maya than in M.
inca (compare Fig. 157 with Fig. 164). The
epigynum of M. maya resembles those of
M. inca and M. comanche because the oval
epigynal openings are formed out of mem-
branous surfaces that are distinctly sepa-
rate from the base of the scape (Figs. 159,
166, 187). The epigynal openings of M.
maya (Fig. 159) are much wider than
those of M. comanche (Fig. 187); and the
posterior lobes on M. maya are thickened
(Fig. 159) but not pointed as in M. inca
(Fig. 166).
Variation. Average body length of five
females examined 5.2 mm, range 3.9 to 6.3
mm. Average body length of four males
examined 2.8 mm, range 2.3 to 3.1 mm.
Natural History. Mature specimens have
been collected in March through August
Metepeira • Piel 57
gressa
(18)
It
maya
(19)
A
r^;''i
164
169^^ 170
inca
(20)
Figures 149-156. Metepeira gressa (Keyserling) (sp. 18; 34°50'S, 56°26'W). 149, male palpus, mesal. 150, epigynum, posterior.
151, epigynum, ventral. 152, male embolic division, ventral. 153, male, dorsal. 154, male, ventral. 155, female, dorsal. 156,
female, ventral.
Figures 157-163. Metepeira maya new species (sp. 19; 16°50'N, 88°5'W). 157, male palpus, mesak 158, epigynum, posterior.
159, epigynum, ventral. 160, male, dorsal. 161, male, ventral. 162, female, dorsal. 163, female, ventral.
Figures 164-170. Metepeira inca new species (sp. 20; 4°30'S, 81°8'W). 164, male palpus, mesal. 165, epigynum, posterior.
166, epigynum, ventral. 167, male, dorsal. 168, male, ventral. 169, female, dorsal. 170, female, ventral.
Scale bars: dorsum and venter figures 1.0 mm.
58 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
(Fig. 317). Habitats range from mangroves
at sea level to pine forests at 1,600 m.
Distribution. Southern Mexico to Costa
Rica (Map 14).
Records Examined. BELIZE Stann Creek: Twin
Cays, Andera Flats, 16°50'N, 88°5'W, 20.iii.l986 (P.
Sierwald, USNM); Twin Cays, North part of West
Pond, West of Swamp Doc, 16°50'N, 88°5'W
28.iii.1986 (P Siei-wald, USNM); Twin Cays, North-
west Point, North Bay, 16°50'N, 88°5'W, 14.iii.l986
(P. Sieiw^ald, USNM); Twin Cays, Northwest Point,
North Bay, ground not flooded, 16°50'N, 88°5'W,
14.iii.l986 (P Sierwald, USNM); Twin Cays, red
mangrove, 16°50'N, 88°5'W, 17.V.1985 (Feller,
USNM); Twin Cays, white mangrove, 16°50'N,
88°5'W, 15.vi.l984 (Feller, USNM), 5.vi.l985 (Erwin,
Mathis, Sims, USNM). COSTA RICA San Jose: San
Jose, 9°56'N, 84°5'W (Tristan & Banks, MCZ). GUA-
TEMALA Chiquimula: Chiquimula, 14°48'N,
89°33'W, 22.vii.1947 (C. & P Vaurie, AMNH). Sa-
catepequez: Antigua, 14°34'N, 90°44'W, 16.viii.l947
(C. & P Vaurie, AMNH). MEXICO Chiapas: Com-
itan de Dominguez, 16°1.5'N, 92°8'W, 19.vii.l950 (C.
J. & M. Goodnight, AMNH); near Rio San Gregorio,
between Comitan and Ocotal, 15°45'N, 92°0'W,
18.vii.l950 (C. J. & M. Goodnight, AMNH); pine for-
est, 15 mi NW Arriaga, 16°25'N, 94°1'W, 27.viii.1966
(Jean & Wilton Ivie, AMNH); Tuxda Gutierrez,
16°45'N, 93°7'W, 10.vi.l964 (Palhster, AMNH). NIC-
ARAGUA Mfltogoipa; Matagaslpa, 12°53'N, 85°57'W
15.vii.l989 (R. Reinbold, JMM).
20. Metepeira inca new species
Figures 164-170, 318; Map 11
Holotype. Male from Pariiias Valley, Piura, Peru,
21.V.1939, D. L. & H. E. Frizzell in CAS. The
specific name is a noun in apposition after the Que-
chuan people who once luled Peru.
Description. Female paratype from Par-
iiias Valley, Piura, Peru. Light reddish
brown carapace, white around eyes, very
faint light mark in center (Fig. 169). Legs
white; slightly darker on dorsal surfaces.
Femur I with row of four to five macro-
setae on anterior side; none on anterov-
entral side. Dorsal folium white; darker on
shoulders and posteriorly (Fig. 169). Ven-
ter grayish browii with wide white median
mark inside U-shaped pattern; pair of
white spots on either side of spiracle (Fig.
170). Sternum white (Fig. 170). Ratio of
eye diameters: posterior medians and an-
terior medians 0.9, anterior laterals 1.0,
posterior laterals 1.1. Anterior median
eyes separated by 1.6 diameters, posterior
median eyes by 1.3, anterior median eyes
separated from anterior laterals by 2.5 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.3 their diameters. To-
tal length 6.8 mm. Carapace 2.9 mm long,
2.3 wide. First femur 3 mm, patella and
tibia 3.3, metatarsus 2.6, tarsus 1. Second
patella and tibia 2.8 mm, third 1.7, fourth
2.5.
Male holotype. Light reddish brown
carapace, white around eyes, faint light tri-
angular mark in center (Fig. 167). Legs
white, save for light reddish brown on dor-
sal surfaces of femora and patellae. Femur
I with row of three macrosetae on anterior
side; two to three on anteroventral side.
Dorsal folium white with wavy black lines
thickening posteriorly (Fig. 167). Venter
and sternum as in female (Fig. 168). Ratio
of eye diameters: posterior inedians and
anterior medians 1.0, anterior laterals 1.3,
posterior laterals 1.1. Anterior median
eyes separated by 1.3 diameters, posterior
median eyes by 0.8, anterior median eyes
separated from anterior laterals by 1.9 di-
ameters of anterior lateral eyes, lateral
eyes almost touching. Total length 3.7 mm.
Carapace 1.9 mm long, 1.4 wide. First fe-
mur 2.9 mm, patella and tibia 2.9, meta-)
tarsus 2.7, tarsus 1. Second patella and tib-
ia 2.4 mm, third 1.3, fourth 1.9.
Diagnosis. Unlike others in the M. in-
crassata species group, M. inca and M.
niaija both have distal embolic apophyses
that project forward, forming rounded
bumps (Figs. 157, 164). The male palps of
these two species are separated by the
thinner flagellae and base on the median
apophysis of M. inca (compare Fig. 164
with Fig. 157). The epigynum of M. inca
resembles those of M. inaija and M. coni-
anche because the oval epigynal openings
are formed out of membranous surfaces
that are distinctly separate from the base
of the scape (Figs. 159, 166, 187). The epi-
gynal openings of M. inca (Fig. 166) are
much wider than those of M. conianche
(Fig. 187); the posterior lobes on M. inca
Metepeira • Piel
59
are pointed (Fig. 166), not just thickened,
as in M. may a (Fig. 159).
Variation. Average body length of six fe-
males examined 5.7 mm, range 5.2 to 6.8
mm. Average body length of four males
examined 3.7 mm, range 2.8 to 4.4 mm.
Coloration varies significantly among lo-
calities, especially in the males, where
some are frequently much darker than
others.
Natural History. Mature specimens
have been collected in April through Oc-
tober (Fig. 318) at elevations between 300
and 600 m.
Distribution. Most northern tip of Peru
(Map 11).
Records Examined. PERU Piiira: 12 mi N Man-
cora, 4°0'S, 80°54'W, ll.xii.l93S; Parinas Valley,
4°30'S, 81°8'W, 3.iv.l939, 8.iv.l939, 16.iv.l939,
7.V.1939, 21. V. 1939, 25.vi.1939, 3.vii.l939, 6.viii.l939,
15.viii.l939, 15.X.1939; Quebrada de Pariiias, 4°32'S,
81°17'W, 14.iv.l939, 7.V.1939, 21.V.1939; Quebrada
MogoUon, 4°32'S, 81°4'W, 30.iv.l939, ll.vi.l939,
18.vi.l939, 21.vi.l939, ll.vii.l939, 16.vii.l939,
24.ix.1939 (all records: D. L. & H. E. Frizzell, CAS).
21 . Metepeira gosoga
Chamberlin and Ivie
Figures 171-177, 322; Map 9
Metepeira gosoga Chamberlin and Ivie, 1935: 21 figs
82-83, 9 . Female holotype from Pilot Knob Valley,
Mohave Desert, California, in the AMNH, exam-
ined. Roewer, 1942: 868. Ronnet, 1957: 2820. Levi,
1977: 200, figs. 28-36.
Description. Female from Baja Califor-
nia Norte, Mexico. Carapace diiiy yellow-
ish brown, lighter anterior half, darker di-
amond-shaped mark behind eyes (Fig.
176). Legs same color as carapace; dark
rings on distal ends of articles. Femur I
with row of three to five inacrosetae on
anterior side; two or three macrosetae on
anteroventral side. Folium lighter than in
other species, darkening posteriorly (Fig.
176). Venter black surrounded by yellow.
Wide white median line with pair of large
white spots on either side of spiracle.
Sometimes with parallel pair of lateral
white lines that join transverse white line
posteriorly (Fig. 177). Sternum black with
wide, white line widening anteriorly, often
broken in center (Fig. 177). Ratio of eye
diameters: posterior medians and anterior
medians 0.9, anterior laterals 1.3, posterior
laterals 1.2. Anterior median eyes separat-
ed by 1.2 diameters, posterior median eyes
by 0.6, anterior median eyes separated
from anterior laterals by 3.4 diameters of
anterior lateral eyes, lateral eyes separated
by 0.4 their diameters. Total length 8.3
mm. Carapace 4 mm long, 3.3 wide. First
femur 4.5 mm, patella and tibia 4.8, meta-
tarsus 4.5, tarsus 1.3. Second patella and
tibia 4.3 mm, third 2.5, fourth 3.8.
Male from Baja California Norte, Mex-
ico. Male carapace, dorsum, venter, ster-
num as in female (Figs. 174, 175). Femur
I with row of five macrosetae on anterior
side; row of nine macrosetae on anterov-
entral side. Ratio of eye diameters: poste-
rior medians and anterior medians 0.8, an-
terior laterals 1.1, posterior laterals 1.3.
Anterior median eyes separated by 1 di-
ameter, posterior median eyes by 0.8, an-
terior median eyes separated from anterior
laterals by 2.1 diameters of anterior lateral
eyes, lateral eyes separated by 0.2 their di-
ameters. Total length 5.5 mm. Carapace
2.8 mm long, 2.3 wide. First femur 4.5
mm, patella and tibia 4.5, metatarsus 4.5,
tarsus 1.4. Second patella and tibia 3.8
mm, third 1.9, fourth 3.
Diagnosis. Within the M. incrassata spe-
cies group, only M. gosoga, M. inca, and
M. maya have projecting distal embolic
apophyses (Figs. 157, 164, 171). Of these,
only M. gosoga has a distinctly pointed one
(Fig. 171). The epigynum of M. gosoga dif-
fers from those of other species in the M.
incrassata group by the small, oval shape
of the openings (Fig. 173).
Variation. Average body length of two
females examined 6.8 mm, range 5 to 8.5
mm. Average body length of two males ex-
amined 4.9 mm, range 4.1 to 5.7 mm.
Natural History. Mature specimens
have been collected in June through Au-
gust (Fig. 322) between 500 and 2,000 m.
Levi (1977) notes that these have been
collected on Opuntia and desert vegeta-
tion.
60
Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
Distribution. Southwestern U.S. and
Baja California Norte, Mexico (Levi, 1977,
map 1; Map 9).
Records Examined. MEXICO BaJa Calif. Norte: 23
mi S Catavina, 29°32'N, 114°57'W, 20.vi.l983 (L.
Strange & R. Miller, FSCA). USA Arizona: Show
Low Lake, 34°12'N, 110°0'W, 15.viii.l966 (MCZ).
22. Metepeira olmec new species
Figures 178-184, 324; Map 11
Holotype. Female from Foitin de las Flores, Vera-
cmz, Mexico, 26.vi.1944, L. L Davis, in AMNH.
The specific name is a noun in apposition after the
ancient Indian people of southern Veracruz and Ta-
basco.
Description. Female holotype. Carapace
reddish brown with white region around
eyes that extends posteriorly along lateral
margins (Fig. 183). Legs ringed on distal
ends of articles. Femur I with row of four
macrosetae on anterior side; none on an-
teroventral side. Dorsum of abdomen with
usual Metepeira folium (Fig. 183). Venter
of abdomen with wide, long white median
line set within surrounding U-shaped
markings. Pair of white spots on either
side of spiracle (Fig. 184). Sternum has
wide median white line widening anteri-
orly with broken constriction in center
(Fig. 184). Ratio of eye diameters: poste-
rior medians and anterior medians 1.0, an-
terior laterals 1.1, posterior laterals 1.5.
Anterior median eyes separated by 1.5 di-
ameters, posterior median eyes by 0.8, an-
terior median eyes separated from anterior
laterals by 2.1 diameters of anterior lateral
eyes, lateral eyes separated by 0.2 their di-
ameters. Total length 5.9 mm. Carapace
2.8 mm long, 2 wide. First femur 2.8 mm,
patella and tibia 2.8, metatarsus 2.5, tarsus
1. Second patella and tibia 2.5 mm, third
1.5, fourth 2.3.
Male paratype from Fortin de las Flo-
res, Veracruz, Mexico. Carapace dark with
light region around eyes and a light mark
anterior to thoracic furrow (Fig. 181).
Legs darker than in female. Macrosetae on
legs variable with spiders size — femur I
with row of three or four macrosetae on
anterior side; one to three on anteroven-
tral side. Dorsum, venter, sternum as in
female, except median white line on ven-
ter shortened to spot and median white
line on sternum often broken (Figs. 181,
182). Ratio of eye diameters: posterior me-
dians and anterior medians 1.0, anterior
laterals 1.3, posterior laterals 1.3. Anterior
median eyes separated by 1.3 diameters,
posterior median eyes by 0.8, anterior me-
dian eyes separated from anterior laterals
by 1.3 diameters of anterior lateral eyes,
lateral eyes separated by 0.1 their diame-
ters. Total length 2.8 mm. Carapace 1.4
mm long, 1.2 wide. First femur 1.7 mm,
patella and tibia 1.6, metatarsus 1.1, tarsus
0.7. Second patella and tibia 1.3 mm, third
0.8, fourth 1.1.
Diagnosis. Female M. olmec are diag-
nosed by their almost perfectly round epi-
gynal depressions with uniform thickness
around the posterior edges (Fig. 180). The
male embolus is relatively short and has a
vestigial distal embolic apophysis that does
not project foi^ward, but instead drops off,
forming a shai^ cuive (Fig. 178). Among
other species in the M. incrassata species
group, this embolus shape is also seen in
M. conianche and M. triangularis: unlike
M. conianche (Fig. 185), the darker, scler-
otized portion of the embolus does not ex-\
tend over the hump of the distal embolic
apophysis; unlike M. triangularis (Fig.
199), the flagellae are thicker than the
base of the median apophysis.
Variation. Average body length of four
females examined 6.2 mm, range 5.6 to 6.7
mm. Average body length of three males
examined 2.3 mm, range 2.2 to 2.6 mm.
Natural History. Mature specimens
have been collected perennially (Fig. 324)
at elevations between 500 and 1,400 m.
Distribution. Montane rain forests from
Veracruz to Panama (Map 11).
Records Examined. COSTA RICA San Jose: San
Antonio de Escanza, 9°59'N, 84°11'W [?], 4.iii.l984
(W. Eberhard, MCZ). MEXICO Veracruz: Foitin de
las Flores, 18°54'N, 97°0'W, 26.vi.1944 (L. I. Davis,
AMNH); Papantla, 20°27'N, 97°19'W, 12.x. 1947 (H.
Wagner, AMNH). PANAMA Chiriqut: Boquete,
8°47'N, 82°26'W, 15.viii.l950 (A. M. Chickering,
Metepeira • Piel 61
gosoga
(21)
184
olmec
(22)
190 ^ 191
Comanche
(23)
Figures 171-177. Metepeira gosoga CnamberWn and Ivie (sp. 21; 29°32'N, 114°57'W). 171, male palpus, mesal. 172, epigynum,
posterior. 173, epigynum, ventral. 174, male, dorsal. 175, male, ventral. 176, female, dorsal. 177, female, ventral.
Figures 178-184. Metepeira olmec new species (sp. 22; 18°54'N, 97°0'W). 178, male palpus, mesal. 179, epigynum, posterior.
180, epigynum, ventral. 181, male, dorsal. 182, male, ventral. 183, female, dorsal. 184, female, ventral.
Figures 185-191. Metepeira comancfie Levi (sp. 23 [185,188,189] 33°22'N, 99°56'W; [186,187] 25°45'N, 10r55'W; [190, 191]
26°41'N, 101°23'W). 185, male palpus, mesal. 186, epigynum, posterior. 187, epigynum, ventral. 188, male, dorsal. 189, male,
ventral. 190, female, dorsal. 191, female, ventral.
Scale bars: dorsum and venter figures 1 .0 mm.
62 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
MCZ), 7.viii.l954 (A. M. Chickering, MCZ). Panama:
Cerro Galera, 8°55'N, 79°38'W, 7.i.l977 (H. W. Levi
& Y. Lubin, MCZ).
23. Metepeira comanche Levi
Figures 185-191, 331; IVlap 11
Metepeira comanche Levi, 1977: 204, figs. 61-69, S,
? . Male holotype from 9.7 km west of O'Brien,
Haskell Co., Texas, in the MCZ, examined. Brig-
noli, 1983: 275.
Description. Female from Gloria, Coa-
huila, Mexico. Carapace dark with light re-
gion around eyes, extending posteriorly
behind posterior lateral eyes (Fig. 190).
Legs ringed. Femur I with row of four ma-
crosetae on anterior side; one or two on
anteroventral side. Dorsal folium high in
contrast; black comma-shaped markings
shadow white fleur-de-lis pattern (Fig.
190). Venter of abdomen with wide white
median line posteriorly set within sur-
rounding U-shaped marking. Pair of white
spots on either side of spiracle (Fig. 191).
Sternum has wide median white line wid-
ening anteriorly (Fig. 191). Ratio of eye
diameters: posterior inedians and anterior
medians 0.9, anterior laterals 1.2, posterior
laterals 1.1. Anterior median eyes separat-
ed by 1.2 diameters, posterior median eyes
by 1.0, anterior median eyes separated
from anterior laterals by 2.7 diameters of
anterior lateral eyes, lateral eyes separated
by 0.2 their diameters. Total length 7.8
mm. Carapace 3.3 mm long, 2.5 wide.
First femur 3.6 mm, patella and tibia 3.9,
metatarsus 3.4, tarsus 1.2. Second patella
and tibia 3.3 mm, third 2, fourth 2.9.
Male from 9.7 km west of O'Brien, Has-
kell Co., Texas. Carapace light around
eyes, darker posteriorly (Fig. 188). Legs
ringed. Macrosetae on femur I variable;
usually row of four macrosetae on anterior
side, six on anteroventral side. Dorsum
with usual Metepeira folium (Fig. 188).
Venter with median oval white mark; pair
of white spots on either side of spiracle
(Fig. 189). Median white line on sternum
usually broken (Fig. 189). Ratio of eye di-
ameters: posterior medians and anterior
medians 1.0, anterior laterals 1.3, posterior
laterals 1.3. Anterior median eyes separat-
ed by 1.4 diameters, posterior median eyes
by 0.9, anterior median eyes separated
from anterior laterals by 1.6 diameters of
anterior lateral eyes, lateral eyes separated
by 0.2 their diameters. Total length 3.1
mm. Carapace 1.7 mm long, 1.3 wide.
First femur 2.4 mm, patella and tibia 2.2,
metatarsus 2, tarsus 0.9. Second patella
and tibia 1.9 mm, third 1.0, fourth 1.5.
Diagnosis. The epigynum of M. com-
anche resembles those of M. may a and M.
inca because the oval epigynal openings
are formed out of membranous surfaces
that are distinctly Separate from the base
of the scape (Figs. 159, 166, 187). The epi-
gynum of M. comanche differs from M.
niaya and M. inca by having much narrow-
er epigynal openings (compare Fig. 187
with Figs. 159, 166). The male embolus is
relatively short and has a vestigial distal
embolic apophysis that does not project
forward, but instead drops off, forming a
sharp curve (Fig. 185). Among other spe-
cies in the M. incrassata species group,
this embolus shape is also seen in M. olmec
and M. triangularis: unlike M. olmec (Fig.
178), the darker, sclerotized portion of the
embolus extends over the hump of the dis-
tal embolic apophysis; unlike M. triangu-
laris (Fig. 199), the flagellae and base of
the median apophysis are wide (Fig. 185).
Natural Histortj. Mature specimens
have been collected from May to Novem-
ber (Levi, 1977, Fig. 331).
Distribution. New Mexico, Texas, and
northern Mexico (Levi, 1977, map 1; Map
11).
Records Examined. MEXICO Coahuila: Gloria,
26°41'N, 101°23'W, 24.viii.1947 (W. J. Gerisch,
AMNH); Paila, 25°45'N, 101°55'W, 21.viii.l947 (W.
J. Gertsch, AMNH).
24. Metepeira pimungan new species
Figures 192-198, 330; Map 9
Holotype. Male from San Miguel Island, Santa Bar-
bara Co., California, USA, 20.vii.l968, M. E.
Thompson, in the MCZ. The specific name is a
noun in apposition after the Indian people who
once lived on the Channel Islands.
Metepeira • Piel
63
Description. Female paratype from San
Miguel Island, California USA. Carapace
with large white eye region and lateral
posterior extensions. White median arrow-
shaped mark extends to thoracic furrow
(Fig. 197). Legs mostly white with rings on
legs III and IV. Femur I with row of four
macrosetae on anterior side; one on anter-
oventral side. Dorsum of abdomen with
usual Metepeira folium (Fig. 197); venter
wide, with long white median line, flanked
by shorter parallel thin white lines on ei-
ther side. Pair of white spots on either side
of spiracle. Sternum black with posterior
drop-shaped white mark (Fig. 198). Ratio
of eye diameters: posterior medians and
anterior medians 1.0, anterior laterals 1.4,
posterior laterals 1.3. Anterior median
eyes separated by 1.6 diameters, posterior
median eyes by 1.0, anterior median eyes
separated from anterior laterals by 3.4 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.3 their diameters. To-
tal length 6.9 mm. Carapace 3.3 mm long,
2.6 wide. First femur 3.5 mm, patella and
tibia 3.5, metatarsus 2.8, tarsus 1.3. Sec-
ond patella and tibia 3 mm, third 1.9,
fourth 2.8.
Male holotype. Carapace, legs, abdo-
men, venter, sternum as in female, though
legs darker and not ringed (Figs. 195,
196). Femur I with row of four to five ma-
crosetae on anterior side; four to five on
antero ventral side: both rows concentrated
distally on femur. Ratio of eye diameters:
posterior medians and anterior medians
1.0, anterior laterals 1.3, posterior laterals
1.3. Anterior median eyes separated by 1.6
diameters, posterior median eyes by 0.8,
anterior median eyes separated froin an-
terior laterals by 1.9 diameters of anterior
lateral eyes, lateral eyes separated by 0.3
their diameters. Total length 4.5 mm. Car-
apace 2.3 mm long, 1.8 wide. First femur
3.1 mm, patella and tibia 3, metatarsus 2.8,
tarsus 1. Second patella and tibia 2.5 mm,
third 1.3, fourth 1.9.
Diagnosis. Like M. atascadero, but un-
like other species in the M. incrassata
group, the posterior end of the sternum of
M. pimungan has a white water drop mark
(Figs. 196, 198). Unlike M. atascadero, M.
pimungan lacks U-shaped white markings
on the venter, just anterior to the spiracle
(compare Fig. 198 with Fig. 221). In con-
trast to other species in the M. incrassata
group, the embolus on male M. pimungan
speciinens is curved to forin a gentle S-
shape (Fig. 192). The female has a squar-
ish-shaped black mark inside each epigyn-
al depression (Fig. 198) which is not seen
in other M. incrassata group species.
Variation. Average body length of four
females examined 7.7 mm, range 6.9 to 8.5
mm. Average body length of five inales ex-
amined 5.3 mm, range 4.6 to 6.4 mm.
Natural History. Mature specimens
have been collected in August (Fig. 330).
Webs are found near the ground, shel-
tered from the perennial strong winds by
Cortjopsis and lupines.
Distribution. Endemic to San Miguel
Island (Map 9).
Records Examined. USA California: San Miguel Is-
land, 34°2'28.6"N, 120°21'13.6"W, ll.viii.l995"(W. H.
Piel, MCZ); 34°2'6.1"N, 120°21'9.1"W, 12.viii.l995
(W. H. Piel, MCZ); 34°2'N, 120°22'W, 20.viii.l968
(M. E. Thompson, MCZ); 34°3'3.1"N, 120°21'53.3"W,
13.viii.l995 (W. H. Piel, MCZ); behind dunes on
beach, 34°2'43.8"N, 120°21'0.8"W, ll.viii.l995 (W. H.
Piel, MCZ).
25. Metepeira triangularis (Franganillo)
Figures 199-205, 337; Map 14
Mangora triangularis Franganillo, 1930: 21—22. Fe-
male holotype from Sierra Maestra, lost. Platnick,
1993: 449.
Metepeira labijrinthea: — Franganillo, 1936: 75. Bry-
ant, 1940: 341. Platnick, 1993: 449.
Metepeira triangidaris: — Archer, 1958: 15, fig. 37, 9 .
Metepeira acostai Archer, 1958: 15, fig. 36, 9 . Female
holotype from the savannas, Agramont, Camagiiey
Province, Cuba, in the AMNH, examined. NEW
SYNONYMY.
Metepeira hani Archer, 1965: 132, figs. 11, 17 9, 13
S . Male and female syntypes from Bani, Domini-
can Republic, in the AMNH, examined. NEW
SYNONYMY. Brignofi, 1983: 275.
Note. Franganillo's collection vials are not la-
beled, making it impossible to find the holotype.
Based on the descriptions by Franganillo (1930),
Archer (1958), the illustrations of Archer (1958,
1965), and material determined by Archer, I con-
clude that there is no evidence to support three
64 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
separate species. For the name M. triangularis, I
designate a female neotype from Camagiiey Prov-
ince, Cuba. 21°30'N, 78°10'W, x.l954 (J. T. Acosta,
AMNH).
Description. Female from La Descu-
bierta, Lago Enriquillo, Isla Cabritos, Do-
minican Republic. Light reddish brown
carapace, white around eyes (Fig. 204).
Legs yellowish white, sometimes ringed
brown at distal ends of articles. Femur I
with row of three, or sometimes four, ma-
crosetae on anterior side; none on anter-
oventral side. Dorsal folium white with
slightly darker markings posteriorly (Fig.
204). In some cases, dark anterior marks
on shoulders of dorsum. Venter of abdo-
men black with wide white median line,
sometimes flanked by thinner white lines
that together form an anchor shape (Fig.
205). Pair of small white spots on either
side of spiracle. Sternum black with me-
dian white line, wider anteriorly (Fig. 205).
Ratio of eye diameters: posterior medians
and anterior medians 1.0, anterior laterals
1.4, posterior laterals 1.4. Anterior median
eyes separated by 1.5 diameters, posterior
median eyes by 1.1, anterior median eyes
separated from anterior laterals by 3.3 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.2 their diameters. To-
tal length 5.9 mm. Carapace 2.7 mm long,
2 wide. First femur 2.6 mm, patella and
tibia 2.9, metatarsus 2.5, tarsus 1. Second
patella and tibia 2.4 mm, third 1.4, fourth
2.2.
Male from La Descubierta, Lago Enri-
quillo, Isla Cabritos, Dominican Republic.
Carapace as in female, though lighter eye
region often extends to cover entire ante-
rior half (Fig. 202). Legs yellowish white.
Distal half of femora often dark. Femur I
with row of four macrosetae on anterior
side; three to five on anteroventral side.
Dorsal folium white witli slightly darker
markings posteriorly (Fig. 202). Venter of
abdomen as in female; sternum black with
median white line (Fig. 203). Ratio of eye
diameters: posterior medians and anterior
medians 1.1, anterior laterals 0.7, posterior
laterals 1.1. Anterior median eyes separat-
ed by 1.6 diameters, posterior median eyes
by 0.8, anterior median eyes separated
from anterior laterals by 1.0 diameters of
anterior lateral eyes, lateral eyes separated
by 0.1 their diameters. Total length 4.2
mm. Carapace 2 mm long, 1.5 wide. First
femur 2.9 mm, patella and tibia 3, meta-
tarsus 2.8, tarsus 1.1. Second patella and
tibia 2.4 mm, third 0.8, fourth 1.3.
Diagnosis. The epigynum of M. trian-
gularis differs from other species by the
thin, sclerotized openings in the shape of
Ray-Ban sunglasses (Fig. 201). The male
embolus has a vestigial distal embolic
apophysis that does not project forward,
but instead drops off, forming a sharp
cui-ve (Fig. 199). Among other species in
the M. incrassata species group, shorter
versions of this embolus shape is also seen
in M. Comanche and M. olmec: unlike M.
Comanche (Fig. 185), the darker, sclero-
tized portion of the embolus does not ex-
tend over the hump of the distal embolic
apophysis; unlike M. olmec (Fig. 178), the
flagellae and base of the median apophysis
are much thinner (Fig. 199).
Variation. Average body length of seven
females examined 5.5 mm, range 4.6 to 6.3
mm. Average body length of five males ex-
amined 3.8 mm, range 3.2 to 4.3 mm. j
Natural History. Mature specimens
have been collected from June to Febru-
ary, although I suspect that like other Ca-
ribbean species, M. triangularis is actually
entirely perennial (Fig. 337).
Distribution. Dominican Republic and
Cuba (Map 14).
Records Examined. CUBA Camagiiey Prov.:
21°30'N, 78°10'W, X.1954 (J. T. Acosta, AMNH). Ma-
tanzas: Matanzas, Parque Watldns, 23°3'N, 81°35'W,
9.viii.l955 (A. F. Archer, AMNH); Pan de Palenque
[?], 23°1'N, 81°43'W, ll.viii.l955 (A. F. Archer,
AMNH). Oriente: Banes, 20°58'N, 75°43'W,
2.viii.l955 (A. F Archer, AMNH); Cuabitas, Santiago,
20°4'N, 75°48'W, 15.xii.l955 (P Alayo, AMNH);
Puerto Boniato, 20°7'N, 75°47'W, 4.xi.l945 (P Alayo,
AMNH). Sierra las Casas: Isla de Pinos, 21°53'N,
82°48'W, 17.viii.l955 (A. F Archer, AMNH). DO-
MINICAN REPUBLIC Azua: Hatillo, 18°24'N,
70°32'W, 30.i.l991 (FelLx E. Del Monte, CAS). Ban-
ahona: Sierra Martin Garcia [?], 18°25'N, 70°30'W,
8.viii.l958 (A. F Archer & E. de Boyrie Moya,
Metepeira • Piel
65
pimungan
(24)
A.
9
triangularis
(25)
212
arizonica
(26)
Figures 192-198. Metepeira pimungan new species (sp. 24; 34°2'N, 120°22'W). 192, male palpus, mesal. 193, epigynum,
posterior. 194, epigynum, ventral. 195, male, dorsal. 196, male, ventral. 197, female, dorsal. 198, female, ventral.
Figures 199-205. Metepeira triangularis (Franganillo) (sp. 25; 18°34'N, 71°44'W). 199, male palpus, mesal. 200, epigynum,
posterior. 201, epigynum, ventral. 202, male, dorsal. 203, male, ventral. 204, female, dorsal. 205, female, ventral.
Figures 206-212. Metepeira arizonica Chamberlin and Ivie (sp. 26; 28°53'N, 1 13°4'W). 206, male palpus, mesal. 207, epigynum,
posterior. 208, epigynum, ventral. 209, male, dorsal. 210, male, ventral. 211, female, dorsal. 212, female, ventral.
Scale bars: dorsum and venter figures 1 .0 mm.
66
Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
AMNH). La Independencia: 2 km SE El Limon, Ji-
mani. S side of road, 18°25'N, 71°45'W, 30.i.l991
(Felix E. Del Monte, MNSD); Isla Cabritos: La Des-
cubierta, Lago Enriquillo, 18°34'N, 71°44'W,
18.vi.l981 (E. Marcano, MNSD). Montecristi: Car-
retera Montecristi, El Morro [?], 19°54'N, 71°39'W,
2.xii.l991 (EelLx E. Del Monte, MNSD), 21.xii.l991
(FelLx E. Del Monte, MNSD). Peravia: Las Dunas,
Prov. Bani [?], 18°25'N, 71°25'W, 23.i.l992 (Felix E.
Del Monte, MNSD). Frov. Tnijillo Valdez: W of
Bani, 18°17'N, 70°22'W, 8.viii.l958 (A. F. Archer,
AMNH). San Juan: 1 km S Las Matas de Farfan,
18°51'N, 71°31'W, 25.viii.1970 (B. Patterson, MCZ).
26. Metepeira arizonica
Chamberlin and Ivie
Figures 206-213, 319; Map 9
Metepeira arizonica Chamberlin and Ivie, 1942: 69,
figs. 182-187, 9,6. Female holotype from Canyon
Lake, Arizona, in the AMNH, examined. Levi,
1977: 200, figs. 12, 13, 39^6, 9,3. Brignoh, 1983:
275.
Description. Female from Isla Partida,
Baja Calif. Norte, Mexico. General color-
ation high in contrast. Carapace dark
brown; large white eye region, white lat-
eral posterior extensions, and short median
posterior extension (Fig. 211). Distinct
black rings on all legs. Femur I with row
of four macrosetae on anterior side; four
on anteroventral side. Dorsum of abdo-
men with usual Metepeira folium, though
generally slightly lighter (Fig. 211); venter
wide, with long white median line set
within surrounding U-shaped markings.
Pair of white spots on either side of spi-
racle (Fig. 212). Wide median white line
widening anteriorly with constriction in
center, set on black sternum (Fig. 212).
Ratio of eye diameters: posterior medians
and anterior medians 1.0, anterior laterals
1.4, posterior laterals 1.2. Anterior median
eyes separated by 1.4 diameters, posterior
median eyes by 0.9, anterior median eyes
separated from anterior laterals by 2.7 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.1 their diameters. To-
tal length 6.2 mm. Carapace 2.8 mm long,
2.2 wide. First femur 3.4 mm, patella and
tibia 3.5, metatarsus 3, tarsus 1.2. Second
patella and tibia 3 mm, third 1.9, fourth
2.8.
Male from Isla Partida, Baja Calif. Nor-
te, Mexico. Carapace, legs, abdoiTien, ven-
ter, sternum as in female, though legs
darker with less distinct rings, and white
line on sternum often broken (Figs. 209,
210). Femur I with row of four macrosetae
on anterior side; five to eight on anterov-
entral side. Ratio of eye diameters: poste-
rior medians and anterior medians 1.1, an-
terior laterals 1.4, posterior laterals 1.2.
Anterior median eyes separated by 1.5 di-
ameters, posterior median eyes by 0.7, an-
terior median eyes separated from anterior
laterals by 1.6 diameters of anterior lateral
eyes, lateral eyes separated by 0.2 their di-
ameters. Total length 4.4 mm. Carapace
2.3 mm long, 1.8 wide. First femur 3.3
mm, patella and tibia 3.3, metatarsus 3.2,
tarsus 1.2. Second patella and tibia 2.8
mm, third 1.5, fourth 2.2.
Diagnosis. Unlike all other species in
the M. incrassata group, the epigynum of
M. arizonica is puffy and swollen, causing
the openings to reduce (from a ventral
view) to crescent-shaped slits (Fig. 187).
The embolus differs from others in the M.
incrassata group because the distal em-
bolic apophysis is reduced to a gentle arch-
ing cui-ve (Fig. 206).
Variation. Average body length of nine
females examined 5.5 mm, range 4.7 to 6.3
mm. Average body length of 13 males ex-
amined 3.9 mm, range 3.1 to 5.1 mm. In
some males a small ventral keel is visible
extending beyond the flagella of the me-
dian apophysis (Levi, 1977, fig. 46).
'Natural History. Mature specimens are
most commonly collected between April
and August (Fig. 319), from dry oak-pine-
juniper woodland, alfalfa, cactus, and de-
sert shrub (Levi, 1977). Localities have a
large range in altitude, from sea level to
2,300 m; however, spiders from southern
regions live at higher elevations than spi-
ders from northern regions (Fig. 213).
Distribution. American southwest to
San Luis Potosi (Levi, 1977, map 1; Map
9).
Records Examined. MEXICO Baja Calif. Norte: 15
Metepeira • Piel
67
o
o
1
2200 •
^^
°
B
1800 -
°
^^^
o
o
1
1400 ■
1000 •
"""V
»^
i
■^^
°
1
600-
200-
— ma — a__-
nn. a. . . . . — ^
Latitude (degrees north)
Figure 213. The elevation of collection localities for M. arizon-
ica at their corresponding latitudes. Species-specific altitudes
appear to decrease with distance from the equator. Elevations
estimated from NOAA database of 5- by 5-minute geographic
tiles.
mi S Punta Prieta, 28°58'N, 114°17'W, 7.vii.l973 (S.
C. Williams & K. B. Blair, CAS); 24 mi S Santa Ines,
29°20'N, 114°20'W, 7.vii.l973 (S. C. Williams & K.
B. Blair, CAS); 26 mi S San Felipe, 30°38'N,
114°50'W, 15.iv.l965 (D. Q. Cavagnaro, C. E. & E.
S. Ross, V. L. Vesterby, CAS); 6 km NW Racho Santa
Ines, 29°43'N, 114°43'W, 28.iii.1981 (Paul E. Blom,
WHO; 6 mi S San Felipe, 30°55'N, 114°52'W
14.iv.l965 (D. Q. Cavagnaro, C. E. & E. S. Ross, V.
L. Vesterby, CAS); 9 km NW Racho Santa Ines,
29°46'N, 114°46'W, 3.vii.l981 (David Crowe, WHC);
Arroyo de Calamajue at carbonated spring, 29°38'N,
114°25'W, 26.vi.1973 (S. C. Williams & K. B. Blair,
CAS); Canon de Cuadalupe, off Laguna Salada,
32°10'N, 115°48'W, ll.i.l958 (V. Roth, AMNH); El
Mayor, 32°5'N, 115°13'W, 15.vi.l952 (M. Cazier, W
Gertsch, & R. Schrammel, AMNH); Isla Mejia,
29°34'N, 113°35'W, 30.ivl921 (J. C. Chamberlin,
MCZ); Isla Partida, 28°53'N, 113°4'W, 2.vii.l921 (J.
C. Chamberlin, MCZ); Isla San Lorenzo, N end,
28°40'N, 112°52'W, 24.vi.1921 (J. C. Chamberlin,
CAS); Isla San Lorenzo, south Tobart, 28°36'N,
112°46'W [?], 9.iv.l921 (J. C. Chamberlin, CAS);
Puerto Refugio, N end Angel de la Guarda Isl. Webs
in cracks in cliff overhanging beach, 29°34'N,
113°32'W, 18.iv.l962 (Howard W. Campbell, CAS);
San Pedro Martir, 30°45'N, 115°13'W, 18.iv.l921 (J.
C. Chamberlin, CAS). Coahuila: 25 mi SE San Pedro,
25°35'N, 102°50'W, 21.viii.l947 (W. J. Gertsch,
AMNH); Cabos, 25°35'N, 101°43'W, 21.viii.l947 (W.
J. Gertsch, AMNH); Gloria, 26°41'N, 10r23'W,
24.viii.1947 (W J. Gertsch, AMNH); Saltillo,
25°25'N, 101°0'W, 23.V.1952 (M. Cazier, W Gertsch,
& R. Schrammel, AMNH). Durango: La Loma,
25°27'N, 103°40'W, 20.viii.l947 (W J. Gertsch,
AMNH); Providencia, 26°44'N, 105°56'W
24.viii.1947 (A. M. Davis, AMNH); San Isidro, 60 mi
NW Durango, 25°1'N, 105°6'W, 19.viii.l947 (W J.
Gertsch, AMNH). San Luis Potosi: 3 km W Pilares,
21°55'34"N, 100°48'6"W, 21.X.1994 (W H. Piel,
MCZ). Sinaloa: Las Saleras Is. (= Isla SaHaca [?]),
25°11'N, 108°20'W [?], 13.vi.l921 (J. C. Chamberlin,
CAS). Sonora: 20 mi N Hermosillo, 29°8'N,
110°58'W, 13.Lx.1966 (Jean & Wilton Ivie, AMNH);
20 mi SW Sonoyta, 31°38'N, 113°4'W, 13.vi.l952 (W
J. Gertsch, AMNH); 22 mi E Hermosillo on the
banks of Rio Sonora, 29°4'N, 110°55'W, 17.viii.l959
(B. A. Branson, AMNH); Los Angeles, 29°27'N,
110°46'W, 4.X.1966 (V. Roth, AMNH); Puerto Pefi-
asco at seashore, 31°20'N, 113°33'W, 3.iv.l968 (D. E.
Bixler, MCZ). Zacatecas: 14 mi S Fresnillo, 23°5'N,
102°45'W, 4.viii.l954 (W J. Gertsch, AMNH); 4 mi
NE Concepcion del Oro, 24°41'N, 101°23'W,
4.vii.l984 (J. B. Woolley AD). USA Anzona: Santa
Catalina Mtns., mile 7.9 of Catalina Highway from
Tuscon to Mt. Lemmon., 32°17'N, 110°48'W,
19.vi.l985 (W Maddison, MCZ); Tucson, 32°15'N,
110°57'W, 5.vii.l991 (W H. Piel & G. S. Bodner,
MCZ). California: Corn Springs, Chuckawalla Mtns.,
10 mi SE Desert Center, 33°33'N, 115°24'W,
29.xi.1963 (D. C. Lorn, MCZ); Manzanita chaparral,
San Gabriel Canyon, Coldbrook Ranger Station,
34°11'N, 117°53'W, 19.viii.l964 (L. Pinter, MCZ).
Texas: Big Bend Nat'l Park, Old Ranch House on St.
Elena Rd., 29°10'N, 103°30'W, 25.V.I967 (E. Sabath,
MCZ).
27. Metepeira atascadero new species
Figures 214-221, 336; Map 14
Holottjpe. Male from San Miguel de Allende, Guana-
juato, Mexico, 25.x. 1982, George Uetz, in MCZ.
The specific name is a noun in apposition after the
Rancho Hotel Atascadero in San Miguel de Allen-
de. This name was coined by George Uetz when
he and his colleagues stayed in Rancho Hotel Atas-
cadero while studying the behavioral ecology of this
species. This name has been in informal use in the
literature (e.g., Uetz and Hodge, 1990).
Description. Female paratype from San
Miguel de Allende, Guanajuato, Mexico.
Reddish brown carapace, white around
eyes, faint light marks extend posteriorly
behind lateral eyes (Fig. 220). Legs yel-
lowish, ringed brown at distal ends of ar-
ticles. Femur I with row of four macro-
setae on anterior side; three on anterov-
entral side. Largest branches of fleur-de-
lis pattern on dorsal folium form large
paired white spots (Fig. 220). Venter of ab-
domen black with wide white median line,
flanked by white U-shape mark (Fig. 221).
Pair of small white spots on either side of
spiracle. Sternum black with white poste-
rior mark (Fig. 221). Ratio of eye diame-
ters: posterior medians and anterior me-
dians 1.0, anterior laterals 1.6, posterior
laterals 1.3. Anterior median eyes separat-
Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
ed by 1.4 diameters, posterior median eyes
by 0.9, anterior median eyes separated
from anterior laterals by 3.7 diameters of
anterior lateral eyes, lateral eyes separated
by 0.4 their diameters. Total length 7.5
mm. Carapace 3.1 mm long, 2.5 wide.
First femur 4 mm, patella and tibia 4,
metatarsus 3.9, tarsus 1.3. Second patella
and tibia 3.3 mm, third 1.8, fourth 2.8.
Male holotype. Reddish brown cara-
pace, lighter around eyes, faint light marks
extend posteriorly behind lateral eyes and
median eyes (Fig. 218). Legs yellowish
white, each article gradually turning red-
dish brown distally. Femur I with row of
four or five macrosetae on anterior side;
eight or nine on anteroventral side. Larg-
est branches of fleur-de-lis pattern on dor-
sal folium form large paired white spots
(Fig. 218). Venter of abdomen black with
wide white median line, flanked by faint
white U-shape mark (Fig. 219). Pair of
small white spots on either side of spiracle.
Sternum black with white posterior mark
(Fig. 219). Ratio of eye diameters: poste-
rior medians and anterior medians 1.0, an-
terior laterals 1.7, posterior laterals 1.4.
Anterior median eyes separated by 1.8 di-
ameters, posterior median eyes by 1.0, an-
terior median eyes separated from anterior
laterals by 3.6 diameters of anterior lateral
eyes, lateral eyes separated by 0.5 their di-
ameters. Total length 5 mm. Carapace 2.5
mm long, 1.8 wide. First femur 4 mm, pa-
tella and tibia 4.2, metatarsus 4.5, tarsus
1.4. Second patella and tibia 3.3 mm, third
1.7, fourth 2.5.
Diagnosis. The male embolus of M.
atascadero has a protruding bump be-
tween the embolus tip and the basal em-
bolic apophysis (Figs. 214, 217) which sep-
arates it from all other Metepeira species.
As with M. incrasssata (Fig. 224) and, to
a lesser degree, M. triangularis (Fig. 201),
the epigynal depressions on either side of
the scape of M. atascadero (Fig. 216) are
sclerotized to create a scooped and slick
quality. The darker epigynal openings in-
side the depressions are hidden anteriorly
under the scapes hood. Metepeira atas-
cadero's epigynum differs from that of M.
incrassata by its much smaller depressions
(compare Fig. 216 with 224).
Variation. Average body length of 12 fe-
males examined 7.9 mm, range 6 to 9.5
mm. Average body length of 12 males ex-
amined 5.7 mm, range 3.5 to 6.9 miTi.
Natural History. Mature specimens
have been collected from the end of Au-
gust to the end of October (Fig. 336). This
species lives solitarily: only 20% of spiders
live in aggregations of two or more indi-
viduals (Uetz and Hodge, 1990). It is
thought that the low level of social behav-
ior in M. atascadero occurs because the
species pursues a risk-prone foraging strat-
egy (Uetz, 1988a,b). Spiders are found be-
tween 1,500 and 2,500 m elevation.
Distribution. Mexican highlands from
Durango to Guerrero (Map 14).
Records Examined. MEXICO Coahuila: Saltillo, 14
mi E in Larrea Desert, 25°25'N, 100°55'W,
28.vii.1944 (AMNH). Durango: El Taseate, 26°12'N,
105°7'W, 27.vii.1947 (W. J. Gertsch, AMNH); Yer-
banis, 80 mi NW Durango, 24°45'N, 103°50'W,
19.viii.l947 (W. J. Gertsch, AMNH). Guanajuato: 20
mi E. Guanajuato, 21°1'N, 100°57'W, 15.ix.l976 (C.
E. Griswold & Jackson, CAS); Guanajuato, 21°1'N,
101°15'W (N. Banks, MCZ); San Miguel de Allende,
20°55'N, 100°45'W, 25.x. 1982 (George Uetz, MCZ),
14.X.1983 (MCZ), 14.X.1985 (MCZ), 17.x.l98^
(MCZ), 26.X.1985 (MCZ). Guerrero: Tecalpuico, 25
km N Iguala, 18°29'N, 99°38'W, l.i.l948 (AMNH).
Hidalgo: Ozumbilla, 20°9'N, 101°16'W, 2.X.1957 (R.
Dreisbach, MCZ). Jalisco: Cyarco Onda, 30 km W
Ojuelos, 21°47'N, 101°53'W (H. Wagner, AMNH).
Michoacan: Hills N of Patzcuaro, 19°45'N, 101°36'W,
24.viii.1959 (A. F. Archer, AMNH); Hwy 110, 4 mi
W Jiquilpan, 19°59'N, 102°47'W, 2.viii.l967 (R. E.
Leech, REL). Zacatecas: East of Guadalupe,
22°46'N, 102°31'W, 21.viii.l959 (A. F. Archer,
AMNH); Guadalupe, 22°45'N, 102°31'W, 16.viii.l947
(W. J. Gertsch, AMNH).
28. Metepeira incrassata
F. O. P. -Cambridge
Figures 222-228, 323; IVIap 9
Epeira Salei KeyserUng, 1864: 93, fig., 6,9. Female
holotype from Oaxaca, Mexico, in BMNH, exam-
ined. Keyserhng, 1892: 196, fig. 145, 9. Roewer,
1942: 851. NEW SYNONYMY.
Metepeira incrassata F. O. R-Cambridge, 1904: 460,
fig. 11, ?. Female holotype from Jalapa, Mexico,
in BMNH. Roewer, 1942: 868. Bonnet, 1957: 2821.
Metepeira • Piel
69
220"^" 221
atascadero
(27)
224
^im^ 227 ' 228
incrassata
(28)
234
Figures 214-221. Metepeira atascadero new species (sp. 27; 20°55'N, 100°45'W). 214, male palpus, mesal. 215, epigynum,
posterior. 216, epigynum, ventral. 217, male embolic division, ventral. 218, male, dorsal. 219, male, ventral. 220, female, dorsal.
221, female, ventral.
Figures 222-228. Metepeira incrassata F. O. P.-Cambridge (sp. 28 [222,225-228] 19°4'N, 97°2'W; [223,224] 18°54'N, 97°0'W).
222, male palpus, mesal. 223, epigynum, posterior. 224, epigynum, ventral. 225, male, dorsal. 226, male, ventral. 227, female,
dorsal. 228, female, ventral.
Figures 229-235. Metepeira ventura Chamberlin and Ivie (sp. 29; 28°5'N, 114°8'W). 229, male palpus, mesal. 230, epigynum,
posterior. 231, epigynum, ventral. 232, male, dorsal. 233, male, ventral. 234, female, dorsal. 235, female, ventral.
Scale bars: dorsum and venter figures 1.0 mm.
70
Bulletin Museum of Comparative Zoology, Vol. 157, No.
Aranea sallei: — F.O.P.-Cambridge, 1904: 519. Roew-
er, 1942: 851.
Araneus incrassata: — Petrunkevitch, 1911: 289.
Araneus sallei: — Petrunkevitch, 1911: 314.
Metepeira salei: — Levi, 1991: 180. Platnick, 1993:
449.
Note. Although M. incrassata is not the oldest
name, it has been cited more tlian 20 times in the
general literature (e.g., Caraco et al. 1995; Hieber
and Uetz, 1990; Hodge and Uetz, 1992, 1995,
1996; Jakob et al., 1996; Rayor, 1996; Rayor and
Uetz, 1990, 1993; Uetz, 1988a,b, 1989, 1991, 1992,
1996; Uetz and Hieber, 1994, 1997; Uetz and Hod-
ge, 1990; Uetz et al., 1994). In contrast, the senior
synonym has not been cited outside of taxonomic
catalogues. In compliance with Article 79 of the
ICZN (1985), I choose to assert the priority of tlie
more popular junior name.
Description. Female from Rancho Chu-
la-Vista, Veracruz, Mexico. Browii cara-
pace, darker around margins, lighter
around eyes and behind lateral eyes (Fig.
227). Legs brown, yellow at base of articles
and on articles distal to the femur for legs
I and II. Femur I with row of three to four
macrosetae on anterior side; one on anter-
oventral side. Dorsal folium darker than
most Metepeira species. White fleur-de-lis
pattern with thin branches (Fig. 227). Ven-
ter dark brown to black with white median
mark that is shorter than it is in most other
species (Fig. 228). Sternum dark brown to
black with one or two small white spots,
usually in the center or anteriorly (Fig.
228). Ratio of eye diameters: posterior me-
dians and anterior medians 0.9, anterior
laterals 1.3, posterior laterals 1.1. Anterior
median eyes separated by 1.5 diameters,
posterior median eyes by 1.1, anterior me-
dian eyes separated from anterior laterals
by 4.1 diameters of anterior lateral eyes,
lateral eyes separated by 0.5 their diame-
ters. Total length 7.8 mm. Carapace 3.6
mm long, 2.9 wide. First femur 3.9 mm,
patella and tibia 4.1, metatarsus 3.5, tarsus
1.4. Second patella and tibia 3.7 mm, third
2.3, fourth 3.3.
Male from Rancho Chula-Vista, Vera-
cruz, Mexico. Carapace, dorsum, venter,
sternum a darker version of female (Figs.
225, 226). Base of femora yellow, remain-
der dark brown; other articles gradually
turning lighter distally Femur I with row
of four to six macrosetae on anterior side;
six to nine on anteroventral side. Ratio of
eye diameters: posterior medians and an-
terior medians 1.0, anterior laterals 1.4,
posterior laterals 1.2. Anterior median
eyes separated by 1.4 diameters, posterior
median eyes by 1.0, anterior median eyes
separated from anterior laterals by 3.7 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.4 their diameters. To-
tal length 7 mm. Carapace 3.3 mm long,
2.7 wide. First femur 4.9 mm, patella and
tibia 5.3, metatarsus 4.8, tarsus 1.7. Sec-
ond patella and tibia 4.3 mm, third 2.2,
fourth 3.4.
Diagnosis. Overall pigmentation of M.
incrassata is darker than all others in its
species group. The thin branches on the
dorsal folium (Figs. 225, 227), the almost
entirely dark sternum, and the shortened
median line on the venter (Figs. 226, 228)
are distinctive. As with M. ohnec (Fig.
178), M. comanche (Fig. 185), and M. tri-
angularis (Fig. 199), the embolus of M. in-
crassata (Fig. 222) curves off sharply from
a distal embolic apophysis that does not
project forward. Compared to M. olniec
(Fig. 178) and M. comanche (Fig. 185), M.
incrassata's embolus tip beyond the distal
apophysis is relatively longer and not as
curved (Fig. 222). Male M. incrassata are
easily separated from M. triangularis by
the shape of the median apophysis (com-
pare Fig. 222 with Fig. 199). As with M.
atascadero (Fig. 216) and, to a lesser de-
gree, M. triangularis (Fig. 201), the epi-
gynal depressions on either side of the
scape of M. incrassata (Fig. 224) are scler-
otized to create a scooped and slick qual-
ity. The darker epigynal openings inside
the depressions are hidden anteriorly un-
der the scape's hood. Metepeira incrassa-
ta's epigynum differs from those of M.
atascadero and M. triangularis by its much
larger and disk-shaped depressions (com-
pare Fig. 224 with Figs. 201, 216).
Variation. Average body length of 31 fe-
males examined 7.6 mm, range 6.4 to 9.1
Metepeira • Piel
71
mm. Average body length of 22 males ex-
amined 6.2 mm, range 4.3 to 8 mm.
Natural History. Mature specimens
have been collected in February through
October but are most frequently found in
July (Fig. 323). This species lives socially:
half of all spiders live in aggregations of
1,000 or more individuals (Uetz and Hod-
ge, 1990). Colonies are frequently found
spanning telephone lines, houses, fences,
and other man-made structures. It is
thought that this high level of social be-
havior occurs because the species pursues
a risk-averse foraging strategy in which the
variance in survival attenuates with in-
creasing colony size (Uetz, 1988a,b). Ele-
vations center around 800 m and range
from about 500 to 1,500 m.
Distribution. Tropical Mexico from San
Luis Potosi to southern Veracruz and Oa-
xaca (Map 9).
Records Examined. MEXICO Hidalgo: 20 mi S of
Jacala, 20°50'N, 99°16'W, 18.iv.l946 (L. I. Davis &
M. Johnston, AMNH); Chapulhuacan, 21°10'N,
98°54'W, 20.V.1952 (M. Cazier, W. Gertsch, & R.
Schrammel, AMNH), 27.vii.1966 (Jean & Wilton
Ivie, AMNH), 16.vii.l969 (S. & J. Peck, MCZ). Mex-
ico: 1 mi S Palomas, 19°50'N, 99°5'W, 28.vii.1950
(AMNH). Oaxaca: Oaxaca, 17°3'N, 96°43'W (Nathan
Banks, MCZ). Puebla: Huauchinango, 20°11'N,
98°3'W, 7..X.1947 (H. Wagner, AMNH); north of Xic-
otepec de Juarez, 20°18'N, 97°57'W 19.ivl967 (W
B. Peck, MCZ). San Luis Potosi: 10km W Xilitla on
rl20, 21°22'N, 99°4'W 10.viii.l991 (W H. Piel & G.
S. Bodner, MCZ). Veracruz: 1 mi SW Tlapacoyan,
19°57'N, 97°14'W 16.vii.l973 (A. Newton, MCZ); 2
mi N Fortin de las Flores, 18°56'N, 97°1'W,
5.viii.l966 (Jean & Wilton Ivie, AMNH); 5 mi E Ori-
zaba, 18°51'N, 97°4'W 25.vii.1956 (W Gertsch & V.
Roth, AMNH); Coatepec, 19°27'N, 96°58'W,
28.vii.1955 (C. & P Vaurie, AMNH), 19.vii.l991 (W.
H. Piel & G. S. Bodner, MCZ); Fortin de las Flores,
18°54'N, 97°0'W, 7.vii.l947 (G. & M. Goodnight,
AMNH), 25.iv.1963 (W. J. Gertsch & W Ivie,
AMNH), 10.vii.l976 (A. Newton, MCZ), 21.X.1982
(George Uetz, MCZ), 17.vii.l991 (W. H. Piel & G. S.
Bodner, MCZ); Jalapa, 19°32'N, 96°55'W, 14.ii.l948
(H. Wagner, AMNH), 15.iii.l948 (H. Wagner,
AMNH), 15.X.1962 (N. L. H. Krauss, AMNH); Los
Naranjos, 18°21'N, 96°10'W 4.iii.l948 (H. Wagner,
AMNH); near Monte Blanco, 18°58'N, 97°1'W
3.viii.l973 (A. Newton, MCZ); Orizaba, 18°51'N,
97°4'W, 6.vii.l963 (D. BLxler, MCZ); Rancho Chula-
Vista, N of Cordoba, 19°4'N, 97°2'W 18.vii.l991 (W
H. Piel & G. S. Bodner, MCZ).
Metepeira ventura Group
Spiders in the M. ventura group {Me-
tepeira ventura, Metepeira revillagigedo ,
Metepeira celestun, Metepeira uncata, Me-
tepeira crassipes, Metepeira chilapae) are
closely related to those in the M. minima
group (M. petatlan, M. minima, M. paci-
fica, M. jamaicensis) . Females in the M.
ventura group have epigynal openings that
are wider than long and shaped as trian-
gles (Fig. 231), ovals (Fig. 252), or squares
(Figs. 259, 266), with their posterior edges
open. The posterior edges are observed to
be open because the posterior lobes at
their distal edge are wider than the distal
end of the scape. In many other species,
the gap between the lobes is narrower
than the scape and therefore hidden from
a ventral view, giving the impression that
the openings form closed shapes (e.g.. Fig.
123). In addition, with the exception of M.
celestun (Fig. 245), the scape is relatively
thin throughout its entire length (Figs.
231, 238, 252, 259, 266). In males, the em-
bolus can be slini and elongated with a
gentle curve (e.g.. Figs. 236, 243), or as
with some species in the M. minima group,
it can be tapering to a shaip bend right at
the tip (e.g.. Figs. 229, 264). In general,
the larger flagellum is a simple tapering
extension off the base (Figs. 229, 243, 250,
257, 264, and to a lesser extent, Fig. 236),
as opposed to being a distinctly separate
structure that abruptly cuives off the base
(e.g.. Fig. 129). Although the flagellae can
be thin, as in the M. minima species
group, they are not set off on a distinctly
narrower stalk (e.g., compare Fig. 264 with
Fig. 278).
29. Metepeira ventura
Chamberlin and Ivie
Figures 229-235, 320; Map 13
Metepeira ensenada Chamberlin and Ivie, 1942: 65,
figs. 166-168, 6. Male holotype from beach near
Ensenada, Mexico, in the AMNH, examined. Syn-
onymized by Levi (1977).
Metepeira ventura Chamberlin and Ivie, 1942: 67,
figs. 175-179, ?. Female holotype, 1 male and 3
female paratypes from between Oxnard and Santa
Monica, California, USA, in the AMNH, examined.
72 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
Levi, 1977: 204, figs. 53-60. Brignoli, 1983: 275.
Note. As first reviser, Levi (1977) preferred to
use the name M. ventura because a larger number
of specimens were available from the type locality.
Description. Female from 10 mi north
of Colonia Guerrero, Baja California Nor-
te, Mexico. Carapace brown with white
eye region, lateral posterior extensions,
and white median line (Fig. 234). Legs yel-
lowish tan with darker rings on fourth pair
of legs. Femur I with row of four macro-
setae on anterior side; two on anteroven-
tral side. Dorsum of abdomen with usual
Metepeira folium, lighter in anterior third
(Fig. 234). Venter of abdomen with wide
white median line posteriorly set within
hint of surrounding U-shaped marking.
Pair of white spots on either side of spi-
racle (Fig. 235). Sternum has wide inedian
white line widening anteriorly, sometimes
fragmented (Fig. 235). Ratio of eye di-
ameters: posterior medians and anterior
medians 1.0, anterior laterals 1.6, posterior
laterals 1.4. Anterior median eyes separat-
ed by 1.4 diameters, posterior median eyes
by 0.7, anterior median eyes separated
from anterior laterals by 2.9 diameters of
anterior lateral eyes, lateral eyes separated
by 0.3 their diameters. Total length 5.5
mm. Carapace 2.8 mm long, 2.2 wide.
First femur 3.3 mm, patella and tibia 3.3,
metatarsus 3, tarsus 1.2. Second patella
and tibia 2.9 mm, third 1.7, fourth 2.5.
Male from 10 mi north of Colonia
Guerrero, Baja California Norte, Mexico.
Carapace, abdomen, venter as in female
(Figs. 232, 233). Ringed legs darker than
in female. Femur I with row of four ma-
crosetae on anterior side; six to seven on
anteroventral side. Median white line on
sternum often broken (Fig. 233). Ratio of
eye diameters: posterior medians and an-
terior medians 0.9, anterior laterals 1.2,
posterior laterals 1.1. Anterior median
eyes separated by 1.6 diaineters, posterior
median eyes by 0.8, anterior median eyes
separated from anterior laterals by 1.9 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.3 their diameters. To-
tal length 4.3 mm. Carapace 2.2 mm long,
1.7 wide. First femur 3.6 mm, patella and
tibia 3.6, metatarsus 3.7, tarsus 1.3. Sec-
ond patella and tibia 2.8 mm, third 1.5,
fourth 2.3.
Diagnosis. Unlike other members of the
M. minima species group, the larger fla-
gellum on the median apophysis of M.
ventura is a tapering extension off the base
(Fig. 229), as opposed to arising from a
distinctly separate stalk (Fig. 286). Like M.
uncata, the epigynal openings of M. ven-
tura (Fig. 231) are somewhat triangular in
shape, in contrast to oval (Fig. 288) or slit-
shaped (Fig. 280) as in other members of
the M. minima species group. The rela-
tively shorter scape and the dark circles
inside the epigynal openings of M. ventura
(Fig. 231) contrast with M. uncata s rela-
tively longer scape and the dark sinuous
shapes inside the epigynal openings (Fig.
252).
Variation. Average body length of eight
females examined 6.7 mm, range 5.6 to 9.7
mm. Average body length of seven males
examined 5.1 mm, range 3.9 to 7 mm.
Natural History. Mature speciinens are
mostly collected April through September
(Fig. 320). Altitudes range from near sea
level to 1,000 m.
Distribution. California to northwestern
Mexico (Levi, 1977, map 1; Map 13).
Records Examined. MEXICO Baja Calif. Not~te: 10
m E El Rosario, 30°11'N, 115°46'W, 8.vii.l973 (S. C.
Wilhams & K. B. Blair, CAS); 10 mi N Col. Guerrero,
28°5'N, 114°8'W, l.b(.1957 (V. Roth, AMNH); 2 mi
SE Erendira, 3ri9'N, 116°19'W, 12.V.1973 (S. C.
Wilhams & K. B. Blair, CAS); Isla Cedros, Gran Cafi-
on, 28°12'N, 115°15'W, 10.iii.l945 (B. F. Osorio Taf-
all, AMNH); Isla de Cedros, Cerro de Cedros,
28°12'N, 115°15'W, l.vii.l983 (V. F. Lee, CAS); Isla
de Cedros, Punta Norte [?], 28°22'N, 115°14'W,
3.vii.l983 (V. F. Lee, CAS); Isla de Cedros, trail to
Cerro de Cedros, at spring, 28°12'N, 115°15'W,
27.Lx.1984 (D. B. Weissman, V. F Lee, CAS); Islas
San Benito, Benitos del Oeste, 28°18'N, 115°35'W,
4.vii.l983 (D. C. Lightfoot & V. F Lee, CAS); Islas
San Benito, Middle Island, 28°19'N, 115°34'W,
9.iv.l981 (Stanley C. Williams, CAS); Islas San Be-
nito, South Island, 28°18'N, 115°35'W, 9.iv.l981
(Stanley C. Williams, CAS); near Consuelo, 6 mi NW
El Rosario, 30°11'N, 115°46'W 18.iv.l965 (D. Q.
Cavagnaro, C. E. & E. S. Ross, V. L. Vesterby, CAS);
Rancho Las Parritas, 10 mi S San Quintin, 30°20'N,
Metepeira • Piel
73
115°57'W, 27.vi.1977 (C. E. Griswold, CAS); Santo
Tomas, 31°33'N, 116°24'W, 8.vii.l953 (W. J. & J. W.
Gertsch, AMNH). Baja Calif. Sun Bahia de los
Muertos, 23°58'N, 109°50'W, 20.xii.l958 (H. B.
Leech, CAS); Desierto del Vizcaino, Laguna Ojo de
Liebre, sobre frutilla, 27°43'N, 114°15'W, ll.xi.l981
(A. Cota & M. Jimenez, MLJ); E edge of Sierra Pla-
ceres, 27°35'N, 114°30'W, 25.iii.1984 (W. J. Pulawsld,
CAS). Sonora: 6 mi E Navojoa, 27°6'N, 109°23'W,
23.viii.1965 (W. J. Gertsch & R. Hastings, AMNH).
USA California: 1 mi NW Winchester (Double
Butte), 33°43'N, 117°6'W, 7.xii.l976 (W. Icenogle,
MCZ); Lompoc, by US 63, 34°38'N, 120°27'W,
17.viii.l966 (L. & P. Pinter, MCZ); Lucia, Monterey
Co., 36°1'N, 121°33'W, 15.ix.l964 (L. Pinter, MCZ);
Manzanita chaparral, San Gabriel Canyon, Coldbrook
Ranger Station, 34°11'N, 117°53'W, 29.V.1965 (L.
Pinter, MCZ); Santa Catalina Island Area near Hay-
press Res, 33°23'N, 118°25'W, 6.ii.l993 (Martin C.
Ramirez & Laura B. Fandino, MCZ); Santa Catalina
Island, Mt. Torquemada, 33°26'N, 118°33'W,
15.viii.l965 (L. Pinter, MCZ); Winchester, 33°42'N,
117°5'W, ll.v.1970 (W. Icenogle, MCZ); Winchester,
Double Butte, 33°42'N, 117°5'W, 19.V.1974 (W. Icen-
ogle, MCZ). Nevada: Quin River Crossing, 41°35'N,
118°27'W, 21.vi.l975 (G. F. Knowlton, MCZ).
30. Metepeira revillagigedo new species
Figures 236-242, 301; IVIap 12
Holotype. Female from south of Isla Socorro, Archi-
pielago de Revillagigedo, State of Colima, Me.xico,
15.xii.l988, M. Jimenez, in MCZ. The specific
name is a noun in apposition after the locality.
Description. Female holotype. Carapace
dark brown with light region in median
eye quadrangle and surrounding lateral
eyes. Pair of darker, elliptic, walnut leaf
shapes on lighter patch near center of car-
apace (see Fig. 241). Darker rings on distal
ends of femora, patellae, tibiae, and prox-
imal dorsal portion of tibia I, II. Femur I
with three to four macrosetae on anterior
side; none on the anteroventral side. Dor-
sum of abdomen with white oak leaf foli-
um, margined with dark markings, partic-
ularly on trailing edges of lobes (Fig. 241).
Coloration has a slightly golden hue. Dark
lateral band wraps around sides of abdo-
men and stretches up over anterior dorsal
portion. Venter with two white spots on
either side of spiracle. Anterior to spiracle,
slight indication of a V-shaped mark with
longitudinal extensions reaching halfway
up abdomen. Wide, white medial longitu-
dinal line to epigynal groove (Fig. 242).
Sternum with wide, white longitudinal
mark widening anteriorly (Fig. 242). Ratio
of eye diameters: posterior medians and
anterior medians 0.8, anterior laterals 1.1,
posterior laterals 1. Anterior median eyes
separated by 1.3 diameters, posterior me-
dian eyes by 1.0, anterior median eyes sep-
arated from anterior laterals by 2.5 diam-
eters of anterior lateral eyes, lateral eyes
separated by 0.3 their diameters. Total
length 7.5 mm. Carapace 3.2 mm long, 2.5
wide. First femur 3.5 mm, patella and tibia
3.8, metatarsus 3.2, tarsus 1.2. Second pa-
tella and tibia 3.2 mm, third 1.8, fourth
2.8.
Male paratype froin Isla Socorro, Coli-
ma. Carapace dirty yellowish brown with
lighter posteriorly pointing acute triangle
in center (Fig. 239). Legs same color as
carapace, except lighter on proximal half
of femora. Femur I with row of four ma-
crosetae on anterior side, row of two ma-
crosetae on anteroventral side. Male ab-
domen lighter and with less contrast than
female (Fig. 239). Venter with a much re-
duced and shorter longitudinal line than
female. Sternum as in female, except lon-
gitudinal line more often broken (Fig.
240). Ratio of eye diameters: posterior me-
dians and anterior medians 1.0, anterior
laterals 1.1, posterior laterals 1.1. Anterior
median eyes separated by 1.5 diameters,
posterior median eyes by 0.8, anterior me-
dian eyes separated from anterior laterals
by 1.4 diameters of anterior lateral eyes,
lateral eyes separated by 0.3 their diame-
ters. Total length 3.2 mm. Carapace 1.7
mm long, 1.1 wide. First femur 2.5 mm,
patella and tibia 2.5, metatarsus 2.3, tarsus
0.9. Second patella and tibia 2 mm, third
1.0, fourth 1.5.
Diagnosis. The epigynum of M. revilla-
gigedo looks veiy different from those of
other species in the M. ventura group. In-
stead of having small, sharply delineated
depressions on either side of the scape
(e.g.. Fig. 259), M. revillagigedo has larger
but more gradual depressions (Fig. 238).
Despite the unique appearance of the epi-
74 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
gynum, the affinity between M. revillagi-
gedo and the M. ventura species group can
be seen in the black comma-shaped marks
inside the epigynal depressions (Fig. 238)
similar to those in, for example, M. Ven-
tura (Fig. 231). The male emboli of M.
revillagigedo (Fig. 236) and M. celestun
(Fig. 243) are veiy similar in shape and
relatively longer than in other species in
the M. Ventura group. Males of M. revil-
lagigedo differ from males of M. celestun
by the thickness and shape of the median
apophysis (compare Fig. 236 with Fig.
243).
Natural History. Specimens have only
twice been collected: once in December
and once in April (Fig. 301). This species
is known to live in fig trees.
Distribution. This species is found on
Isla Socorro in the Pacific Ocean (Map
12), the island being one of several that
form the Archipielago de Revillagigedo. It
is well isolated, situated about 465 km
south of Baja California Sur and 588 km
west of the Jalisco coastline.
Records Examined. MEXICO Colinia: Archipiela-
go de Revillagigedo: Isla Socorro, 18°45'N, 110°57'W,
26.iv.1932 (Templeton, Crocker Exped., CAS); Ar-
chipielago de Revillagigedo: Sur de la Isla Socorro,
18°44'N, 110°57'W, 15.xii.l988 (M. Jimenez, MCZ).
31 . Metepeira celestun new species
Figures 243-249, 326; IVlap 13
Holotype. Male from Celestun, Yucatan, Mexico,
24.vii.1991, W. H. Piel & G. S. Bodner, in MCZ.
The specific name is a noun in apposition after the
locality.
Note. Since males of this species were never
found witli females, one cannot be sure absolutely
that they are conspecific. Several facts argue for
conspecificity: all specimens were collected in sim-
ilar habitats and during the same season; dorsal fo-
lium of males and females share similar patterns
and both have hint of gold coloration; based on
parallel correspondence widi other species, die
male palp is arguably compatible with the female s
epigynum.
Description. Female paratype from
Edzna, Campeche, Mexico. Carapace yel-
lowish tan with lighter area around eyes
(Fig. 248). Legs yellowish tan with darker
rings on fourth pair of legs. Femur I with
row of four macrosetae on anterior side;
sometimes four on anteroventral side.
Dorsal folium as in other Metepeira, ex-
cept speckled an unusual reddish gold col-
or (Fig. 248). Venter wide white median
line with pair of large white spots on either
side of spiracle (Fig. 249). Sternum has
wide median white line widening anteri-
orly with constriction in center (Fig. 249).
Ratio of eye diameters: posterior medians
and anterior medians 1.0, anterior laterals
1.3, posterior laterals 1.2. Anterior median
eyes separated by 1.5 diameters, posterior
median eyes by 0.9, anterior median eyes
separated froin anterior laterals by 2.4 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.2 their diameters. To-
tal length 5.2 mm. Carapace 2.5 mm long,
1.9 wide. First femur 2.6 mm, patella and
tibia 2.8, metatarsus 2.2, tarsus 0.9. Sec-
ond patella and tibia 2.4 mm, third 1.4,
fourth 2.
Male holotype. Carapace dark with light
region around eyes and a light triangular
mark anterior to thoracic furrow (Fig.
246). Coxae and proximal third of femora
white, remaining two-thirds black; other
articles similarly ringed. Femur I with row
of four macrosetae on anterior side; three
on anteroventral side. Dorsal folium with
slight gold iridescence. Fleur-de-lis pat-
tern leafy and pinnate (Fig. 246). Venter,
sternum as in female, except median white
line on sternum often broken (Fig. 247).
Ratio of eye diameters: posterior medians
and anterior medians 1.1, anterior laterals
1.4, posterior laterals 1.3. Anterior median
eyes separated by 1.6 diameters, posterior
median eyes by 0.9, anterior median eyes
separated from anterior laterals by 1.4 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.2 their diameters. To-
tal length 2.8 mm. Carapace 1.4 mm long,
1.1 wide. First femur 1.8 mm, patella and
tibia 1.8, metatarsus 1.4, tarsus 0.8. Sec-
ond patella and tibia 1.5 mm, third 0.8,
fourth 1.2.
Diagnosis. The overall coloration of M.
celestun in alcohol has an unusually golden
quality that is rare among preserved Me-
Metepeira • Piel 75
Figures 236-242. Metepeira revillagigedo new species (sp. 30; 18°44'N, 110°57'W). 236, male palpus, mesal. 237, epigynum,
posterior. 238, epigynum, ventral. 239, male, dorsal. 240, male, ventral. 241, female, dorsal. 242, female, ventral.
Figures 243-249. Metepeira celestun new species (sp. 31 [243,246,247] 20°56'N, 90°21'W; [244,245,248,249] 19°35'N,
90°15'W). 243, male palpus, mesal. 244, epigynum, posterior. 245, epigynum, ventral. 246, male, dorsal. 247, male, ventral.
248, female, dorsal. 249, female, ventral.
Figures 250-256. Metepeira uncata F. O. P.-Cambridge (sp. 32 [250,251,253-256] 14°40'N, 92°9'W; [252] 14°49'N, 91°31'W),
250, male palpus, mesal. 251, epigynum, posterior. 252, epigynum, ventral. 253, male, dorsal. 254, male, ventral. 255, female,
dorsal. 256, female, ventral.
Scale bars: dorsum and venter figures 1.0 mm.
76 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
tepeira. Unlike all other members of the
M. Ventura species group, the scape of M.
celestun is very wide at its base, thereby
partly concealing the epigynal openings
(Fig. 245). However, if the scape is dis-
carded, the exposed openings have black
comma-shaped marks similar to those of
M. Ventura (Fig. 231), thereby confirming
the species' affinities with other members
in the M. ventura species group. Unlike
other members, the posterior lobes nearly
touch each other near their base (Fig.
244). The emboli of M. celestun and M.
revillagigedo are very similar in shape and
relatively longer than in other species in
the M. ventura group (Fig. 243). Metepei-
ra celestun differs from M. revillagigedo by
the thickness and shape of the median
apophysis (compare Fig. 243 with Fig.
236).
Variation. Average body length of four
feinales examined 5.6 mm, range 5.2 to 6.4
mm. Average body length of three males
examined 2.7 mm, range 2.2 to 2.9 mm.
Female scape sometimes wider than it ap-
pears in Fig. 245, resembling Metepeira
arizonica. Legs and carapace color vary
from yellowish tan to orange-red.
Natural History. Mature specimens
have been collected in July (Fig. 326) in
forested clearings, roadside bushes, palms,
and swampy areas near the beach.
Distribution. Yucatan peninsula (Map
13).
Records Examined. MEXICO Campeche: Edzna,
19°35'N, 90°15'W, 22.vii.1991 (W. H. Piel & G. S.
Bodner, MCZ). Quintana Roo: Kohunlich ruins, 9 km
S Franciso Villa, 18°26'N, 88°48'W, 15.vii.l983 (R. S.
Anderson, MCZ). Yucatan: 3km S San Felipe,
21°32'N, 88°14'W, 25.vii.1991 (W. H. Piel & G. S.
Bodner, MCZ); 4 km N Xocenpich, 12 km N Piste,
on road to Dzitas, 20°47'N, 88°34'W, 20.vii.l983 (W.
Maddison, MCZ); Balankanche Cave, 2 km E Chi-
chen Itza, 20°40'N, 88°33'W, 19.vii.l983 (W. Mad-
dison, MCZ); beach north of Celestun, 20°56'N,
90°21'W, 24.vii.1991 (W. H. Piel & G. S. Bodner,
MCZ); Chichen Itza ruins on HWY 180, seasonal for-
est, 20°40'N, 88°34'W, 19.vii.l983 (W. Maddison &
R. S. Anderson, MCZ); Uxmal, 20°22'N, 89°46'W,
23.vii.1991 (W. H. Piel & G. S. Bodner, MCZ).
32. Metepeira uncata
F. O. P. -Cambridge
Figures 250-256, 329; Map 12
Metepeira uncata F. O. P.-Cambridge, 1903: 459, fig.
8, 6. Male holotype from Santa Ana, Guatemala,
in BMNH. Roewer, 1942: 868. Bonnet, 1957: 2823.
Araneus uncatus: — Petrunkevitch, 1911: 321.
Description. Female from Ayutla, San
Marcos, Guatemala. Carapace dark brown
with light region surrounding the eyes and
extending posteriorly behind the lateral
eyes. Pair of darker feather-shaped patches
anterior of thoracic furrow, touching pos-
teriorly to create a U-shape (see Fig. 255).
Dark rings on distal half of leg articles, an-
terior side of patellae, and dark markings
on dorsal, proximal end of tibia I and II.
Femur I with four macrosetae on anterior
side; one on anteroventral side. Dorsum of
abdomen with white fleur-de-lis folium;
this pattern thinner than in most other
species. Fleur-de-lis pattern on a dark
background, outlined on either side by
thin white stripe (Fig. 255). Dark lateral
band follows sides of abdomen and
stretches up over anterior dorsal portion.
Venter with two white spots on each side
of spiracle; wide, white, medial longitudi-
nal line starts anterior to the colulus and
ends posterior to the epigynal groove.
Thin, faint lines run parallel to wide me-
dian one anterior to white spots. White
patch separates epigynum and dark de-
pression posterior to pedicel (Fig. 256).
Sternum black with white dewdrop-
shaped mark at posterior end (Fig. 256).
Ratio of eye diameters: posterior medians
and anterior medians 1.0, anterior laterals
1.3, posterior laterals 1.2. Anterior median
eyes separated by 1.6 diameters, posterior
median eyes by 1.1, anterior median eyes
separated froin anterior laterals by 4 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.4 their diameters. To-
tal length 7.8 mm. Carapace 3.7 mm long,
2.9 wide. First femur 4 mm, patella and
tibia 4.3, metatarsus 3.8, tarsus 1.4. Sec-
ond patella and tibia 3.6 mm, third 2.3,
fourth 3.2.
Metepeira • Piel 77
Male from Ayutla, San Marcos, Guate-
mala. Carapace dirty yellowish brown with
lighter mark in center (Fig. 253). Saine
color as carapace except lighter on proxi-
mal half of femora. Femur I with row of
nine macrosetae on anterior side; five on
anteroventral side. Male abdomen similar
to female, but leaves of fleur-de-lis pattern
thinner (Fig. 253). Venter with a shorter
longitudinal line, as compared with female
(Fig. 254). Sternum with posterior white
dewdrop as in female, and additional an-
terior white mark near labium (Fig. 254).
Ratio of eye diameters: posterior medians
and anterior medians 0.9, anterior laterals
1.3, posterior laterals 1.3. Anterior median
eyes separated by 1.5 diameters, posterior
median eyes by 1.0, anterior median eyes
separated from anterior laterals by 2 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.3 their diameters. To-
tal length 4.5 mm. Carapace 2.3 mm long,
1.8 wide. First femur 3.4 mm, patella and
tibia 3.4, metatarsus 3.2, tarsus 1.1. Sec-
ond patella and tibia 2.7 mm, third 1.4,
fourth 2.
Diagnosis. The generally darker pig-
mentation (Fig. 255) and the small white
mark on the sternum (Fig. 256) separate
M. uncata from all other members of the
M. Ventura species group. Like M. ventura
(Fig. 231) but unlike other members of
the M. ventura species group, the open-
ings to the epigynum are more triangular
to oval (Fig. 252), rather than square (Figs.
259, 266). Also distinctive are the sinuous
black lines that form upside-down U-
shapes inside each epigynal opening (Fig.
252), in contrast to the comma-shaped
marks in M. ventura or M. chilapae (Figs.
231, 266). Like M. chilapae but unlike oth-
er members of the M. ventura group, the
median apophysis and flagellae of M. un-
cata are slimmer and of uniform thickness
(Figs. 250, 264). The palp of M. uncata
differs from that of M. chilapae by the
more basal position of the bend in the em-
bolus tip (compare Fig. 250 with Fig. 264).
Variation. Average body length of five
females examined 8.2 mm, range 7.5 to 8.8
mm. Average body length of four males
examined 5.6 mm, range 4.5 to 6.4 mm.
Natural Historij. Mature specimens
have been collected in August (Fig. 329).
Distribution. Southwestern Guatemala
to northern Costa Rica (Map 12) at alti-
tudes ranging from 100 to 3,000 m.
Records Examined. COSTA RICA Cordillera: 20
km N Siquires, 10°9'N, 84°17'W, 15.viii.l980 (W.
Eberhard, MCZ). GUATEMALA El Quiche: Chichi-
castenango, 14°56'N, 91°7'W, 6.viii.l947 (C. & P.
Vaurie, AMNH). Huehuetenango: Todos Santos Cu-
chumatan, 15°31'N, 91°37'W, 16.viii.l979 (C. E.
Griswold, CAS). Quetzaltenango: El Baul, 1 km S
Quetzaltenango, 14°49'N, 91°31'W, 14.viii.l979 (T C.
Meikle & C. E. Griswold, CAS); Quetzaltenango,
14°50'N, 91°31'W, 16.viii.l950 (C. J. & M. Good-
night, AMNH). San Marcos: Ayutla, 14°40'N, 92°9'W,
19.viii.l947 (AMNH).
33. Metepeira crassipes
Chamberlin and Ivie
Figures 257-263, 309; Map 12
Metepeira josepha Chamberlin and Ivie, 1942; 64, fig.
165, 9 . Female holotype from Kings Mtn. near
Palo Alto, California in the AMNH. Synonymized
by Levi (1977).
Metepeira crassipes Chamberlin and Ivie, 1942: 66,
figs. 171-173, 9, S. Male holotype, female, male
paratypes from Laguna Beach, California in the
AMNH. Levi, 1977: 202, figs. 47-52, 9,6.
Note. As first reviser, Levi (1977) preferred to
use name M. crassipes.
Description. Female from Isla San Pe-
dro Nolasco, Sonora, Mexico. Carapace
with large white eye region and lateral
posterior extensions (Fig. 262). Legs
ringed at distal ends of articles — though
sometimes only lightly. Femur I with four
macrosetae on anterior side aligned in
straight row; one to four light setae on an-
teroventral side. Dorsum of abdomen with
usual Metepeira folium (Fig. 262); venter
wide, with long white median line set
within U-shaped thinner white lines. Pair
of white spots on either side of spiracle
(Fig. 263). Sternum black with wide me-
dian white line widening anteriorly (Fig.
263). Ratio of eye diameters: posterior me-
dians and anterior medians 1.1, anterior
laterals 1.4, posterior laterals 1.3. Anterior
median eyes separated by 1.4 diameters.
78 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
posterior median eyes by 0.8, anterior me-
dian eyes separated from anterior laterals
by 3 diameters of anterior lateral eyes, lat-
eral eyes separated by 0.1 their diameters.
Total length 5.7 mm. Carapace 2.8 mm
long, 2.1 wide. First femur 3.2 mm, patella
and tibia 3.1, metatarsus 2.8, tarsus 1. Sec-
ond patella and tibia 2.7 mm, third 1.6,
fourth 2.5.
Male from Winchester, California. Car-
apace as in female except often with me-
dian white mark. Femur I with row of four
macrosetae on anterior side; about four to
eight on anteroventral side. Dorsum, ven-
ter as in female (Fig. 260); median white
line on sternum more often broken (Fig.
261). Ratio of eye diameters: posterior me-
dians and anterior medians 1.0, anterior
laterals 1.6, posterior laterals 1.4. Anterior
median eyes separated by 1.2 diameters,
posterior median eyes by 0.6, anterior me-
dian eyes separated from anterior laterals
by 2 diameters of anterior lateral eyes, lat-
eral eyes separated by 0.3 their diameters.
Total length 4 mm. Carapace 2.1 mm long,
1.5 wide. First femur 3.2 mm, patella and
tibia 3.2, metatarsus 3.2, tarsus 1.1. Sec-
ond patella and tibia 2.5 mm, third 1.3,
fourth 2.
Diagnosis. Unlike other species in the
M. Ventura species group, the openings to
the epigynum of M. crassipes resemble
those of M. chilapae because they are
shaped like squares with rounded edges
(compare Fig. 259 with Fig. 266). How-
ever, compared to M. chilapae, the epigyn-
al openings of M. crassipes are relatively
smaller and the scape is relatively longer.
The embolus of M. crassipes is distin-
guished by its gentle cui'vature (Fig. 257),
in contrast to much more abi"upt cui"vature
seen in other species in the M. ventura
group (e.g.. Fig. 264), with the exception
of M. revillagigeclo. The median apophysis
of M. crassipes (Fig. 257) is slimmer than
that of M. revillagigedo (Fig. 236).
Variation. Average body length of four
females examined 6 mm, range 5 to 6.5
mm. Average body length of two males ex-
amined 4.4 mm, range 4.1 to 4.7 mm.
Sometimes the scape is greatly swollen,
adding to its relative width. In such cases
the scape can cover the epigynal openings,
which may mislead the investigator to con-
fuse it for Metepeira arizonica.
Natural Histortj. Levi (1977) reports
that males in the U.S. have been collected
from April to October, primarily in Cali-
fornia buckwheat and sage. Mexican re-
cords expand this seasonality to include
the entire year (Fig. 309).
Distribution. From northern California
in the U.S. to Baja California Sur, Mexico
(Levi, 1977, map 1; Map 12).
Records Examined. MEXICO Baja Calif. Norte:
Santo Tomas, 31°33'N, 116°24'W, 8.vii.l953 (W. J. &
J. W. Gertsch, AMNH), 12.xi.l976 (S. C. Williams &
K. B. Blair, CAS). Baja Calf. Sun 26 mi S Loreto,
25°37'N, 111°17'W, l.i.l977 (C. E. Griswold & L.
Vincent, CAS); Isla Magdalena, Puerto Magdalena,
24°38'N, 112°9'W, 16.iii.l957 (R. Zweifel, AMNH);
Isla San Francisco, South Side, 24°50'N, 110°35'W,
19.V.1970 (S. C. Williams & V. F. Lee, CAS); Sierra
San Nicolas, 26°32'N, 111°36'W [?] (Eisen & Vaslit,
MCZ). Sonora: Isla San Pedro Nolasco, 27°58'N,
111°25'W, 17.iv.l921 (G. C. Chamberlin, MCZ); Si-
erra de Alamos, 30°51'N, 112°2'W [?], 19.i.l968 (V.
Roth, AMNH). USA California: 1 mi NW Winches-
ter (Double Butte), in web between Artemisia cali-
fornica bushes, 33°43'N, 117°6'W, 4.xii.l976 (W.
Icenogle, MCZ).
34. Metepeira chilapae
Chamberlin and Ivie
Figures 264-270, 333; Map 13
Metepeira chilapae Chamberlin and Ivie, 1936: 45,
figs. 119-121, S. Male holotype from Chilapa,
Guerrero, Mexico, in the ZMB. Roewer, 1942: 868.
Metepeira chilapica: — Bonnet, 1957: 2820. Unjusti-
fied eiuendation.
Note. Examination of Chamberlin and Ivies
(1936) and Levi's (personal illustrations) figures of
the holotype was sufficient to identify M. chilapae
accurately.
Description. Female from Cocoyoc,
Morelos, Mexico. Brownish black cara-
pace, paler around and just posterior to
lateral eyes (Fig. 269). Distal halves of
ventral leg articles black, elsewhere yel-
lowish. Femur I with row of four macro-
setae on anterior side; zero to three setae
on anteroventral side. Dorsal folium a
white fleur-de-lis marking set on golden-
Metepeira • Piel 79
257
crassipes
(33)
260 261
269
chilapae
(34)
270
276
277
272
petatlan
(35)
Figures 257-263. Metepeira crassipes Chamberlin and Ivie (sp. 33 [257-261] 33°43'N, 117°6'W; [262,263] 27°58'N, 1 1 1°25'W).
257, male palpus, mesal. 258, epigynum, posterior. 259, epigynum, ventral. 260, male, dorsal. 261, male, ventral. 262, female,
dorsal. 263, female, ventral.
Figures 264-270. Metepeira cliilapae Chamberlin and Ivie (sp. 34 [264] 17°39'N, 99°22'W; [265-270] 17°39'N, 99°22'W). 264,
male palpus, mesal. 265, epigynum, posterior. 266, epigynum, ventral. 267, male, dorsal. 268, male, ventral. 269, female, dorsal.
270, female, ventral.
Figures 271-277. Metepeira petatlan new species (sp. 35 [271,274,275] 17°14'N, 100°53'W; [272,273,276,277] 17°17'32"N,
101°2'40"W). 271, male palpus, mesal. 272, epigynum, posterior. 273, epigynum, ventral. 274, male, dorsal. 275, male, ventral.
276, female, dorsal. 277, female, ventral.
Scale bars: dorsum and venter figures 1.0 mm.
80
Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
brown speckled pattern over white. Foli-
um darkens posteriorly (Fig. 269). Ante-
rior shoulders black. Venter black with
wide median white line and pair of white
spots on eidier side of spiracle (Fig. 270).
Sternum brownish black with wide, white
line widening anteriorly (Fig. 270). Ratio
of eye diameters: posterior medians and
anterior medians 1.0, anterior laterals 1.1,
posterior laterals 1.1. Anterior median
eyes separated by 1.3 diameters, posterior
median eyes by 0.8, anterior median eyes
separated from anterior laterals by 2.2 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.2 their diameters. To-
tal length 5.8 mm. Carapace 2.7 mm long,
2.3 wide. First femur 2.9 mm, patella and
tibia 3, metatarsus 2.6, tarsus 1. Second
patella and tibia 2.4 mm, third 1.6, fourth
2.3.
Male from 6 mi northeast of Tixtla de
Guerrero, Guerrero, Mexico. Carapace,
dorsum, venter, sternum darker and more
contrasty version of female (Figs. 267,
268). Distal halves of ventral leg articles
black, elsewhere yellowish. Femur I with
row of four macrosetae on anterior side;
three on an tero ventral side. Ratio of eye
diameters: posterior medians and anterior
medians 1.0, anterior laterals 1.3, posterior
laterals 1.3. Anterior median eyes separat-
ed by 1.7 diameters, posterior median eyes
by 0.9, anterior median eyes separated
from anterior laterals by 1.9 diameters of
anterior lateral eyes, lateral eyes separated
by 0.3 their diameters. Total length 3.5
mm. Carapace 1.8 mm long, 1.4 wide.
First femur 2.3 mm, patella and tibia 2.1,
metatarsus 1.8, tarsus 0.8. Second patella
and tibia 1.8 mm, third 0.9, fourth 1.4.
Diagnosis. The openings to the epigyn-
um resemble those of M. crassipes be-
cause they are shaped like squares with
rounded edges (compare Fig. 266 with
Fig. 259). However, the dark, comma-
shaped mark inside each opening resem-
bles that of M. Ventura (compare Fig. 266
with Fig. 231). As in M. ventura (Fig. 229)
but unlike all other species in the M. ven-
tura species group, the embolus tapers
strongly then curves sharply up and inward
right at the tip (Fig. 264). The bend in the
embolus in other M. ventura group species
is less pronounced (Figs. 236, 257) or not
so near the tip (Figs. 243, 250). Unlike M.
ventura, the median apophysis in M. chi-
lapae is slimmer (Fig. 264) and the sepa-
ration between the embolus and the basal
embolic apophysis is relatively greater.
Variation. Average body length of eight
females examined 6.5 ixim, range 5.7 to 8.3
mm. Average body length of three males
examined 2.9 mm, range 2.3 to 3.5 mm.
Natural History. Mature specimens
have been collected in July through Oc-
tober (Fig. 333).
Distribution. Southern Mexican states
between Nayarit and Oaxaca; elevations
from 1,000 to 2,000 m (Map 13).
Records Examined. MEXICO Guerrero: 6 mi NE
Tixtla de Guerrero, 17°39'N, 99°22'W, 16.vii.l984 (J.
B. Woolley, ADC). Morelos: Cocoyoc, 18°52'N,
98°59'W, 28.vii.1956 (W. Gertsch & V. Roth, AMNH);
Cuemavaca, 18°55'N, 99°15'W, 15.X.1944 (N. L. H.
Krauss, AMNH). Nayarit: 30 mi SE Tepic, 21°12'N,
104°33'W, 23.xi.1948 (E. S. Ross, CAS); 5 mi NW
Tepic, 21°32'N, 104°57'W, 13.V.1963 (W. J. Gertsch
& W. Ivie, AMNH). Oaxaca: 6 mi NE Mitla, 16°59'N,
96°21'W, 20.viii.l985 (J. Woolley & G. Zolnerowich,
ADC); Huajuapan, 17°48'N, 97°46'W, l.x.1946 (H.
Wagner, AMNH); Oaxaca, 17°3'N, 96°43'W,
17.vii.l955 (C. & R Vaurie, AMNH), 12.viii.l991 (W.
H. Piel & G. S. Bodner, MCZ); San Bait. Chichica-
pan, 16°45'N, 96°29'W, 4.viii.l991 (W. H. Piel & G.
S. Bodner, MCZ).
Metepeira minima Group
Female spiders in the M. minima group
(Metepeira petatlan, Metepeira uninima,
Metepeira pacifica, Metepeira jamaicensis)
have a thin scape with epigynal openings
that are shaped from longitudinal slits
(Figs. 273, 280, 281) to ovals that are lon-
ger than wide (Figs. 288, 295). In males,
the flagellae on the median apophysis are
set off on a distinctly narrower stalk (Figs.
271, 278, 286, 293).
35. Metepeira petatlan new species
Figures 271-277, 334; IVIap 10
Holotype. Male from 50 km southeast of Petatlan,
Guerrero, Mexico, 14.viii.l984, J. B. Woolley, in
Metepeira • Piel
81
MCZ. The specific name is a noun in apposition
after the locality.
Description. Female paratype from Pa-
panoa, Guerrero. Light region around pos-
terior eye row; carapace darkens posteri-
orly, then lightens under overhang of ab-
domen (Fig. 276). Slight rings on patella
and tibia. Femur I with three macrosetae
on anterior side, none on anteroventral
side. Dorsum of abdoinen with typical fo-
lium, except that white oak leaf pattern
narrower than most species and set on a
narrow, remarkably darker brownish dove-
tail (Fig. 276). Dark lateral band wraps
around abdomen and stretches up over an-
terior dorsal portion. Venter with two spots
on either side of spiracle and slight indi-
cation of a transverse bar. Wide, white lon-
gitudinal mark ends at the epigynal groove
(Fig. 277). Sternum with wide, white lon-
gitudinal mark v^dening anteriorly (Fig.
277). Ratio of eye diameters: posterior me-
dians and anterior medians 1.0, anterior
laterals 1.3, posterior laterals 1.2. Anterior
median eyes separated by 1.1 diameters,
posterior median eyes by 0.7, anterior me-
dian eyes separated from anterior laterals
by 1 diameter of anterior lateral eyes, lat-
eral eyes aliTiost touching. Total length 5
mm. Carapace 2.2 mm long, 1.7 wide.
First femur 2.3 mm, patella and tibia 2.5,
metatarsus 2, tarsus 0.9. Second patella
and tibia 2 mm, third 1.3, fourth 1.9.
Male holotype. Carapace uniform
brown with a lighter median streak just an-
terior to thoracic depression (Fig. 274).
Weakly ringed on tibia; distal half of fem-
ora dark. Feinur I with two macrosetae on
anterior side. Abdomen is a lighter version
of the female (Fig. 274). Venter with a
much reduced and shorter longitudinal
line, compared to the female (Fig. 275).
Sternum as in female. Ratio of eye diam-
eters: posterior medians and anterior me-
dians 1.0, anterior laterals 1.4, posterior
laterals 1.4. Anterior median eyes separat-
ed by 1.1 diameters, posterior median eyes
by 0.7, anterior median eyes separated
from anterior laterals by 1.1 diameters of
anterior lateral eyes, lateral eyes separated
by 0.2 their diameters. Total length 2 mm.
Carapace 1 mm long, 0.7 wide. First fe-
mur 1.2 inm, patella and tibia 1.0, meta-
tarsus 0.8, tarsus 0.5. Second patella and
tibia 0.9 mm, third 0.5, fourth 0.8.
Diagnosis. The female's dorsum can be
distinguished from other species by the
unusually dark brown color of the dovetail-
shaped mark (Fig. 276). Both this mark
and the white oak leaf pattern have nar-
rower margins that are more parallel and
not as wedge-shaped as they are in other
species (Fig. 276). The epigynum resem-
bles that of M. uninima by the narrow
scape and the parallel slitlike openings on
either side. These openings differ from
those of M. minima because they have
conspicuous sclerotized spheres just ante-
rior to the lateral edge of the slits (Fig.
273) but which are much farther away in
M. minima (Fig. 280). The male is unusu-
ally small and its palp is unique and easily
distinguished from other species. The lon-
ger flagellum on the median apophysis is
needlelike over its entire length, not grad-
ually tapering (compare Fig. 271 with Fig.
278). In addition, the long needlelike fla-
gelluni has a sharper elbow. At the elbow
it projects away from the palp for a short
distance and then strongly curves around
the palp (Fig. 271).
Natural History. The female paratype
was found at eye level on a hot, dry, wood-
ed hillside overlooking the Pacific Ocean
at an altitude of about 200 m. Her web
differed considerably from the usual Me-
tepeira web: instead of an elaborate barrier
web, it consisted of a much reduced Y-
shaped structure with a curled leaf in the
center. The leaf served as a retreat for the
spider and protected and hid three egg
sacs. The retreat was suspended very near
the hub, as opposed to farther away and
above it, as found in most other species.
Mature specimens have been collected in
August through October (Fig. 334).
Distribution. Western Mexico: Guerrero
and Sinaloa (Map 10).
82 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
Records Examined. MEXICO Guerrero: 32 mi SE
Petatlan, 17°14'N, 100°53'W, 14.viii.l984 (J. B. Wool-
ley, MCZ); Microondas Tamarindos, S. Papanoa,
17°17'32"N, 101°2'40"W, 18.x. 1994 (W. H. Piel,
MCZ). Sinaloa: 2 mi S Elota, 23°55'N, 106°48'W,
ll.Lx.1966 (Jean & Wilton Ivie, AMNH).
36. Metepeira minima Gertsch
Figures 278-285, 328; Map 15
Metepeira minima Gertsch, 1936: 10, fig. 31, 6 . Male
hoiotype from Edinburg, Texas, in the AMNH, ex-
amined. Roewer, 1942: 869. Chamberlin and Ivie,
1942: 67, fig. 174, 9. Levi, 1977: 206, fig. 70-73,
76-77, 9 ; 74-75, 3 . Bonnet, 1957: 2822. Brignoli,
1983: 275.
Description. Female from Celestun, Yu-
catan, Mexico. Carapace dark brown with
light region surrounding the eyes, some-
times extending posteriorly behind the lat-
eral eyes. Faint pair of darker feather-
shaped patches in center of carapace (Fig.
284). Legs same color as carapace; darker
on distal ends of articles. Femur I with
row of four macrosetae on anterior side;
rarely any on anteroventral side. Dorsum
of abdomen with usual Metepeira folium
(Fig. 284); venter wide, with long white
median line surrounded by faint, thin,
white U-shaped line (Fig. 285). Pair of
large white spots on either side of spiracle.
Sternum black with wide, white, uneven
median line (Fig. 285). Ratio of eye di-
ameters: posterior medians and anterior
medians 1.0, anterior laterals 1.2, posterior
laterals 1.1. Anterior median eyes separat-
ed by 1.2 diameters, posterior median eyes
by 0.9, anterior median eyes separated
from anterior laterals by 1.9 diameters of
anterior lateral eyes, lateral eyes separated
by 0.2 their diameters. Total length 6.5
mm. Carapace 2.7 mm long, 2 wide. First
femur 2.8 mm, patella and tibia 2.8, meta-
tarsus 2.5, tarsus 0.9. Second patella and
tibia 2.4 mm, tliird 1.5, fourth 2.
Male from Celestun, Yucatan, Mexico.
Carapace dark with light mark anterior to
thoracic furrow (Fig. 282). Femur I with
row of four macrosetae on anterior side;
row of two to five macrosetae on anter-
oventral side. Distal two-thirds same color
as carapace, proximal third white. Folium
of abdomen as in female, except posterior
half darker than anterior half (Fig. 282).
Venter of abdomen and sternum as in fe-
male, except median white line of sternum
often broken (Fig. 283). Ratio of eye di-
ameters: posterior medians and anterior
medians 1.1, anterior laterals 1.4, posterior
laterals 1.1. Anterior median eyes separat-
ed by 1.2 diameters, posterior median eyes
by 0.8, anterior median eyes separated
from anterior laterals by 1.5 diameters of
anterior lateral eyes, lateral eyes separated
by 0.1 their diameters. Total length 3 mm.
Carapace 1.5 mm long, 1.2 wide. First fe-
mur 2 mm, patella and tibia 1.9, metatar-
sus 1.8, tarsus 0.8. Second patella and tibia
1.6 mm, third 0.8, fourth 1.3.
Diagnosis. Unlike other members of the
M. niininia species group, the openings to
the epigynum of M. minima are narrow
slits around a scape of variable size (Figs.
280, 281). The openings of M. petatlan are
only slightly wider than M. minima, but
the internal darker sclerotized spheres in
M. minima sit much farther apart (com-
pare Fig. 280 with 273). Unlike M. pacifica
(Fig. 286) and M. jamaicensis (Fig. 293),
the bent embolus on M. minima tapers
strongly and is therefore not like a needle
(Fig. 278). Unlike M. petatlan, the longer
flagellum on M. minima tapers (Fig. 278)
and is not thin over its entire length (Fig.
271).
Variation. Average body length of 13 fe-
males examined 5.4 mm, range 3.7 to 6.5
mm. Average body length of 17 males ex-
amined 3.4 mm, range 2.5 to 4.5 mm. The
epigyna of females from the Yucatan pen-
insula differ noticeably from all others, but
the males hardly show any distinguishing
features. These differences can be seen in
the much wider scape and the greater sep-
aration between openings in a specimen
from the Yucatan (Fig. 281), compared to
a specimen from Tamaulipas (Fig. 280) or
one from Texas (Levi, 1977: 209, fig. 71).
In addition, the posterior view of the epi-
gynum from a Yucatan specimen shows
the lobes converging to form a V-shape
(Fig. 279), whereas females outside of the
Metepeira • Piel
83
278
minima
(36)
s 291
292
289 ^ 290
pacifica
(37)
298
297
jamaicensis
(38)
Figures 278-285. Metepeira minima Gertsch (sp, 36 [278,279,281-285] 20°56'N, 90°21'W; [280] 22°30'N, 99°4'W). 278, male
palpus, mesal. 279, epigynum, posterior. 280,281, epigynum, ventral. 282, male, dorsal. 283, male, ventral. 284, female, dorsal.
285, female, ventral.
Figures 286-292. Metepeira pacifica new species (sp. 37; 10°27'N, 85°9'W). 286, male palpus, mesal. 287, epigynum, posterior.
288, epigynum, ventral. 289, male, dorsal. 290, male, ventral. 291, female, dorsal. 292, female, ventral.
Figures 293-299. Metepeira jamaicensis Archer (sp. 38; 18°17'N, 76°48'W). 293, male palpus, mesal. 294, epigynum, posterior.
295, epigynum, ventral. 296, male, dorsal. 297, male, ventral. 298, female, dorsal. 299, female, ventral.
Scale bars: dorsum and venter figures 1 .0 mm.
84 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
Yucatan show largely parallel lobes (Levi,
1977: 209, fig. 71).
Natural History. Adults have been ob-
served on tree trunks and on bushes at and
above 150 cm. They can be found in shad-
ed areas, which is unusual for Mexican
Metepeira. Most mature specimens have
been collected in May through September
(Fig. 328). Elevations range from sea level
to just under 2,000 m.
Distribution. Southern Texas to Hon-
duras (Map 15).
Records Examined. HONDUFIAS Copdn: 14°50'N,
89°9'W, 7.iii.l939 (AMNH); Ruinas, 14°50'N,
89°9'W, 7.iii.l939 (AMNH). MEXICO Campeche:
Becan, 18°33'N, 89°30'W, 31.vii.l991 (W. H. Piel &
G. S. Bodner, MCZ); Campeche, 19°51'N, 90°32'W,
14.vii.l948 (C. J. Goodnight, AMNH); Champoton
beach, 19°21'N, 90°43'W, 22.vii.1991 (W. H. Piel &
G. S. Bodner, MCZ); Seybaplaya, 19°39'N, 90°40'W,
2.vlii.l949 (C. J. Goodnight, AMNH). Chiapas: 16 mi
W Cintalpa on rt 190. 16°36'N, 93°51'W, 15.vi.l982
(F. Coyle, MCZ). Nuevo Leon: 20 km E Montemo-
relos, Gamine Q. Rayones, 25°16'N, 99°41'W,
22.vi.1981 (L. Stange, FSCA); Los Cristales, 25°33'N,
100°12'W, 15.viii.l972 (A. F. Archer, AMNH); Villa
de Santiago, Hacienda Vista Hermosa, 25°25'N,
100°9'W, 19.vi.l940 (H. Hoogstraal, MCZ); Villa San-
tiago, Horsetail Falls, 25°25'N, 100°9'W, 19.vi.l940
(H. Hoogstraal, MCZ). San Luis Potosi: 20 km W Cd.
Valles, 21°58'N, 99°11'W, 18.iii.l972 (J. A. L. Cooke,
AMNH); Cd. Valles, 21°59'N, 99°1'W, 6.vii.l940 (R
Ran, MCZ), 19.vli.l956 (W. J. Gertsch & V. Roth,
AMNH), 15.vii.l959 (L. Steude, AMNH); Cd. Valles,
Hotel Covadonga, 21°59'N, 99°1'W (L. Steude,
AMNH); Huichichuyan, 21°25'N, 98°55'W, 19.V.1952
(M. Cazier, W. Gertsch, & R. Schrammel, AMNH);
Medina, 24°1'N, 100°24'W, 9.Lx.l956 (A. F. Archer,
AMNH); N section of San Luis Potosi, 22°13'N,
100°58'W, 8.ix.l956 (A. F. Archer, AMNH); Venado
Arroyo, 22°56'N, 101°5'W [?], 27.vii.1934 (MCZ). So-
nora: 8 mi W Alamos, 29°13'N, 110°10'W,
23.viii.1965 (W. J. Gertsch & R. Hastings, AMNH).
Tamaulipas: 60 km N Cd Valles, 22°30'N, 99°4'W,
10.viii.l991 (W. H. Piel & G. S. Bodner, MCZ); Cd.
Victoria, 23°44'N, 99°8'W, 17.V.1952 (M. Cazier, W.
Gertsch, & R. Schrammel, AMNH); Laredo road
near Cd. Victoria, 23°44'N, 99°8'W, 20.viii.l947 (C.
J. & M. Goodnight, AMNH); rlOl 26km S Tula,
22°49'N, 99°55'W, 8.Lx.l991 (W. H. Piel & G. S. Bod-
ner, MCZ); Sisal, 15 mi S Cd. Victoria, 23°38'N,
99°12'W, 22.vii.1966 (Jean & Wilton Ivie, AMNH).
Veracruz: 15 mi E Panuco, 22°10'N, 98°3'W,
29.xi.1941 (A. M. & L. I. Davis, AMNH); Plan del
Rio, 19°6'N, 96°6'W [?], 26.vii.1956 (W. J. Gertsch &
V. Roth, AMNH). Yucatan: 20 km E Valladolid,
20°41'N, 88°2'W, 26.vii.1991 (W. H. Piel & G. S.
Bodner, MCZ); 3 km S San Felipe, 21°32'N,
88°14'W, 25.vii.1991 (W. H. Piel & G. S. Bodner,
MCZ); 5 mi E Sisal salt flat, 21°9'N, 90°5'W, 9.i.l984
(V. & B. Roth, CAS); Balankanche Cave, 2 km E Chi-
chen Itza, 20°40'N, 88°33'W, 19.vii.l983 (W. Mad-
dison, MCZ); beach north of Celestun, 20°56'N,
90°21'W, 24.vii.1991 (W. H. Piel & G. S. Bodner,
MCZ); Chichen Itza, 20°40'N, 88°34'W (C. J. Good-
night, AMNH), 15.vii.l981 (C. Gold, CAS); Cordil-
leria Mayapan, 20°28'N, 89°11'W, 6.Lx.l952 (J. & D.
Pallister, AMNH); Uxmal, 20°22'N, 89°46'W,
7.ix.l970 (A. F. Archer, AMNH), 23.vii.1991 (W. H.
Piel & G. S. Bodner, MCZ). USA Texas: 29 mi S
Sarita, 26°47'N, 97°47'W, 14..xi.l958 (A. Brady,
MCZ); 1 mi S Pharr, 26°10'N, 98°11'W, 14.xi.l958
(A. Brady, MCZ); 1 mi S Pharr on U.S. HW 281,
26°10'N, 98°11'W, 14..xi.l95S (A. Brady, MCZ).
37. Metepeira pacifica new species
Figures 286-292, 302; Map 10
Holotijpe. Male from La Pacifica, Guanacaste, Costa
Rica, I.ii.l975-2.iii.l975, R. E. Coville, in MCZ.
The specific name is a noun in apposition after the
locality.
Description. Female paratype from La
Pacifica, Guanacaste, Costa Rica. Carapace
tan, lighter around eyes. Legs white. Fe-
mur I with row of three macrosetae on an-
terior side; none on anteroventral side.
Dorsum of abdomen white with faint, in-
distinct folium, darker distally (Fig. 291).
Gravid females often slightly marbled.
Venter wide, with long white median line
surrounded by faint, thin, white U-shaped
line on black. Pair of large white spots on
each side of spiracle (Fig. 292). Sternum
black with wide, white, uneven median
line (Fig. 292). Ratio of eye diameters:
posterior medians and anterior medians
1.0, anterior laterals 1.2, posterior laterals
1.2. Anterior median eyes separated by 1.2
diameters, posterior median eyes by 0.9,
anterior median eyes separated from an-
terior laterals by 1.8 diameters of anterior
lateral eyes, lateral eyes separated by 0.2
their diameters. Total length 4.5 mm. Car-
apace 2.2 mm long, 1.7 wide. First femur
2.3 mm, patella and tibia 2.4, metatarsus
1.9, tarsus 0.8. Second patella and tibia 2
mm, third 1.2, fourth 1.8.
Male holotype. Carapace, legs, abdo-
men as in female, though often darker
(Figs. 289, 290). Femur I with row of four
Metepeira • Piel
85
300 Dec
Nov
301
gressa, n = 26
Sep
Dec
Jan
Nov ^
<
z
'^Mar
V
-r*
W""
Aug^
\
Julyl
June
revillagigedo, n = 2
304 Dec
Nov
Apr Sep
galatheae, n = 304
tarapaca, n = 23
July
rectangula, n = 1 1
desenderi, n = 46
308 De':
Nov
calamuchita, n = 15
Nov
■\
\"^
OaA
1
Am
'"V
\y
V^
Aug
y^ ^
May
July
rjune
crassipes, n
May
July I June
nigriventris, n = 44
Aug NC^ ^ jP\ ^^y
July I June
compsa, n = 279
datona, n = 66
jamaicensis, n = 24
glomerabilis, n = 33
cajabamba, n = 1 1
Aug "V^ 1 Js/ May
July June
roraima, n = 11
maya, n = 1 3
Aug ^yl ■" \^ May
July June
inca, n = 29
Aug ^^ ^1 J^ May
July ' June
arizonica, n = 45
Figures 300-319. Circular liistograms depicting relative seasonal abundance of collecting events for mature spiders. 300-308.
Primarily collected during the northern hemisphere winter and spring seasons. 300, Metepeira gressa; 301, Metepeira revillagi-
gedo; 302, Metepeira pacifica; 303, Metepeira karkii; 304, Metepeira galatheae; 305, Metepeira tarapaca; 306, Metepeira rec-
tangula; 307, Metepeira desenderi; 308, Metepeira calamuchita. 309-314. Generally collected throughout. 309, Metepeira cras-
sipes; 310, Metepeira nigriventris; 311, Metepeira compsa; 312, Metepeira datona; 313, Metepeira jamaicensis; 314, Metepeira
glomerabilis. 315-319. Primarily collected during the Northern Hemisphere summer season. 315, Metepeira cajabamba; 316,
Metepeira roraima; 317, Metepeira maya; 318, Metepeira inca; 319, Metepeira arizonica.
86
Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
macrosetae on anterior side; two to three
on antero ventral side. Median white line
on sternum often broken (Fig. 290). Ratio
of eye diameters: posterior medians and
anterior medians 1.0, anterior laterals 1.6,
posterior laterals 1.6. Anterior median
eyes separated by 1.1 diameters, posterior
median eyes by 0.6, anterior median eyes
separated from anterior laterals by 1.4 di-
ameters of anterior lateral eyes, lateral
eyes separated by 0.3 their diameters. To-
tal length 3.4 mm. Carapace 1.7 mm long,
1.2 wide. First femur 2.2 mm, patella and
tibia 2, metatarsus 1.7, tarsus 0.8. Second
patella and tibia 1.6 mm, third 0.9, fourth
1.3.
Diagnosis. Unlike other members of the
M. minima species group, M. pacifica and
M. jamaicensis share very light pigmenta-
tion, and the embolus in both species is
needle-shaped (Figs. 286, 289, 291, 293,
296, 298). Unlike M. jamaicensis, the fla-
gellae on the median apophysis of M. pa-
cifica are set off on a short, wide stalk (Fig.
286) rather than a long, thin one (Fig.
293). A ventral view of the epigynum in
M. pacifica shows a ridge under the scape
that is more V-shaped (Fig. 288), com-
pared to a straighter line (Fig. 295).
Variation. Average body length of three
females examined 5.3 mm, range 4.5 to 5.8
mm. Average body length of five males ex-
amined 3 mm, range 2.8 to 3.4 mm. Most
specimens appear whitish, but the loss of
pigment is variable, especially among
males.
Natural History. Specimens have been
collected from the wasp nests of Trtjpar-
gilum nitidum, T. tenoctitlan, and T. hen-
soni. In Costa Rica, mature specimens
have been collected in November through
February; Honduras and Nicaragua in July
(Fig. 302). Altitudes range from 100 to
2,600 m.
Distribution. Costa Rica, Honduras, and
Nicaragua (Map 10).
Records Examined. COSTA RICA Guanacaste: 4
km NW Canas, La Pacifica, prey of Trypargilum ben-
soni, 10°27'N, 85°9'W, 29.1.1975 (R. E. Coville,
MCZ); 4 km NW Canas, La Pacifica, prey of Trypar-
gilum nitidum, 10°27'N, 85°9'W, I.ii.l975-2.iii.l975
(R. E. Coville, MCZ); 4 km NW Caiias, La Pacifica,
prey of Trypargilum tenoctitlan, 10°27'N, 85°9'W
l.ii.l975 (R. E. Coville, MCZ); 4 km NW Canas, La
Pacifica, wasp collected, 10°27'N, 85°9'W, 16.ii.l975
(R. E. Coville, MCZ); Bagaces, Palo Verde, 10°21'N,
85°21'W 19.1.1978 (W. Eberhard, MCZ); ca. Canas,
10°25'N, 85°7'W, 15.xi.l982 (W Eberhard, MCZ);
Canas, 10°25'N, 85°7'W 15.xl.l982 (W Eberhard,
MCZ). Puntarena.s: Elnca San Miento-Sialas, 10°9'N,
84°54'W, 5.11.1976 (Roth & Schroepfer, AMNH). San
Jose [?]: Santa Maria, 9°39'N, 83°58'W, 15.1.1930
(Dodge, MCZ). HONDURAS Tegucigalpa: Teguci-
galpa, 14°6'N, 87°13'W, l.vii.l948 (Clarke, AMNH).
NICARAGUA Managua: Laguna de Jiloa, SW Ma-
nagua, campsite, 12°13'N, 86°19'W 8.vil.l970 (S.
Riechert, SR).
38. Metepeira jamaicensis Archer
Figures 293-299, 313; Map 15
Metepeira jamaicensis Archer, 1958: 16, fig. 33, 9 .
Female holotype from Port Henderson, St. Cath-
erine Parish, Jamaica, in the AMNH, examined.
Metepeira rmnima: — Levi, 1977: 206, 208. BrignoH,
1983: 275. Erroneous synonymy.
Description. Female from Saint Mary's
Parish, Strawberry Fields near Robin s Bay
and Green Castle, Jamaica. Carapace dirty
brown, white around eyes, central w^hite
wedge (Fig. 298). Legs whitish yellow;
slight rings on legs II and III. Femur I
with row of three to four macrosetae on
anterior side; none or only a few very fine
setae on anteroventral side. Dorsal folium
lighter than in most species; fleur-de-lis
white on speckled light gray (Fig. 298).
Venter brownish gray with white margins.
Wide median white line with pair of large
white spots on either side of spiracle (Fig.
299). Sternum brownish black with wide,
white line widening anteriorly, sometimes
broken in center (Fig. 299). Ratio of eye
diameters: posterior medians and anterior
medians 1.0, anterior laterals 1.2, posterior
laterals 1.1. Anterior median eyes separat-
ed by 1.4 diameters, posterior median eyes
by 0.7, anterior median eyes separated
from anterior laterals by 2 diameters of an-
terior lateral eyes, lateral eyes separated by
0.2 their diameters. Total length 5.1 mm.
Carapace 2.1 mm long, 1.5 wide. First fe-
mur 2.1 mm, patella and tibia 2.2, meta-
Metepeira • Piel
87
Aug NC^ ^t \^ May
July I June
Ventura, n = 35
Aug N4/ 1 J\ May
July I June
grandiosa grandiosa, n = 4
22
Dec J
Jan
Nov ,
Feb
OctN
-^Mar
Sepi^
J*,
Aug
July
June
•'^^May
gosoga, n = 2
Aug 7^^ J\ May
July I June
incrassata, n = 33
Aug N/ W\ \; ^^"^
July I June
grandiosa alpina, n = 5
326
Dec
Jan
Nov
7^
Feb
Oct /^
^
Am^
Sep\
J^
Jap.
Aug
\
May
July
June
327
Dec
Jan
Nov
7^
Feb
Oct A
■V
"^Mar
Sepi^
^
/ Apr
Aug^
\
May
July
June
celestun, n = 8
vigilax, n = 10
28
Dec J
Jan
NOV/
\
Feb
Oct?^
f^
\ Mar
Sep\>
V
\ / ^^'
Aug^
/r
May
juijn
June
329
Sep
Dec
Nov J\
Jan
^\^Mar
^
J*"
AugNc^^
\
May
J^
\lune
30
Nov X
Dec
Jan
Feb
Oc.?^
^
-^Mar
Sep^
K
-^Apr
AugV
;4
May
July
June
minima, n = 39
uncata, n = 6
pimungan, n = 6
Comanche, n = 2
332
Dec
Jan
Nov
y^
^Feb
OoN
V
-A Mar
.pi^
^
J Apr
Aug
\
May
Julyl
June
333
Dec J
Jan
Nov
\
Feb
Oct /^
j^
•^A Mar
Sep\
\t
Ja.
Aug
YJ
May
July
Fjune
34
Nov X
Dec
Jan
Feb
o„/V
J^
'^A Mar
sS^
V
■\ / ^P'
Aug^
^
May
July
Ijune
335
Dec
Jan
Nov
\
^Feb
OctN
1^
-^Mar
'"U
|L
J Apr
Aug
^
/
May
jiiiTl
rTune
lacandon, n = 10
chilapae, n = 15
petatlan, n = 3
spinipes, n = 1 16
336
Dec J
Jan
Nov ,
Feb
Oct r^
•^A Mar
Sep y .
V4
^Apr
Aug
July
June
May
atascadero, n = 1 7
triangularis, n = 19
Figures 320-337. Circular histograms depicting relative seasonal abundance of collecting events for mature spiders. 320-331,
Primarily collected during the northern hemisphere summer season. 320, Metepeira ventura; 321, Metepeira grandiosa gradiosa;
322, Metepeira gosoga; 323, Metepeira ventura; 324, Metepeira olmec; 325, Metepeira grandiosa alpina; 326, Metepeira celes-
tun; 327, Metepeira vigilax; 328, Metepeira minima; 329, Metepeira uncata; 330, Metepeira pimungan; 331, Metepeira comanche.
332-337. Primarily collected during the northern hemisphere fall season. 332, Metepeira lacandon; 333, Metepeira ctiilapae;
334, Metepeira petatlan; 335, Metepeira spinipes; 336, Metepeira atascadero; 337, Metepeira triangularis.
88 Bulletin Museum of Comparative Zoology, Vol. 157, No.
tarsus 1.8, tarsus 0.8. Second patella and
tibia 1.9 mm, third 1.2, fourth 1.7.
Male from same locality as female. Male
carapace, dorsum, venter, sternum darker
and more contrasty version of female
(Figs. 296, 297). All legs ringed. Femur I
with row of three macrosetae on anterior
side; none on anteroventral side. Ratio of
eye diameters: posterior medians and an-
terior medians 0.9, anterior laterals 1.2,
posterior laterals 1.2. Anterior median
eyes separated by 1.3 diameters, posterior
median eyes by 0.9, anterior median eyes
separated from anterior laterals by 1.2 di-
ameters of anterior lateral eyes, lateral
eyes separated by 2.3 their diameters. To-
tal length 2.3 mm. Carapace 1.2 mm long,
0.9 wide. First femur 1.4 mm, patella and
tibia 1.3, metatarsus 1.0, tarsus 0.5. Sec-
ond patella and tibia 1.2 mm, third 0.6,
fourth 0.9.
Diagnosis. Unlike other members of the
M. minima species group, M. jamaicensis
and M. pacifica share very light pigmen-
tation and the embolus in both species is
needle-shaped (Figs. 286, 289, 291, 293,
296, 298). Unlike M. pacifica, the flagellae
on the median apophysis of M. jamaicensis
are set off on a long, thin stalk (Fig. 293)
rather than a short, wide one (Fig. 286). A
ventral view of the epigynum in M. jamai-
censis shows a ridge under the scape that
almost forms a straight line (Fig. 295),
compared to a V-shape (Fig. 288).
Variation. Average body length of elev-
en females examined 5 mm, range 4.2 to
6.1 mm. Average body length of four
males examined 2.4 mm, range 2.3 to 2.6
mm.
Natural History. Mature specimens
have been collected in July through March
(Fig. 313).
Distribution. Primarily in Jamaica and
Haiti (Map 15), near sea level.
Records Examined. BRITISH WEST INDIES
Grand Cayman Island: 19°20'N, 81°10'W, 15.ii.l960
(R. A. Lewin, MCZ). HAITI Departement de
L'Ouest: Port-au-Prince, 18°32'N, 72°20'W,
19.vii.l955 (A. F. Archer, AMNH), 20.vii.l955 (A. F.
Archer, AMNH). Dept. de L'Artibonite: Saint-Marc,
19°7'N, 72°42'W, 15,i.l913 (W. M. Mann, MCZ). JA-
MAICA Cornwall: Montego Bay, 18°28'N, 77°55'W,
l.iii.l984 (L. E. Schulten Jr, MCZ). Middlesex: 3 mi
E Old Harbor, 17°56'N, 77°10'W, 21.X.1957 (A. M.
Chickering, MCZ); Christiana, 18°10'N, 77°29'W,
13.xi.l957 (A. M. Chickering, MCZ), 15.vii.l960 (C.
& P Vaurie, AMNH), 17.vii.l960 (C. & P Vaurie,
AMNH); Strawberry Fields near Robin's Bay and
Green Casde, 18°17'N, 76°48'W, 23.iii.1972 (H. W.,
L. & F Levi, MCZ), 25.iii.1972 (H. W., L. & F. Levi,
MCZ), 26.iii.1972 (H. W., L. & F Levi, MCZ). Saint
Ann: Roaring River, 18°24'N, 77°9'W [?], 8.ii.l946
(B. Heineman, AMNH); Saint Ann's Bay, 18°26'N,
77°8"W, 20.xi.l959 (A. M. Nadler, AMNH). Saint
Catherine: E Green Harbour, S slope of Healdishire,
17°53'N, 76°51"W, 12.viii.l958 (A. F. Archer,
AMNH). St. Andrews: Ferry, 9/10 mi on Spanishtown
Road, 18°2'N, 76°53"W, 26.vii.1955 (A. F Archer,
AMNH). Surrey: Kingston, Mona Road, pasture,
17°59'N, 76°24"W, lO.x.1957 (A. M. Chickering,
MCZ); Roselle Falls, 24 mi E Kingston, 17°59'N,
76°24"W, 29.X.1957 (A. M. Chickering, MCZ); Saint
Andrew, 18°4'N, 76°45'W, 7.X.1957 (A. M. Chicker-
ing, MCZ). Trelawny: Falmouth, 18°30'N, 77°39"W,
20.vii.l960 (C. & P. Vaurie, AMNH). Westinorland:
Negril, 18°16'N, 78°21'W, 24.iii.1955 (A. M. Nadler,
AMNH); 'Whitehouse, 18°4'N, 77°58"W, 26.iii.1955
(A. M. Nadler, AMNH).
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INDEX
Valid taxon names are printed in italics. Page numbers in bold refer to illustrations; those in italics refer to
species descriptions.
acostai, Metepeira 63
Aculepeira 8
alpina, Metepeira 23, 24
Amazonepeira 8
Arachnidornyia 10
Araneus 8
arizonica, Metepeira 9, 19, 65, 66
atascadero, Metepeira 9, 16, 18, 67, 69
bani, Metepeira 63
biogeography 9, 10
cajabamba, Metepeira 9, 12, 17, 26, 31
calamuchita, Metepeira 14, 18, 39, 42
cancriforrnis, Gasteracantha 10
celestun, Metepeira 9, 10, 16, 19, 74, 75
cereicola, Metepeira 43
chilapae, Metepeira 9, 16, 19, 78, 79
chilapica, Metepeira 78
Comanche, Metepeira 14, 19, 60, 61
compsa group, Metepeira 47
compsa, Aranea 48
compsa, Metepeira 7, 9, 10, 11, 14, 17, 48, 49
crassipes, Metepeira 7, 9, 10, 16, 19, 77, 79
Cyrtophora 8, 9
dakota, Metepeira 23, 24
daytona, Metepeira 6, 7, 8, 9, 10, 12, 17, 20
desenderi, Metepeira 6, 10, 12, 17, 19, 21, 25
digital photography 4
dominicana, Metepeira 30
dorsal folium 6
douglasi, Metepeira 34
ensenada, Metepeira 71
epigynum 8
foxi group, Metepeira 19
foxi, Metepeira 8, 9, 10, 12, 17
galatheae, Araneus 43
galatheae, Epeira 43
galatheae, Metepeira 9, 14, 18, 43, 45
glomerabilis, Araneus 28
glomerabilis, Epeira 28
glomerabilis, Metazygia 28
glomerabilis, Metepeira 9, 12, 17, 28, 31
gosoga, Metepeira 14, 18, 59, 61
grandiosa alpina, Metepeira 9, 12, 17, 24, 25
grandiosa grandiosa, Metepeira 9, 10, 12, 17, 23, 25
grandiosa palustris, Metepeira 9, 12, 17
grandiosa, Metepeira 23
gressa, Epeira 54
gressa, Metazygia 54
gressa, Metepeira 9, 18, 54, 57
gressus, Araneus 54
grinnelli, Metepeira 34
habitats 9
inca, Metepeira 6, 14, 17, 18, 57, 58
incrassata group, Metepeira 55
incrassata, Metepeira 8, 9, 10, 11, 16, 19, 68, 69
inerma, Metepeira 20
jamaicensis, Metepeira 9, 17, 19, 83, 86
josepha, Metepeira 77
Kaira 6, 8
karkii, Araneus 46
karkii, Metepeira 9, 13, 18, 45, 46
koepckeonim, Araneus 6
92 Bulletin Museum of Comparative Zoology, Vol. 157, No. 1
labyrinthea grinnelli, Aranea 34
labyrinthea grinnelli, Epeira 34
labyiinthea grinnelli, Metepeira 34
labyrintliea group, Metepeira 33
labyrinthea, Epeira 21
labtjrinthea, Metepeira 7, 9, 10, 14, 17, 32, 34, 46, 48
labyrintheus, Araneus 48
lacandon, Metepeira 9, 14, 17, 35, 37
latigyna, Metepeira 48
lindae, Arachnidoijia 10
maija, Metepeira 9, 14, 18, 56, 57
measurements 5
Mecynogea 8, 9
median apophysis 8
Metepeira 5
minima group, Metepeira 80
minima, Metepeira 9, 10, 12, 16, 19, 81, 83, 86
nigriventris group, Metepeira 38
nigriventris, Araneus 38
nigriventris, Epeira 38
nigriventris, Metepeira 9, 10, 11, 12, 18, 38, 39
ocosingo, Mecynogea 10
olmec, Metepeira 9, 16, 19, 5.9, 61
pacifica, Metepeira 16, 19, 83, 84
palomara, Metepeira 23
palp 8
palustris, Metepeira 23
perezi, Metepeira 48
petatlan, Metepeira 16, 19, 79, 80
pimungan, Metepeira 8, 9, 16, 19, 62, 65
predation 10
rayorae, Arachnidomyia 10
rectangula, Epeira 32
rectangula, Metepeira 5, 6, 9, 12, 17, 32, 35
rectangulata, Metepeira 32
revillagigedo, Metepeira 16, 19, 73, 75
roraima, Metepeira 9, 12, 18, 49, 53
Salei, Epeira 68
salei, Metepeira 70
sallei, Aranea 70
sallei, Araneus 70
santa, Aranea 28
scitulus, Araneus 54
seditiosa, Epeira 54
seditiosa, Eustala 54
seditiosa, Metepeira 54
seditiosus, Araneus 54
Singa 6
species groups 10
spinipes, Araneus 34
spinipes, Metepeira 5, 6, 8, 9, 14, 17, 34, 35
suspended retreat 8
tarapaca, Metepeira 7, 9, 14, 18, 39, 40
triangularis, Metepeira 9, 15, 18, 63, 65
Trypargilum 10
uncata, Metepeira 9, 16, 19, 75, 76
uncatus, Araneus 76
vaurieomm, Metepeira 48
Ventura group, Metepeira 71
Ventura, Metepeira 10, 16, 19, 69, 71
vigilax group, Metepeira 26
vigilax, Araneus 30
vigilax, Epeira 29
vigilax, Metepeira 9, 10, 12, 17, 26, 30, 31
virginensis, Metepeira 48
web 8
Zygiella 6
OF THE
seum
(US ISSN 0027-4100)
Type Specimens of Recent Mammals
in the Museum of Comparative Zoology
K. M. HELGEN ANDT. L. McFADDEN
1V1CZ
LIBRARY
JUL 3 2001
HA^VARO
y N L V ii:^ R -^ ^"^ '
HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS, U.S.A.
VOLUME 157, NUMBER 2
8 JUNE 2001
(US ISSN 0027-4100)
PUBLICATIONS ISSUED
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Breviora 1952-
bulletin 1863-
Memoirs 1865-1938
JoHNSONiA, Department of Mollusks, 1941-1974
Occasional Papers on Mollusks, 1945-
SPECIAL PUBLICATIONS.
1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and
Evolution of Crustacea. 192 pp.
2. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere-
dinidea (Mollusca: Bivalvia). 265 pp.
3. Sprinkle, J-., 1973. Morphology and Evolution of Blastozoan Echino-
derms. 284 pp.
4. Eaton, R. J., 1974. A Flora of Concord from Thoreau's Time to the
Present Day. 236 pp.
5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology
and Evolutionary Biology: Essays in Honor of Ernest E. Williams.
725 pp.
6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp.
Other Publications.
Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine.
Reprinted 1964.
Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of
Insects. {Bulletin of the M. C. Z, Vol. 108.) Reprinted 1971.
Creighton, W. S., 1950. The Ants of North America. Reprinted 1966.
Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In-
ternational Symposium on Natural Mammalian Hibernation. {Bulletin
of the M. C. Z, Vol 124.)
Orinthological Gazetteers of the Neotropics (1975-).
Peter's Check-list of Birds of the World, vols. 1-16.
Proceedings of the New England Zoological Club 1899-1947. (Complete
sets only.)
Proceedings of the Boston Society of Natural History.
Price list and catalog of MCZ publications may be obtained from Publica-
tions Office, Museum of Comparative Zoology, Harvard University, Cambridge,
Massachusetts 02138, U.S.A.
This publication ha': been printed on acid-free permanent paper stock.
©The President and Fellows of Harvard College 2001.
TYPE SPECIMENS OF RECENT MAMMALS IN THE MUSEUM OF
COMPARATIVE ZOOLOGY
K. M. HELGEN AND T. L McFADDEN^
CONTENTS
Abstract - 94
Introduction 94
Authors of Type Descriptions 94
Definitions and Organization 96
Taxa Included in This Catalogue .._ 97
Abbreviations 98
Acknowledgments 98
Paralectotype Series 98
Accounts of Type Specimens 99
Order Didelphimoi"phia 99
Family Caluromyidae 99
Family Marmosidae 99
Family Didelphidae 100
Order Dasyuromoqohia 100
Family Dasyuridae 100
Order Peramelia 100
Family Peroiyctidae 100
Order Cingulata 101
Family Dasypodidae 101
Order Afrosoricida 101
Family Chiysochloridae 101
Family Tenrecidae 101
Order Rodentia 103
Family Aplodontidae 103
Family Sciuridae 103
Family Castoridae 109
Family Geomyidae 110
Family Heteromyidae 111
Family Dipodidae 112
Family Muridae 112
Subfamily AiAdcolinae 112
Subfamily Cricetinae 117
Subfamily Cricetomyinae 117
Subfamily Dendromurinae 117
Subfamily Gerbillinae 117
Subfamily Murinae 117
Subfamily Nesomyinae 120
Subfamily Otomyinae 120
Subfamily Sigmodontinae 121
Family Pedetidae 128
^ Mammal Department, Museum of Comparative
Zoology, Harvard University, Cambridge, Massachu-
setts 02138.
Family Myoxidae 129
Family Bathyergidae 129
Family Erithizontidae 130
Family Dasyproctidae 130
Family Agoutidae 130
Family Octodontidae 130
Family Echimyidae 131
Family Capromyidae 131
Order Lagomoqiha 133
Family Ochotonidae 133
Family Leporidae 133
Order Scandentia 134
Family Tupaiidae 134
Order Primates 135
Family Indridae 135
Family Daubentoniidae 135
Family Galagonidae 135
Family Cebidae 135
Family Hylobatidae 136
Family Hominidae 136
Order Lipotyphla 136
Family Nesophontidae 136
Family Solenodontidae 137
Family Soricidae 137
Family Talpidae 140
Order Chiroptera 140
Family Pteropodidae 140
Family Emballonuridae 142
Family Nycteridae 142
Family Rhinolophidae 142
Family Mormoopidae 143
Family Phyllostomidae 143
Family Molossidae 145
Family Vespertilionidae 145
Family Thyi^opteridae 147
Order Artiodactyla 147
Family Tayassuidae 147
Family Monodontidae 148
Family Phocoenidae 148
Family Cervidae 148
Family Bovidae 149
Order Carnivora 149
Family Canidae 149
Family Ursidae 151
Bull. Mus. Comp. ZooL, 157(2): 93-181, June, 2001 93
94 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Family Procyonidae 151
Family Mustelidae 152
Family Mephitidae 154
Family Viverridae 155
Family Heipestidae 155
Family Felidae 155
Order Cimolesta 156
Family Manidae 156
References 157
Index 171
Abstract. The Mammal Department at the Mu-
seum of Comparative Zoology houses name-bearing
types of 342 species-group taxa of Recent inammals.
The type collection consists of 327 holotypes, 2 lec-
totypes, 4 complete syntype series, and 9 partial syn-
type series. This catalogue notes information on the
type locality, collector, date of collection, present con-
dition, original publication, and synonyms for all
name-bearing types in the Mammal Department.
Comments on the taxonomic and historical impor-
tance of many type specimens are included. Lecto-
types for a number of taxa are designated for pur-
poses of taxonomic consistency.
INTRODUCTION
"Some of them are more or less historic
specimens," wrote the great mainiTialogist
Glover M. Allen in 1931, "whose location
has undoubtedly in many cases been lost
sight of, so that it may be of value to make
the present record" (1931: 230). Allen,
then Curator of Mammals at the Museum
of Comparative Zoology at Harvard Uni-
versity, was referring to the type speci-
mens of inammals in the museum, the
subject of a listing he published that year.
Allen's words resound with even greater
truth today than when they were written
70 years ago. Since that time, the inuse-
um s holdings of mainmal type specimens
have grown considerably. Furthermore,
type specimens of ixiamiTials housed in the
MCZ have sometimes been credited to the
collections of other institutions or consid-
ered to no longer exist, making the need
for a new catalogue obvious.
Type specimens are biological reference
points that lend objectivity to taxonomy
and are thus of critical importance in sys-
teinatic investigations; accordingly, "eveiy
institution in which name-bearing types
are deposited should publish lists of name-
bearing types in its possession or custody"
(International Commission on Zoological
NoiTienclature, 1999: 79). Our duty in this
regard is thus long overdue, and this cat-
alogue should sei've to fulfill that obliga-
tion. Without doubt, it will alleviate con-
fusion in the scientific coiximunity as to the
w^hereabouts of certain specimens, thought
to be lost, especially those type speciinens
that were acquired from Guillaume Gran-
didier. In addition, it should bring to light
information, fonnerly unavailable, on this
museum s veiy notable holdings of name-
bearing types.
All taxonoinic judgments in this work,
including new name combinations and lec-
totype designations, reflect the decision of
the first author (KMH) alone and should
be cited accordingly.
AUTHORS OF TYPE DESCRIPTIONS
Four maminalogists authored the over-
whelming majority of names based on type
specimens in the MCZ: Glover Allen, Out-
ram Bangs, Guillaume Grandidier, and
Barbara Lawrence. Following are brief bi-
ographies of these four outstanding mam-
malogists.
Glover M. Allen (1879-1942) |
Curator of Mammals at the MCZ from
1924 until his death in 1942, Glover Allen
began his work in the Mammal Depart-
ment in 1907. He was known as a careful,
dependable researcher and an outstand-
ing, patient teacher. His goal in all en-
deavors was to increase the sum total of
knowledge about the mammals of the
world. Although much of his career was
spent in the museum studying, as he put
it, "the dried remains of animals" (Barbour
et al, 1943: 300), he also traveled widely
throughout the world collecting specimens
as well as observing and learning about liv-
ing animals. He once commented that the
actual knowledge of living creatures could
all too often be summed up by saying
"when we found it, it ran like hell, where-
upon we shot it!" (Barbour et al., 1943:
300). He wrote prolifically; his bibliogra-
phy of publications is 81 pages long, the
Type Specimens of Recent Mammals • Helgen and McFadden 95
first of which he pubHshed when he was
only 11 years old (Lawrence 1947a: 1).
Holotypes for 96 taxa, which Allen de-
scribed alone or with colleagues, are de-
posited in the MCZ.
Outram Bangs (1863-1932)
Outram Bangs, Curator of Mammals
from 1899 to 1924, "was one of those for-
tunate mortals, born with a love of nature
and the outdoors which rule their entire
lives" (Peters 1933: 265). Bangs authored
135 MCZ mammal names (only one co-
authored!). He and brother Edward began
collecting as boys using slingshots and
horsehair nooses. He was an early ecolo-
gist, saving Microtus breweri by killing the
feral cat population on Massachusetts'
Muskeget Island and restocking the mouse
from a tiny islet across the channel. Al-
though his greatest passion was the natural
history of birds, he was also fascinated with
mammals and served as curator of both
departments in the MCZ. He decided to
systematically collect the mammals of east-
ern North America in about 1890. He be-
gan by trapping in New England and later
made trips to the southeastern United
States and Canada. Other collectors assist-
ed Bangs, expanding the collection area to
western North America and south to Cen-
tral America. His precision and organiza-
tion were legendaiy, and today his mam-
mal collection of more than 10,000 speci-
mens, donated to the Museum in 1899, re-
mains one of the best curated and most
informative in the department. A complete
list of Bangs' scientific publications was
compiled by Porter (1943).
Guillaume Grandidier (1873-1957)
French explorer and scientist Guillaume
Grandidier, son of naturalist Alfred Gran-
didier, authored and coauthored descrip-
tions of 13 taxa whose types were donated
to the MCZ in 1947, along with his exten-
sive personal collection of Malagasy mam-
mals. The collection was purchased by
Robert Barbour and donated to the MCZ
in honor of his brother, MCZ director
Thomas Barbour. Between 1898 and 1902,
Grandidier explored the center and south-
ern portions of Madagascar, collecting
specimens and describing the geography
of the area. Through his writings he
brought the unique fauna of this remote
region to the attention of the scientific
world. He was known for his devotion to
the careful acquisition of knowledge,
whether it was geographical, historical, or
scientific. His work was honored by both
the scientific community and the French
government (Chapus, 1953).
Barbara Lawrence (1909-97)
After graduating from Vassar College in
1931, Barbara Lawrence became a volun-
teer at the MCZ. She was encouraged by
Dr. Glover Allen to do her own research,
and in the late 1930s she made field trips
to the Philippines and Sumatra to collect
mammals. In 1952 she was appointed Cu-
rator of Mammals, a position she held un-
til her retirement in 1976. Her areas of
scientific interest were many, ranging from
echolocation in w^hales to zooarchaeology,
as well as the more traditional mammalog-
ical pursuit of taxonomy. "She once wrote
'. . . to know and love a bit of the world
so well that you can give it to someone else
... is a rare talent' " (Rutzmoser, 1999:
1049), certainly a talent Barbara Lawrence
had in abundance. She authored or coau-
thored 20 MCZ names.
The specimens described by these au-
thors and others are the result of world-
wide collecting by numerous expeditions
and individuals, listed in the accounts that
follow. A handful of type specimens in the
MCZ were formerly in the collection of
the National Museum of Natural Histoiy
in Washington, D.C. According to Glover
Allen, these were obtained via exchanges
"at a time when 'duplicates' were more
freely disposed of, [and would] prove to be
cotypes or even actual types" (1931: 229).
The collections of the Boston Society of
Natural Histoiy, once contained in the
Boston Museum of Science, were trans-
96
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
ferred to the Museum of Comparative Zo-
ology over the course of the 20th century.
Several types are among these specimens,
which consist primarily of mammals from
the New England region.
DEFINITIONS AND ORGANIZATION
Type Categories
Type specimens are categorized as one
of the following:
Holotype. The single specimen desig-
nated as name-bearer in the original pub-
lished description of the species-group tax-
on (Article 73.1, International Commission
on Zoological Nomenclature, 1999: 79).
Syntype. One of multiple specimens on
which a species-group name is equally
based, when no holotype is specified in the
original description and no subsequent
designation of a lectotype has been pub-
lished (Article 73.2, International Com-
mission on Zoological Nomenclature,
1999: 81).
Lectotype. One of multiple specimens
upon which a species-group name is orig-
inally based, designated in a publication
subsequent to the original description to
become the unique name-bearer (Article
74, International Commission on Zoologi-
cal Nomenclature, 1999: 82).
Neotype. A specimen chosen as the
name-bearer of a species-group taxon, "if
no holotype, lectotype, syntype, or prior
neotype is believed to exist" (Article 75,
International Commission on Zoological
Nomenclature, 1999: 84). There are no
neotypes in the MCZ mainmal collection.
Paratype. A specimen other than the
holotype (if designated originally) that is
mentioned in the original description of a
species-group taxon. Paratypes of species-
group taxa whose holotypes are housed in
other institutions are not mentioned in this
catalogue, although inany such specimens
exist in this museum.
Paralectotype. If no holotype is desig-
nated, a specimen other than the lectotype
(if designated subsequently) that is men-
tioned in the original description of a spe-
cies-group taxon. Paralectotypes of spe-
cies-group taxa whose lectotypes are
housed in other institutions are discussed
below, before the accounts for name-bear-
ing types.
Locality
This categoiy includes country, second-
level geopolitical division (state, depart-
ment, territory, province, or district), col-
lection site, and altitude where available.
The geographic name given for a type lo-
cality is that found in the original publi-
cation. When the description of the col-
lection locality does not include current
geopolitical divisions, that information is
provided in parentheses. If an original
name is no longer used, an equals sign ( = )
is included within the parentheses to des-
ignate an equivalent modern name. Where
altitude was originally given in feet, it has
been converted to meters and included in
parentheses.
Sources used for the current names are
from the most current available Gazetteers
of the United States Board on Geographic
Names, the 10th comprehensive edition of
the Times Atlas of the World, the Colum-
bia Gazetteer of the World, and the Or-
nithological Gazetteers of South America.
Account Organization
The type locality, collector, date of col-
lection, and present condition of each
specimen are noted. The publication of
the original description is cited for each
specimen. Many names have changed in
rank or synonymy since their origin; in
these cases, the name by which a taxon is
known today is noted, with a citation of the
publication in which that name combina-
tion was first employed for that taxon.
Comments are offered for most entries to
provide additional information or to dispel
potential sources of confusion.
The format of this catalogue is largely
borrowed from the most recent type cat-
alogue of inammals in the American Mu-
seum of Natural History (Lawrence,
1993). For systematic consistency, the tax-
Type Specimens of Recent Mammals • Helgen and McFadden 97
onomic judgments at the species-level by
the authors of the chapters in Mammal
Species of the World (Wilson and Reeder,
1993) are largely adhered to. However, in
many cases alternate views are explored,
and subsequent work by other authors is
noted. The sequence of mammalian orders
presented here is as follows: Didelphimor-
phia, Dasyuromorphia, Peramelia, Cingu-
lata, Afrosoricida, Rodentia, Lagomoi-pha,
Scandentia, Primates, Lipotyphla, Chirop-
tera, Artiodactyla, Carnivora, and Cimoles-
ta. This sequence represents the ongoing
understanding of higher mainmalian rela-
tionships being produced by research in
molecular phylogenetics (Waddell et al.,
1999) as well as paleontological studies
(McKenna and Bell, 1997). A number of
these names are rather nontraditional; use
of the names Cingulata (for armadillos)
and Cimolesta (for pangolins) at ordinal
rank follows McKenna and Bell (1997). Af-
rosoricida is used for an order including
tenrecs and golden-moles (following Stan-
hope et al, 1998: 9971-9972). We use the
ordinal name Lipotyphla in a restricted
sense to refer to the Recent families Ne-
sophontidae, Solenodontidae, Soricidae,
and Talpidae (others have used the term
"Eulipotyphla," e.g., Waddell et al, 1999).
Cetaceans are included here within the or-
der Artiodactyla (an assemblage often re-
ferred to as "Cetartiodactyla" in recent lit-
erature; see Graur et al., 1997).
To avoid unnecessaiy complexity, or-
ders, families, and genera are the only
ranks above the level of species that are
listed, except for the large family Muridae,
for which subfamilial distinctions are pro-
vided, in alphabetical order. Within each
order, the sequence of families generally
follows Simpson (1945), but Wilson and
Reeder's (1993) order of rodent families
and Simmons' (1998: 12) arrangement of
the bats are observed. Within each genus,
taxa are presented in alphabetical order by
original name.
The format of this catalogue is as fol-
lows, with all the following information
provided when possible:
Original binomen. Name of describer, date
of description.
Citation of original publication.
= Presently used name, if different from
original. Citation of publication in which
this name combination was initially used
for this taxon.
Type Category. Number of specimen.^
Preparation of specimen (skin, skull, alco-
hol, etc.), age and sex.
Locality. Type locality. Date of collec-
tion.
Collector. Name of collector. Original
number of specimen.
Condition. Current condition of the
type material.
Type Series. Any paratypes, paralecto-
types, or additional syntypes in existence
are mentioned, with their preparation, sex,
and age.
Comments. Additional comments re-
garding the systematic status or the histoiy
of the specimen.
TAXA INCLUDED IN THIS CATALOGUE
Unlike Allen's original type catalogue, fos-
sil mammals are not considered here, but
type specimens of Recent mammals known
only from subfossil remains are discussed.
Several type specimens of Recent mam-
mals that were included in Allen's cata-
logue are not considered here as name-
bearing types. A number of syntype series
in the MCZ have since Allen's time been
rendered paralectotypes by the designa-
tion of a lectotype preserved in another
institution; these are discussed in the sec-
tion below on paralectotypes. Additionally,
MCZ 14929, listed as a "cotype" of Nyc-
teris revoili Robin, 1881 by G. M. Allen
(1931: 235), is a paratype rather than a
syntype and is not considered here.
- Mammal specimens in the MCZ bear any of three
kinds of numbers. A number preceded by "MCZ" can
be found in the general collection in the Mammal
Department. A number preceded by a "B" is part of
the collection of E. A. and O. Bangs, also housed in
the Mammal Department. A number preceded by
"VP" designates that the specimen is stored in the
Vertebrate Paleontology Department.
98
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
One other enigmatic specimen in the col-
lection deseives some mention. When it was
received by die Mammal Department, Guil-
laume Grandidiers personal collection con-
tained specimens marked as types for 16
taxa, described by either Grandidier or M.
L. Lavauden. Fifteen of these holotypes are
discussed witliin this catalogue in separate
accounts. The remaining specimen, a fioiit
bat, MCZ 45073, is marked exactly like the
odier type specimens, and both its sldn and
its skull tags bear the name "Eidolon saka-
lava nov. spec, G. Grandidier" in die script-
ed handwriting typical of Grandidiers spec-
imens. The locality given for the specimen
is Ankavandra, in west-central Madagascar.
Though this specimen has been curated as
a type specimen, I (KMH) can find no pub-
lished description of diis taxon or any ref-
erence to diis name in the literature. This
name is dierefore invalid, barring a discov-
eiy in the future diat it was indeed pub-
lished and has since been overlooked. What-
ever die published status of the name, it ap-
pears to me diat diis specimen should be
considered as a young specimen of Eidolon
dupreanum rather dian a distinct species,
and discussion of this mysterious binomial
here should in no way be constiaied as a
formal description of a new taxon.
ABBREVIATIONS
Abbreviations are used in the text to
designate the following institutions.
AMNH American Museum of Natural
Histoiy, New York
BMNH Natural Histoiy Museum, Lon-
don
BSNH Boston Society of Natural His-
tory, Boston
FMNH Field Museum of Natural His-
tory, Chicago
MNHN Museum National d'Histoire
Naturelle, Paris
MVZ Museum of Vertebrate Zoology,
Berkeley, California
RMNH Rijksmuseum van Natuurlijke
Historic, Leiden
USNM
YPM
National Museum of Natural
Histoiy Washington, D.C.
Yale Peabody Museum of Nat-
ural History, New Haven, Con-
necticut
ACKNOWLEDGMENTS
We would like to thank the faculty and
staff of the Mammal Department at the
MCZ — A. W. Crompton, Andrew Biewe-
iier, Judith Chupasko, Jane Harrison, and
especially Maria Rutzmoser for access to
the type specimens. Maiy Sears, Timothy
McNeece, and Ronnie Broadfoot at the
Ernst Mayr Libraiy, HaiA^ard University,
were extremely helpful in pinning down
difficult references. We also thank Alison
Pine for allowing us to make use of the
MCZ Ornithology Department locality
references; Carolyn Kirdahy of the Boston
Museum of Science, w^lio granted us ac-
cess to the archives of the Boston Society
of Natural History; and the many individ-
uals associated with other museums who
assisted by answering any questions that
arose. Finally, we are thankful for the sug-
gestions we received from two anonymous
reviewers, which helped us improve this
catalogue.
PARALECTOTYPE SERIES
Hesperomys eremicus Baird, 1858 =Peromyscus er-
emicus eremicus (Baird, 1858). Lectotype, USNM
2575, designated by Osgood (1909: 241). MCZ
4310 and 5273 are paralectotypes.
Neotoma fuscipes Baird, 1858 =Neotoinafuscipesfus-
cipes Baird, 1858. Lectotype, USNM 22026, des-
ignated by Lyon and Osgood (1909: 99). MCZ 4336
and 5264 are paralectotypes.
Mus bairdii Hoy and Kennicott, 1857 =Peromyscus
maniculatus bairdii (Hoy and Kennicott, 1857).
Lectotype, number 750 in the Collection of the
Academy of Natural Sciences of Philadelphia, des-
ignated by Osgood (1909: 80). MCZ 8073 is a par-
alectotype.
Pteropus lanigera [sic] H. Allen, 1890 =Pteropus in-
stdaris Hombron and Jacquinot, 1842. The cor-
rected spelling of the original name is Pteropus lan-
iger (see Andersen 1912: 297). H. Allen based the
description of Pteropus laniger on two syntypes,
USNM 19066 (skin)/37815 (skull) and MCZ 7023,
a skin. Andersen (1912: 297-298) was unaware of
Type Specimens of Recent Mammals • Helaen and McFadden
99
the whereabouts of this latter specimen and based
his evaluation of the systematic status of P. laniger
solely on the USNM specimen in his taxonomic
review of the Megachiroptera. Because it is rep-
resented by both a skin and a skull in good con-
dition and because it has been used in past taxo-
nomic treatments, USNM 19066/37815 is hereby
designated as the lectotype of Pteropus laniger to
ensure consistency between past and future taxo-
nomic treatments of this name. MCZ 7023 is thus
a paralectotype. The type locality of laniger is the
Caroline Islands, as emended by Andersen (1912:
298), not Samoa, as originally described.
Sciunis castanotus Baird, 1855 and Sciurus castan-
onotus Baird, 1858 =Scmnis aherti aherti Wood-
house, 1853. Baird (1858: 266) noted that the name
castanotus, used in his original description of this
taxon, was a misprint for castanonotus. Lectotype,
USNM 121/1107, designated by Lyon and Osgood
(1909: 183). Though not explicitly stated by Lyon
and Osgood, this specimen should serve as a lec-
totype for both names (Sciunis castanotus Baird,
1855 and Sciunis castanonotus Baird, 1858), for
ttixonomic consistency. MCZ 4692 is a paralecto-
type.
Spennophiliis obsoletus Kennicott, 1863 =Spenno-
philus spilosoma obsoletus Kennicott, 1863. Lec-
totyjoe, USNM 3222/27998, designated by A. H.
Howell (1938: 130). See G. M, Allen (1931: 252)
for a list of paralectotypes (then considered synty-
pes) in the MCZ.
Spennophiliis parnji van kodiacensis J. A. Allen, 1874
= Spennophiliis parnji kodiacensis J. A. Allen,
1874. Lectotype, USNM 9242/38543, designated
by A. H. Howell (1938: 103). See G. M. Allen
(1931: 252) for a list of paralectotypes (then con-
sidered syntypes) in the MCZ.
Spennophiliis tridecemlineatus van pallidus ]. A. Al-
len, 1874 = Spennophiliis tridecemlineatus pallidus
J. A. Allen, 1874. Lectotype, USNM 16237, des-
ignated by A. H. Howell (1938: 112). See G. M.
Allen (1931: 253) for a Ust of paralectotypes (then
considered syntypes) in the MCZ.
Tamias quadrivittatus var. pallidus J. A. Allen, 1874
= Tamias minimus pallidus J. A. Allen, 1874. Lec-
totype, USNM 11656/38311, designated by Gary
(1906: 88). G. M. Allen (1931: 255) provided a hst
of paralectotypes.
ACCOUNTS OF NAME-BEARING TYPE
SPECIMENS
Order DIDELPHIMORPHIA Gill, 1872
Family CALUROMYIDAE Kirsch and Reig,
1977
Genus CALUROMYSJ. A. Allen, 1900
Philander cicur Bangs, 1 898k
Proc. Biol. Soc. Washington, 12: 161, 10
August.
= Caluromys lanatus cicur {Bangs, 1898).
See Cabrera (1958: 2).
Holotype. B8114. Skin and skull. Adult female.
Locality. Colombia: (Magdalena), Santa Marta
Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 27
March 1898.
Collector. W. W. Brown, Jr. Original number 123.
Condition. Skin and skull complete. Mandible dis-
articulated.
Tijpe Series. 3 paratypes; B8036, skin and skull,
adult male; B8115, skin and skull, adult male;
B8116, skin and skull, adult male.
Family MARMOSIDAE Hershkovitz, 1992
Genus MARMOSA Gray, 1821
Marmosa robinsoni Bangs, 18981
Proc. Biol. Soc. Washington, 12: 95, 30
April.
= Marmosa robinsoni robinsoni Bangs,
1898. See Cabrera (1958: 24).
Holotype. B7749. Skin and skull. Adult male.
Locality. Venezuela: (Nueva Esparta), Margarita
Island. 12 July 1895.
Collector. W. Robinson. Original number 506.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 2 paratypes, both in the USNM;
USNM 63209, skin and skull, adult male; USNM
63210, skin and skull, adult female.
Comments. M. robinsoni was considered a valid
species by Gardner (1993: 18) and Nowak (1999:
21).
IVIarmosa mitis Bangs, 1 898k
Proc. Biol. Soc. Washington, 12: 162, 10
August.
= Marmosa robinsoni robinsoni Bangs,
1898. See Cabrera (1958: 24).
Holotype. B8123. Skin and skull. Adult male.
Locality. Colombia: (Magdalena), Santa Marta
Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 25
March 1898.
Collector. W. W. Brown, Jr. Original number 91.
Condition. Skin and skull complete. Mandible dis-
articulated.
Tijpe Series. 26 paratypes; B8117-B8122, B8124-
B8143; all represented by skin and skull, 15 fe-
males and 11 males. 5 paratypes are no longer in
the MCZ (B8118 is at Wellesley College, B8124
and B8141 are at FMNH, and B8136 and B8138
are at USNM).
100
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Marmosa fulviventer Bangs, 1901b
Amer. Nat., 35: 632, 22 August.
= Marmosa robinsoni fulviventer Bangs,
1901. See Handley (1966: 775).
Holotype. B8435. Sldn and skull. Adult male.
Locality. Panama: (Panama), Gulf of Panama, San
Miguel Island. 28 April 1900.
Collector. W. W. Brown, Jr. Original number 123.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; B8436, skin and skull, fe-
male (exchanged to FMNH in 1931); B8437, skin
and skull, female; B8438, skin and skull, male.
Comments. M. r. fulviventer was retained as a valid
subspecies by Hall (1981: 14) and O'Connell (1983:
1).
Family DIDELPHIDAE Gray, 1821
Genus D/DELPH/S Linnaeus, 1758
Didelphis marsupialis particeps Goldman,
1917
Proc. Biol. Soc. Washington, 30: 107, 23
May.
Holotype. B8439. Skin and skull. Adult male.
Locality. Panama: (Panama), Gulf of Panama, San
Miguel Island. 8 May 1900.
Collector. W. W. Brown, Jr. Original number 165.
Condition. Skin and skull complete.
Type Series. 1 paratype; B8440, skin and skull,
adult female.
Comments. Retained as a valid subspecies by Hall
(1981: 4).
Didelphis virginiana pigra Bangs, 1 898b
Proc. Boston Soc. Nat. Hist., 28: 172, 15
March.
Holotype. B3500. Sldn and skull. Adult female.
Locality. (United States): Florida, Brevard Gounty,
Oak Lodge, East Peninsula opposite Micco. 31 Jan-
uaiy 1895.
Collector. O. Bangs.
Condition. Skin and skull complete.
Type Series. 11 paratypes; all represented by skin
and skull, most still in the MGZ.
Comments. Retained as a valid subspecies by Hall
(1981: 5) and McManus (1974: 1). The type de-
scription lists 31 January 1896 as the date of col-
lection, but the date is written as "January 31,
1895" on the original specimen label and in Bangs"
accession catalogue.
Order DASYUROIVIORPHIA Gill, 1872
Family DASYURIDAE Goldfuss, 1820
Genus ANTECHINUS Macleay, 1841
Antecfiinus may eh misim Tate, 1947
Bull. Amer. Mus. Nat. Hist., 88: 130, 20
February.
= Antecfiinus naso misim Tate, 1947. See
Laurie and Hill (1954: 7).
Holotype MGZ 29924. Skin and skull. Adult male.
Locality. Papua New Guinea: Morobe Province,
Mount Misim ( = Missim), 5,850 ft (1,784 m). 14
April 1933.
Collector H. Stevens.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; MGZ 29923, sldn and
skull, adult female.
Comments. Tate, in the original description, erro-
neously listed 24 April 1933 as the date of collec-
tion. A. n. misim was retained as a valid subspecies
by Flanneiy (1995a: 80), who also noted that the
New Guinean species assigned to the genus Ante-
chinus are not closely related to the Australian spe-
cies of that genus and will be reassigned at the
generic level pending a full taxonomic revision of
the group.
Genus IVIYOICTIS Gray, 1858
Myoictis melas wavicus Tate, 1 947
Bull. Amer. Mus. Nat. Hist., 88: 140, 20
February.
= Myoictis melas wallacei Gray, 1858. See
Flanneiy (1990: 56).
Holotijpe. MGZ 28082. Skin and skull. Adult male.
Locality. Papua New Guinea (S. Morobe), Wau,
3,800 ft (1,159 m). 27 March 1932.
Collector H. Stevens. Original number 1.
Condition. Sldn and skull complete.
Type Series. Holot}'pe only.
Comments. Stevens' original field label bears the
following lament — "To my intense anguish I failed
to retrieve the female, shot on recumbent, decayed
log in undergrowth."
Order PERAMELIA Ameghino, 1889
Family PERORYCTIDAE Groves and
Flannery, 1990
Genus ECHYMIPERA Lesson, 1842
Suillomeles hispida G. M. Allen and
Barbour, 1909
Proc. New England Zool. Club, 4: 44, 12
July.
= Echymipera kalubu kalubu (Fischer,
1829). See Laurie and Hill (1954: 11).
Holotype. MGZ 7006. Skin and skull. Adult.
Type Specimens of Recent Mammals • Helaen and McFadden
101
Localitij. (Indonesia): Dutch New Guinea ( = Ii-ian
Jaya), Doreh Bay, Manokwari, "not far from the
foot of Mt. Arfak." 23 Februaiy 1907.
Collector. T. Barbour.
Condition. Skin complete. Skull partial (occipital
region and two posterior upper molars on each side
missing).
Type Serie.s. Holotype only.
Comments. S. hispida is the type species of Suil-
lomeles G. M. Allen and Barbour, 1909.
Echymipera rufescens austral is Tate, 1948
Bull. Amer. Mus. Nat. Hist., 92: 334, 25
November.
Holotype. MCZ 29214. Skin and skull. Adult male.
Locality. Australia: Queensland, Cape York, near
Coen, east slope of Mcllwraith Ranges. Rocky Riv-
er, "Rocky Scrub." 20 June 1932.
Collector. R J. Darlington, Jr., Hanard Australian
Expedition. Original number 209.
Condition. Skin complete, but tip of tail worn.
Skull complete.
Type Series. Holotype only.
Comments. Tate (1952: 582) reported that the ex-
act type locality is "in the dense rain forests of the
upper Nesbit River on the east slopes of the
Mcllwraith Range." Retained as a valid subspecies
by Flanneiy (1995a: 111). Darlington noted in his
field notebook, regarding this specimen, "Animal
about the fattest I have skinned — I shall dream of
it!"
Order CINGULATA llliger, 1911
Family DASYPODIDAE Gray, 1821
Genus D/ASVPL/S Linnaeus, 1758
Dasypus novemcinctus hoplites G. M.
Allen, 1911a
Bull. Mus. Comp. Zool., 54: 195, July.
Holotype. MCZ 8116. Skin, skull, and postcranial
skeleton. Adult female.
Locality. Grenada: hills back of Gouyave. 7 Sep-
tember 1910.
Collector G. M. Allen. Original number 26.
Condition. Skin, skull, and postcranial skeleton
complete.
Type Series. 2 paratypes; MCZ 8117, skin and skull,
adult male; MCZ 8118, skin and skull, adult male.
Comments. Retained as a valid subspecies by Hall
(1981: 283) and McBee and Baker (1982: 1).
Order AFROSORICIDA Stanhope et a!.,
1998
Family CHRYSOCHLORIDAE Gray, 1825
Genus CHRYSOCHLORIS Lacepede,
1799
Chlorotalpa tropicalis G. M. Allen and
Loveridge, 1927
Proc. Boston Soc. Nat. Hist., 38: 418, 23
December.
= Chrysochloris stuhlmanni tropicalis (G.
M. Allen and Loveridge, 1927). See
Meester (1974: 3).
Holotype. MCZ 22435. Skin and skull. Adult female.
Locality. Tanganyika Territory (=Tanzania): Ulu-
gui"u Mountains, Bagilo. 5 October 1926.
Collector A. Loveridge.
Condition. Skin complete. Skull slightly damaged
(coronoid processes missing from both mandibular
rami). Mandible disarticulated.
Type Series. Holotyj^De only.
Comments. Included in Chrysochloris stuhlmanni
by Meester (1974: 3) but recognized as distinct by
Simonetta (1968: 42). Hutterer (1993: 75) noted
that the systematic status of tropicalis merits fur-
ther study.
Family TENRECIDAE Gray, 1821
Genus GEOGALE Milne-Edwards and A.
Grandidier, 1872
Cryptogale australis G. Grandidier, 1928
Bull. Mus. Hist. Nat. Paris, 34: 64, 26
Januaiy.
= Geogale aurita Milne-Edwards and A.
Grandidier, 1872. See Genest and Petter
(1975: 3).
Holotype. MCZ 45057. Skull fragments.
Locality. Madagascar: (Toliary), south of Fort Dau-
phin (=Tolanaro), Andrahomana grotto. 1927.
Collector R. Decary.
Condition. MCZ 45047 includes 18 partial crania
and 8 mandibular rami. Fragmentary.
Type Series. All the type material of C australis
bears a single accession number.
Comments. C. australis is the type species of the
genus Cn/pto^ale G. Grandidier, 1928.
Geogale aurita orientalis G. Grandidier
and Petit, 1930
Faune des Colonies Frangaises, 4: 446.
Holotype. MCZ 45660. Body in alcohol, cranium sep-
arate.
102 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Locality. Madagascar: (Toamasina), east coast, Fe-
iierive ( = Fenoarivo Atsinanana). April 1928.
Collector. R. Decaiy. Original number 12.
Condition. Alcoholic, cranium complete.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Ge-
nest and Fetter (1975: 3). The holotype of G. a.
orientalis seems to be the only record of Geogale
aurita from the east coast of Madagascar.
Genus /W/CROG/\/.E Thomas, 1882
Microgale decaryi G. Grandidier, 1928
Bull. Mus. Hist. Nat. Paris, 34: 69, 26
January.
= Microgale principula Thomas, 1926. See
MacPhee (1987a: 9).
Holotype. MCZ 45049. Cranium. Adult.
Locality. Madagascar: (Toliary), south of Fort Dau-
phin (=Tolanaro), Andrahomana caves. 1926.
Collector. R. Decary.
Condition. Cranium partial (back of cranium miss-
ing posterior to parietals).
Type Series. Paratype material consists of MCZ
45408, which includes 3 partial crania and 5 man-
dibular rami, and MAD- 1649, a partial skull in the
collections of the Institut de Paleontologie,
MNHN.
Microgale drouhardi G. Grandidier, 1934
Bull. Mus. Hist. Nat. Paris, 6: 474, 29
November.
Holotype. MCZ 45034. Body in alcohol, skull extract-
ed. Juvenile female.
Locality. Madagascar: (Antsiranana), east coast, Di-
ego-Suarez (= Antsiranana). May 1934.
Collector M. E. Drouhard. Original number A.
Condition. Alcoholic, skull complete.
Type Series. 6 paratypes; MCZ 46007-MCZ 46012,
all in alcohol. MCZ 46017, represented by a skull
and postcranial skeleton, was collected at the same
time and place as the type series but is not men-
tioned in the original description.
Comments. MCZ 45034 represents an immature
animal (MacPhee 1987a: 7), not an adult as claimed
in the original description. MacPhee (1987a: 9)
synonyinized drouhardi with Microgale cowani, an
approach followed by Hutterer (1993: 71), but Jen-
kins et al. (1997: 6) argued that M. droidiardi is a
distinct species.
Microgale parvula G. Grandidier, 1934
Bull. Mus. Hist. Nat. Paris, 6: 476, 29
November.
Holotype. MCZ 45465. Body in alcohol, skull extract-
ed. Juvenile male.
Locality. Madagascar: (Antsiranana), east coast, Di-
ego-Suarez (= Antsiranana). May 1934.
Collector. M. Drouhard.
Condition. Alcoholic, skull complete.
Tijpe Series. Holotype only.
Comments. MCZ 45465 represents an immature
animal (MacPhee 1987a: 7), not an adult as claimed
in the original description. The holotype of Micro-
gale pulla Jenkins, 1988 actually represents an
adult specimen of M. parvula (Jenkins et al., 1996:
204). Considered a valid species by Hutterer (1993:
71) and Nowak (1999: 190).
Microgale prollxacaudata G. Grandidier,
1937
Bull. Mus. Hist. Nat. Paris, 9: 348, 25
November.
= Microgale longicaudata Thomas, 1882.
See MacPhee (1987a: 9).
Holotype. MCZ 45035. Body in alcohol, skull extract-
ed. Juvenile.
Locality. Madagascar: (Antsiranana), east coast, Di-
ego-Suarez (= Antsiranana). May 1934.
Collector M. Drouhard.
Condition. Alcoholic; skull partial (left tympanic
bulla missing). Mandible disarticulated.
Type Series. 1 paratype; MCZ 46020, in alcohol.
Comments. This specimen represents an immature
animal (MacPhee 1987a: 8), not an adult as claimed
in the original description.
Paramlcrogale occldentalls G. Grandidier
and Petit, 1931
Bull. Soc. Zool. France, 56: 129, 15 June.
= Microgale brevlcaudata G. Grandidier,
1899. See MacPhee (1987a: 9).
Holotype. MCZ 45047. Body in alcohol, skull extract-
ed. Juvenile male.
Locality. Madagascar: (Antananarivo), northwest of
Maintirano, Andriafeuelo. 1930.
Collector M. A. de la Rue.
Condition. Alcoholic, skull complete.
Type Series. Holotype only, but see comments.
Comments. P. occidentalis is the type species of the
genus Paramicrogale G. Grandidier and Petit,
1931. This specimen represents an immature ani-
mal (MacPhee 1987a: 7), not an adult as claimed
in the original description. The original description
mentions only a single specimen but describes ex-
tensively the skeleton of P. occidentalis. This is puz-
zling, as the skeleton has not been extracted from
MCZ 45047 (MacPhee 1987a: 7).
Type Specimens of Recent Mammals • Helgen and McFadden 103
Genus SETIFER Fronep, 1806
Dasogale fontoynonti G. Grandidier,
1930a
Bull. Acad. Malgache, n. sen, 11: 85 (for
1928).
= Setifer setosus {Schreber, 1777). See
Poduschka and Poduschka (1982: 261).
Holotype. MCZ 45016. Skin, skull, and postcranial
skeleton. Juvenile.
Localitij. Madagascar: east coast. 1917.
Collector. Received by G. Grandidier from the
Academic Malgache in 1917.
Condition. Sldn represented by a small patch of
lur. Skull partial (occipital region missing). Postcra-
nial skeleton complete, partially articulated.
Tijpe Series. 1 paratype; MCZ 45532, in alcohol.
Comments. Walker (1975: 110) commented that
"the only specimen known of D. fontoynonti ... is
in the Paris Museum"; actually, the only material
attributed to Dasogale is in the MCZ, a fact first
noted in publication by Poduschka and Poduschka
(1982: 253). Dasogale was often considered to be
an extremely rare or recently extinct species until
Poduschka and Poduschka (p. 261) and MacPhee
(19S7b: 135) demonstrated that the holotype is
probably a juvenile Setifer .setosus. D. fontoynonti
is the type species of the genus Dasogale G. Gran-
didier, 1930.
Order RODENTIA Bowdich, 1821
Family APLODONTIDAE Brandt, 1855
Genus APLODONTIA Richardson, 1829
Aplodontia californica columbiana Taylor,
1916
Univ. California Publ. Zool., 12: 499, 6
May.
= Aplodontia rufa rainieri Merriam, 1899.
See Dalquest (1948: 369).
Holotype. B1899. Sldn and skull. Adult male.
Locality. (Canada): British Columbia, Hope,
Roabs Ranch. 14 June 1894.
Collector W. C. Colt. Original number 479.
Condition. Skin and skull complete.
Type Series. 8 paratypes; B1892-B1898, B1900; all
represented by skin and skull, 3 females and 5
males.
Family SCIURIDAE Fischer de Waldheim,
1817
Genus CALLOSCIURUS Gray, 1867
Callosciurus baluensis medial is G. M.
Allen and Coolidge, 1940
Bull. Mus. Comp. Zool., 87: 156, 31
December.
Holotype. MCZ 22265. Skin and skull. Adult female.
Locality. (Indonesia): Dutch Borneo, (Kalimantan),
Mount Tibang (possibly =Bukit Tungun). 1925.
Collector. E. Mjoberg. Original number 6.
Condition. Skin complete. Skull partial (right pa-
rietal, right tympanic bulla, and lachiymal bro-
ken — bulla present). Mandible disarticulated.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Med-
way (1977: 90).
Callosciurus ferruginous primus G. M.
Allen and Coolidge, 1940
Bull. Mus. Comp. Zool, 87: 157, 31
December.
= Callosciurus erythraeus (Pallas, 1778).
See Corbet and Hill (1992: 283).
Holotype. MCZ 35352. Skin and skull. Adult female.
Locality. Siam (=Thailand): Mae Wan River near
Doi, Mount Souket ( = Saket), 1,500 ft (458 m). 20
Febiaiary 1937.
Collector J. A. Griswold, Jr., Asiatic Primate Ex-
pedition. Original number 5.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; MCZ 35353, skin and
skull, adult male.
Sciurus castaneoventris haemobaphes G.
M. Allen, 1912c
Proc. Biol. Soc. Washington, 25: 177, 24
December.
= Callosciurus erythraeus haemobaphes
(G. M. Allen, 1912). See Hayman and
Holt (1940: 359).
Holotype. MCZ 13693. Skin and skull. Male.
Locality. China: southeastern Yunnan, Chih-ping
( = Shiping). 26 Februaiy 1911.
Collector Kobayashi Collection. Original number
46.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Only the holotype is mentioned in the
original publication, but Allen had four other spec-
imens of these squirrels at the time of description
(MCZ 13692, 13694-13696; all represented by skin
and skull, 3 males, 1 female).
104 Bulletin Museum of Comparative Zoolog^y, Vol. 157, No. 2
Genus DREMOMYS Heude, 1 898
Dremomys pernyi flavior G. M. Allen,
1912c
Proc. Biol. Soc. Washington, 25: 178, 24
December.
Holottjpe. MCZ 13691. Skin and skull. Male.
Locality. China: southeastern Yunnan, Mongtz
( = Mengzi). 1911.
Collector. Kobayashi Collection. Original number
6/8.
Condition. Skin complete. Skull partial (squamo-
sals missing, supraoccipital damaged, right jugal
missing, palatine missing, tympanic bulla dam-
aged).
Type Series. Holoty]^)e only.
Dremomys senex G. M. Allen, 1912b
Mem. Mus. Comp. ZooL, 40: 229,
August.
= Dremomys pernyi senex G. M. Allen,
1912. See Hayman and Holt (1940: 382).
Holotype. MCZ 7582. Skin and skull. Adult female.
Locality. China: Hupeh ( = Hubei), Ichanghsien,
Nantou. 5 Februaiy 1909.
Collector W. R. Zappey. Original number .373.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paraty^je; MCZ 7583, skin and skull,
adult male.
Genus FUNISCIURUS Trouessart, 1880
Funisciurus pyrrhopus victoriae G. M.
Allen and Loveridge, 1942
Bull. Mus. Comp. Zool., 8: 180,
February.
= Funisciurus pyrriiopus al<l<a De Winton,
1899. See Amtmann (1975: 8).
Holotype. MCZ 39199. Skin and skull. Adult male.
Locality. Uganda: Toro, Kibale Forest, 4,200 ft
(1,281 m). 16 December 1938.
Collector A. Loveridge.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Genus GLAUCOIVIYS Thomas, 1908
Sciuropterus alpinus bangsi Rhoads, 1897
Proc. Acad. Nat. Sci. Philadelphia, 1897,
p. 321 (footnote), July.
^Glaucomys sabrinus bangsi (Rhoads,
1897). See A. H. Howell (1918: 38).
Holotype. B6959. Sldn and skull. Adult male.
Locality. (United States): Idaho, Idalio County.
March 8, 1897.
Collector Harbison and Bargamin.
Condition. Skin and skull complete.
Type Series. 1 paratype; B6960, skin and skull,
adult male.
Comments. G. s. bangsi was retained as a valid sub-
species by Hall (1981: 450).
Sciuropterus alpinus lascivus Bangs,
1899J
Proc. New England Zool. Club, 1: 69, 31
July.
= Glaucomys sabrinus lascivus (Bangs,
1899). See A. H. Howell (1918: 55).
Holotype. B9186. Skin and skull. Adult female.
Locality. (United States): California, El Dorado
County, Tallac. 28 August 1898.
Collector W. VV. Price and P. O. Simons. Original
number 1722.
Condition. Sldn and skull complete.
Type Series. 2 paratypes, B9187, skin and skull, fe-
male; B9188, sldn and skrdl, female.
Comments. G. s. la.scivns was retained as a valid
subspecies by Hall (1981: 451).
Sciuropterus sabrinus makkovikensis
Sornborger, 1900
Ottawa Nat., 14: 48, 6 June.
= Glaucomys sabrinus makkovikensis
(Sornborger, 1900). See A. H. Howell
(1918: 34).
Syntypes. MCZ 10476: Skin and skull. Adult. MCZ
10477: Skin and skull. Adult. MCZ 10478: Skin and
skull. Adult.
Locality. (Canada): Labrador Peninsula, Makkovik.
1899.
Collector. W. W. Perrett. Original number 1540.
Condition. MCZ 10476: Skin complete. Skull par-
tial (base of skull missing). Mandible disarticulated.
MCZ 10477: Skin partial (tail broken but present).
Skull partial (base of skull missing). Mandible dis-
articulated. MCZ 10478; Skin partial (missing left
hind foot and tail). Skull partial (occipital chipped).
Mandible disarticulated.
Type Series. 3 syntypes only.
Comments. Sornborger's original description was
based on three specimens, original numbers 1540,
1541, and 1542, now MCZ 10476, 10477, 10478,
respectively. G. .$. makkovikensis was retained as a
valid subspecies by Hall (1981: 453).
Sciuropterus silus Bangs, 1896J
Proc. Biol. Soc. Washington, 10: 163, 28
December.
= Glaucomys volans volans (Linnaeus,
1758). See A. H. Howell (1918: 20).
Holotype. B4931. Sldn and skull. Adult male.
Locality. (United States): West Virginia, Greenbri-
Type Specimens of Recent Mammals • Helpen and McFodden
105
er County, White Sulphur Springs, top of Katis
Mtn. 3,200 ft (976 m). 2 September 1895.
Collector. T. Surber. Original number 19.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Sciuropterus volans querceti Bangs, 1 896j
Proc. Biol. Soc. Washington, 10: 166, 28
December.
= Glaucomys volans querceti (Bangs,
1896). See A. H. Howell (1918: 26).
Holotype. B2451. Skin and skull. Adult female.
Locality. (United States): Florida, Citrus County,
Citronelle. 17 September 1894.
Collector F. L. Small. Original number 1363.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. 2 paratypes; B2452, skin and skull,
adult male; B2453, skin and skull, adult female.
Comments. G. v. querceti was retained as a valid
subspecies by Hall (1981: 449).
Genus HYLOPETES Thomas, 1 908
Pteromys (Hylopetes) alboniger orinus G.
M. Allen, 1940
The Mammals of China and Mongolia,
Natural Histoiy of Central Asia, 11: 723,
3 September.
= Hylopetes alboniger alboniger (Hodgson,
1836). See Ellerman and Morrison-Scott
(1951: 469).
Holotype. MCZ 28086. Skin and skull. Adult female.
Locality. China: Yunnan, Likiang Range ( = Li-
jiang), 7,800 ft (2,379 m). December 1931.
Collector J. F Rock.
Condition. Skin complete. Skull partial (most of
right tympantic bulla missing, supraoccipital dam-
aged). Mandible disarticulated.
Type Series. Allen examined 10 specimens in ad-
dition to the type, including 4 specimens from the
BMNH, 3 from AMNH, and MCZ 28087, skin and
skull, an unsexed adult.
Pteromys phayrei anchises G. M. Allen
and Coolldge, 1940
Bull. Mus. Comp. Zool., 87: 153, 31
December.
= Hylopetes phayrei anchises (G. M. Allen
and Coolldge, 1940). See Ellerman and
Morrison-Scott (1951: 469).
Holotype. MCZ 35776. Skin and skull. Adult male.
Locality. Siam (=Thailand): (Chiang Mai) Mount
Angka ( = Doi Inthanon), 4,300 ft (1,312 m). 27
February 1937.
Collector. J. A. Griswold, Jr., Asiatic Primate Ex-
pedition. Original number 24.
Condition. Skin and skvdl complete. Mandible dis-
articulated.
Type Series. 3 paratypes; MCZ 35775, skin and
sk-ull, adult female; MCZ 35777, skin and skull,
adult male; MCZ 35778, skin and skull, subadult
male.
Genus MARMOTA Blumenbach, 1779
Arctomys flaviventer avarus Bangs, 1899]
Proc. New England Zool. Club, 1: 68, 31
July.
= Marmota flaviventris avara (Bangs,
1899). See A. H. Howell (1915: 41).
Holotype. B7299. Skin and skull. Juvenile female.
Locality. (Canada): British Columbia, Okanagan.
17 July 1897.
Collector A. C. Brooks. Original number 969.
Condition. Skin and skadl complete.
Type Series. 2 paratypes; B7298, skin and skull, ju-
venile male; B7300, skin and skull, juvenile female.
Comments. M. f. avara was retained as a valid sub-
species by Hall (1981: 371).
Arctomys ignavus Bangs, 1899d
Proc. New England Zool. Club, 1: 13, 28
February.
= Marmota monax ignava (Bangs, 1899).
See A. H. Howell (1915: 29).
Holotype. B7971. Skin and skull. Adult male.
Locality. (Canada): Labrador Peninsula, Black Bay.
13 July 1898.
Collector. E. Doane.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 5 paratypes; B7968-B7970, B7972,
B7973 (juvenile); all represented by skin and skull,
4 females and 1 male.
Comments. M. m. ignava was retained as a valid
subspecies by Hall (1981: 370) and Kwiecinski
(1998: 1).
Genus MICROSCIURUS J. A. Allen, 1895
Sciurus (Microsciurus) browni Bangs,
1902b
Bull. Mus. Comp. Zool., 39:24, April.
= Microsciurus alfari browni (Bangs, 1902).
See J. A. Allen (1914:151).
Holotype. MCZ 10404. Skin and skull. Adult male.
Locality. Panama: Chiriqui, Bogaba, 600 ft (183
m). 15 July 1901.
106 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Collector. W. W. Browii, Jr. Original number 631.
Condition. Sldn and skull complete.
Type Series. 4 paratypes; MCZ 10405, adult fe-
male, 10406, adult female, 10407, adult female,
10408, juvenile male; all represented by sldn and
skull.
Comments. M. a. broivni was retained as a valid
subspecies by Hall (1981: 439).
Genus PARAXERUS Forsyth Major, 1893
Aethosciurus byatti laetus G. M. Allen and
Loverldge, 1933
Bull. Mus. Comp. Zool, 75: 96,
Februaiy.
= Paraxerus vexillarius byatti (Kershaw,
1923). See Amtmann (1975: 11).
Holotype. MCZ 26198. Skin and skull. Adult male.
Locality. Tanganyika Territory (=Tanzania): north
end of Lake Nyasa, Ukinga Mountains, Madehani,
7,000 ft (2,135 m). 22 Februaiy 1930.
Collector. A. Love ridge.
Condition. Sldn and skull complete.
Type Series. 9 paratypes; MCZ 26196, 26197,
26199-26202, 26204-26206; all represented by
skin and skull, 4 females and 5 males.
Genus SCIUROTAMIAS M\\\er, 1901
Sciurotamias davidanus thayeri G. M.
Allen, 1912b
Mem. Mus. Comp. Zool., 40: 231,
August.
= Sciurotamias davidianus consobrinus
(Milne-Edwards, 1868). See Moore and
Tate (1965: 308).
Holotype. MCZ 8008. Skin and skull. Adult male.
Locality. China: western Szechwan ( = Sichuan),
Washan (=Wushan), 6,000 ft (1,830 m). 17 May
1908.
Collector W. R. Zappey. Original number 163.
Condition. Sldn complete. Skull partial (basioccip-
ital and left maxilla damaged). Mandible disartic-
ulated.
Type Series. Holotype only.
Comments. G. M. Allen (1912b: 231) noted in the
original description of thayeri that "unfortunately,
tlie skull of the type was lost." The sldn and skull
have been revmited subsequently.
Genus SC/L/RL/S Linnaeus, 1758
Sciurus (Guerlinguetus) aestuans
cfiiriquensis Bangs, 1902b
Bull. Mus. Comp. Zool., 39: 22, April.
= Sciurus granatensis chiriquensis Bangs,
1902. See Hershkovitz (1947: 7).
Holotype. MCZ 10044. Skin and skull. Adult male.
Locality. Panama: Chiriqui, Divala. 18 November
1900.
Collector. W. W. Brown, Jr. Original number 10.
Condition. Skin and skull complete.
Type Series. There is a large series of paratypes in
the MCZ.
Comments. S. g. chiriquensis was retained as a val-
id subspecies by Hall (1981: 437) and Nitikman
(1985: 1).
Sciurus carolinensis extimus Bangs, 1896j
Proc. Biol. Soe. Washington, 10: 158, 28
December.
Holotype. B4519. Sldn and skull. Adult female.
Locality. (United States): Florida, Dade County,
Miami. 12 March 1895.
Collector. L. Brownell. Original number 59.
Condition. Skin complete (with bald spots on ven-
trum). Skull complete.
Type Series. 7 paratypes; B3406, B4517, B4518,
(all represented by skin and skull), B4520-B4523
(skins only); 4 females and 3 males.
Comments. Retained as a valid subspecies by Hall
(1981: 417) and Koprowskd (1994: 1).
Sciurus carolinensis var. yucatanensis J.
A. Allen, 1877
In Coues and J. A. Allen, Monogr. N.
Amer. Rodentia, U.S. Geol. Geograph.
Suivey Terr., Rep. 11: 705, August.
= Sciurus yucatanensis J. A. Allen, 1877.
See Elliot (1896: 80).
Syntype. MCZ 5398. Skin. Adult male.
Locality. (Mexico): Yucatan, Merida. March 1865.
Collector. A. Schott. Original number 228. For-
merly USNM 8502.
Condition. Sldn complete.
Type Series. J. A. Allen referred to "four specimens
of this variety before me" in the original descrip-
tion (1877: 705). Three specimens are mentioned
by number: USNM 8502 (now MCZ 5398), 8503,
and 8505. A juvenile referred to in the description
(but not by number) corresponds to USNM 8504.
Comments. S. yucatanensis was considered a valid
species by Hoffman et al. (1993: 443) and Nowak
(1999: 1265). The syntype of yucatanensis in the
MCZ was received from the USNM in March
1877. Poole and Schantz (1942: 554) state that
USNM 8505 is no longer in the USNM.
Sciurus ludovicianus vicinus Bangs, 1896j
Proc. Biol. Soc. Washington, 10: 150, 28
December.
= Sciurus niger vulpinus Gmelin, 1896.
See Bark-alow (1954: 25).
Holotype. B5215. Skin and skull. Adult female.
Locality. (United States): West Virginia, Greenbri-
Type Specimens of Recent Mammals • Helgen and McFodden 107
ei" County, White Sulphur Springs. 29 Januaiy
1896.
Collector. T. Surber. Original number 55.
Condition. Sldn and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Sciurus nesaeus G. M. Allen, 1902b
Proc. Biol. Soc. Washington, 15: 93, 25
April.
= Sciurus granatensis nesaeus G. M.
Allen, 1902. See Hershko\atz (1947: 37).
Holotijpe. MCZ 10744. Skin. Adult female.
Locality. Venezuela: (Nueva Esparta), Margarita
Island, El Valle. 8 July 1901.
Collector. A. H. Clark. Original number 619.
Condition. Skin complete. The lower incisors are
present in the sldn.
Type Series. Holotype only.
Comments. S. g. nesaeus was retained as a valid
subspecies by Cabrera (1961: 367) and Nitikman
(1985: 1).
Sciurus variabilis morulus Bangs, 1 900d
Proc. New England Zool. Club, 2: 43, 20
September.
= Sciurus granatensis morulus Bangs,
1900. See Miller and Kellogg (1955:
257).
Holotype. B8420. Skin and skull. Adult female.
Locality. Panama: Canal Zone, Loma del Leon. 13
March 'l900.
Collector W. W. Brown, Jr. Original number 5.
Condition. Skin complete. Skull slightly damaged
(left bulla broken).
Type Series. 5 paratypes; B8418, B8419, B8421-
B8423 (juvenile); all represented by skin and skull,
3 females and 2 males.
Comments. S. g. mondus was retained as a valid
subspecies by Hall (1981: 437) and Nitikman
(1985: 1).
Sciurus variabilis saltuensis Bangs, 1898o
Proc. Biol. Soc. \\'ashington, 12: 185, 16
November.
= Sciurus granatensis saltuensis Bangs,
1898. See Hershkovitz (1947: 15).
Holotype. B8144. Skin and skull. Adult female.
Locality. Colombia: Magdalena, Santa Marta
Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 26
March 1898.
Collector. W. W. Brown, Jr. Original number 112.
Condition. Sldn and skull complete.
Type Series. 2 paratypes; B8145, skin and skull,
adult male; B8244, skin and skull, adult female.
Comments. S. g. saltuensis was retained as a valid
subspecies by Cabrera (1961: 368) and Nitikman
(1985: 1).
Genus SPERMOPHILUS F. Cuvier, 1825
Citellus obscurus siccus G. M. Allen, 1925
Amer. Mns. Novitates, 163: 3, 2 April.
= Spermophilus alashanicus Buchner,
1888. See Hoffmann et al. (1993: 444).
Holotype. MCZ 19924. Skin and skull. Adult female.
Locality. China: Shansi ( = Slienxi), 10 miles (16.1
km) west of Taiyuanfu. August 1921.
Collector. F. R. Wulsin. Original number 146.
Condition. Skin complete. Skull partial (parietals
and basiooccipital missing, right tympanic bulla
broken but present). Mandible disarticulated.
Ti/pe Series. Holotype only.
Spermophilus armatus Kennicott, 1863
Proc. Acad. Nat. Sci. Philadelphia, 15:
158, June.
Sipitypes. MCZ 297: Skull. Male. Collected 11 April
' 1858. Original number 167, formerly USNM 4799.
MCZ 4793: Skin and skull. Male. Collected 2 April
1858. Original number 140, formerly USNM 3478
(3373). MCZ 4790: Skin. Female. Collected 26
May 1858. Original number 455, formerly USNM
3470 (3470). MCZ 4794: Skull only (skin spoiled in
repreparation, discarded). Male. Collected April
14, 1858. Original number 215, formerly USNM
3474 (3373).
Locality. (United States): Utah (now Wyoming),
(Uinta County), near Fort Bridger, foothills of the
Uinta Mountains, Camp Scott.
Collector C. Drexler.
Type Series. See comments below.
Comments. Considered a valid species by Hoff-
mann et al. (1993: 444) and Nowak (1999: 1254).
Kennicott does not designate a type in the original
descidption; thus, "all the specimens from Fort
Bridger collected by C. Drexler and in the collec-
tions [of the USNM] prior to 1863 are evidently
cotypes [ = syntypes] of this species" (Lyon and Os-
good 1909: 163). The syntypes of annatus in the
MCZ were received from the USNM in January
1874. These specimens possess USNM labels that
bear numbers in discrepancy with the USNM cat-
alogue; the numbers listed above are taken from
the catalogue of the USNM, followed in parenthe-
ses by the number on the specimen label. A com-
plete list of syntypes, most of the remainder of
which are in the USNM, can be found in Poole
and Schantz (1942: 504-505). The skin of MCZ
297, which formerly bore tlie number USNM
3472, is not to be found in the collections of either
108 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Spermophilus elegans Kennicott, 1863
Proc. Acad. Nat. Sci. Philadelphia, 15:
158, June.
Syntijpes. MCZ 4791: Skin. Female. Collected 11
April 1858. Original number 168. Formerly USNM
3468. MCZ 4792: Skin. Male. Collected 11 April
1858. Original number 166. Formerly USNM
3473.
Locality. (United States): Utah (now Wyoming),
(Uinta County), Fort Bridger
Collector. C. Dre.-der.
Condition. Following their receipt, these speci-
mens were spoiled in repreparation and discarded.
Type Series. See comments below.
Comments. These specimens are no longer in ex-
istence. Kennicott does not designate a type in the
original description; thus, "all the specimens from
Fort Bridger collected by C. Drexler and in the
collections [of the USNM] prior to 1863 are evi-
dently cotypes [ = syntypes] of this species" (Lyon
and Osgood 1909: 163). These syntypes oi elegans
were received from the USNM in January 1874. A
complete list of syntypes can be found in Lyon and
Osgood (1909: 166). G. M. Allen (1931: 251) er-
roneously included MCZ 4791 in a list of the type
series of Spennophihis annatus. S. elegans is con-
sidered a valid species by Hoffmann et al. (199.3:
446) and Nowak (1999: 1254).
Spermophilus (Ictidomys) tridecemlineatus
badius Bangs, 1899c
Proc. New England Zool. Club, 1: 1, 8
Februaiy.
= Spermophilus tridecemlineatus texensis
Merriam, 1898. See Hall and Kelson
(1959: 347).
Holotype. B1682. Skin and skull. Adult male.
Locality. (United States): Missouri (Vernon Coun-
ty), Stotesbury. 17 April 1894.
Collector. T. Surber. Original number 81.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 2 paratypes; B1683, skin and skull,
adult male; B5609, skin and skull, adult male.
Comments. A. H. Howell (1938: 110) first synon-
yinized badius with texensis under die genus Ci-
tellus.
Genus SYNTHEOSCIURUS Bangs, 1902
Syntheosciurus brochus Bangs, 1 902b
Bull. Mus. Comp. Zool., 39: 25, April.
Holotype. MCZ 10402. Sldn and skull. Adult male.
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo-
quete), 7,000 ft (2.135 m). 30 April 1901.
Collector W. W. Brown, Jr Original number 415.
Condition. Skin complete. Skull partial (left supra-
occipital damaged)
Type Series. 1 paratype; MCZ 10403, sldn and
skull, adult female.
Comments. Type species of the genus Syntheo.sciu-
nis Bangs, 1902. Considered a valid species by
Hoffman et al. (1993: 452) and Nowak (1999:
1268). Following a discussion with a collector em-
ployed by W. W. Brown, Jr., and a biological survey
of Boquete, Enders (1953b: 509) recommended
that the type locality be considered "the Cordillera
about 8 miles [12.9 km] north of Boquete and not
at Boquete which is on the lower slopes of El Vol-
can de Chiriqui."
Genus LAMMS llliger, 1811
Tamias cooperi Baird, 1855
Proc. Acad. Nat. Sci. Philadelphia, 7:
334, 24 April 24.
= Tamias townsendii cooperi Baird, 1855.
See Baird (1858: 737).
Syntype. MCZ 4754. Skin only. Adult, unsexed.
Locality. (United States): Washington (Skagit
County), Clickitat ( = Klickitat) Pass, Cascade
Mountains, 4,500 ft (1,373 m). July 1853. See com-
ments.
Collector J. G. Cooper. Formerly USNM 211/
1182.
Condition. Sldn complete.
Type Series. The other syntype is USNM 212/1183,
skin and skull, unsexed adult.
Comments. No type is designated in the original
description, but Baird subsequently ex-plained that
the two specimens listed above were those on
which cooperi was based (1858: 301). Cooper
(1869: .531) emended the type locality. The syntype
of cooperi in the MCZ was received from the
USNM in Januarv' 1874.
Tamias dorsalis Baird, 1855
Proc. Acad. Nat. Sci. Philadelphia, 7:
332, 24 April.
= Tamias dorsalis dorsalis Baird, 1855.
See Hayman and Holt (1940: 435).
Syntype. MCZ 4759. Skin and mandible.
Locality. (United States): New Me.xico (Grant
County), "Fort Webster, Coppermines of die Mim-
bres" (near present Georgetown). 32°47'N,
108°41'W. 1851. See A. H. Howell (1929: 131).
Collector J. H. Clark. Formerly USNM 119/3151.
Condition. Skin complete. Mandible partial (pos-
terior of right mandibular ramus broken off and
missing).
Type Series. The other syntype is USNM 120, skull
within skin.
Comments. No type is designated in the original
description, but Baird subsequently listed the two
Type Specimens of Recent Mammals • Helaen and McFadden
109
specimens above as those on which dorsalis was
based (1858: 300). The syntype of dorsalis in the
MCZ was received from the USNM in January
1874.
Tamias quadrivittatus neglectus J. A.
Allen, 1890
Bull. Amer. Mus. Nat. Hist., 3: 106,
June.
= Tamias minimus neglectus (J. A. Allen,
1890). See Hayman and Holt (1940:
430).
Holotype. MCZ 1575. Sldn and cranium.
Locality. (Canada): (Ontario), eastern end of Lake
Superior (near mouth of Montreal River). 5 July
1848.
Collector. L. Agassiz.
Condition. Sldn complete. Cranium partial (zygo-
matic arches, parietals, left tympanic bulla, and
mandible missing).
Type Series. The description mentions 6 paratypes,
including MCZ 1567, skin and skull; and 4 others
at USNM.
Comments. Following A. H. Howell (1929: 54),
Hayman and Holt (1940: 430) considered T. m.
neglectus a synonym of T. m. borealis. However,
neglectus was retained as a valid subspecies of min-
imus by Hall (1981: 346) under the genus Euta-
mias.
Tamias striatus venustus Bangs, 1 896h
Proc. Biol. Soc. Washington, 10: 137, 28
December.
Holotype. B5478. Skin and skull. Adult male.
Locality. (United States): Indian Territory
( = Oklahoma) (Adair County), Stilwell. 13 August
1896,
Collector T. Surber. Original number 63.
Condition. Skin complete. Skull partial (condyle of
left mandibular ramus inissing).
Type Series. 2 paratypes; B5479, skin and skull,
adult female; B5605, sldn and skull, adult male.
Comments. Retained as a valid subspecies by Hall
(1981: 340).
Genus r>4/WMSC/L/f?L/S Trouessart, 1880
Sciurus iiudsonicus gymnicus Bangs,
18991
Proc. New England Zool. Club, 1: 28, 31
March.
= Tamiasciurus Iiudsonicus gymnicus
(Bangs, 1899). See Osgood (1938: 438).
Holotype. B4914. Skin and skull. Adult female.
Locality. (United States): Maine, Piscataquis
County, Greenville, near Moosehead Lake. 1 De-
cember 1895.
Collector C. H. Goldthwaithe. Original number 2.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. T. h. gymnicus was retained as a valid
subspecies by Hall (1981: 442) and Steele (1998:
1).
Sciurus hudsonicus loquax Bangs, 1 896j
Proc. Biol. Soc. Washington, 10: 161, 28
December.
= Tamiasciurus hudsonicus loquax
(Bangs, 1896). See A. H. Howell (1936:
1).
Holotype. B4270. Skin and skull. Adult male.
Locality. (United States): Connecticut, New Lon-
don County, Liberty Hill. 24 December 1895.
Collector. O. Bangs. Original number 3.
Condition. Skin and skull complete.
Type Series. There is a large series of paratypes in
the MCZ.
Comments. T. h. loquax was retained as a valid sub-
species by Hall (1981: 443) and Steele (1998: 1).
Sciurus hudsonicus orarius Bangs, 18971
Proc. Biol. Soc. Washington, 11: 281, 30
December.
= Tamiasciurus douglasii mollipilosus
(Audubon and Bachman, 1841). See
Hayman and Holt (1940: 347).
Holotype. B4978. Skin and skull. Adult female.
Locality. (United States): California, Mendocino
County, Philo. 9 December 1895.
Collector. C. A. Allen. Original number 887.
Condition. Skin complete. Skull partial (right tym-
panic bulla broken).
Type Series. 13 paratypes; B4832, B4979-B4989,
B5462; all represented by sldn and skull, 9 feinales
and 4 males.
Family CASTORIDAE Hemprich, 1820
Genus C/^SrOR Linnaeus, 1758
Castor caecator Bangs, 1 91 3
Bull. Mus. Comp. Zool., 54: 513, July.
= Castor canadensis caecator Bangs,
1913. See G. M. Allen (1942: 62).
Holotype. B6979. Skull. Adult male.
Locality. (Canada): Newfoundland, near Bay St.
George. 1896.
Collector. E. Doane.
Condition. Skull complete. Mandible disarticulat-
ed.
110 Bulletin Museum of Comparative Zoologi/, Vol. 157, No. 2
Type Series. Holotype only.
Comments. C. c. caecator was retained as a valid
subspecies by Hall (1981: 602).
Family GEOMYIDAE Bonaparte, 1845
Genus GEOMVS Rafinesque, 1817
Geomys colonus Bangs, 1898b
Proc. Boston Soc. Nat. Hist., 28: 178, 15
March.
= Geomys pinetis pinetis Rafinesque,
1817. See Williams and Genoways (1980:
444).
Holotijpe. B5001. Skin and skull. Advilt male.
Locality. (United States): Georgia, Camden Coun-
ty, Arnot Plantation, about 4 miles (6.4 km) west
of St. Marys. 21 March 1896,
Collector. O. Bangs. Original number 8.
Condition. Sldn and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. Hall (1981: 505) recognized Geomys
colonus as a valid species.
Geomys cumberlandius Bangs, 1898b
Proc. Boston Soc. Nat. Hist., 28: 180, 15
March.
= Geomys pinetis pinetis Rafinesque,
1817. See Wihiams and Genoways (1980:
444).
Holotype. B5016. Skin and skull. Adult male.
Locality. (United States): Georgia, Camden Coun-
ty, Cumberland Island, Stafford Place. 17 April
1896.
Collector O. Bangs. Original number 1.
Condition. Sldn and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. In the original description. Bangs lists
B5015 as the type and gives its data. The data given
lit B5016, not B5015. B5016 has "Type" written
after the entiy in Bangs' catalogue and should be
considered the holotype, rather than B5015. Hall
(1981: 505) and Laerm (1981: 150) supported the
specific status of Geomys cumherlandius.
Geomys floridanus austrinus Bangs,
1898b
Proc. Boston Soc. Nat. Hist., 28: 177, 15
March.
= Geomys pinetis pinetis Rafinesque,
1817. See WilHams and Genoways (1980:
444).
Holotype. B6983. Skin and skull. Adult male.
Locality. (United States): Florida, Pinellas County,
Belleair. 3 August 1897.
Collector. W. S. Dickinson.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype, in the USNM.
Geomys tuza gfof// Sherman, 1944
Proc. New England Zool. Club, 23: 38,
30 August.
= Geomys pinetis goffi Slierman, 1944.
See Harper (1952: 37).
Holotype. B7222. Skin and skull. Adult male.
Locality. (United States): Florida, Brevard County,
Eau Gallie. 18 March 1897.
Collector. O. Bangs. Original number 1.
Condition. Sldn and skull complete.
Type Series. 12 paratypes; B7212-B7217, B7219-
B7221; all represented by sldn and skull, 5 females
and 7 males.
Comments. Williams and Genoways (1980: 444)
synonymized gojfi with G. pinetis pinetis; however.
Hall (1981: 504) maintained G, p. goffi as a valid
subspecies. The lUCN designates goffi as extinct.
Genus ORTHOGEOMYS Merriam, 1895
IVIacrogeomys cavator Bangs, 1902b
Bull. Mus. Conip. Zool., 39: 42, April.
= Orttiogeomys cavator cavator (Bangs,
1902). See Russell (1968a: 532).
Holotijpe. MCZ 10381. Skin and sk-ull. Adult male.
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo-
quete), 4,800 ft (1,464 m). 9 March 1901.
Collector W. W. Brown, Jr. Original number 212.
Condition. Skin and skull complete. Mandible dis-
articvdated.
Type Series. There is a large series of paratypes in
the MCZ.
Comments. O. cavator was considered a valid spe-
cies by Patton (1993: 470) and Nowak (1999: 1314).
Macrogeomys pansa Bangs, 1902b
Bull. Mus. Comp. Zool., 39: 44, April.
= Ortiiogeomys cavator pansa (Bangs,
1902). See Russell (1968a: 532).
Holotype. MCZ 10364. Skin and skull. Adult female.
Locality. Panama: Chiriqui, Bogaba, 600 ft (183
m). 6 July 1901.
Collector W. W. Brown, Jr Original number 567.
Condition. Skin complete. Skull partial (right tym-
panic bulla missing).
Type Series. There are 7 paratypes; MCZ 10362—
10363, 10365-10369, 4 females and 3 males, all
represented by skin and skull. MCZ 10362 and
10366 are now in the FMNH, and MCZ 10365 is
in the USNM.
Comments. O. c. pansa was retained as a valid sub-
species by Hall (1981: 513).
Type Specimens of Recent Mammals • Helaen and McFadden 111
Orthogeomys grandis pluto Lawrence,
1933a
Proc. New England Zool. Club, 13: 66, 8
May.
Holotype. MCZ 29040. Skin and skull. Adult female.
Locality. Honduras: Francisco Morazan, north of
Tegucigalpa, Cerro Cantoral. 20 July 1932.
Collector. C. F. Underwood. Original number
1100.
Condition. Skin and skull complete.
Type Series. 2 paratypes; MCZ 29038, skin and
sk-ull, juvenile female; MCZ 29039, skin and skull
unsexed juvenile.
Coninient.s. Retained as a valid subspecies by Hall
(1981: 509).
Genus PAPPOGEOMYS Mernam, 1895
Cratogeomys castanops rubellus Nelson
and Goldman, 1934a
Proc. Biol. Soc. Washington, 47: 147, 13
June.
= Pappogeomys castanops rubellus
(Nelson and Goldman, 1934). See Russell
(1968b: 682).
Holotype. MCZ 20507. Skin and skrdl. Adult male.
Locality. Mexico: San Luis Potosi, near San Luis
Potosi, Soledad, 6,400 ft (1,952 m). 1 August 1923.
Collector W. W. Browai, Jr. Original number 2.31.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a large paratype series in the
MCZ.
Comments. P. a. nibellus was retained as a valid
subspecies by Hall (1981: 520).
Family HETEROMYIDAE Gray, 1868
Genus DIPODOMYS Gray, 1841
Dipodomys californicus pallidulus Bangs,
1899J
Proc. New England Zool. Club, 1: 65, 31
July.
= Dipodomys californicus californicus
Merriam, 1890. See Kelt (1988: 1).
Holotype. B9147. Skin and skull. Adult female.
Locality. (United States): California, Colusa Coun-
ty, Sites. 27 June 1896.
Collector P. O. Simons. Original number 222.
Condition. Skin complete. Skull partial (left jugal
missing).
Type Series. 1 paratype; B9148, sldn and skull,
adult female.
Dipodops ordii palmeri J. A. Allen, 1891
Bull. Amer. Mus. Nat. Hist., 3: 276, 30
June.
= Dipodomys ordii palmeri {y}. A. Allen,
1891). See Grinnell (1921: 96).
Syntypes. MCZ 5886: Skin and skull. Juvenile male.
MCZ 5887: Skin and skull. Unsexed juvenile.
Locality. Mexico: San Luis Potosi, San Luis Potosi.
1 May 1878.
Collector E. Palmer.
Condition. MCZ 5886: Sldn complete. Skull partial
(part of parietal, right occipital condyle, and tym-
panic bulla missing). MCZ 5887: Skin complete.
Skull partial (left squamosal process missing). Man-
dible disarticulated.
Type Series. 2 syntypes only.
Comments. Allen did not specify a holotype in the
original description, which he based on the two
specimens above. D. o. palmeri was retained as a
vahd subspecies by Hall (1981: 569).
Genus THOMOMYS Wied-Neuwied, 1839 Genus HETEROMYS Desmarest, 1817
Heteromys repens Bangs, 1902b
Bull. Mus. Comp. Zool, 39: 45, April.
^Heteromys desmarestianus repens
Bangs, 1902. See Goldman (1920: 115).
Thomomys umbrinus atrodorsalis Nelson
and Goldman, 1934b
J. Mammal., 15: 111, 15 May.
Holotype. MCZ 20487. Skin and skull. Adult male.
Locality. Mexico: San Luis Potosi, Alvarez, 8,000
ft (2,440 m). 7 November 1923.
Collector W. W. Brown, Jr. Original number 338.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a large series of paratypes in
the MCZ.
Comments. Retained as a valid subspecies by Hall
(1981: 476).
Holotype. MCZ 10356. Skin and skull. Adult female.
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo-
quete), 4,000 ft (1,220 m). 8 April 1901.
Collector W. W. Brown, Jr. Original number 264.
Condition. Sldn complete. Skull partial (left jugal
and process of squamosal missing).
Type Series. 5 paratypes; MCZ 10359, adult male,
10355, adult male, 10358, adult male, 10361, ju-
venile female, 10360, juvenile female.
Comments. H. d. repens was retained as a valid
subspecies by Hall (1981: 597).
112 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Genus LIOMYS Uernam, 1902
Heteromys alleni Coues, 1881
In J. A. Allen, Bull. Mus. Comp. Zool., 8:
187, March.
= Liomys irroratus alleni (Coues, 1881).
See Goldman (1911: 56).
Holotype. MCZ 5889. Skin and skull Male.
Locality. (Mexico): San Luis Potosi, Rio Verde, Ha-
cienda Angostura. 26 Febniaiy 1878.
Collector. E. Palmer.
Condition. Sldn complete, skull withiji skin.
Type Series. Holotype only.
Comments. L. i. alleni was retained as a valid sub-
species by Hall (1981: 590).
Genus PEROGNATHUSW\e6-NeuW\e6,
1839
Perognathus longimembris bangs! Mearns,
1898
Bull. Amer. Mus. Nat. Hist., 10: 300, 31
August.
Holotype. B5304. Skin and skull. Adult female.
Locality. (United States): California, (Riverside
County), Colorado Desert, Palm Springs, 450 ft
(137 m). 13 April 1896.
Collector. E. C. Thurber. Original number 644.
Condition. Sldn complete. Skull partial (teeth sep-
arate from skull). Mandible disaiUculated.
Type Series. 2 paratypes; B5302, adult male, skin
and sloill; B5303, adult female, skin and skull.
Comments. Retained as a valid subspecies by Hall
(1981: 537).
Family DIPODIDAE Fischer de Waldheim,
1817
Genus ZAPUS Coues, 1875
Zapus hudsonius hardy! Batchelder, 1 899
Proc. New England Zool. Club, 1: 5, 8
February.
=Zapus hudsonicus acadlcus (Dawson,
1856). See Krutzsch (1954: 432).
Holotype. MCZ 41681. Skin and skull. Adult female.
Locality. (United States): Maine, Hancock County,
Mount Desert Island. 24 August 1898.
Collector C. F. Batchelder. Original number 1597.
Condition. Skin and skull complete.
Tijpe Series. There is a series of paratypes in tlie
MCZ.
Zapus hudsonius ladas Bangs, 1 899d
Proc. New England Zool. Club, 1: 10, 28
February.
Holotype. B4169. Skin and skull. Adult female.
Locality. (Canada): Labrador, Hamilton Inlet, Ri-
goulette. 18 July 1895.
Collector. C. H. Goldthwaite. Original number 2.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a large series of paratypes,
most of which are in the MCZ.
Comments. Retained as a valid subspecies by Hall
(1981: 843).
Zapus orahus Preble, 1899
N. Amer. Fauna, 15: 29, 8 August.
=Zapus trinotatus orarlus Preble, 1899.
See Hooper (1944: 67).
Holotype. B250. Skin and skull. Adult male.
Locality. (United States): California, Marin Coun-
ty, Point Reyes. 14 May 1893.
Collector C. A. Allen. Original number 618.
Condition. Skin and skull complete.
Type Series. 3 paratypes, not in MCZ; mentioned
by locality in original description.
Comments. Z. t. orarius was retained as a valid sub-
species by Hall (1981: 846) and Gannon (1988: 1).
Family MURIDAE llliger, 1811
Subfamily ARVICOLINAE Gray, 1821
Genus CH/OA/O/V/yS Miller, 1908
Hypudaeus nivlcola Schinz, 1845
Syst. Verzeichniss Saugethiere Synopsis
Mammalium, 2: 236.
= Chionomys nivalis (Martins, 1842). See
Musser and Carleton (1993: 507).
Syntype. MCZ 1291. Skin. Juvenile female.
Locality. Switzerland: St. Gotthard.
Collector Received from L. Agassiz,
Condition. Sldn complete.
Type Series. Schinz wrote that he had e.xamined 16
specimens, "junge und alte."
Comments. G. M. Allen discussed the somewhat
uncertain tyjie status of this specimen (1931: 263).
The original label bears the following: "Hypodaeus
[sic] nivicola juv. 9 sp. nov. du St. Gotthard et du
Faulhorn." Louis Agassiz, a student of Schinz's, de-
posited the specimen in the MCZ.
Type Specimens of Recent Mammals • Helgen and McFadden 113
Genus CLETHRIONOMYSJWes'wJs, 1850 Genus EOTHENOMYS Miller, 1896
Evotomys proteus Bangs, 1 897g
In Bailey, Proc. Biol. Soc. Washington,
11: 137, 13 May.
= Clethrionomys gapperi proteus (Bangs,
1897). See Jackson (1938: 433).
Holotype. B4081. Sldn and skull. Adult female.
Locality. (Canada): Labrador, Hamilton Inlet. 27
August 1895.
Collector. C. H. Goldthwaite. Original number 10.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a large series of paratypes in
tlie MCZ.
Comments. C. g. proteus was retained as a valid
subspecies by Hall (1981: 783).
Genus DICROSTONYX Gloger, 1841
Dicrostonyx chionopaes G. M. Allen,
1914b
Proc. New England Zool. Club, 5: 62, 9
April.
= Dicrostonyx torquatus chionopaes G. M.
Allen, 1914. See Ognev (1948: 507).
Holotype. MCZ 15263. Skin and skull. Adult male.
Locality. U.S.S.R. ( = Russian Federation): eastern
Siberia, Nijni Kolymsk (Nizhnekolymsk), near
mouth of Kolyma River. 15 October 1911.
Collector J. Koren. Original number 257.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Dicrostonyx exsuKj. M. Allen, 1919a
Bull. Mus. Comp. Zool., 62: 532,
February.
Holotype. MCZ 11885. Skin and skull. Adult male.
Locality. (United States): (Alaska), St. Lawi-ence Is-
land. 24 June 1913.
Collector J. Dixon. Original number 3267.
Condition. Skin complete. Skull partial (basioccip-
ital and right tympanic bulla missing). Mandible
disarticulated.
Type Series. 3 paratypes; MCZ 11883, skin and
skull, female; MCZ 11884, skin and skull, female;
USNM 232007, skin and skull, female.
Comments. Considered a valid species by Musser
and Carleton (1993: 510) but included in D. groen-
landicus by Nov^^ak (1999: 1479).
Craseomys aquilus G. M. Allen, 1912b
Mem. Mus. Comp. Zool, 40: 216,
August.
= Eotlienomys eva eva (Thomas, 1911).
See G. M. Allen (1940: 837).
Holotype. MCZ 7190. Skin and skull. Adult male.
Locality. China: Hupeh ( = Hubei), Showlungtan.
17 May 1907.
Collector W. R. Zappey Original number 10.
Condition. Sldn complete. Skull partial (hole in left
parietal, occiput missing).
Type Series. 5 paratypes; MCZ 7189, 7191-7194,
7196; all represented by sldn and skull, all female.
Microtus (Eothenomys) aurora G. M.
Allen, 1912b
Mem. Mus. Comp. Zool., 40: 211,
August.
= Eotlienomys melanogaster aurora (G. M.
Allen, 1912). See Hinton (1923: 149).
Holotype. MCZ 7788. Skin and skull. Male.
Locality. China: Hupeh ( = Hubei), Changyangh-
sieh. 2 Februaiy 1909.
Collector W. R. Zappey. Original number 372.
Condition. Skin and skull complete.
Type Series. 3 paratypes; MCZ 7185, sldn and skull,
female; MCZ 7186, skin and skull, male; MCZ
7188, skin and skull, male.
Comments. E. ni. aurora was retained as a valid
subspecies by Zhang et al. (1997: 229).
Microtus (Eotlienomys) mucronatus G. M.
Allen, 1912b
Mem. Mus. Comp. Zool., 40: 214,
August.
= Eotiienomys melanogaster melanogaster
(Milne-Edwards, 1871). See G. M. Allen
(1940: 806).
Holotype. MCZ 7789. Sldn and skull. Aduk female.
Locality. China: western Szechwan ( = Sichuan),
Tachiao, 12,000 ft (3,660 m). 11 August 1908.
Collector. W. R. Zappey. Original number 269.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; MCZ 7790, sldn and skull,
adult female (exchanged to the BMNH); MCZ
7791, skin and skuU, adult female; MCZ 7803, skin
and skuU, juvenile female.
114 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Genus LEMMUS Link, 1795
Lemmus paulus G. M. Allen, 1914b
Proc. New England Zool. Club, 5: 60, 9
April.
= Lemmus sibiricus chrysogasterJ. A.
Allen, 1903. See Ellerman and Morrison-
Scott (1951: 656).
Holotijpe. MCZ 15268. Skin and skull. Adult male.
Locality. (Russian Federation): northeastern Sibe-
ria, Kalascliowo, near mouth of Kolyma River. 22
June 1912.
Collector. J. Koren. Original number 152.
Condition. Skin complete. Skull partial (left man-
dibular ramus missing).
Ti/pe Series. 1 paratype; MCZ 15267, skin, female.
Genus M IC ROTUS Schrank, 1798
Arvicola breweri Bah6, 1858
Mammals, in Repts. Explor. Surveys
Railr. to Pacific, 8(1): 525, 14 July.
= Microtus breweri {Bai'cd, 1858). See
Miller (1896: 83).
Syntype. MCZ 4365. Body in alcohol. Formerly
USNM 2833.
Locality. (United States); Massachusetts, Nantuck-
et County, off Nantucket Island, Muskeeget ( = Mu-
skeget) Island. July 1856.
Collector. T. M. Brewer.
Condition. Alcoholic.
Type Series. There are 5 other syntypes (see Poole
and Schantz, 1942: 271-272), all of which were at
one time in the USNM.
Comments. Poole and Schantz (1942: 271) noted
that USNM 2829, a svntype of breweri, could not
be found.
Arvicola riparia var. longipilis Baird, 1858
Mammals, in Repts. Explor. Sin-vevs
Railr. to Pacific, 8(1): 524, 14 July'
= Microtus pennsylvanicus pennsylvanicus
(Ord, 1815). See Bailey (1900: 16).
Syntype. MCZ 5292. Skin and skull.
Locality. (United States): Illinois, (Cook County),
West Northfield. Spring 1855.
Collector R. Kennicott. Formerly USNM 745.
Condition. Sldn complete. Skull partial (part of pa-
rietal, both tympanic bullae and palatine missing).
Mandible disarticulated.
Tijpe Series. See comments.
Comments. G. M. Allen (1931: 260), mentioned
that "although no individuals are mentioned [in
Baird's description], . . . [tliis] specimen was un-
doubtedly among those examined by Baird in pre-
paring his diagnosis and, therefore, is a cotype
[ = syntype]." Baird's specimens were collected by
Kennicott at West Northfield, Illinois, and by Hoy
at Racine, Wisconsin, and were apparently depos-
ited in the USNM. Lyon and Osgood (1909) and
Poole and Schantz (1942) make no mention of oth-
er sxii types.
Arvicola rufidorsum Baird, 1858
Repts. Explor. Surveys Railr. to Pacific,
8(1): 526, 14 July.
= Microtus pennsylvanicus pennsylvanicus
(Ord, 1815). See Bailey (1900: 16).
Holotype. MCZ 54372. Skin and skull.
Locality. (United States): Massachusetts, (Duke's
County), Martha's Vineyard, Holmes Hole.
Collector D. J. Wyman.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. This specimen has formerly been
BSNH 1949 and USNM 901.
Arvicola terraenovae Bangs, 1894a
Proc. Biol. Soc. Washington, 9: 129, 27
July.
= Microtus pennsylvanicus terraenovae
(Bangs, 1894). See Davis (1936: 290).
Holotype. B1104. Skin and skull. Adult male.
Locality. (Canada): Newfoundland, Codroy. 27 No-
vember 1893.
Collector E. Doane. Original number 4.
Condition. Sldn and sk-ull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. M. p. terraenovae was retained as a val-
id subspecies bv Hall (1981: 796).
Microtus chrotorrhinus ravus Bangs,
1898p
Proc. Biol. Soc. Washington, 12: 188, 16
November.
Holotype. B7951. Skin and skull. Adult male.
Locality. (Canada): Labrador, Strait of Belle Isle,
Black Bay. 15 July 1898.
Collector. E. Doane. Original number 4.
Condition. Skin and skull coiuplete. Mandible dis-
aiticulated.
Type Series. Paratype mateiial consists of Bangs'
series from the type locahty; B7952-7967, all rep-
resented by skin and skull, 10 females, 3 males, and
3 unsexed individuals. B7953 and B7955 v^ere ex-
changed to MVZ in 1937; B7964 was exchanged to
USNM in 1922.
Comments. Retained as a valid subspecies by Hall
(1981: 811) and Kirkland and Jannett (1982:' 1).
Type Specimens of Recent Mammals • Helaen and McFadden
115
Microtus enixus Bangs, 1896g
Amer. Nat., 30: 1051, 1 December.
= Microtus pennsylvanicus enixus Bangs,
1896. See Davis (1936: 290).
Holotype. B3973. Skin and skull. Adult female.
Locality. (Canada): Labrador, Hamilton Inlet. 15
July 1895.
Collector. C. H. Goldthwaite. Original number 4.
Condition. Skin and skull complete.
Type Series. There is a large series of parat\'j3es in
the MCZ.
Comments. M. p. enixus was retained as a valid
subspecies by Hall (1981: 79.3).
Microtus fontigenus Bangs, 1896d
Proc. Biol. Soc. Washington, 10: 48, 9
March.
= Microtus pennsylvanicus fontigenus
Bangs, 1896. See Miller (1897: 14).
Holotype. B3837. Skin and skull. Adult female.
Localitij. (Canada): Quebec, Lake Edward. 28 Sep-
tember 1895.
Collector. E. A. and O. Bangs. Original number 9.
Condition. Skin and skull complete. Mandible dis-
articulated.
Tijpe Series. 7 paratypes; B3838-B3844; all repre-
sented by sldn and skull, 2 females and 5 males.
Comments. M. p. fontigenus was retained as a valid
subspecies by Hall (1981: 793).
Microtus l<oreni G. M. Allen, 1914b
Proc. New England Zool. Club, 5: 64, 9
April.
= Microtus oeconomus koreni G. M. Allen,
1914. See Ellerman and Morrison-Scott
(1951: 706).
Holotype. MCZ 15213. Skin and skull. Adult female.
Locality. (Russian Federation): northeastern Sibe-
ria, Nijni Kolymsk ( = Nizhnekolymsk), near mouth
of Kolyma River. 1 November 1911.
Collector J. Koren. Original number 132.
Condition. Skin complete. Skull partial (left tym-
panic bulla broken, right mandibular ramus miss-
ing).
Type Series. There is a large series of paratypes in
the MCZ.
Microtus pennsilvanicus [sic] sliattucl<i
Howe, 1901
Proc. Portland Soc. Nat. Hist., 2: 201, 31
December.
= Microtus pennsylvanicus shattucki
Howe, 1901. See Wyman (1922: 162).
Holotype. MCZ 10011. Sldn and skull. Adult female.
Locality. (United States): Maine, Penobscot Bay,
near Long Island, Tvmible Down Dick Island. 10
July 1900.
Collector R. H. Howe and G. C. Shattuck. Original
number 31.
Condition. Skin and skull complete.
Type Series. There is a small series of paratypes in
the MCZ.
Comments. Wyman (1922: 166) considered M. p.
shattucki a synonym of M. p. pennsylvanicus, but
Hall (1981: 796) retained it as a separate subspe-
cies.
Microtus pennsylvanicus acadicus Bangs,
1897c
Amer. Nat., 31: 239, 1 March.
Holotype. B2155. Skin and skull. Adult female.
Locality. Canada: Nova Scotia, Digbv. 22 |uly
1894.
Collector O. Bangs.
Condition. Sldn and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. Retained as a valid subspecies by Hall
(1981: 792).
Microtus provectus Bangs, 1 908
Proc. New England Zool. Club, 4: 20, 6
March.
= Microtus pennsylvanicus provectus
Bangs, 1908. See Chamberlain (1954:
589).
Holotype. B9794. Skin and skull. Adult female.
Locality. (United States): Rhode Island, Block Is-
land. 5 August 1899.
Collector O. Bangs. Original number 11.
Condition. Sldn complete. Skull partial (tympanic
bullae broken). Mandible disarticulated and angu-
lar process chipped.
Type Series. There is a series of paratypes in the
MCZ.
Comments. M. p. provectus was retained as a valid
subspecies by Hall (1981: 796).
Genus MYOPUSmWer, 1910
Myopus thayeri G. M. Allen, 1914b
Proc. New England Zool. Club, 5: 58, 9
April.
^Myopus schisticolor thayeri G. M. Allen,
1914. See Ognev (1948: 526).
Holotype. MCZ 15264. Sldn and skull. Adult male.
Locality. (Russian Federation): northeastern Sibe-
ria, Yakutsk, Nijni Kolyinsk ( = Nizhnekolyinsk),
near mouth of Kolyma River 28 March 1912.
Collector J. Koren. Original number 264.
Condition. Skin and skull complete. Mandible dis-
articulated.
116 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Comment.^. 1 paratype; MCZ 15265, skin and skull,
male.
Genus ONDATRA Link, 1795
Fiber obscurus Bangs, 1894b
Proc. Biol. Soc. Washington, 9: 133, 15
September.
= Ondatra zibethicus obscurus (Bangs,
1894). See Cameron (1959: 85).
Holotype. B1155. Skin and skull. Adult female.
Locality. (Canada): Newfoundland, Codroy. 14
May 1894.
Collector. E. Doane. Original number 3.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a small series of paratypes in
the MCZ.
Comments. O. z. obscurus was retained as a valid
subspecies by Hall (1981: 827).
Fiber zibethicus aquilonius Bangs, 1899d
Proc. New England Zool. Club, 1: 11, 28
February.
= Ondatra zibettiicus aquilonius (Bangs,
1899). See Miller (1912:231).
Holotype. B3957. Skin and skull. Adult male.
Locality. (Canada): Labrador, Hamilton Inlet, Ri-
goulette. 15 August 1895.
Collector. C. H. Goldthwaite. Original number 11.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; B7974, sldn and skull,
adult female; B7975, sldn and skull, juvenile;
B8704, skull, juvenile.
Comments. O. z. aquilonius was retained as a valid
subspecies by Hall (1981: 825).
Fiber zibethicus rivalicius Bangs, 1895b
Proc. Boston Soc. Nat. Hist., 26: 541, 31
= Ondatra zibethicus rivalicius (Bangs,
1895). See Davis and Loweiy (1940:
212).
Holotype. B2719. Sldn and skull. Adult male.
Locality. (Canada): Louisiana, Plaquemines Parish,
Burbridge. 31 January 1895.
Collector. F. L. Small. Original number 1556/165.
Condition. Sldn and skull complete.
Type Series. 4 paratypes; B2720, skin and skull,
adult female; B2721, skin and skull, adult male;
B2883, skin and skull, male; B2884, sldn and skull,
male.
Comments. O. z. rivalicius was retained as a valid
subspecies by Hall (1981: 827).
Genus PHENACOMYS Mernam, 1889
Phenacomys celatus crassus Bangs,
1900c
Proc. New England Zool. Club, 2: 39, 20
September.
= Phenacomys ungava crassus Bangs,
1900. See A. B. Howell (1926: 27).
Holotype. B3959. Skin and skull. Adult male.
Locality. (Canada): Labrador, Hamilton Inlet, Ri-
goulette. 15 August 1895.
Collector C. H. Goldthwaite. Original number 1.
Condition. Sldn and skull complete.
Type Series. There is a small series of paratypes,
most of which are no longer in the MCZ.
Comments. In the original description, "3946" is a
misprint for 3964.
Genus SYNAPTOMYS Ba\r6, 1858
Synaptomys fatuus Bangs, 1 896d
Proc. Biol. Soc. Washington, 10: 47, 9
March.
= Synaptomys cooperi cooperi Baird,
1858. See Wetzel (1955:8).
Holotype. B3857. Skin and skull. Adult female.
Locality. (Canada): Quebec, Lake Edward. 28 Sep-
tember 1895.
Collector E. A. and O. Bangs. Original number 3.
Condition. Sldn and skull complete.
Type Series. 8 paratypes; B3854-B3856, B3858-
B3862; all represented by sldn and skull, 3 females
and 5 males.
Synaptomys (Mictomys) innuitus
medioximus Bangs, 1 900c
Proc. New England Zool. Club, 2: 40, 20
September.
= Synaptomys borealis medioximus
Bangs, 1900. See A. B. Howell (1927: 9).
Holotype. B8852. Skin and skull. Adult male.
Locality. (Canada): Labrador, Strait of Belle Isle,
Lance ( = Lanse) an Loup. 15 April 1899.
Collector E. Doane. Original number 7.
Condition. Sldn complete. Skull slightly damaged
(hole in left parietal).
Type Series. 1 paratype; B3972, sldn and skull,
adult male.
Comments. S. b. medioximus was retained as a val-
id subspecies by Hall (1981: 834).
Type Specimens of Recent Mammals • Helaen and McFadden 117
Subfamily CRICETINAE Fischer de
Waldheim, 1817
Genus CANSUMYS G. M. Allen, 1928
Cansumys canus G. M. Allen, 1928
J. Mammal., 9: 245, 9 August.
Holotype. MCZ 23779. Skin and skull. Adult female.
Locality. China: southern Kansu (=Xinjiang),
Choni. 9 December 1925.
Collector. R. B. Ekvall. Original number 79.
Condition. Skin complete. Skull partial (left jugal
and occiput missing).
Type Series. 1 paratype; MCZ 23780, skin and
skull, juvenile male.
Comments. Type species o( Cansumys G. M. Allen,
1928. Considered a valid species by Musser and
Carleton (1993: 537) and Nowak (1999: 1423).
Known only by the type series and one additional
specimen, FMNH 36067.
Subfamily CRICETOMYINAE Roberts,
1951
Genus SACCOSTOMUS Peters, 1846
Saccostomus cricetulus G. M. Allen and
Lawrence, 1936
Bull. Mus. Comp. Zool., 79: 100, January.
= Saccostomus mearnsiHeWer , 1910. See
Hubert (1978: 51).
Holotype. MCZ 31475. Skin and skaill. Subadult male.
Locality. Uganda: Sabei District, due north of
Mount Elgon, south bank of Greek River ( = Kelim
River), 3,000 ft (915 m). 5 December 1933.
Collector. A. Loveridge.
Condition. Skin complete. Skull partial (right tym-
panic bulla missing). Mandible disarticulated.
Type Series. 1 paratype; MCZ 31474, skin and
skull, adult female.
Subfamily DENDROMURINAE G. M.
Allen, 1939
Genus S7E/\ rO/WVS Peters, 1846
Steatomys pratensis nyasae Lawrence
and Loveridge, 1953
Bull. Mus. Comp. Zool, 110: 39, June.
Holotype. MCZ 44213. Skin and skull. Adult male.
Locality. Nyasaland ( = Malawi): foot of Mulanje
Mountain, Likabula River, 2,100 ft (641 m). 29 July
1948.
Collector. A. Loveridge.
Condition. Sldn and skull complete.
Type Series. 14 paratypes; MCZ 44214-44216,
44218-44228; all represented by skin and skull, 9
males and 5 females.
Comments. Retained as a valid subspecies by An-
sell (1978: 77).
Subfamily GERBILLINAE Gray, 1825
Genus TATERA Lataste, 1882
Tatera flavipes G. M. Allen, 1914d
Bull. Mus. Comp. Zool., 5S: 331, July.
= Tatera valida /cemp/ Wroughton, 1906.
See Bates (1988: 277).
Holotype. MCZ 14491. Skin and skull. Adult female.
Locality. Sudan: Blue Nile, north of (Er) Roseires,
Aradeiba. 22 Januaiy 1913.
Collector G. M. Allen and J. C. PhilHps. Original
number 69.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Tatera sororG. M. Allen, 191 4d
Bull. Mus. Comp. Zool, 58: 333, July
= Tatera valida /cemp/ Wroughton, 1906.
See Bates (1988: 277).
Holotype. MCZ 14492. Skin and skull. Adult female.
Locality. Sudan: Blue Nile, Fazogli. 16 January
1913.
Collector CM. Allen and J. C. Phillips. Original
number 53.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; MCZ 14493, skin and
skull, juvenile male.
Genus TATERILLUS Thomas, 1910
Taterillus melanops G. M. Allen, 1912a
Bull. Mus. Comp. Zool., 54: 446, April.
= Taterillus harringtoni (Thomas, 1906).
See Musser and Carleton (1993: 563).
Holotype. MCZ 8132. Skin and skull. Male.
Locality. British East Africa ( = Kenya): arid plains
by the Meru River. 11 August 1909.
Collector G. M. Allen. Original number 108.
Condition. Skin and skull complete.
Tijpe Series. Holotype only.
Subfamily MURINAE llliger, 1811
Genus APODEMUS Kaup, 1829
Apodemus mystacinus euxinus G. M.
Allen, 1915b
Bull. Mus. Comp. Zool, 59: 11,
Februaiy.
= Apodemus mystacinus mystacinus
(Danford and Alston, 1877). See Corbet
(1978: 133).
Holotype. MCZ 14887. Skin and skull. Male.
Locality. Asia Minor (=Turkey): (Trabzon), near
118 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Trebizond (=Trabzon), Scalita, 1,000 m. 25 No-
vember 1905.
Collector. A. Robert. Original number 2189.
Condition. Skin complete. Skull partial (right and
left jugal missing, right tympanic bulla separated
but present, right and left squamosal missing).
Mandible disarticulated.
Tijpe Series. HolotN-pe only.
Genus DASYMYS Peters, 1875
Dasymys incomtus alleni Lawrence and
Loveridge, 1953
Bull. Mus. Comp. ZooL, 110: 53, June.
Holotijpe. MCZ 26322. Skin and skull. Adult male.
Locality. Tanganyika Territoiy (=Tanzania):
(Mbeya), Ilolo, near Rungwe Mountains, 4,600 ft
(1,403 m). 31 March 1930.
Collector. A. Loveridge.
Condition. Skin and skull complete. Mandible dis-
articulated and right coronoid process chipped.
Type Series. 11 paratypes, all represented by sldn
and skull, in the MCZ.
Comments. Retained as a valid subspecies by An-
sell (197S: S3).
Genus GRAMMOMYS Jhomas, 1915
Thamnomys ochraceus G. M. Allen,
1912a
Bull. Mus. Comp. ZooL, 54: 442, April
1912.
= Grammomys macmillani (Wroughton,
1907). See Musser and Carleton (1993:
594).
Holotype. MCZ 8126. Skin and skull. Adult male.
Locality. British East Africa ( = Kenya): Meru Riv-
er, near junction with northern Guaso Nyiro
( = Ewaso Ngiro). 8 August 1909.
Collector. G. M. Allen. Original number 103.
Condition. Skin and skull complete.
Type Series. Holotype only.
Genus HyLO/WySCL/S Thomas, 1926
Hylomyscus alleni simus G. M. Allen and
Coolidge, 1930
7/7 Strong, The African Republic of
Liberia and the Belgian Congo, 2: 599,
October.
= Hylomyscus alleni (Waterhouse, 1838).
See Heim de Balsac and Aellen (1965:
722).
Holotype. MCZ 24028. Skin and skull. Adult male.
Locality. Liberia: Merikay. 13 September 1926.
Collector G. M. Allen and H. J. Coolidge, Jr. Orig-
inal number 97.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a series of paratypes in the
MCZ.
Genus MELO/W/S Thomas, 1922
Melomys levipes stevensi Rummler, 1935
Z. Saugetierkunde, 10: 109, 31
December.
= Melomys mollis Thomas, 1913. See
Flanneiy (1990: 226).
Holotype. MCZ 29890. Skin and skull. Adult male.
Locality. Papua New Guinea: Morobe, Mount Mis-
im ( = Missim), 6,700 ft (2,044 m). 17 April 1933.
Collector H. Stevens.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 11 paratypes; MCZ 29889, 29892-
29899, 29901; all represented by sldn and skull, 4
females and 7 males.
Melomys moncktoni alleni Rummler, 1935
Z. Saugetierkunde, 10: 112, 31
December.
= Melomys rubex alleni Rummler, 1935.
See Tate (1951: 299).
Holotype. MCZ 29902. Skin and skull. Adult female.
Localiti/. Papua New Guinea: Morobe, Mount Mis-
im ( = Missim), 6,700 ft (2,044 m). 17 April 1933.
Collector H. Stevens.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratvpe; MCZ 29900, skin and
skull, adult male.
Comments. Flannery (1995a: 308—309) did not list
subspecies of M. nibex, noting, "It is a veiy variable
species, and a taxonomic revision is needed in or-
der to clarify the taxonomic status of many popu-
lations."
Genus MUS Linnaeus, 1758
Leggada bufo ablutus G. M. Allen and
Loveridge, 1942
Bull. Mus. Comp. ZooL, 89: 199,
Februaiy.
= Mus bufo (Thomas, 1906). See Musser
and Carleton (1993: 622).
Holotype. MCZ 40745. Skin and skull. Adult male.
Locality. Belgian Congo ( = Democratic Republic
of Congo): Kivu, Lake Kivu, Idj\\d Island, Upper
Mulinga, 6,500 ft (1,983 m). 24 February 1939.
Collector A. Loveridge.
Type Specimens of Recent Mammals • Helgen and McFadden 119
Condition. Skin and skull complete. Mandible dis-
articulated.
Tijpe Series. 2 paratypes; MCZ 40746, skin and
skull (no longer in MCZ); MCZ 40747, skin and
skull, female.
Leggada gerbillus G. M. Allen and
Loveridge, 1933
Bull. Mus. Comp. ZooL, 75: 112,
Febriiaiy.
= Mus tenellus (Thomas, 1903). See
Musser and Carleton (1993: 629).
Holotype. MCZ 26586. Skin and skull. Adult male.
Locality. Tanganyika Territoiy (= Tanzania): Do-
doma, Ugogo, 3,700 ft (1,129 m). 23 December
1929.
Collector. A. Loveridge.
Condition. Sldn complete. Skull partial (right jugal
missing). Mandible disarticulated and angular pro-
cess of right ramus chipped.
Type Series. Holotype only.
Mus bactrianus tantillus G. M. Allen, 1927
Amer. Mus. Novitates 270: 9, 31 May.
= Mus musculus tantillus G. M. Allen,
1927. See Marshall (1977: 214).
Holotype. MCZ 23476. Skin and skull. Adult female.
Locality. China: Sze-chuan ( = Sichuan), Wanhsien
(=Wanxian). 14 November 1921.
Collector W. W. Granger.
Condition. Skin complete. Skull partial (6 frag-
ments). Mandible disarticulated.
Type Series. There is a series of paratypes in the
AMNH, and 1 paratype in the MCZ; MCZ 23475
(formerly AMNH 56403), skin and skull, male.
Comments. This specimen was formerly AMNH
56416. The original description lists AMNH 56413
as the holotype, but this specimen is a juvenile, not
an adult female, as Allen stated. The collection date
and measurements of the holotype in the original
description fit MCZ 23476 rather than any other
specimens in the type series at the AMNH, which
are all immatures (Lawrence 1993: 136).
Genus /W/O/W/S Thomas, 1915
Praomys fumatus oweni Setzer, 1 956
Proc. U.S. Nat. Mus., 106: 525, 28
November.
= Myomys fumatus (Peters, 1878). See
Musser and Carleton (1993: 631).
Holotype. MCZ 45883. Skin and skull. Adult male.
Locality. Anglo-Egyptian Sudan ( = Sudan): Equa-
toria Province, Torit District, Murukurun, 50 miles
(80.5 km) east of Torit, 2,000 ft (610 m). 9 May
1950.
Collector J. S. Owen. Original number 1030.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Genus A//l//\/EA/rE/? Marshall, 1976
Epimys zappeyi G. M. Allen, 1912b
Mem. Mus. Comp. ZooL, 40: 225,
August.
^Niviventer confucianus (Milne-Edwards,
1871). See Musser and Carleton (1993:
633).
Holotype. MCZ 7607. Skin and skull. Adult male.
Locality. China: western Szechwan ( = Sichuan),
Washan Mountains (=Wu Shan), 9,000 ft (2,745
m). 26 October 1908.
Collector W. R. Zappey. Original number 305.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Genus OEA/OM/S Thomas, 1904
Oenomys hypoxanthus talangae Setzer,
1956
Proc. U.S. Nat. Mus., 106: 505, 28
November.
Holotype. MCZ 45315. Skin and skull. Adult female.
Locality. Anglo- Egyptian Sudan ( = Sudan): Equa-
toria Province, Imatong Mountains, Talanga Foi-
est, 3,000 ft (915 m). 10 July 1950.
Collector J. S. Owen. Original number 1338.
Condition. Skin and skull complete.
Type Series. 1 parats-pe; MCZ 45284, skin and
skuU, male.
Genus PR/A 0/WVS Thomas, 1915
Allen
Praomys tullbergi melanotus G.
and Loveridge, 1933
Bull. Mus. Comp. ZooL, 75: 106,
February.
= Praomys delectorum (Thomas, 1910).
See Musser and Carleton (1993: 642).
Holotype. MCZ 26287. Skin and skull. Adult male.
Locality. Tanganyika Territoiy (=Tanzania): north-
west end of Lake Nyasa, Poroto Mountains, Nyam-
wanga, 6,400 ft (1,952 m). 21 March 1930.
Collector. A. Loveridge.
Condition. Skin complete. Skull partial (left tym-
panic bulla missing). Mandible disarticulated.
Type Series. 23 paratypes; MCZ 26259, 26285,
26286, 26288-26293, 26295-26297, 26387-26394,
26411, 26497, 26498; all represented by skin and
skull.
120 Bulletin Museum of Comparative Zoologi/, Vol. 157, No. 2
Comments. Van der Straeten and Dieterlen (1987:
9) and Van der Straeten and Dudu (1990: 81) treat-
ed melanotus as a species within the Praomijs de-
lectorum species complex.
Genus PSEUDOHYDROMYS Rummler,
1934
Pseudohydromys murinus Rummler, 1 934
Z. Saugetierkunde, 9: 48, 30 December.
Holotype. MCZ 29904. Sldn and skull. Adult male.
Locality. Papua New Guinea, Morobe, Mount Mis-
im ( = Missim), 7,000 ft (2,135 m). 8 March 1933.
Collector. H. Stevens.
Condition. Sldn complete, with small bald spots.
Skull complete. Mandible disarticulated.
Type Series. Holotype only.
Comments. Type species of Pseudohydromys Rii-
mmler, 1934. Considered a valid species by Flan-
nery (1995a: 255), Musser and Carleton (1993:
644), and Nowak (1999: 1612).
Genus STENOMYS Ihomas, 1910
Stenomys niobe stevensi Rummler, 1935
Z. Saugetierkunde, 10: 117, 31
December.
= Stenomys niobe niobe (Thomas, 1906).
See Flamieiy (1995a: 339).
Holotype. MCZ 29915. Skin and skull. Adult male.
Locality. Papua New Guinea: Morobe, Mount Mis-
im ( = Missim), 7,000 ft (2,135 m). 16 Januaiy 1933.
Collector. H. Stevens. Original number 8.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. Taylor et al. (1982: 193) first synony-
mized .stevensi with niobe, under the genus Rattus.
Stenomys verecundus mollis Rummler,
1935
Z. Saugetierkunde, 10: 116, 31
December.
Holotype. MCZ 29905. Skin and skull. Adult female.
Locality. Papua New Guinea: Morobe, Mount Mis-
im ( = Missim), 5,850 ft (1,784 m). 14 April 1933.
Collector. H. Stevens.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Flan-
neiy (1995a: 346).
Genus THAMNOMYS Thomas, 1 907
Thamnomys venustus kivuensis G. M.
Allen and Loverldge, 1942
Bull Mus. Comp. Zool, 89: 192,
Februaiy.
Holotype. MCZ 39151. Skin and sk-ull. Adult female.
Locality. Belgian Congo ( = Democratic Republic
of Congo): (Sud-Kivu), Lake Kivu, Idjwi Island,
Upper Mulinga, 6,500 ft (1,983 m). 2 March 1939.
Collector. A. Loveridge.
Condition. Skin complete. Skull partial (separated
between nasal and frontal). Right and left angular
processes of mandible chipped.
Tiipe Series. Holotype only.
Subfamily NESOMYINAE Forsyth Major,
1897
Genus NESOMYS Peters, 1870
Nesomys lambertoni G. Grandidier, 1930c
Bull. Acad. Malgache, n. sen, 11: 95 (for
1928).
Holotype. MCZ 45941. Skull and postcranial skele-
ton. Adult inale.
Locality. Malagasy Republic ( = Madagascar): west
coast, vicinity of Maintirano.
Collector No collection data available. Received by
G. Grandidier from Academic Malgache, 1928.
Condition. Skull partial (left zygomatic arch miss-
ing, right jugal missing, several small holes). Post-
cranial skeleton complete (small foot bones pre-
svHTiably preserved in skin).
Type Series. 2 paratopes, MCZ 45933 and 45934,
poorly prepared skins (■with partial skulls within),
both male.
Comments. The skin belonging to the holotype is
in the MNHN, Paris (M. Carleton, personal com-
munication). Considered a valid species by Nowak
(1999: 1434).
Subfamily OTOMYINAE Thomas, 1897
Genus OTOMYS F. Cuvier, 1824
Otomys anchietae lacustris G. M. Allen
and Loveridge, 1933
Bull. Mus. Comp. Zool., 75: 120,
Februaiy.
= Otomys lacustris G. M. Allen and
Loveridge, 1933. See Dieterlen and Van
der Straeten (1992: 385).
Holotype. MCZ 26358. Skin and sk-ull. Adult female.
Locality. Tanganyika Territoiy (=Tanzania): north
end of Lake Nyasa, Ukinga Mountains, Madehani,
7,000 ft (2,135 m). 21 Februaiy 1930.
Type Specimens of Recent Mammals • Helgen and McFadden 121
Collector. A. Loveridge.
Condition. Skin complete. Skiill partial (left tym-
panic bulla damaged). Mandible disarticulated and
left coronoid process chipped.
Type Series. 17 paratypes; MCZ 26326, 26344-
26351, 26353-26357, 26359, 26654, 26658; all rep-
resented by skin and skull.
Comments. Included in O. anchietae by Musser
and Carleton (1993: 680) but treated as a full spe-
cies by Nowak (1999: 1439).
Otomys barbouri Lawrence and Loveridge,
1953
BulL Mus. Comp. ZooL, 110: 63, June.
Holotijpe. MCZ 31369. Skin and skull. Adult male.
Locality. Uganda: Mount Elgon, Kaburomi,
1°14'N, 34°31'E, 10,500 ft (3,203 m). 28 December
1933.
Collector. A. Loveridge.
Condition. Skin and skull complete.
Type Series. 9 paratypes; MCZ 31371-31372,
31376, 31421-31425, 31438 (4 males, 4 females,
and 1 unsexed individual).
Comments. Considered a valid species by Dieter-
len and Van der Straeten (1992: 385) and Nowak
(1999: 1439) but included in O. anchietae by Mus-
ser and Carleton (1993: 680).
Otomys uzungwensis Lawrence and
Loveridge, 1953
Bull Mus. Comp. ZooL, 110: 61, June.
= Otomys typus uzungwensis Lawrence
and Loveridge, 1953. See Hanney (1965:
626).
Holotype. MCZ 26645. Sldn and skull. Adult female.
Locality. Tanganyika Territory (=Tanzania): Iringa
District, Uzungwa Mountains, Dadaga, 6,000 ft
(1,830 m). 31 December 1929.
Collector. A. Loveridge.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. O. t. uzungwensis was retained as a val-
id subspecies bv Ansell (1978: 78).
Subfamily SIGMODONTINAE Wagner,
1843
Genus ISTHMOMYS Hooper and IVIusser,
1964
Megadontomys flavidus Bangs, 1902b
Bull Mus. Comp. ZooL, 39: 27, ApriL
= lsthmomys flavidus (Bangs, 1902). See
Hooper and Musser (1964: 12).
Holotype. MCZ 10331. Skin and skull. Adult male.
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo-
quete), 4,000 ft (1,220 m). 12 April 1901.
Collector W. W. Brown, Jr. Original number 28.
Condition. Skin complete, widi small bald spots.
Skull complete. Mandible disarticulated.
Type Series. There is a large series of paratypes in
the MCZ.
Comments. Type species oihthmomys Hooper and
Musser, 1964. I. flavidus was considered a valid
species by Musser and Carleton (1993: 706) and
Nowak (1999: 1357).
Genus NECTOMYS Peters, 1861
Nectomys squamipes amazonicus
Hershkovitz, 1944
Misc. Pub., Mus. ZooL, Univ. Mich., 58:
47, 4 Januaiy.
Holotype. MCZ 30820. Skin and skull. Male.
Locality. Brazil: (Para), Rio Tapajos, Tauary (=Tau-
ari). 23 January 1934.
Collector A. M. Olalla. Original number 7312.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Hershkovitz examined 28 specimens,
in the MCZ, AMNH, and FMNH.
Comments. Retained as a valid subspecies by Er-
nest (1986: 1).
Genus NEOTOMA Say and Ord, 1825
Neotoma abbreviata Goldman, 1909
Proc. BioL Soc. Washington, 22: 140, 25
June.
= Neotoma lepida abbreviata Goldman,
1909. See Burt (1932: 182).
Holotype. MCZ 12260. Skin and skull. Adult male.
Locality. (Mexico): Lower ( = Baja) California, San
Francisco Island. 22 February 1909.
Collector. W. W. Brown, Jn Original number 22.
Condition. Skin and skull complete.
Type Series. 9 paratypes; MCZ 12256-12259,
12261—12265; all represented by skin and skull, 5
females and 4 males. MCZ 12265 is now in the
USNM.
Comments. N. I. abbreviata was retained as a valid
subspecies by Hall (1981: 755).
Neotoma bella Bangs, 1899]
Proc. New England ZooL Club, 1: 66, 31
July.
= Neotoma lepida lepida Thomas, 1893.
See Goldman (1932: 62).
Holotype. B5308. Skin and skull. Adult male.
Locality. (United States): California, Riverside
County, Palm Springs. 12 April 1896.
Collector. E. C. Thurber Original number 623.
Condition. Skin and skull complete. Mandible dis-
articulated and left airgular pr'ocess chipped.
122 Bulletin Museum of Comparative Zoologi/, Vol. 157, No. 2
Type Series. Holotype only.
Neotoma distincta Bangs, 1903a
Proc. Biol. Soc. Washington, 16: 89, 25
June.
= Neotoma mexicana distincta Bangs,
1903. See Hall (1955: 329).
Holotype. B9819. Skin and skull. Adult male.
Locality. Mexico: Vera Cruz, near Jalapa, Texolo. 8
March 1899.
Collector. S. N. Rhoads. Original number 295.
Condition. Skin and skull complete.
Type Series. 4 paratypes; B9818, sldn and skull,
male, B9820, skin and skull, female; B9821, skin
and skull, male (exchanged to USNM); B9822, skin
and skull, male.
Comments. N. m. distincta was retained as a valid
subspecies by Hall (1981: 761) and Cornely and
Baker (1986: 1).
Neotoma floridana rubida Bangs, 1 898b
Proc. Boston Soc. Nat. Hist., 28: 185, 15
March.
Holotype. B2872. Skin and skull. Adult male.
Locality. (United States): Louisiana, Terrebonne
Parish, Gibson. 4 April 1895.
Collector F. L. Small. Original number 1751.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 2 specimens are mentioned by num-
ber in the original description: B2871, sldn and
skull, adult male, and B2873, skin and skull, adult
female.
Comments. Retained as a valid subspecies by Hall
(1981: 749).
Genus NYCTOMYS 6e Saussure, 1860
Nyctomys nitellinus Bangs, 1 902b
Bull. Mus. Comp. ZooL, 39: 30, April.
= Nyctomys sumichirasti nitellinus Bangs,
1902. See Goldman (1916: 155).
Holotype. MCZ 10249. Skin and skull. Adult female.
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo-
quete), 4,000 ft (1,220 m). 8 Februaiy 1901.
Collector. W. W. Brown, Jr. Original number 119.
Condition. Skin and skull complete.
Type Series. 5 paratypes; MCZ 10245, skin and
skull, adult male; MCZ 10246, skin and skull, adult
male; MCZ 10247, adult female, skin and skull;
MCZ 10248, skin and skull, adult female; MCZ
10250, sldn and skull, juvenile female.
Comments. N. s. nitellinus was retained as a valid
subspecies by Hall (1981: 630).
Genus OECOMYS Thomas, 1906
Oecomys trabeatus G. M. Allen and
Barbour, 1923
Bull. Mus. Comp. ZooL, 65: 262,
Februaiy.
= Oecomys bicolor trabeatus G. M. Allen
and Barbour, 1923. See comments.
Holotype. MCZ 19837. Skin and skull. Male.
Locality. Panama: Rio Jesusito. 10 April 1922.
Collector. T. Barbour and W. S. Brooks. Original
number 2027.
Condition. Skin complete. Skull partial (right zy-
gomatic arch broken). Mandible disarticulated.
Type Sei-ies. Holotype only.
Comments. Following Hershkovitz (1960: 533), O.
b. trabeatus was retained as a valid subspecies by
Hall (1981: 619) under the genus Oryzomys. The
use of Oecomys as a generic name follows Musser
and Carleton (1993: 715).
Oryzomys flavicans illectus Bangs, 1898k
Proc. Biol. Soc. Washington, 12: 164, 10
August.
= Oecomys flavicans illectus (Bangs,
1898k). See comments.
Holotype. B8101. Sldn and skull. Adult female.
Locality. Colombia: Magdalena, Santa Marta
Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 24
March 1898.
Collector W. W. Bi^own, Jr. Original number 89.
Condition. Skin complete. Skull partial (right jugal
missing).
Tt/pe Series. There is a small series of paratypes in
the MCZ.
Comments. The use of Oecomys as a generic name
follows Musser and Carleton (1993: 715).
Genus OLIGORYZOMYS Bangs, 1 900
Oryzomys navus Bangs, 1899a
Proc. Biol. Soc. Washington, 13: 9, 31
januaiy.
= Oligoryzomys fulvescens (Saussure,
1860). See Carleton and Musser (1989:
70).
Holotype. B8107. Skin and skull. Adult male.
Locality. Colombia: Magdalena, Santa Marta
Mountains, Pueblo Viejo, 8,000 ft (2,440 m). 26
March 1898.
Collector W. W. Browni, Jr. Original number 154.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 9 paratypes; B8223-B8231; all repre-
sented by sldn and skull, except B8228 (skin only);
2 females and 7 males. B8224 is now in the USNM,
Type Specimens of Recent Mammals * Helpen and McFadden
123
and B8223, B8226, B8230, and B8231 are now in
the AMNH.
Coininents. Oryzoini/s navus is the t)'pe species of
Oligoryzomijs Bangs, 1899.
Oryzomys (Oligoryzomys) vegetus Bangs,
1902b
Bull. Mus. Comp. ZooL, 39: 35, April.
= Oligoryzomys vegetus (Bangs, 1902).
See Musser and Carleton (1993: 718).
Holotype. MCZ 10298. Skin and skull. Adult female.
Locality. Panama: Chiriqui, Volcan de Chiriqui,
Boquete, 4,000 ft (1,220 m). 16 April 1901.
Collector. W. W. Brown, Jr. Original number 304.
Condition. Skin complete. Skull partial (left jugal
missing). Mandible disarticulated.
Type Series. 12 paratypes; MCZ 10295 (now in
USNM), 10297, 10300-10306, 10308-10310, 2 fe-
males and 10 males, all represented by skin and
skull except for 10308 (skin only).
Comments. Considered a valid species by Musser
and Carleton (1993: 718) and Nowak (1999: 1368).
Carleton and Musser (1995) provided distribution-
al information and support of specific status for O.
vegetus.
Genus OR VZOM VS Baird, 1858
Oryzomys devius Bangs, 1902b
Bull. Mus. Comp. ZooL, 39: 34, April.
Holotype. MCZ 10324. Skin and skull. Adult female.
Locality. Panama: Chiriqui, Volcan de Chiriqui,
Boquete ( = Bajo Boquete), 5,000 ft (1,525 m). 29
January 1901.
Collector. W. W. Brown, jr. Original number 73.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 3 paraty^aes; MCZ 10325, skin and
skull, adult female; MCZ 10326, skin and skull,
adult female; MCZ 10340, sldn and skull, adult
male.
Comments. Considered a valid species by Musser
and Carleton (1993: 722) and Nowak (1999: 1366).
Oryzomys palustris coloratus Bangs,
1898b
Proc. Boston Soc. Nat. Hist., 28: 189, 15
March.
= Oryzomys palustris natator Chapman,
1893. See Wolfe (1982: 1).
Holotype. B4470. Skin and skull. Adult male.
Localitij. (United States): Florida, Monroe County,
Cape Sable. 17 April 1895.
Collector. C. L. Brownell. Original number 148.
Condition. Sldn and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Oryzomys rostratus carrorum Lawrence,
1947b
Proc. New England ZooL Club, 24: 101,
29 May.
Holotype. MCZ 41280. Skin, skull, and postcranial
skeleton. Adult male.
Locality. Mexico: Tamaulipas, Rio Soto la Marina,
Rancho Santa Ana, 8 miles (12.9 km) southwest of
Padilla. 21 December 1941.
Collector. G. H. Pournelle, Third University of
Florida Mexican Expedition. Original number 196.
Condition. Skin and skull complete. Mandible dis-
articulated. Postcranial skeleton partial (right fore-
limb, hindlimbs, pelvis, ribs, vertebral column).
Type Series. 2 paratypes; MCZ 41281, skin, skull,
and postcranial skeleton, female; MCZ 41282, sldn,
skull, and postcranial skeleton, male.
Comments. Retained as Oryzomys melanotis car-
rorum by Hall (1981: 614). Musser and Carleton
(1993: 724) regarded O. rostratus as distinct from
O. melanotis.
Genus PEROMYSCUS G\oger, 1841
Hesperomys gossypinus LeConte, 1853
Proc. Acad. Nat. Sci. Philadelphia, 6:
411, 25 October.
= Peromyscus gossypinus gossypinus
(LeConte, 1853). See Rhoads (1896b:
189).
Lectotype. MCZ 5275. Skin and skull. Adult male.
Locality. (United States): Georgia, Liberty County,
Riceboro, probably on the LeConte Plantation near
Riceboro. 13 September 1847.
Collector J. E. LeConte. Formerly USNM 546.
Condition. Skin complete. Skull partial (basioccip-
ital and tympanic bullae broken). Mandible disar-
ticulated.
Type Series. Poole and Schantz (1942: 320) listed
potential "cotypes," meaning syntypes. Osgood
(1909: 136) mentioned that a specimen, number
752 in the collection of the Academy of Natural
Sciences in Philadelphia, possibly possessed type
status, but this specimen was not mentioned by
Koopman (1976).
Comments. In the original description of gossypi-
nus, measurements were given for a single speci-
men only; however, in addition to this specimen,
LeConte made reference to the coloration of
"younger indixdduals." The measureinents given
correspond to MCZ 5275 rather than to any other
of LeConte "s specimens still in the USNM (see G.
M. Allen, 1931: 262; Poole and Schantz, 1942:
320). In addition, MCZ 5275 was the only speci-
men of Peromyscus gossypinus from Georgia men-
tioned in Baird's Mammals of North America
(1858: 469). Because LeConte did not specify a ho-
lotype, MCZ 5275, which seems to be the speci-
124 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
men discussed and described by measurement in
the original publication and the only adult speci-
men mentioned, is here designated as lectotype. P.
gossijpinus was considered a valid species by Mus-
ser and Carieton (1993: 730) and Nowak (1999:
1361).
Hesperomys leucopus arcticus Mearns,
1890
Bull. Amer. Mus. Nat. Hist., 2: 285, 21
Februaiy.
Name preoccupied by Hesperomys
arcticus Coues, 1 877
Peromyscus maniculatus borealis Mearns,
1911
Proc. Biol. Soc. Washington, 24: 102, 15
May. (Replacement name for
Hesperomtjs leucopus arcticus Mearns,
1890)
Holotype. MCZ 5555. Skin and skull. Adult male.
Locality. Canada: Northwest Territories, Mac-
Kenzie, Fort Simpson, 7 September 1859.
Collector. R. Kennicott. Original number 157. For-
merly USNM 4531.
Conclition. Skin complete. Skull partial (left jugal
and process of squamosal missing, right process of
squamosal missing, basioccipital and palatine miss-
ing). Mandible disarticulated and both angular pro-
cess missing.
Type Series. Holot)'pe only.
Comments. P. m. borealis was retained as a valid
subspecies by Hall (1981: 672).
Hesperomys sonoriensis nebrascensis
Coues, 1877
In Coues and J. A. Allen, Monographs N.
Amer. Rodentia, U.S. Geol. Geogr.
Survey Terr. Rep. Washington, 11: 79,
August.
= Peromyscus maniculatus nebrascensis
(Coues, 1877). See Osgood (1909: 75).
Sijntype. MCZ 5528. Skin and sk-ull.
Locality. Nebraska (now Wyoming): (Converse
County), Deer Creek. 19 Januaiy 1860.
Collector F. V. Hayden. Original no. 80. Formerly
USNM 4310.
Condition. Sldn complete. Skull partial (posterior
missing from parietal to palatine; both jugal and
process of squamosal broken). Left angular process
of mandible chipped, right side broken at cheek
teeth.
Type Series. 1 other syntype, present location un-
known, formerly in die USNM.
Comments. Hesperomys sonoriensis nebrascensis
Baird, 1858 is a nomen nudum. Osgood (1909: 78)
and Mearns (1911: 102) designated Deer Creek,
Nebraska, as the type locaHty. Jones (1958: 107-
111) reviewed the systematic histoiy oi nebrascen-
sis, and Jones and Mursaloglu (1961: 101-103) re-
ported the discovery of one of the two original syn-
types in the MCZ. The location of the other syn-
tN-pe, if it still exists, is unknown.
Peromyscus anastasae Bangs, 1898b
Proc. Boston Soc. Nat. Hist., 28: 195, 15
March.
= Peromyscus gossypinus anastasae
Bangs, 1898. See Osgood (1909: 141).
Holotype. B7179. Skin and skull. Adult female.
Locality. (United States): Florida, St. Johns Coun-
ty, Anastasia Island, Point Romo. 15 February
1897.
Collector O. Bangs. Original number 2.
Condition. Sldn complete, skull partial (premaxilla
to frontal in fragments). Right mandible missing,
left present.
Type Series. There is a small series of paratypes in
the MCZ.
Comments. P. g. anastasae was retained as a valid
subspecies by Hall (1981: 689).
Peromyscus bellus Bangs, 1896h
Proc. Biol. Soc. Washington, 10: 137, 28
December.
= Peromyscus attwateri ^. A. Allen, 1895.
See Schmidly (1973: 125).
Holott/pe. B5483. Skin and skull. Adult male.
Locality. (United States): Indian Territory
( = Oklahoma), (Adair County), Stilwell. 15 August
1896.
Collector T Surber. Original number 67.
Condition. Skin and sk-ull complete. Mandible dis-
articulated.
Type Series. 1 paratype; B5484, skin and skull,
adult female.
Peromyscus cacabatus Bangs, 1 902b
Bull. Mus. Comp. Zool, 39: 29, April.
= Peromyscus mexicanus (Saussure,
1860). See Huckaby (1980: 15).
Holotype. MCZ 10225. Sldn and skull. Adult female.
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo-
quete), 5,000 ft (1,525 m). 22 April 1901.
Collector. W. W. Brown, Jr. Original number 331.
Condition. Skin complete. Skull partial (left jugal
missing). Mandible disarticulated.
Type Series. There is a very large series of paraty-
pes in the MCZ.
Type Specimens of Recent Mammals • Helgen and McFadden 125
Peromyscus canadensis abietorum Bangs,
1896d
Proc. Biol. Soc. Washington, 10: 49, 9
March.
= Peromyscus maniculatus abietorum
Bangs, 1896. See Osgood (1909: 45).
Holotype. B2205. Skin and sknll. Adult female.
Locality. (Canada): Nova Scotia, James River. 8
August 1894.
Collector. C. H. Goldthw^aite.
Condition. Sldn complete. Skull partial (right jugal
missing).
Type Series. There is a series of paratypes in the
MCZ.
Comments. P. m. abietorum was retained as a valid
subspecies by Hall (1981: 670).
Peromyscus canadensis argentatus
Copeland and Church, 1906
Proc. Biol. Soc. Washington, 19: 122, 6
September.
= Peromyscus maniculatus argentatus
Copeland and Church, 1906. See Osgood
(1909: 46).
Holotype. MCZ 54627. Skin and skull. Adult male.
Locality. (Canada): New Brunswick, Grand Manan
Island, Grand Harbor. 19 September 1905.
Collector M. L. Church and M. Copeland. Original
number 168.
Condition. Skin and skull complete.
Type Series. There is a large series of paratypes in
the MCZ.
Comments. P. m. argentatus was retained as a valid
subspecies by Hall (1981: 671).
Peromyscus crinitus scitulus Bangs, 1899j
Proc. New England Zool. Club, 1: 67, 31
July.
= Peromyscus crinitus crinitus (Merriam,
1891). See Osgood (1909: 229).
Holotype. B9175. Skin and skull. Adult male.
Locality. (United States): Nevada, Douglas County,
Gardnerville. 13 July 1898.
Collector W. W. Price and R O. Simons. Original
number 1571.
Condition. Skin complete. Skull partial (right and
left jugal missing).
Type Series. 7 paratypes; B9177-B9180, B9183-
B9185; all represented by skin and skull, 4 females
and 3 males.
Peromyscus gossypinus nigriculus Bangs,
1896e
Proc. Biol. Soc. Washington, 10: 124, 5
November.
= Peromyscus gossypinus gossypinus
(LeConte, 1853). See Osgood (1909: 136).
Holotype. B2731. Sldn and skull. Adult female.
Locality. (United States): Louisiana, Plaquemines
Parish, Burbridge. 30 Januaiy 1895.
Collector F. L. Small. Original number 1547/156.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a veiy large series of paraty-
pes in the MCZ.
Peromyscus gossypinus palmarius Bangs,
1896e
Proc. Biol. Soc. Washington, 10: 124, 5
November.
Holotype. B3224. Skin and skull. Adult female.
Locality. (United States): Florida, Brevard County,
Oak Lodge on East Peninsula opposite Micco. 23
Februaiy 1895.
Collector O. Bangs.
Condition. Skin and skull conaplete.
Type Series. There is a very large series of paraty-
pes in the MCZ.
Comments. Retained as a valid subspecies by Hall
(1981: 690).
Peromyscus insulanus Bangs, 1898b
Proc. Boston Soc. Nat. Hist., 28: 196, 15
March.
= Peromyscus gossypinus anastasae
Bangs, 1898. See Osgood (1909: 141).
Holotype. B6438. Skin and skull. Adult male.
Locality. (United States): Georgia, Camden Coun-
ty, Cumberland Island, north end. 10 April 1897.
Collector W. W. Brown, Jr. Original number 804.
Condition. Skin complete. Skull partial (right jugal
missing) .
Type Series. There is a large series of paratypes in
the MCZ.
Peromyscus leucopus ammodytes Bangs,
1905a
Proc. New England Zool. Club, 4: 14, 28
Februaiy.
Holotype. B828. Skin and skull. Adult male.
Locality. (United States): Massachusetts, Barnsta-
ble County, Monomoy Island. 28 December 1893.
Collector G. S. Miller, Jr. and O. Bangs. Original
number 8.
Condition. Skin and skull complete.
126 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Type Series. There is a series of paratypes in the
MCZ.
Comments. Retained as a vaUd subspecies by Hall
(1981: 685).
Peromyscus leucopus fusus Bangs, 1 905a
Proc. New England Zool. Club, 4: 13, 28
Februaiy.
Holotijpe. B9737. Skin and skull. Adult male.
Localitij. (United States): Massachusetts, Dukes
County, Martha's Vineyard, W. Tisbury. 17 June
1899.
Collector. O. Bangs. Original number 1.
Condition. Skin and skull complete.
Type Series. There is a small series of paratypes in
the MCZ.
Comments. Retained as a valid subspecies by Hall
(19S1: 686).
Peromyscus oreas Bangs, 1 898f
Proc. Biol. Soc. Washington, 12: 84, 24
March.
Holotype. B3696. Skin and skull. Adult female.
Locality. Canada: British Columbia, near boundaiy
of Whatcom County, Washington, 49th parallel.
Mount Baker Range, 6,500 ft (1,983 m). 29 August
1896.
Collector A. C. Brooks. Original number 745.
Condition. Sldn and skull complete.
Type Series. There is a large series of paratypes in
the MCZ.
Comments. Considered a valid species by Musser
and Carleton (1993: 734) and Nowak (1999; 1361).
Peromyscus phasma Bangs, 1898b
Proc. Boston Soc. Nat. Hist., 28: 199, 15
March.
= Peromyscus poloniotus phasma Bangs,
1898. See Osgood (1909: 107).
Holotype. B7175. Skin and skull. Adult female.
Locality. (United States): Florida, St. Johns Coun-
ty, Anastasia Island, Point Romo. 14 Februaiy
1897.
Collector O. Bangs. Original number 3.
Condition. Skin complete. Skull partial (parietal,
tympantic bulla, left jugal, and process of squa-
mosal broken). Mandible disarticulated.
Type Series. There is a series of paratypes in the
MCZ.
Comments. P. p. phasma was retained as a valid
subspecies by Hall (1981: 668).
Peromyscus subgriseus arenarius Bangs,
1898b
Proc. Boston Soc. Nat. Hist., 28: 202,
March.
Name preoccupied by Peromyscus
eremicus arenarius Mearns, 1 896.
Peromyscus subgriseus baliolus Bangs,
1898n
Science, n. sen, 8: 215, 19 August.
(Replacement name for Peromyscus
subgriseus arenarius Bangs, 1898)
= Peromyscus polionotus polionotus
(Wagner, 1843). See Osgood (1909: 104).
Holotype. B5925. Skin and skull. Adult male.
Locality. (United States): Georgia, Scriven County,
Hursman's Lake (Savannah River), near Bascom.
15 December 1896.
Collector W. W. Brown, Jr Original number 60.
Condition. Skin and skull complete.
Type Series. There is a large series of paratypes in
the MCZ.
Peromyscus subgriseus rhoadsi Bangs,
1898b
Proc. Boston Soc. Nat. Hist., 28: 201, 15
March.
= Peromyscus polionotus riioadsi Bangs,
1898. See Osgood (1909: 107).
Holotype. B6980. Skin and skull. Adult male.
Locality. (United States): Florida, Hillsborough
County, head of Anclote River. 23 May 1895.
Collector. W. S. Dickinson.
Condition. Sldn and skvdl complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. P. p. rhoadsi was retained as a valid
subspecies by Hall (1981: 669).
Peromyscus texanus saturatus Bangs,
1897a
Amer. Nat. 31: 75, 1 Januaiy.
= Peromyscus maniculatus saturatus
Bangs, 1897. See Osgood (1909: 61).
Holotype. B2581. Skin and skull. Adult male.
Locality. Canada: British Columbia, Saturna Is-
land. 31 Januaiy 1894.
Collector W. C. Colt. Original number 128.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. P. m. saturatus was retained as a valid
subspecies by Hall (1981: 682).
Type Specimens of Recent Mammals • Helpen and McFadden
127
Genus REITHRODONTOMYS Giglioli,
1873
Reithrodontomys australis vulcanius
Bangs, 1902b
Bull. Mus. Comp. Zool, 39: 38, April.
= Reithrodontomys sumichrasti vulcanius
Bangs, 1902. See Hooper (1952: 83).
Holotijpe. MCZ 10281. Skin and skull. Adult male.
Locality. Panama: Chiriqui, Volcan de Chiriqui,
10,300 ft (3,142 m). 26 May 1901.
Collector. W. W. Brown, Jr. Original number 500.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. R. s. vulcanius was retained as a valid
subspecies by Hall (1981: 644).
Reittirodontomys creper Bangs, 1902b
Bull. Mus. Comp. Zool, 39: 39, April.
Holotype. MCZ 10284. Skin and skull. Adult female.
Locality. Panama: Chiriqui, Volcan de Chiriqui,
11,000 ft (3,355 m). 2 June 1901.
Collector. W. W. Brown. Original number 504.
Condition. Skin and skull complete. Mandible dis-
articulated. Left lower incisor missing.
Type Series. Holotype only.
Comments. Considered a valid species by Musser
and Carleton (1993: 740) and Nowak (1999: 1364).
Reithrodontomys lecontii impiger Bangs,
1898m
Proc. Biol. Soc. Washington, 12: 167, 10
August.
= Reithrodontomys humulis humulis
(Audubon and Bachman, 1841). See Hall
and Kelson (1959: 584).
Holotype. B7784. Skin and skull. Adult male.
Locality. (United States): West Virginia, Green-
briar County, White Sulphur Springs, 2,000 ft (610
m). 27 Febmaiy 1898.
Collector. T. Surber. Original number 466.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. 2 paratypes; B6931 (not B6932, as
stated in the original description), skin and skull,
adult male; B7785, skin and skull, adult female.
Genus SCOT/A/OM/S Thomas, 1913
AlKodon teguina apricus Bangs, 1 902b
Bull. Mus. Comp. Zool, 39: 40, April.
= Scotinomys teguina apricus (Bangs,
1902). See Thomas (1913: 409).
Holotype. MCZ 10236. Skin and skull. Adult female.
Locality. Panama: Chiriqui, Boquete ( = Baio Bo-
quete), 4,000 ft (1,220 m). 24 February 1901.
Collector. W W. Brown, Jr. Original number 192.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 4 paratypes; MCZ 10234, skin and
skull, adult male; MCZ 10235, skin, adult female;
MCZ 10237, skin and skull, adult female; MCZ
10238, sldn and skull, adult female.
Comments. S. t. apricus was retained as a valid sub-
species by Hooper (1972: 21) and Hall (1981: 734).
AI<odon xerampelinus Bangs, 1 902b
Bull. Mus. Comp. Zool., 39: 41, April.
= Scotionomys xerampelinus (Bangs,
1902). See Thomas (1913: 409).
Holotype. MCZ 10240. Skin and skull. Adult male.
Locality. Panama: Chiriqui, Volcan de Chiriqui,
10,300 ft (3,142 m). 26 May 1901.
Collector W W Brown, Jr. Original number 501.
Condition. Sldn complete. Skull partial (right jugal
missing). Mandible disarticulated.
Type Series. 2 paratypes; MCZ 10239, skin and
skull, adult male; MCZ 10241, skin and skull, adult
male.
Comments. S. xerampelinus was considered a valid
species by Musser and Carleton (1993: 746) and
Nowak (1999: 1355).
Genus SIGMODON Say and Ord, 1825
Sigmodon austerulus Bangs, 1 902b
Bull. Mus. Comp. Zool, 39: 32, April.
= Sigmodon hispidus borucae J. A. Allen,
1897. See Hall and Kelson (1959: 672).
Holotype. MCZ 10288. Sldn and skull. Adult male.
Locality. Panama: Chiiiqui, Volcan de Chiriqui,
10,000 ft (3,050 m). 1 June 1901.
Collector W W Brown, Jr. Original number 503.
Condition. Skin complete. Skull partial — according
to original description, "unfortunately it [the skull]
was broken by the trap directly across the orbits."
Tijpe Series. Holotype only.
Comments. For comments regarding the type lo-
cality, see Enders (1953a: 508-509), who suggested
that a native helper of W. W Brown, Jr, transplant-
ed a specimen of S. [h.] borucae, trapped during a
trip to his home in the lowlands near Boruca, to
the top of the Volcan, "after Brown had promised
a bottle of wine to any man who captured a Sig-
modon at that altitude."
Sigmodon hispidus exsputus G. M. Allen,
1920b
J. Mammal., 1: 236, 4 December.
Holotype. MCZ 18100. Skin and skull. Adult male.
Locality. (United States): Florida, Monroe County,
Big Pine Key 16 April 1920.
Collector W S. Brooks. Original number 1936.
128 Bidletm Museum of Comparative Zoology, Vol. 157, No. 2
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paraty][De; MCZ 18101, skin and
skull, adult male.
Comments. Retained as a valid subspecies by Hall
(1981: 737).
Sigmodon hispidus furvus Bangs, 1 903b
Bull. Mus. Comp. Zool., 39: 158, July.
Holottjpe. MCZ 10665. Skin and skull. Male.
Locality. Honduras: Atlantida, La Ceiba. 16 Janu-
ary 1902.
Collector. W. W. Brown, Ji". Original number 4.
Condition. Skin and skull complete.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Hall
(1981: 738). Sigmodon hispidus fei'vidus Lydekker,
1904 (p. 34) was a misspelling and thus accidental
renaming of Sigmodon hispidus furvtis Bangs,
1903.
Sigmodon hispidus spadicipygus Bangs,
1898b
Proc. Boston Soc. Nat. Hist., 28: 192, 15
March.
Holotype. B4477. Skin and skull. Adult female.
Locality. (United States): Florida, Monroe County,
Cape Sable. 18 April 1895.
Collector C. L. Brownell. Original number 153.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a series of paratypes in the
MCZ.
Comments. Retained as a valid subspecies by Hall
(1981: 739).
Sigmodon sanctae-martae Bangs, 1898q
Proc. Biol. Soc. Washington, 12: 189, 30
December.
= Sigmodon hispidus hirsutus (Burmeister,
1854). See Cabrera (1961: 508).
Holotype. B8105. Skin and skull. Adult male.
Locality. Colombia: (Magdalena), (Santa Marta
Mountains), Pueblo Viejo (not far from the source
of Rio Ancho), 8,000 ft (2,440 m). 23 March 1898.
Collector. W. W. Brown, Jr. Original number 73.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; B8250, skin and skull,
adult male.
Genus THOIVI ASOIVIYS Coues, 1884
Oryzomys (Erioryzomys) monochromos
Bangs, 1900b
Proc. New England Zool. Club, 1: 97, 23
February.
= Thomasomys monochromos (Bangs,
1900). See Gardner and Patton (1976:
26).
Holotype. B8348. Sldn and skull. Adult male.
Locality. Colombia: Magdalena, Santa Marta
Mountains, Paramo de Macotania, 11,000 ft (3,355
m). 7 March 1899.
Collector W. W. Brown, Jr. Original number 93.
Condition. Skin complete. Skull partial (right jugal
missing). Majidible disarticulated.
Type Series. 4 paratypes; B8248, juvenile male,
B8345, adult female, B8346, adult female, B8347,
adult male; all represented by sldn and skull.
Comments. Considered a valid species by Musser
and Carleton (1993: 750) and Nowak (1999: 1362).
Paramo de Macotama is in La Guajira (Paynter
1997: 261).
Genus ZYGODONTOMYS ^. A. Allen,
1897
Zygodontomys seorsus Bangs, 1901b
Amer. Nat., 35: 642, 22 August.
= Zygodontomys brevicauda cherriei (J. A.
Allen, 1895). See Voss (1991: 59).
Holotype. B8490. Skin and skull. Adult male.
Locality. Panama: Gulf of Panama, San Miguel Is-
land, Isla del Rey 5 May 1900.
Collector W. W. Brown, Jr Original number 147.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. Bangs based the description on a se-
ries of 68 specimens, most of which are still in the
MCZ.
Family PEDETIDAE Gray, 1825
Genus PEDETES llliger, 1811
Pedetes cafer taborae G. M. Allen and
Loveridge, 1927
Proc. Boston Soc. Nat. Hist., 38: 438, 23
December.
= Pedetes capensis (Forster, 1778). See
Misonne (1974: 8).
Holotype. MCZ 23080. Skin and skull. Adult female.
Locality. Tanganyika Territory (=Tanzania): Ta-
bora, Unyamwezi, 4,000 ft (1,220 m). 18 November
1921.
Collector A. Loveridge. Original number R6915.
Type Specimens of Recent Mammals • Helgen and McFadden 129
Condition. Sldn complete, with small bald spots on
dorsum. Skull complete.
Type Series. Holotype only.
Family MYOXIDAE Gray, 1821
Genus GRAPHIURUS Smuts, 1832
Aethoglis hueti argenteus G. M. Allen,
1936
J. Mammal, 17: 293, 14 August.
= Graphiurus hueti argenteus (G. M. Allen,
1936). See Hayman and Holt (1940:
608).
Holotype. MCZ 17920. Skin and skull. Adult female.
Locality. Cameroons ( = Cameroon): (Kribi), Lolo-
dorf. 17 March 1911.
Collector. G. Schwab. Original number 1.
Condition. Sldn and skull complete.
Type Series. 2 paratypes; MCZ 17607, sldn and
skull, adult female; and a specimen in the USNM,
number 125434.
Comments. Rosevear considered G. h. argenteus
"possibly valid as a race" (1969: 501).
Claviglis soleatus collaris G. M. Allen and
Loveridge, 1933
Bull. Mus. Comp. Zool., 75: 122,
Februan'.
= Graphiurus lorraineus Dollman, 1910.
See Holden (1993: 764).
Holotype. MCZ 26373. Skin and skull. Adult female.
Locality. Tanganyika Territory (=Tanzania): (Irin-
ga), north end Lake Nyasa, Uldnga Mountains, Ma-
dehani, 7,000 ft (2,135 m). 24 Februaiy 1930.
Collector. A. Loveridge.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 2 paratypes; MCZ 26374, skin and
skull, male; MCZ 26375, skin and skull, female.
Graphiurus microtis griseus G. M. Allen,
1912a
Bull. Mus. Comp. Zool, 54: 440, April.
= Grapfiiurus murinus griseus G. M. Allen,
1912. See Hayman and Holt (1940: 610).
Holotype. MCZ 8244. Skin and skull. Adult male.
Locality. British East Africa ( = Kenya): northern
Guaso Nyiro River ( = Ewaso Ngiro), 4,000 ft (1,220
m). 25 July 1909.
Collector G. M. Allen. Original number 51.
Condition. Skin and skull complete.
Type Series. 2 paratypes; MCZ 8248, skin and skull,
male; MCZ 8249, skin and skaiU, adult female.
Grapiiiurus scfiwabi G. M. Allen, 1912a
Bull. Mus. Comp. Zool., 54: 441, April.
= Grapiiiurus surdus Dollman, 1912. See
Holden (1993: 765).
Holotype. MCZ 8607. Skin and skull. Juvenile.
Locality. Cameroun ( = Cameroon), Kribi. 1911.
Collector G. Schwab.
Condition. Skin and skull complete.
Type Series. Holotype only.
Family BATHYERGIDAE Waterhouse,
1841
Genus CRYPTOIVIYS Gmy, 1864
Cryptomys tiottentotus occlusus G. M.
Allen and Loveridge, 1933
Bull. Mus. Comp. Zoo)., 75: 125,
Februaiy.
= Cryptomys tiottentotus wfiytei (Thomas,
1897). See Honeycutt et al. (1991: 51).
Holotype. MCZ 26557. Skin and skull. Adult male.
Locality. Tanganyika Territoiy (=Tanzania): (Irin-
ga), Ugungwe Mountains, Kigogo, 6,000 ft (1,830
m). 18 January 1930.
Collector. A. Loveridge.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 15 paratypes; MCZ 26558-26572, all
represented by skin and skull.
Genus HETEROCEPHALUS Ruppell,
1842
l-ieterocepiiaius stygius G. M. Allen,
1912a
Bull. Mus. Comp. Zool, 54: 444, April.
= l-leterocepiiaius giaber Ruppell, 1842.
See HoUister (1919: 160).
Holotype. MCZ 12470. Body in alcohol, skull e.xtract-
ed. Adult female.
Locality. British East Africa ( = Kenya): northern
Guaso Nyiro ( = Ewaso Ngiro), Neumann's Boma. 6
August 1909.
Collector G. M. Allen.
Condition. Alcoholic. Skull complete. Mandible
disarticulated.
Type Series. Holotype only.
130 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Family ERITHIZONTIDAE Bonaparte,
1845
Genus ERITHIZONF. Cuvier, 1822
Erethizon dorsatus picinus Bangs, 1900c
Proc. New England Zool. Club, 2: 37, 20
September.
= Erithizon dorsatum dorsatum (Linnaeus,
1758). See Hai-per (1961: 90).
Holotype. B8839. Sldn and skull. Adult male.
Locality. (Canada): Labrador, Strait of Belle Isle,
Lance ( = Lanse) au Loup. 16 Februaiy 1899.
Collector. E. Doane.
Condition. Skin and skull complete.
Type Series. 1.5 paratypes; B8832-B8838, B8840-
B8847; all represented by skin and skull, 8 females
and 7 males. Bangs had also a skull of picimis
(B8831) from Black Bay, Labrador, at the time of
description, not mentioned in the original publi-
cation.
Family DASYPROCTIDAE Gray, 1825
Genus DASYPROCTA llliger, 1811
Dasyprocta callida Bangs, 1 901 b
Amer. Nat., 35: 635, 22 August.
= Dasyprocta punctata callida Bangs,
1901. See Kellogg (1946: 59).
Holotype. B8443. Skin and skull. Adult male.
Locality. Panama: (Panama), San Miguel Island. 8
May 1900.
Collector. W. W. Brown, Jr Original number 171.
Condition. Skin and skull complete.
Type Series. 5 paratypes; B8442, B8444-B8447; all
represented by skin and skull, 3 females and 2
males.
Comments. D. p. callida was retained as a valid
subspecies by Hall (1981: 860).
Dasyprocta colombiana Bangs, 1 898k
Proc. Biol. Soc. Washington, 12: 163, 10
August.
= Dasyprocta punctata colombiana (Gray,
1898). See Cabrera (1961: 589).
Holotype. B8008. Skin and skull. Adult female.
Locality. Colombia: Magdalena, Santa Marta. 6
January 1898.
Collector. W. W. Brown, Jr. Original number 37.
Condition. Sldn and skull complete.
Type Series. 1 paratype; B8113, skin and skull, ju-
venile male.
Comments. D. p. colombiana was retained as a val-
id subspecies by Cabrera (1961: .589).
Dasyprocta noblei G. M. Allen, 1914e
Proc. New England Zool. Club, 5: 69, 7
October.
= Dasyprocta leporina noblei G. M. Allen,
1914. See Woods (1993: 781).
Holotype. MCZ 15936. Sldn and skull (and atias).
Adult female.
Locality. Guadeloupe Island: Goyave. 22 August
1914.
Collector G. K. Noble.
Condition. Sldn complete. Skull partial (bone miss-
ing from nasal and frontal).
Type Series. 1 paratype; MCZ 15937, sldn and
skull, subadult female.
Dasyprocta punctata nuchalis Goldman,
1917
Proc. Biol. Soc. Washington, 30: 113, 23
May.
Holotype. MCZ 1008 L Skin and skull. Adult female.
Locality. Panama: Chiriqui, Divala. 30 November
1900.
Collector W. W. Brown, Jr. Original number 17.
Condition. Skin and skull complete.
Type Series. 4 paratypes; MCZ 10080, skin and
skull, adult male; 10084, skull; 10175, skin and
skull, adult female; 10176, sldn and skull, adult
male.
Comments. Retained as a valid subspecies by Hall
(1981: 860).
Family AGOUTIDAE Gray, 1821
Genus AGOUTI Lacepede, 1799
Agouti paca virgatus Bangs, 1902b
Bull. Mus. Comp. Zool., 39: 47, April.
Holotype. MCZ 10079. Sldn and skull. Adult male.
Locality. Panama: Chiriqui, Divala. 16 December
1900.
Collector W. W. Brown, Jr Original number 21.
Condition. Sldn and skull complete.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Hall
(1981: 8.58) and Perez (1992: 1).
Family OCTODONTIDAE Waterhouse,
1839
Genus TYMPANOCTOMYS Yepes, 1941
Octomys barrerae Lawrence, 1941
Proc. New England Zool. Club, 18: 43,
28 Januaiy.
= tympanoctomys barrerae (Lawrence,
1941). See Yepes (1942: 75).
Holotype. MCZ 39716. Skin and skull. Aduk male.
Locality. Argentina: Mendoza Province, La Paz.
April 1939.
Type Specimens of Recent Mammals • Helpen and McFadden
131
Collector. J. M. de la Barrera. Original number
A31.
Condition. Sldn and skull complete.
Type Series. Holotype only.
Comments. Type species of Tyinpanoctomys Yepes,
1941. T. barrerae was considerd a valid species by
Woods (1993: 789) and Nowak (1999: 1683).
Family ECHIMYIDAE Gray, 1825
Genus BOROMYS M\\\er, 1916
Boromys torrei G. M. Allen, 1917a
Bull. Mus. Comp. Zool., 61: 6, Januaiy.
Holotype. VP 9601. Palate.
Locality. Cuba: Matanzas Province, cave in Sierra
de Hato Nuevo.
Collector C. de la Torre.
Condition. Condition as described in original —
"palate with root of right z}'gomatic arch, pni^ and
alveolar row of right side, i)i' and posterior part of
alveolar row of left side."
Type Series. Paratype material consists of 8 lower
jaws and 2 separate lower molars; in the MCZ.
Comments. Considered a valid species by Woods
(1993: 799) and Nowak (1999: 1703) but almost
certainly extinct. This specimen is stored in the
Vertebrate Paleontology Department of tlie MCZ.
Genus DIPLOMYS Thomas, 1916
Loncheres labilis Bangs, 1901b
Amer. Nat., 35: 638, 22 August.
= Diplomys labilis (Bangs, 1901). See
Thomas (1916: 296).
Holotype. B8480. Skin and skull. Adult male.
Locality. Panama: (Panama), Gulf of Panama, San
Miguel Island. 26 April 1900.
Collector. W W Brown, Jr. Original ninuber 103.
Condition. Skin and skull complete.
Type Series. There is a small series of paratypes in
the MCZ and FMNH.
Comments. D. labilis was considered a valid spe-
cies by Woods (1993: 791) and Nowak (1999:
1695).
Genus PROECHIMYS ^. A. Allen, 1899
Proechimys burrus Bangs, 1901b
Amer. Nat., 35: 640, 22 August.
= Proechimys semispinosus burrus Bangs,
1901. See Goldman (1920: 120).
Holotype. B8458. Skin and skull. Adult male.
Locality. Panama: (Panama), Gulf of Panama, San
Miguel Island. 30 April 1900.
Collector WW. Bro\\ai, Jr. Original number 130.
Condition. Skin and skull complete. Bight dentary
broken at pnij.
Type Series. There is a large series of paratypes in
the MCZ.
Comments. P. s. burnis was retained as a valid sub-
species bv Hall (1981: 873).
Proechimys gorgonae Bangs, 1905b
Bull. Mus. Comp. Zool., 46: 89, June.
Holotype. MCZ 10828. Sldn and skull. Adult male.
Locality. Colombia: (Cauca), Gorgona Island. 2
July 1904.
Collector W. W Brown, Jr. Original number 25.
Condition. Skin and skull complete.
Tijpe Series. 6 paratypes; MCZ 10829-10834, all
represented by skin and skull, 1 female and 5
males. MCZ 10830 and 10834 are now in the
FMNH.
Comments. Treated as a separate species by Woods
(1993: 796) but included in P. cayennensis by No-
wak (1999: 1689).
Proechimys guyannensis hyleae Moojen,
1948
Univ. Kansas Publ. Mus. Nat. Hist., 1:
361, 10 December.
= Proechimys cayennensis hyleae Moojen,
1948. See comments.
Holotype. MCZ 30887. Skin and skull. Adult male.
Locality. Brazil: Para, Porto de Moz, Rio Tapajoz
(=Tapajos), Tauari (=Tauaiy), 87 km south of San-
tarem. 19 January 1934.
Collector A. M. Olalla. Original number 7288.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 20 paratypes (19 at MCZ and 1 at
FMNH), represented by sldn and skull.
Comments. Retained as a valid subspecies by Ca-
brera (1961: 521). The use of the specific name
cayennensis over guyannensis follows Woods
(1993: 795).
Family CAPROMYIDAE Smith, 1842
Genus CAPROMYS Desmarest, 1822
Capromys pilorides relictus G. M. Allen,
1911a
Bull. Mus. Comp. Zool, 54: 207, July.
Holotype. MCZ 10996. Skin and skull. Adult male.
Locality. Cuba: Isle of Pines ( = Isla de Pinos), Nue-
va Gerona, Casas Mountains. 10 March 1902
Collector. W. R. Zappey.
Condition. Sldn complete. Skull partial (bone miss-
ing from palatine to occipital).
Type Series. 1 paratype; MCZ 10997, sldn and
skull.
Comments. Retained as a valid subspecies by Hall
(1981: 863).
132 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Genus GEOCAPROMYS Chapman, 1901
Geocapromys cubanus G. M. Allen, 1917a
Bull. Mus. Comp. Zool. 61: 9, Januaiy.
= Geocapromys columbianus (Chapman,
1892). See G. M. Allen (1918b: 145).
Holotype. VP 9602. Part of right lower ramus. Juve-
nile.
Locality. Cuba: Matanzas Province, cave in Sierra
de Hato Neuvo.
Collector. C. de la Torre.
Condition. Portion of ramus includes incisor and
three anterior cheekteeth.
Type Series. Allen mentions in the original descrip-
tion, "Five palates with teeth, about 15 jaw frag-
ments mostly with teeth, and numerous other frag-
ments." This material is in the MCZ.
Comments. The type material o{ cubanus is stored
in the Vertebrate Paleontology Department of the
MCZ.
Geocapromys ingrahami abaconis
Lawrence, 1934
Occas. Pap. Boston See. Nat. Hist., 8:
190, 7 November.
Holotype. VP 2108. Left lower ramus. Adult.
Locality. Bahamas: Great Abaco Island, Hole in
the Wall, Imperial Lighthouse Caves. 1934.
Collector F. Rainey.
Condition. Ramus intact.
Type Series. The parats'pe series is a collection of
cranial and mandibular fragments, described in the
original description and housed in the MCZ.
Comments. G. i. abaconis was retained as a valid
subspecies by Hall (1981: 866) under the genus
Capromys. Woods (1993: 800) employed the genus
Geocapromys. G. i. abaconis is extinct. The type
material of abaconis is stored in the Vertebrate Pa-
leontology Department of the MCZ.
Geocapromys ingrahami irrectus
Lawrence, 1934
Occas. Pap. Boston Soc. Nat. Hist., 8:
190, 7 November.
Holotype. VP 2107. Right lower ramus. Adult.
Locality. Bahamas: Crooked Island, Gordon Hill
Caves, "Burial Cave No. 1." 1934.
Collector. F. Rainey.
Condition. Ramus intact.
Tijpe Series. The paratype series is a collection of
cranial and mandibular fragments, described in the
original description and housed in the MCZ.
Comments. G. i. irrectus was retained as a valid
subspecies by Hall (1981: 866) vmder the genus
Capromys. Woods (1993; 800) employed tlie genus
Geocapromys. G. i. irrectus is extinct. The type ma-
terial of irrectus is stored in the Vertebrate Pale-
ontology Department of tlie MCZ.
Genus MESOCAPROMYS Marona, 1970
Capromys nana G. M. Allen, 1917b
Proc. New England Zool. Club, 6: 54, 28
March.
= Mesocapromys nanus (G. M. Allen,
1917). See Kratochvil et al. (1978: 15).
Holotype. VP 9864. Right mandible.
Locality. Cuba: Matanzas, cave in Sierra de Hato
Nuevo. March 1917.
Collector. T. Barbour.
Condition. Mandible partial — coronoid and angu-
lar processes broken off.
Tijpe Series. A series of paratype material is stored
at the MCZ.
Comments. G. M. Allen supplemented his descrip-
tion of Capromys nanus after Thomas Barbour
provided him with a freshly killed specimen in
1918 (1918b: 140-145). Type species of the sub-
genus Paracapromys Kratochvil, Rodriguez, and
Baiois, 1978. M. nanus was considered a valid spe-
cies by Woods (1993: 801) and Nowak (1999:
1703). The type series is stored in the Vertebrate
Paleontology Department of the MCZ.
Genus PLAGIODONTA F. Cuvier, 1836
Plagiodonta araeum Ray, 1 964
Breviora, Mus. Comp. Zool., 203: 2, 10
April.
Holotype. VP 7675. Left upper cheektooth, originally
described as "almost certainly the fourth (decidu-
ous) premolar."
Locality. Dominican Republic: San Rafael Prov-
ince, Hondo Valle Municipality, unnamed cave 2
km southeast of Rancho de La Guardia. April 1963.
Collector R. Allen and C. E. Ray.
Condition. Condition of tooth as originally de-
scribed — "damaged slighdy along the posterolabial
wall."
Type Series. Holotype only.
Comments. Considered a valid species by Woods
(1993: 804) and Nowak (1999: 1708). Complete
cranial and dentaiy material of araeum has been
collected in Haiti and deposited in the Florida Mu-
seum of Natural Histoiy (Woods 1993: 804). This
species is extinct. This specimen is stored in the
Vertebrate Paleontology Department of the MCZ.
Type Specimens of Recent Mammals • Helaen and McFadder
133
Order LAGOMORPHA Brandt, 1855
Family OCHOTONIDAE Thomas, 1897
Genus OCHOTONA Link, 1795
Ochotona cuppes Bangs, 1899g
Proc. New England Zool. Club, 1: 40, 5
June.
= Ochotona princeps cuppes Bangs, 1899.
See A. H. Howell (1924: 27).
Holotype. B7389. Skin and skull. Adult male.
Locality. (Canada): British Columbia, Gold Range,
Monishee Divide, 4,000 ft (1,220 m). 2 August
1897.
Collector. A. C. Brooks. Original number 10.30.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; MCZ 7390-7392; all rep-
resented by sldn and skull, all female. MCZ 7390
is now at FMNH.
Comments. O. p. cuppes was retained as a valid
subspecies by Hall (1981: 289) and Smith and Wes-
ton (1990: 1).
Ochotona saxatilis Bangs, 1 899g
Proc. New England Zool. Club, 1: 41, 5
June.
= Ochotona princeps saxatilis Bangs,
1899. See A. H. Howell (1924: 27).
Holotype. MCZ 2703. Skin. Adult male.
Locality. (United States): Colorado, Park County,
Snowy Range, near Mount Lincoln, Montgomery.
27 July 1871.
Collector. J. A. Allen, Rocky Mountain Expedition.
Original number 945.
Condition. Skin complete.
Type Series. Large paratype series; MCZ 209, 243-
263, skulls; B41 and B42, each represented by sldn
and skull; MCZ 2673-2703, skins.
Comments. O. p. saxatilis was retained as a valid
subspecies by Hall (1981: 291) and Smith and Wes-
ton (1990: 1).
Family LEPORIDAE Fischer de Waldheim,
1817
Genus LEPL/S Linnaeus, 1758
Lepus (Macrotolagus) alleni palitans
Bangs, 1900a
Proc. New England Zool. Club, 1: 85, 23
February.
Holotype. B9096. Skin and skull. Adult female.
Locality. Mexico: Sinaloa, Aguacaliente (about 40
miles [64.4 km] southeast of Mazatlan). 7 August
1897.
Collector P. O. Simons. Original number 157.
Condition. Skin and skull complete. Mandible dis-
articulated; left ramus chipped; two holes in right
ramus.
Type Series. 1 paratype; B9097, skin, adult male.
Comments. Retained as a valid subspecies by Hall
(1981: 332) and Best and Heniy (1993: 1).
Lepus americanus struthopus Bangs,
1898e
Proc. Biol. Soc. Washington, 12: 81, 24
March.
Holotype. B2025. Skin and skull. Adult female.
Locality. (Canada): Nova Scotia, Digby. 4 August
1894.
Collector O. Bangs. Original number 9.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. Retained as a valid subspecies by Hall
(1981: 318).
Lepus arcticus bangsii Rhoads, 1896a
Amer. Nat., 30: 236, 6 March.
Holotype. B3752. Skin and skull. Adult female.
Locality. (Canada): Newfoundland, Codroy. 3 Au-
gust 1895.
Collector E. Doane. Original number 1.
Condition. Sldn and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. Retained as a valid subspecies by Hall
(1981: 320).
Lepus bairdii Hayden, 1869
Amer. Nat., 3: 115, May.
= Lepus americanus bairdii Hayden, 1869.
See J. A. Allen (1875: 431).
Syntype. MCZ 5501. Sldn and skull. Adult.
Locality. (United States): Wyoming, (Fremont
County), summit of Wind River Mountains, near
Fremont Peak. 4 June 1860.
Collector. F. V. Hayden. Original number 90. For-
merly USNM 4264.
Condition. Sldn and skull complete.
Tijpe Series. 2 other adult syn types; USNM 4262/
38001, skin and skull, male; USNM 4263/4273,
sldn and skidl; see Poole and Schantz (1942: 210).
Comments. No type is designated in the original
description, which is based on 6 individuals col-
lected by Hayden — 3 adults and 3 juveniles, origi-
nally deposited in the USNM (Poole and Schantz
1942: 210).
134 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Lepus bairdi cascadensis Nelson, 1907
Proc. Biol. Soc. Washington, 20: 87, 11
December.
= Lepus americanus cascadensis Nelson,
1907. See Racey and Cowan (1936:
HIS).
Holotype. B1886. Skin and skull. Adult male.
Locality. Canada: British Columbia, near Hope,
Roab's Ranch. 12 June 1874.
Collector. W. C. Colt. Original number 476.
Condition. Sldn complete. Skull slightly damaged
(supraoccipital and occipital chipped).
Type Series. Holotype only.
Comments. L. a. cascadensis was retained as a valid
subspecies by Hall (1981: 315).
Genus Sy/.WMGL/S Gray, 1867
Lepus (Tapeti) incitatus Bangs, 1901b
Amer. Nat., 35: 633, 22 August.
= Sylvilagus brasiliensis incitatus (Bangs,
1901). See Hershkovitz (1950: 352).
Holotype. B8441. Skin and skull. Adult female.
Locality. Panama: (Panama), Bay of Panama, San
Miguel Island. 30 April 1900.
Collector W. W. Browi:i, Jr. Original number 127
Condition. Skin and skull complete.
Type Series. Holotype only.
Comments. S. b. incitatus was retained as a valid
subspecies by Hall (1981: 296).
Lepus paludicola Miller and Bangs, 1894
Proc. Biol. Soc. Washington, 9: 105, 9
June.
= Sylvilagus palustn's paludicola (Miller
and Bangs, 1894). See Nelson (1909:
269).
Holotype. B1451. Sldn and skull. Adult female.
Locality. (United States): Florida, (Citrus County),
Fort Island, near Crystal River. 28 Januaiy 1894.
Collector F. L. Small. Original number 11.50.
Condition. Skin and skull complete.
Type Series. 3 paratypes; B1452, skin and skull,
adult male; B1453, sldn and skull, adult male;
B1454, skin and skull, adult female.
Connnents. S. p. paludicola was retained as a valid
subspecies by Hall (1981: 299) and Chapman and
Willner (1981: 1).
Lepus sylvaticus alacer Bangs, 1 896h
Proc. Biol. Soc. Washington, 10: 136, 28
December.
= Sylvilagus floridanus alacer (Bangs,
1896). See Lyon (1904: 336).
Holotype. B5480. Sldn and skull. Adult female.
Locality. (United States): Indian Territory
(=Oklahoma), (Adair County), Stilwell. 14 August
1896.
Collector. T. Siu-ber. Original number 65.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; B1677, skin and skull,
B1678, skin and skull, adult male; B5481, skin and
skull, adult female.
Comments. S. f. alacer was retained as a valid sub-
species by Hall (1981: 301) and Chapman et al.
(1980: 1).
Lepus sylvaticus transitionalis Bangs,
1895a
Proc. Boston Soc. Nat. Hist., 26: 405, 31
January.
= Sylvilagus transitionalis (Bangs, 1895).
See Nelson (1909: 195).
Holottjpe. B2407. Skin and skull. Adult female.
Locality. (United States): Connecticut, New Lon-
don County, Libeity Hill. 6 November 1894.
Collector O. Bangs.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. There is a series of paratypes in the
MCZ.
Comments. S. transitionalis was considered a valid
species by Hoffman (1993: 827) and Nowak (1999:
1727).
Order SCANDENTIA Wagner, 1855
Family TUPAIIDAE Gray, 1825
Genus TUPAIA Raffles, 1822
Tana tana griswoldi CooMge, 1938
Proc. New England Zool. Club, 17: 45, 6
May.
= Tupaia tana paitana (Lyon, 1913). See
Medway (1965: 76).
Holotype. MCZ 36416. Sldn, skaill, and baculum.
Subadult male.
Locality. (Malaysia): British North Borneo ( = Sa-
bah). Mount Kinabalu, Kenokok River, Kiau
( = Kampong Kiau), 3,300 ft (1,007 m). 9 August
1937.
Collector J. A. Griswold, Jr., Asiatic Primate Ex-
pedition. Original number 684.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Type Specimens of Recent Mammals • Helpen and McFadden
135
Order PRIMATES Linnaeus, 1758
Family INDRIDAE Burnett, 1828
Genus PROPITHECUS BenneW, 1832
Propithecus perrieri Lavauden, 1931
Compt. Rend. Acad. Sci. Paris, 193: 77, 6
July.
= Propithecus diadema perrieri Lavauden,
1931. See W. C. O. Hill (1953: 568).
Holotype. MCZ 44857. Skin, skull, and postcranial
skeleton. Adult male.
Locality. Malagasy Republic (Madagascar): (Toli-
aiy), Ifotaka, vicinity of Ambovombe. Probably
1928.
Collector. M. Perrier. Original number 1. Grandi-
dier Collection.
Conclition. Skin and skull complete. Postcranial
skeleton partial (right humerus broken, foot bones
left in sldn).
Type Series. 3 paratypes; MCZ 44858, skin, skull,
and postcranial skeleton, adult female; MCZ
44859, skin and skull, juvenile; MCZ 46001, skin
and leg bones, adult.
Comments. P. d. perrieri was retained as a valid
subspecies by Tattersall (1982: 103).
Family DAUBENTONIIDAE Gray, 1863
Genus DAUBENTONIA E. Geoffrey, 1795
Ctieiromys madagascariensis lanigerG.
Grandidier, 1930d
Bull. Acad. Malgache, n. sen, 11: 106
(for 1928).
= Daubentonia madagascariensis (Gmelin,
1788). See G. M. Allen (1939: 134).
Holotype. MCZ 45947, Skull and postcranial skele-
ton. Adult male.
Locality. Madagascar: Forest of the East,
Collector. Received by G, Grandidier from the
Academic Malgache, 1927-28.
Condition. Skull complete. Postcranial skeleton in-
complete (hands and feet missing, presumably in-
tact within mounted skin).
Type Series. Holotype only.
Comments. The mounted skin of the holotype is in
the collection of the Academic Malgache (Anta-
nanarivo Museum). Grandidier named laniger as a
new subspecies on account of its woolly pelage, but
it is probably just a molting individual (Schwarz
1931: 428).
Family GALAGONIDAE Gray, 1825
Genus GALAGOIDES A. Smith, 1833
Galago demidovii orinus Lawrence and
Wasliburn, 1936
Occ. Pap. Boston Soc. Nat. Hist., 8: 259,
8 January.
= Galagoides orinus (Lawrence and
Washburn, 1936). See Honess (1996: 58).
Holotype. MCZ 22453, Skin and skull. Adult male.
Locality. Tanganyika Territoiy (=Tanzania): Mor-
ogoro, Uluguru Mountains, Bagilo, 5,000 ft (1,525
m). 17 September 1926,
Collector A. Love ridge.
Condition. Skin and skull complete.
Type Series. Holotype only.
Family GEBIDAE Bonaparte, 1831
Genus ALOUATTA Lacepede, 1799
Alouatta palliata luctuosa Lawrence,
1933b
Bull. Mus. Comp. Zool., 75: 337,
November.
= Alouatta pigra Lawrence, 1933. See
Smith (1970: 363).
Holotype. MCZ 24059. Skin and skTill. Adult male.
Locality. British Honduras ( = Belize): Cayo Dis-
trict, Mount Cow. 12 April 1928.
Collector. O. L. Austin, Jr. Original number 723,
Condition. Skin and skull complete,
Tijpe Series. HolotyjDe only,
Alouatta palliata trabeata Lawrence,
1933b
Bull. Mus. Comp. Zool., 75: 328,
November.
Holotype. MCZ 29545. Skin and skull. Adult male.
Locality. Panama: Herrera Province, Capina,
March 1933.
Collector T Barboui\ Original number 4,
Condition. Skin and skull complete,
Tijpe Series. Lawrence mentions examining 19
specimens of trabeata and lists them by locality
(1933: 330), 9 of these are in the MCZ; other tlian
the holotype, they are MCZ 27784, 27785, 28735,
28736, 29543, 29544, 29546, and 29548, 1 paraty-
pe, MCZ 29547, has been exchanged to the Museu
Palista, Sao Paolo, Brazil.
Comments. Retained as a valid subspecies by Hall
(1981: 263),
136 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Genus CEBUS Erxleben, 1777
Cebus curtus Bangs, 1905b
Bull. Mus. Comp. ZooL, 46: 91, June.
= Cebus capucinus curtus Bangs, 1905.
See Cabrera (1917: 240).
Holotype. MCZ 10824. Skin and skull. Adult male.
Locality. Colombia: (Cauca), Gorgona Island. 2
July 1904.
Collector. W. W. Brown, Jr. Original number 27.
Condition. Skin and skull complete.
Type Series. 1 paratype; MCZ 10825, skin and
skull, adult female.
Comments. C. c. curtus was retained as a valid sub-
species by Cabrera (1958: 169) and Napier (1976:
36).
Family HYLOBATIDAE Gray, 1870
Genus HYLOBATES llliger, 1811
Hylobates lar carpenteri Groves, 1968
Proc. Biol. Soc. Washington, 81: 625, 30
December.
Holotype. MCZ 41430. Skin and skeleton. Adult
male.
Locality. Thailand: Chiengmai District, Mount
Angka ( = Doi Inthanon), 3,400 ft (1,037 m). 14
March 1937.
Collector. H. J. Coolidge, Jr., Asiatic Primate Ex-
pedition. Original numbers 21 and 185.
Condition. Skin, skull, and skeleton complete.
Type Series. Groves examined a series of 144 skins,
skulls, and skeletons of this subspecies that were
collected by the Harvard Asiatic Primate Expedi-
tion in 1937 from Chiengmai District, Thailand,
"mostly in the Museum of Comparative Zoology,
Hai'vard, but a few of the osteological specimens
in the Anthropology Department, University of
California at Berkeley." He examined also 18 spec-
imens in the USNM and 1 in the AMNH, all rep-
resented by skin and skull.
Comments. Retained as a valid subspecies by Jen-
kins (1990: 15) and Geissman (1995: 474).
Family HOMINIDAE Gray, 1825
Genus GORILLA I. Geoff roy, 1853
Troglodytes gorilla Savage and Wyman,
1847
Boston J. Nat. Hist., 5: 417, December.
= Gorilla gorilla gorilla (Savage and
Wyman, 1847). See Rothschild (1923:
176).
Lectotype. MCZ 9587. Skull and postcranial skeleton.
Adult male.
Locality. Gabon: Gabon Estuaiy, Mpongwe coun-
tiy. 1847.
Collector T. S. Savage. Original Jiumber 28 (of J.
Wyman).
Condition. Postcranial skeleton partial (includes
pelvis, sacrum, scapulae, humeri, radii, left ulna,
femora, left tibia, and 7 vertebrae: 2 cervical, 3 dor-
sal, 2 lumbar). Head of right humerus and left fe-
mur bisected.
Type Series. The original description mentions
"four skulls, two males and two females, one of
each in a perfect condition, and all of them adult;
a male and female peKis, the long bones of the
upper and lower extremities, and a few vertebrae
and ribs." However, measurements and illustrations
of only two of these specimens, an adult male and
female, each represented by a skull and partial
postcranial skeleton, are provided. These two, pre-
sumably the specimens originally described as be-
ing "in a perfect condition," are the only specimens
of the original four to be noted in Wymans per-
sonal notebook of osteology (unpublished, now in
tlie libraiy of the Boston Museum of Science) and
die only original specimens of which there is any
record at all. These two specimens were trans-
ferred from the Boston Society of Natural History
to the MCZ in 1915-16. They bear MCZ numbers
9587 (male) and 9311 (female). The skull of the
female was sawed in half (hemisected), probably by
Wyman. The adult male specimen, MCZ 9587,
possesses an intact skull. MCZ 9587 is the only in-
tact specimen of the syntype series known to exist,
and its measurements and an illustration of its skull
are included in the original publication. Addition-
ally, it is the only syntype whose measurements are
included in Coolidge s rex-ision of the genus Gorilla
(1929: 325), although it is erroneously listed in that
work as MCZ 9586. For these reasons, MCZ 9587
is hereby designated as lectotype of Troglodytes go-
rilla Savage and Wyman, 1847; this should ensure
taxonomic consistency between past and future
treatments of this name.
Comments. Type species of the genus Gorilla I.
Geoffroy, 1853. The original description is occa-
sionally attributed in error to Wyman (1847). G.
gorilla was considered a valid species by Groves
a993: 276) and Nowak (1999: 618).
Order LIPOTYPHLA Haeckel, 1866
Family NESOPHONTIDAE Anthony, 1916
Genus /VESOPHOA/7ES Anthony, 1916
Nesophontes micrus G. M. Allen, 1917a
Bull. Mus. Comp. ZooL, 61: 5, Januaiy.
Holotype. VP 9600. Right ramus.
Locality. Cuba: Matanzas Province, cave in Sierra
de Hato Neuvo.
Collector C. de la Torre.
Type Specimens of Recent Mammals • Helgen and McFadden 137
Condition. Condition as originally described —
"posterior half of the right ramus, containing a part
o£ pm^, m„ m.2, and the roots of 111.."
Type Series. Holotype only.
Comments. Considered a valid species by Hutterer
(1993: 70) and Nowak (1999: 201); almost certainly
extinct. This specimen is stored in the Vertebrate
Paleontology Department of the MCZ.
Family SOLENODONTIDAE Gill, 1872
Genus SOL EA/ODOA/ Brandt, 1833
Antillogale marcanoi Patterson, 1962
Breviora, Mus. Comp. ZooL, 165: 3, 22
August.
= Solenodon marcano/ (Patterson, 1962).
See Varona (1974: 8).
Holotype. VP 7261. Right lower ramus.
Locality. Dominican Republic: San Rafael Prov-
ince ( = Elias Pina), Hondo Valle Municipality, un-
named cave 2 km southeast of Rancho La Guardia.
Summer 1958.
Collector. C. E. Ray and A. S. Rand.
Condition. Condition as originally described — "in-
complete right ramus of mandible with P3— M, and
alveoli of other teeth."
Type Series. 5 paratypes; 7262, left ramus, 7263,
right humerus, 7264, left humerus, 7265, right
ulna, 7266, left ramus, juvenile; all partial.
Comments. A. marcanoi is the type species of the
genus Antillogale Patterson, 1962. Antillogale was
first synonymized with Solenodon by Van Valen
(1967: 255). Considered a valid species by Hutterer
(1993: 69) and Nowak (1999: 199). S. marcanoi is
probably extinct. This specimen is stored in the
Vertebrate Paleontology Department of the MCZ.
Solenodon poeyanus Barbour, 1944
Proc. New England Zool. Club, 23: 6,
March 7.
= Solenodon cubanus poeyanus Barbour,
1944. See Aguayo (1950: 131).
Holotype. MCZ 6597. Mounted skin and skull.
Locality. Cuba: Oriente ( = Holguin), near Nipi Bay
( = Nipe Bay).
Collector Bought by A. Agassiz from H. A. Ward,
1891.
Condition. Skin complete. Skull partial (basioccip-
ital plate and tympanic bullae missing, parietals
damaged). Mandible disarticulated.
Type Series. In the original description, Barbour
includes a photograph of a "living example of So-
lenodon poeyanus now in Zoological Garden, Ha-
vana, Cuba, from vicinity of Baracoa."
Comments. The number of this specimen is 6597,
not 6957 as stated in the original description. S. c.
poeyanus was retained as a valid subspecies by Hall
(1981: 22).
Family SORICIDAE Fischer von
Waldheim, 1817
Genus BLARINA Gray, 1838
Blarina brevicauda aloga Bangs, 1902a
Proc. New England Zool. Club, 3: 76, 31
March.
Holotype. B9727. Skin and skull. Adult male.
Locality. (United States): Massachusetts, Dukes
County, Marthas Vineyard, West Tisbury. 25 June
1899.
Collector O. Bangs. Original number 2.
Condition. Skin and skrdl complete. Mandible dis-
articulated.
Type Series. 9 paratypes, B9725, B9726, B9728-
B9734, all represented by skin and skull, 3 females
and 6 feinales.
Comments. Retained as a valid subspecies by Hall
(1981: 54) and George et al. (1986: 1).
Blarina brevicauda compacta Bangs,
1902a
Proc. New England Zool. Club, 3: 77, 31
March.
Holotype. B9705. Skin and skull. Adult male.
Locality. (United States): Massachusetts, (Nan-
tucket County), Nantucket (Island). 10 July 1899.
Collector O. Bangs. Original number 3.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 6 paratypes; B9701-B9704, B9706,
B9708; all represented by skin and skull, 4 females
and 2 males.
Comments. Retained as a valid subspecies by Hall
(1981: 56) and George et al. (1986: 1).
Genus CROCIDURAVJagner, 1832
Crocidura bicolor tephragaster Setzer,
1956
Proc. U.S. Nat. Mus., 106: 458, 28
November.
= Crocidura fuscomurina (Heuglin, 1865).
See Hutterer (1983: 223).
Holotype. MCZ 44773. Skin and skull. Adult male.
Locality. Anglo-Egyptian Sudan ( = Sudan): (East-
ern) Equatoria, Torit. 25 April 1950.
Collector. J. S. Owen. Original number 1158.
Condition. Sldn and skull complete.
Type Series. Setzer mentions that he examined 18
specimens of tephragaster, 8 of which are in the
MCZ.
138 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Crocidura hildegardeae phaios Setzer,
1956
Proc. U.S. Nat. Mus., 106: 460, 28
November.
Holotype. MCZ 45S55. Skin and skull. Adult female.
Locality. Anglo-Egyptian Sudan ( = Sudan): Equa-
toria, Imatong Mts, Gilo, 6,500 ft (1,983 m). 12
June 1950.
Collector. J. S. Owen. Original number 1266.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; 1 from the MCZ; MCZ
45856, skin and skull, male.
Comments. Retained as a valid subspecies, C gra-
cilipes phaios, by Heim de Balsac and Meester
(1977: 16); included in Crocidura hildegardeae by
Hutterer (1993: 87).
Genus CRYPTOTIS Pomel, 1848
Cryptotis avia G. M. Allen, 1923a
Proc. New England Zool. Club, S: 37, 12
February.
= Cryptotis thomasi {Mernam, 1897). See
Woodman (1996: 414).
Holotype. MCZ 20091. Skin and skull. Adult.
Locality. Colombia: (probably Cundinamarca), El
Verjon (see coments). October 1922.
Collector N. Maria.
Condition. Sldn complete. Skull partial (most of
skull from frontals to occiput missing). Left ramus
of mandible missing.
Type Series. Holotype only.
Comments. Considered a valid species by Hutterer
(1993: 108) and Nowak (1999: 209) but synony-
mized as noted above. The locality "El Verjon" is
not shown on any maps available to us but is pos-
sibly equivalent to "Paramo Cruz Verde" (Paynter
1997: 463).
Genus MYOSOREX Gray, 1838
Crocidura maurisca geata G. M. Allen and
Loveridge, 1927
Proc. Boston Soc. Nat. Hist., 38: 417, 23
December.
= Myosorex geata (G. M. Allen and
Loveridge, 1927). See Heim de Balsac
(1967: 610).
Holotype. MCZ 22447. Skin and skull. Adult male.
Locality. Tanganyika Territoiy (= Tanzania): Mor-
ogoro, Uluguru Mountains, Nyingwa, 7,500 ft
(2,288 m). 19 October 1926.
Collector A. Loveridge.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; MCZ 22448, skin and
skull, adult female.
Comments. M. geata was considered a valid species
by Hutterer (1993; 99) and Nowak (1999: 217).
Genus SOREX Linnaeus, 1758
Neosorex palustris acadicus G. M. Allen,
1915a
Proc. Biol. Soc. Washington, 28: 15, 12
February.
Name preoccupied by Sorex acadicus
Gilpin, 1867.
Sorex palustris gloveralleni Jackson, 1 926
J. Mammal., 7: 57, 15 February.
(Replacement name for Neosorex
palustris acadicus G. M. Allen, 1915)
Holotype. B2046. Skin and skull. Adult female.
Locality. (Canada): Nova Scotia, Digby. 26 July
1894.
Collector O. Bangs. Original number 3.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; B2053, exchanged to the
USNM in 1922.
Comments. Retained as a valid subspecies by Hall
(1981: 41) and Beneski and Stinson (1987: 1).
Sorex araneus ultimus G. M. Allen, 1914b
Proc. New England Zool. Club, 5: 51, 9
April.
= Sorex tundrensis Merriam, 1900. See
Hutterer (1993: 121).
Holotype. MCZ 15000. Skin and skull. Adult male.
Locality. (Russian Federation): northeastern Sibe-
ria, Nijni Kolymsk ( = Nizhnekolymsk), near mouth
of Kolyma River. 6 November 1911.
Collector. J. Koren. Original number 136.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. There is a series of paratypes, most of
which are housed in the MCZ.
Sorex macropygmaeus koreni G. M. Allen,
1914b
Proc. New England Zool. Club, 5: 56, 9
April.
= Sorex caecutiens koreni G. M. Allen,
1914. See Yudin (1989: 281).
Holotype. MCZ 15085. Skin and skiill. Adult female.
Locality. (Russian Federation): northeastern Sibe-
ria, Nijni Kolymsk ( = Nizhnekolymsk), near mouth
of Kolyma River. 19 October 1911.
Collector J. Koren. Original number .50.
Type Specimens of Recent Mammals • Helaen and McFadden
139
Tijpe Series. 5 paratypes; MCZ 15003-15007, all
represented by sldn and skull; 4 males, 1 female.
MCZ 15004, a male, was exchanged to the FMNH
in 1931.
Condition. Skin and skull complete. Mandible dis-
articulated.
Sorex macrurus Batchelder, 1896
Proc. Biol. Soc. Washington, 10: 133, 8
December.
Name preoccupied by Sorex macrourus
Lehmann, 1822.
Sorex dispar Batchelder, 191 1
Proc. Biol. Soc. Washington, 24: 97, 15
May. (Replacement name for Sorex
macrurus Batchelder, 1896)
Holotype. MCZ 41744. Skin and skaill. Adult male.
Locality. (United States): New York, Essex County,
Keene Heights, Beede's (see comments). 9 Sep-
tember 1895.
Collector. C. F. Batchelder. Original number 1384.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; MCZ 41745, skin and
skull, adult male.
Comments. S. dispar was considered a valid species
by Hutterer (1993: 114) and Nowak (1999: 205).
The type loctility was redescribed by Martin (1966:
131) as "0.6 mile south and 0.5 mile east of Saint
Huberts, Essex County, New York, lat. 44°09', long.
73°46'."
Sorex personatus miscix Bangs, 1899d
Proc. New England Zool. Club, 1: 15, 28
Februaiy.
= Sorex cinereus miscix Bangs, 1899. See
Jackson (1925: 56).
Holotype. B8651. Skin and skull. Adult male.
Locality. (Canada): Labrador, Black Bay. 10 Octo-
ber 1898.
Collector E. Doane. Original number 1.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. Bangs based his description on 39
specimens; corresponding to B7931-B7950 and
B8651-B8669; all represented by skin and skull.
Comments. S. c. miscix was retained as a valid sub-
species by Hall (1981: 29).
Sorex sanguinidens G. M. Allen, 1914b
Proc. New England Zool. Club, 5: 54, 9
April.
= Sorex daptiaenodon sanguinidens G. M.
Allen, 1914. See Yudin (1989: 198).
Holotype. MCZ 15012. Skin and skull. Adult female.
Locality. (Russian Federation): northeastern Sibe-
ria, Nijni Kolymsk ( = Nizhnekolymsk), near mouth
Kolyma River. 11 December 1911.
Collector J. Koren. Original number 221.
Condition. Tail separate from sldn. Skull complete.
Mandible disarticulated.
Type Series. There is a large series of paratypes,
most of which are still in the MCZ.
Sorex virG. M. Allen, 1914b
Proc. New England Zool. Club, 5: 52, 9
April.
= Sorex roboratus Hollister, 1913. See
Hoffman (1985: 17).
Holotype. MCZ 15068. Skin and skull. Adult female.
Locality. (Russian Federation): northeastern Sibe-
ria, Nijni Kolymsk ( = Nizhnekolymsk), near mouth
of Kolyma River. 19 December 1911.
Collector J. Koren. Original number 230.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a large series of paratypes,
most of which are still in the MCZ.
Genus SL/A/CL/S Ehrenberg, 1832
Suncus aterMedway, 1965
Mammals of Borneo, J. Malay. Branch R.
Asiat. Soc, 36: 38, December.
Holotype. MCZ 36574. Skin, skull, and postcranial
skeleton. Adult female.
Locality. Malaysia: northern Borneo, Sabah,
Mount Kinabalu, Lumu Lumu, 5,500 ft (1,678 m).
7 July 1937.
Collector J. A. Griswold, Jr. Original number 462.
Condition. Skin complete. Skull partial (tympanic
bullae missing), and mandible disarticulated. Post-
cranial skeleton complete.
Type Series. Holotype only.
Comments. Considered a valid species by Hutterer
(1993: 101) and Nowak (1999: 223).
Suncus varilla minor G. M. Allen and
Loveridge, 1933
Bull. Mus. Comp. Zool, 75: 57,
Februaiy.
Holotype. MCZ 26754. Skin and skaill. Adult female.
Locality. Tanganyika Territory (=Tanzania): Urun-
gu, Kitungulu, 4,500 ft (1,373 m). 14 May 1930.
Collector. A. Loveridge.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotyj^e only.
Comments. Retained as a valid subspecies by Heim
de Balsac and Meester (1977: 6).
140 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Family TALPIDAE Fischer de Waldheim,
1817
Genus NEUROTRICHUS Gm\her, 1880
Neurotrichus gibbsi hyacinthinus Bangs,
1897d
Amer. Nat., 31: 240, 1 March.
Holotype. B1240. Sldn and skull. Adult female.
Locality. U.S.A. (United States): California, Marin
County, Nicasio. 10 March 1894.
Collector. C. A. Allen. Original number 694.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; B1241, skin and skull,
adult male.
Comments. Retained as a valid subspecies by Hall
(1981: 67) and Carraway and Verts (1991: 1).
Genus SCALOPUS Desmarest, 1 804
Scalops anastasae Bangs, 1898b
Proc. Boston Soc. Nat. Hist., 28: 212, 15
March.
= Scalopus aquaticus anastasae (Bangs,
1898). See Jackson (1915: 39).
Holotype. B7192. Skin and skull. Adult male.
Locality. U.S.A. (United States): Florida, St. Johns
County, Anastasia Island, Point Romo. 16 Febioiary
1897.
Collector O. Bangs. Original number 10.
Condition. Skin and skaiU complete. Mandible dis-
articulated.
Tijpe Series. 4 paratypes; B719.3-B7196; all repre-
sented by sldn and skull, 2 females and 2 males.
Comments. S. a. anastasae was retained as a valid
subspecies by Hall (1981: 72).
Scalops texanus aereus Bangs, 1896li
Proc. Biol. Soc. Washington, 10: 138, 28
December.
= Scalopus aquaticus aereus (Bangs,
1896). See Miller (1912: 8).
Holotype. B547.5. Skin and skull. Adult female.
Locality. (United States): Indian Territoiy
( = Oklalioma) (Adair County), Stilwell. 13 August
1896.
Collector. T. Surber. Original number 64.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. S. a. aereus was retained as a valid sub-
species by Hall (1981: 72).
Genus SCAPANUS Pome\, 1848
Scapanus californicus minusculus Bangs,
1899J
Proc. New England Zool. Club, 1: 70, 31
Jnly.
= Scapanus latimanus minusculus Bangs,
1899. See Grinnell and Swarth (1912:
133).
Holotype. B9189. Skin and skull. Adult female.
Locality. (United States): California, El Dorado
County, Fyffe. 10 June 1897.
Collector W. W. Price and E. M. Nutting. Original
number 15.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. S. I. minusculus was retained as a valid
subspecies by Hall (1981: 70).
Order CHIROPTERA Blumenbach, 1779
Family PTEROPODIDAE Gray, 1821
Genus AETHALOPS Thomas, 1923
Aethalops aequalis G. M. Allen, 1938b
J. Mammal., 19: 497, 14 November.
= Aethalops alecto aequalis G. M. Allen,
1938. See Hill (1961: 639).
Holotype. MCZ 36582. Skin, skull, and postcranial
skeleton. Adult female.
Locality. (Malaysia): British North Borneo ( = Sa-
bali). Mount Kinabalu, Luma Luma, 5,500 ft (1,678
m). 12 July 1937.
Collector. J. A. Griswold, Jr., Asiatic Primate E.x;-
pedition. Original number 510.
Condition. Sldn, skull, and postcranial skeleton
complete.
Type Senes. 3 paratypes; MCZ 36583, sldn, skull
and skeleton, female; MCZ 36584, skin and skull,
female; MCZ 36586, skin and skull, female.
Comments. A. a. aequalis was retained as a valid
subspecies by Koopman (1994: 35).
Genus EONYCTERIS Dobson, 1873
Eonycteris spelaea glandifera Lawrence,
1939
7/7 Barbour, Lawrence, and Peters, Bull.
Mus. Comp. Zool., 86: 38, November.
Holotype. MCZ 35159. Sldn and skull. Adult male.
Locality. Philippines: Luzon, (Nueva Ecija), Rizal,
Montalban caves near Manila. 27 Febiniary 1937.
Collector. B. Lawrence. Original number 253.
Condition. Skin and skull complete.
Type Series. In the original description, Lawrence
Type Specimens of Recent Mammals • Helaen and McFadden
141
implies the existence of several specimens in ad-
dition to the holotype; she collected 18 specimens
of glandifera during her 1936—37 expedition to the
Philippines, all of which are in the MCZ.
Comments. Retained as a valid subspecies by Ma-
haradatunkamsi and Kitchener (1997: 59).
Genus HAPLONYCTERIS Lawrence,
1939
Haplonycteris fischeri Lawrence, 1939
In Barbour, Lawrence, and Peters, BulL
Mus. Comp. ZooL, 86: 33, November.
Holotype. MCZ 35258. Skin and skull. Adult male.
Locality. Philippines: Mindoro (Oriental), Mouirt
Halcon, Bignay. 26 April 1937.
Collector F. S. Rivera. Original number BL 502.
Condition. Skin and skull complete.
Type Series. Holotype only.
Comments. Type species of the genus Haplonyc-
teris Lawrence, 1939. Considered a valid species
by Koopman (1993: 142) and Nowak (1999: 292).
Genus PTEROPUS Erxleben, Mil
Pteropus anetianus aorensis Lawrence,
1945
Proc. New England ZooL Club, 23: 66,
26 March.
Holotype. MCZ 42183. Skin and skull. Adult male.
Locality. (Vanuatu): New Hebrides, off southwest
corner of Espiritu Santo Island, Aore Island. 8
April 1944.
Collector O. L. Austin, Jr. Original number 5.
Condition. Skin complete. Skull partial (basioccip-
ital missing), right ramus of mandible broken.
Type Series. 1 paratype; MCZ 42182, skin and
skull, adult male.
Comments. Retained as a valid subspecies by
Koopman (1994: 25) and Flanneiy (1995b).
Pteropus arielG. M. Allen, 1908
Bull. Mus. Comp. ZooL, 52: 28, July.
= Pteropus giganteus ariel G. M. Allen,
1908. See Hill (1958: 5).
Holotype. MCZ 10565. Skin and skull. Adult male.
Locality. Maldive Islands: Male Atoll. 24 Decem-
ber 1901.
Collector. H. B. Bigelow, A. Agassiz E.xpedition.
Condition. Right wing of sldn damaged. Skull in-
tact, with 2 small holes in braincase.
Type Series. 1 paratype; MCZ 10566, skin and
skull, juvenile female.
Comments. P. g. ariel was retained as a valid sub-
species by Koopman (1994: 26).
Pteropus austini Lawrence, 1945
Proc. New England Zool. Club, 23: 59,
26 March.
= Pteropus woodfordi Thomas, 1888. See
Sanborn and Beecher (1947: 389).
Holotype. MCZ 42166. Skin and skull. Subadult fe-
male.
Locality. Solomon Islands: Florida Island (Nggela
Group). 20 Februaiy 1944.
Collector O. L. Austin, Jr. Original number 2.
Condition. Skin and skull complete.
Type Series. 1 paratype; MCZ 42167, skin and
skull, subadult male.
Comments. Lawrence referred to the holotype as
an adult in the original description, but Sanborn
and Beecher (1947: 389) recognized it as a sub-
adult.
Pteropus rayneri monoensis Lawrence,
1945
Proc. New England Zool. Club., 23: 63,
26 March.
Holotype. MCZ 42191. Skin and skull. Aduh male.
Locality. Solomon Islands: Treasuiy (Mono) Island.
11 October 1944.
Collector O. L. Austin, Jr. Original number 27.
Condition. Skin and skull complete.
Type Series. 2 paratypes; MCZ 42192, skin and
skull, adult male; MCZ 42193, skin and skull, adult
male.
Comments. Retained as a valid subspecies by
Koopman (1994: 24) and Flannery (1995b: 285).
Genus ROUSETTUS Gray, 1821
Rousettus madagascariensis G.
Grandidier, 1930b
Bull. Acad. Malgache, n. sen, 11: 91 (for
1928).
Holotype. MCZ 45432. Alcoholic and skull. Adult
male.
Locality. Malagasy Republic (Madagascar): (Anta-
nanarivo), between Tananarive ( = Antananarivo)
and Andevoranto, Grand forest de Est, near Be-
forona.
Collector Received by G. Grandidier from the
Academic Malgache, 1917.
Conditio72. Alcoholic, skull complete.
Type Series. Holotype only.
Comments. Considered a valid species by Koop-
man (1993: 153) and Nowak (1999: 261). In his
review of known material of jR. madagascariensis,
Bergmans (1977: 67) commented in error that the
holotype was in the Academic Malgache, Antana-
narivo, Madagascar.
142 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Family EMBALLONURIDAE Gervais, 1855
Genus PEROPTERYX Peters, 1867
Peropteryx canina phaea G. M. Allen,
1911a
Bull. Mus. Comp. ZooL, 54: 222, July.
= Peropteryx macrotis phaea G. M. Allen,
1911. See Sanborn (1937: 342).
Holotijpe. MCZ 8101. Skin and skull. Adult female.
Locality. Grenada: Point Saline(s). 29 August 1910.
Collector. G. M. Allen. Original number 15.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. There is a series of paratypes in the
MCZ.
Comments. P. m. phaea was retained as a valid sub-
species by Hall (1981: 82) and Koopman (1994:
47).
Genus RHYNCHONYCTERIS Peters,
1867
Rhynchiscus naso priscus G. M. Allen,
1914c
Proc. Biol. Soc. Washins^ton, 27: 109, 10
July.
= Rhynchonycteris naso (Wied-Neuwied,
1820). See Sanborn (1937: 326).
Holotijpe. MCZ 13208. Skin and skull. Adult.
Locality. Me.\ico: Quintana Roo, Xcopen. 18 Feb-
ruary 1912.
Collector. J. L. Peters. Original number 13.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; MCZ 13209, skin and
skuU, adult male; MCZ 14637, alcoholic, adult
male; MCZ 14638, alcoholic, adult female.
Family NYCTERIDAE Van der Hoeven,
1855
Genus NYCTERIS G. Cuvier and E.
Geoffroy, 1795
Nycteris madagascariensis G. Grandidier,
1937
Bull. Mus. Nat. Hist. Paris, 9: 353, 25
November.
Holotype. MCZ 45433. Body in alcohol skull extract-
ed.
Locality. Madagascar: (Antsiranana), Diego-Suarez
( = Antsiranana), Valley of the Rodo, north of Pir-
kana near the Ankarana, 12°5'-13°0'S, 49°5'E. June
1910.
Collector. Grandidier collection.
Condition. Alcoholic, skull complete.
Type Series. 1 paratype, MCZ 45434, in alcohol,
skull extracted, female.
Comments. N. madagascariensis was included in N.
macrotis by Koopman (1993: 162) but retained as
a valid species by Peterson et al. (1995; 6.3).
Nycteris nana tristis G. W\. Allen and
Lawrence, 1936
Bull. Mus. Conip. Zool., 79: 47, January.
^Nycteris nana (Andersen, 1912). See
Hayman and Hill (1971: 19).
Holotype. MCZ 31156. Skin and skull. Adult female.
Locality. Kenya: (W. Nyanza), Kakamega District,
Kaimosi. 13 Februaiy 1934.
Collector A. Loveridge.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Family RHINOLOPHIDAE Gray, 1825
Genus HIPPOSIDEROSGmy, 1831
Hipposideros curtus G. M. Allen, 1921
Rev. Zool. Africaine, 9: 194, December.
Holotype. MCZ 19305. Body in alcohol. Female.
Locality. Cameroons ( = Cameroon): (Littoral), Sak-
bayeme. 1920.
Collector G. Schwab.
Condition. Alcoholic. Skull extracted but not
cleaned (still in alcohol).
Type Series. Holotype only.
Comments. H. curtus was considered a valid spe-
cies by Koopman (1993: 172) and Nowak (1999:
333). '
Hipposideros erigens Lawrence, 1939
In Barbour, Lawrence, and Peters, Bull.
Mus. Comp. Zool., 86: 56, November.
= l-lipposideros bicolor erigens Lawrence,
1939. See Hill (1963: 28).
Holotype. MCZ 35197. Skin and skull. Adult male.
Locality. Philippines: Mindoro, (Oriental) Tabucala
cave near Calapan, northern base of Mount Hal-
con. 7 March 1937.
Collector B. Lawrence. Original number 307.
Condition. Skin complete. Skull partial (occiput
missing), and mandible disarticulated.
Type Series. 3 paratypes; MCZ 35195, skin and
skull, adult female; MCZ 35196, sldn and skull,
adult male; MCZ 35198, skin and skull, adult fe-
male.
Comments. H. b. erigens was retained as a valid
subspecies by Koopman (1994; 61).
Type Specimens of Recent Mammals • Helaen and McFadden
143
Hipposideros turpis Bangs, 1901a
Amer. Nat., 35: 561, 31 July.
= Hipposideros turpis turpis Bangs, 1901.
See Hill (1963: 94).
Holotijpe. MCZ 10003. Skin and skull. Adult female.
Locoliti/. (Japan): Ryaikyu Islands, southern group
of Liu kin Islands, Ishigaki Island. 10 May 1899.
Collector. I. Zensaku.
Condition. Sldn complete. Skull partial (occipital
region missing).
Type Series. 2 paratypes; MCZ 10002, skin and
skull, adult female; MCZ 10004, skin and skvill,
adult male.
Comments. H. tuiyis was considered a valid species
by Koopman (1993: 175) and Nowak (1999: 334).
Genus RHINOLOPHUS Lacepe6e, 1799
Riiinoloplius megaphyllus igniter G. M.
Allen, 1933
J. Mammal., 14: 149, 15 May.
= Rtiinolophus megapfiyilus megapiiyllus
Gray, 1834. See Koopman (1984: 9).
Holotijpe. MCZ 29078. Skin and skull. Adult male.
Locality. Australia: Queensland, Cape York, Coen.
12 June 1932.
Collector P. J. Darlington, Jr., Hai-vard Australian
Expedition. Original number 185.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. 1 paratyj^jc; MCZ 29079, skin and
skull, adult male.
Rtiinolophus ptiilippinensis alleni
Lawrence, 1939
In Barbour, Lawrence, and Peters, Bull.
Mus. Comp. Zool., 86: 46, November.
Holotype. MCZ 35097. Sldn and skull. Adult female.
Locality. Philippines: Mindoro, (Oriental) Tabucala
cave near Calapan, northern base of Mount Hal-
con. 7 March 1937.
Collector. B. Lawrence. Original number 302.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 2 pai'atypes; MCZ 35098, skin and
sk-ull, adult female; MCZ 35099, skin and skull,
adult female.
Comments. Retained as a valid subspecies by
Koopman (1994: 57).
Genus TRIAENOPS Dobson, 1871
Triaenops aurita G. Grandidier, 1912
Bull. Mus. Hist. Nat. Paris, 18: 8, 25
Januaiy.
Holotype. MCZ 45080. Mummy.
Locality. Malagasy Republic ( = Madagascar): (An-
tsiranana), Diego-Suarez (=Antsiranana). 1910.
Collector. Dr. Mazieres.
Condition. Mummy (dried carcass), complete.
Type Series. Holotype only.
Comments. Traditionally included in Triaenops fur-
culus, as in G. M. Allen (1939: 82) and Koopman
(1993: 175), but retained as distinct by Peterson et
al. (1995: 81) pending further material from the
area of the type locality. Known only from the ho-
lotype.
Family MORMOOPIDAE de Saussure,
1860
Genus PTERONOTUS Gray, 1838
Ctiilonycteris parnellii pusillus G. M. Allen,
1917c
Proc. Biol. Soc. Washington, 30: 168, 23
October.
= Pteronotus parnellii pusillus (G. M. Allen,
1917). See Smith (1972: 67).
Holotype. MCZ 16468. Skin and skull. Female.
Locality. Dominican Republic: Santo Domingo,
Arroyo Salado. 7 March 1916.
Collector J. L. Peters. Original number 227.
Condition. Skin complete. Skull partial (left wall of
braincase broken, left tympanic bulla missing).
Type Series. 2 paratypes; MCZ 16599, female;
MCZ 16600, female; both in alcohol.
Comments. P. p. pusillus was retained as a valid
subspecies by Hall (1981: 92) and Koopman (1994:
71).
Ctiilonycteris torrelG. M. Allen, 1916a
Proc. New England Zool. Club, 6: 4, 8
Februai-v.
= Pteronotus quadridens quadridens
(Gundlach, 1840). See Silva-Taboada
(1976: 7).
Holotype. MCZ 11672. Body in alcohol, skull extract-
ed. Adult female.
Locality. Cuba: (Guantanama), Baracoa, La Cueva
de la Majana. 15 June 1915.
Collector V. J. R. Verrier. Presented to the MCZ
by Carlos de la Torre.
Condition. Alcoholic, skull complete.
Type Series. 2 paratypes; MCZ 11670, male; MCZ
11671, female; both in alcohol.
Family PHYLLOSTOMIDAE Gray, 1825
Genus AMETRIDA Gray, 1847
Ametrida minor H. Allen, 1894
Proc. Boston Soc. Nat. Hist., 26: 240, 16
May.
= Ametrida centurio Gray, 1847. See
Peterson (1965: 5).
Holotype. MCZ 11274. Body in alcohol, skull extract-
ed. Adult male.
144
Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Locality. Suriname: Paramaribo. Collected some-
time between 1832 and 1839.
Collector. F. W. Cragin.
Condition. Alcoholic, skull complete.
Type Series. Holotype only.
Comments. For comments on the type locality and
date of acquisition, see G. M. Allen (1902a: 88). A.
minor actually represents the male specimens of A.
centurio.
Genus ARTIBEUS Leach, 1821
Artibeus femurvillosum Bangs, 1899k
Proc. New England Zool. Club, 1: 73, 24
November.
= Artibeus lituratus palmarum J. A. Allen
and Chapman, 1897. See Hershkovitz
(1949: 445).
Holotype. B8314. Skin and skull. Adult male.
Locality. Colombia: (La Guajira), La Concepcion,
3,000 ft (915 m). 21 March 1899.
Collector W. W. Brown, Jr. Original number 31.
Condition. Skin and skull complete.
Ti/pe Series. Holotype only.
Genus EROPHYLLA Miller, 1906
Erophylla sezekomi syops G. M. Allen,
1917c
Proc. Biol. Soc. Washington, 30: 167, 23
October.
Holotype. MCTj 13713. Body in alcohol, skull extract-
ed. Adult male.
Locality. Jamaica: (St. James), Montego Bay. 14
March 1912.
Collector J. A. Cushman.
Condition. Alcoholic, skull complete.
Type Series. 7 paratypes; MCZ 13709-13712,
13714-13716, all alcoholic. MCZ 13709 and 13712
were sent in e.xchange to the USNM.
Comments. Retained as a valid subspecies by Hall
(1981: 171) and Koopman (1994: 79).
Genus GLOSSOPHAGA E. Geoffrey,
1818
Glossophaga longirostris Miller, 1898
Proc. Acad. Nat. Sci. Philadelj^hia, 1898,
p. 330, 2 August.
= Glossophaga longirostris longirostris
Miller, 1898. See Miller (1913: 422).
Holotype. B8046. Skin and skull. Adult female.
Locality. Colombia: (Magdalena), Santa Marta
Mountains, near Santa Marta. 10 February 1898.
Collector W. W. Brown, Jr. Original number 60.
Condition. Skin complete. Skull partial (left zygo-
matic arch missing).
Type Series. Holotype only.
Comments. G. longirostris was considered a valid
species by Koopman (1993: 184) and Nowak (1999:
368).
Genus LONCHOPHYLLA Thomas, 1903
Lonchophylla hesperia G. M. Allen, 1908
Bull. Mus. Comp. Zool, 52: 35, July.
Holotype. MCZ 7011. Body in alcohol, skull extract-
ed. Adult male.
Locality. Peru: (Contralmirante Villar), Tumbes,
Zorritos.
Collector. F. H. Bradley.
Condition. Alcoholic, skull complete.
Type Series. 2 paratypes; in the YPM; 1034 and
1035; both in alcohol.
Comments. Considered a valid species by Koop-
man (1993: 181) and Nowak (1999: 372). This rare-
ly collected bat is known by only two museum
specimens in addition to the type series: USNM
283177 and LSUMZ 14121 (Gardner 1976: 5).
Genus PLATYRRHINUS 6e Saussure,
1860
Vampyrops umbratus Lyon, 1902
Proc. Biol. Soc. Washington, 15: 151, 20
June.
= Platyrrhinus umbratus (Lyon, 1902). See
Koopman (1993: 191).
Holotype. B8180. Sldn and skull. Adult male.
Locality. Colombia: (LaGuajira), San Miguel. 8
June 1898.
Collector W. W. Brown, Jr. Original number 234.
Condition. Sldn and skull complete.
Type Series. 2 paratypes; B8300, skin, male; B8301,
sldn, male.
Comments. P. umbratus was considered a valid spe-
cies by Koopman (1993: 191) and Nowak (1999:
389). Platyrrhinus has priority over the genus name
Vampyrops (Gardner and Ferrell 1990: 501-503).
Vampyrops zarhinus H. Allen, 1891
Proc. Acad. Nat. Sci. Philadelphia, 1891,
p. 400, 22 September.
= Platyrrhinus helleri {Peters, 1866). See
Hall and Kelson (1959: 131).
Holotype. MCZ 3211. Body in alcohol, skull extract-
ed. Adult female, pregnant.
Locality. Panama: Canal Zone, Obispo. 1872. See
comments.
Collector Hassler Expedition.
Condition. Alcoholic, skull complete.
Type Series. Holotype only.
Type Specimens of Recent Mammals • Helgen and McFadden 145
Comments. In the original description, H. Allen re-
port:ed that this specimen had been collected in
Brazil by the Thayer expedition. G. M. Allen
emended this apparently erroneous locality to
Obispo, Panama, in accordance ^A-ith the accession
catalogue of the MCZ (1931: 236-237). In support
of Aliens decision, Rouk and Carter (1972: 4) stat-
ed, after examining the holotype of zarhinus, that
it is "quite like specimens of [Platyrrhiniis] helleri
from Mexico and Central America, and unlikely to
have come from Brazil." For the use of the genus
Platyrrhinus over Vampyrops, see Gardner and
Ferrell (1990: 501-503).
Genus VAMPYRODES Thomas, 1 900
Vampyrodes major G. M. Allen, 1908
Bull. Mus. Comp. ZooL, 52: 38, July.
= Vampyrodes caraccioli major G. M.
Allen, 1908. See Handley (1966: 766).
Holotype. MCZ 6756. Body in alcohol. Adult female.
Locality. Panama: San Pablo (now covered by Ga-
tun Lake). Date unrecorded.
Collector. A. Lesley.
Condition. Alcoholic.
Type Series. Holotype only.
Comments. V. c. major was retained as a valid sub-
species by Koopman (1994: 88).
Family MOLOSSIDAE Gervais, 1855
Genus MOPS Lesson, 1842
Chaerephon leucostigma G. M. Allen,
1918a
Bull. Mus. Comp. ZooL, 61: 513,
Februaiy.
= Mops condylurus leucostigma (G. M.
Allen, 1918). See Koopman (1994: 141).
Holotype. MCZ 16344. Skin and skull. Aduh female.
Locality. Malagasy Republic (Madagascar): (Anta-
nanarivo), Tananarive (= Antananarivo). December
1915.
Collector. F. R. Wulsin.
Condition. Skin partial (bare spot on ventrum).
Skull damaged (right and left zygomatic arch miss-
ing; supraoccipital chipped).
Type Series. 1 paratype; MCZ 16345, skin and
skull, male.
Comments. Peterson et al. (1995: 168) used the
name Tadarida leucostigma.
Mops angolensis orientis G. M. Allen and
Loveridge, 1942
Bull. Mus. Comp. ZooL, 89: 166,
Februaiy.
= Mops condylurus orientis G. M. Allen
and Loveridge, 1942. See Koopman
(1994: 141).
Holotype. MCZ 38829. Skin and skull. Adult male.
Locality. Tanganyika Territoiy (= Tanzania): Mtwa-
ra, Ruviuna River, Kitaya, 300 ft (92 m). 3 April
1939.
Collector A. Loveridge.
Condition. Skin and skull complete.
Tijpe Series. 9 paratypes; MCZ 38826-38828,
38830-38835, all represented by skin and skull, 4
females and 5 males.
Genus OTOMOPS Thomas, 1913
Otomops papuensis Lawrence, 1948
J. Mammal., 29: 413, 31 December.
Holotype. MCZ 45769. Body in alcohol, skull extract-
ed. Adult female.
Locality. Papua New Guinea: Vailala River.
Collector. Bought from Ward's Natural Science Es-
tabhshment, April 1948.
Condition. Alcoholic, skull partial (right zygomatic
arch missing).
Type Series. Holotype only.
Comments. Considered a valid species by Koop-
man (1993: 239) and Nowak (1999: 482). Accord-
ing to Flanneiy (1995a: 481), O. papuensis has
been collected on only two occasions and, other
than the holotype, is known by only 10 specimens;
2 in the BMNH, the remainder in the biological
collections of the University of Papua New Guinea.
Family VESPERTILIONIDAE Gray, 1821
Genus EPTES/CL/S Rafinesque, 1820
Eptesicus darlingtoni G. M. Allen, 1933
J. Mammal., 14: 150, 15 May.
Holotype. MCZ 29113. Skin and skull. Adult female.
Locality. Australia: Queensland, Queensland Na-
tional Park, MacPherson Ranges, 3,000 ft (915 m).
10 March 1932.
Collector P. J. Darlington, Jr., Harvard Australiaji
Expedition. Original number 30.
Condition. Skin and skull complete.
Type Series. 1 paratype; MCZ 29120 (now Queens-
land Museum J 5476), skin and skull, adult female.
Comments. McKean et al. (1978: 533) and Koop-
man (1993: 203) included darlingtoni in Eptesicus
pumilus. However, Koopman also used the name
Pipistrellus darlingtoni (1994: 116). Hoye (1995:
146 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
537) considered darlingtoni to be a valid species of
VespadeJus, to which he gave full generic rank.
Genus HARPIOCEPHALUS Gray, 1842
Harpiocephalus rufulus G. M. Allen, 1913
Proc. Biol. Soc. Washington, 26: 214, 20
December.
= Harpiocephalus harpia rufulus G. M.
Allen, 1913. See Ellerman and Morrison-
Scott (1951: 187).
Holotijpe. MCZ 14206. Skin and skull. Adult male.
Localitij. Vietnam: Tonkin, Lao-Kai ( = Lao Cai). 3
Januaiy 1912.
Collector. Kobayashi Collection. Original number
14.
Condition. Skin complete. Skull partial (parietals
broken).
Type Serie.s. Holotype only.
Comments. H. h. nifuliis was retained as a valid
subspecies by Koopman (1994: 133).
Genus IDIONYCTERIS Anthony, 1923
Corynorhinus phyllotis G. M. Allen, 1916b
Bull. Mns. Comp. Zool., 60: 352, April.
= Idionycteris phyllotis phyllotis (G. M.
Allen, 1916). See Tumlison (1993: 418).
Holotype. MCZ 5943. Skin and skull. Adult.
Locality. Mexico: San Luis Potosi. 24 March 1878.
Collector. E. Paliner.
Condition. Skin and skull complete.
Tijpe Series. Holotype only.
Comments. I. phyllotis was considered a valid spe-
cies by Koopman (1993: 205) and Nowak (1999:
457).
Genus LASIURUS Gray, 1838
Atalapha brachyotisJ. A. Allen, 1892
Bull. Amer. Mus. Nat. Hist., 4: 47, 25
March.
= Lasiurus borealis brachyotis (J. A. Allen,
1892). See Niethammer (1964: 595).
Holotype. MCZ 11143. Body in alcohol. Male.
Locality. (Ecuador), Galapagos Islands: Chatham
Island. 23 June 1891.
Collector G. Baur.
Condition. Alcoholic. The specimen was received
without a skull.
Type Series. Holotype only.
Comments. Lasiunis brachyotis has often been ac-
corded specific status, as in Nowak (1999: 451). In-
cluded in L. borealis as a valid subspecies by Koop-
man (1994: 129).
Genus /W/OT/S Kaup, 1829
Myotis abbotti nugax G. M. Allen and
Coolidge, 1940
Bull. Mus. Comp. Zool., 87: 137, 31
December.
= Myotis muricola nugax G. M. Allen and
Coolidge, 1940. See Koopman (1994:
104).
Holotype. MCZ 36076. Skin and skull. Adult male.
Locality. Malaysia: north Borneo, Sabah, Mount
Kinabalu, Bundutuan, 3,500 ft (1,068 m). 25 July
1937.
Collector J. A. Griswold, Jr., Asiatic Primate Ex-
pedition. Original number 626.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. 16 paratypes, MCZ 36072-36075,
36077-36080, 36082-83, 36085-89, 36091; all rep-
resented by skin and skull, 12 females and 4 males.
Myotis albicinctus G. M. Allen, 1919b
J. Mammal., 1: 2, 28 November.
= Myotis lucifugus carissima Thomas,
1904. See Miller and G. M. Allen (1928:
50).
Holotype. MCZ 11747. Sldn and skull. Adult male.
Locality. (United States): California, (Tulare Coun-
ty), Mount Whitney, 11,000 ft (3,355 m). 14 July
1915.
Collector G. M. Allen. Original number 1.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratyj^^e, probably at USNM.
Comments. The skull of the holotype, which had
been mislaid at the time that albicinctus was de-
scribed, has subsequently been found and reunited
with its skin.
Myotis sodalis Miller and G. M. Allen,
1928
Bull. U.S. Nat. Mus., 144: 130, 25 May.
Holotype. MCZ 10988. Skin and skull. Adult female.
Locality. (United States): Indiana, (Crawford
County), Wyandotte cave. 7 March 1904.
Collector J. O. Sibert.
Condition. Skin and skull complete. Mandible dis-
articulated.
Comments. Considered a valid species by Koop-
man (1993: 215) and Nowak (1999: 419).
Type Series. Miller and Allen based their descrip-
tion on an examination of 443 specimens, described
by locality in the original description (1928: 133).
30 paratypes are in the MCZ, the others are in tlie
FMNH, USNM, AMNH. and BMNH.
Type Specimens of Recent Mammals • Hehen and McFadden 147
Genus A/VCT/CE/L/S Rafinesque, 1819
Nycticeius africanus G. M. Allen, 1911b
Bull. Mus. Comp. Zool., 54: 328,
December.
= Nycticeius sctilieffeni albiventer Thomas
and Wroughton, 1908. See Hayman and
Hill (1971: 36).
Holotype. MCZ 8272. Skin and skull. Male.
Locality. British East Africa ( = Kenya): Meru Riv-
er, effluent of northern Guaso N^'iro ( = Ewaso Ngi-
ro). 11 August 1909.
Collector. G. M. Allen. Original number 113.
Condition. Skin and skull complete.
Type Series. Holotype only.
Genus PIPISTRELLUS Kaup, 1829
Eptesicus phasma G. M. Allen, 1911b
Bull. Mus. Comp. Zool, 54: 327,
December.
= Pipistreilus rendalii phasma (G. M. Allen,
1911). See Koopman (1994: 117).
Holotype. MCZ 8279. Skin and skull. Male.
Locality. British East Africa ( = Kenya): Meru Riv-
er, effluent of northern Guaso Nyiro ( = Ewaso Ngi-
ro). 6 August 1909.
Collector. G. M. Allen. Original number 94.
Condition. Skin and skull complete.
Type Series. There is a small series of paratypes in
the MCZ.
Scabrifer notiusG. M. Allen, 1908
Bull. Mus. Comp. Zool., 52: 46, July.
= Pipistreilus capensis notius (G. M. Allen,
1908). See Koopman (1994: 117).
Holotype. MCZ 4555. Alcoholic, skull extracted.
Adult male.
Locality. South Africa: (Western Cape), Cape
Town.
Collector Received from E. L. Lavard, August
1864.
Condition. Alcoholic, skull partial (right and left
zygomatic arches missing; supraoccipital chipped).
Type Series. Holotyjje only.
Genus PLECOTUS E. Geoffrey, 1813
Plecotus sacrimontis G. M. Allen, 1908
Bull. Mus. Comp. Zool., 52: 50, July
= Piecotus auritus sacrimontis G. M. Allen,
1908. See Ognev (1928: 607).
Holotype. MCZ 6932. Body in alcohol. Adult male.
Locality. Japan: (Honshu), Mount Fuji. 4 Decem-
ber 1906.'
Collector. A. Ovs'ston.
Condition. Alcoholic.
Type Series. Holotype only.
Comments. P. a. sacrimontis was retained as a valid
subspecies by Koopman (1994: 110).
Genus SCOTOPHILUS Leach, 1821
Scotopiiiius altilis G. M. Allen, 1914d
Bull. Mus. Comp. Zool., 58: 350, July
= Scotopiiiius leucogaster (Gretzschmar,
1826). See Koopman (1993: 227).
Holotype. MCZ 14463. Skin and skull. Adult male.
Locality. Sudan: Blue Nile, north of (Er) Roseires,
Aradeiba. 22 January 1913.
Collector G. M. Allen, Phillips Sudan Expedition.
Original number 7.3.
Condition. Skin and skull complete.
Type Series. 3 paratypes; MCZ 14462, skin and
skull, male, exchanged to FMNH; and 14610 and
14611, both males in alcohol.
Family THYROPTERIDAE Miller, 1907
Genus THYROPTERA Sp\x, 1823
Thyroptera tricolor albigula G. M. Allen,
1923c
Proc. New England Zool. Club, 9: 1, 10
December.
= Thyroptera tricolor albiventer (Tomes,
1856). See Dunn (1931: 430).
Holotype. MCZ 20143. Body in alcohol, skull extract-
ed. Adult female.
Locality. Panama: Gutierrez, 25 miles (40.2 km)
inland from Chiriquiscito on trail from Chiriqui La-
goon, Bocas del Toro to Boquete, Chiriqui. August
1923.
Collector E. R. Dunn and C. B. Dvuyea.
Condition. Alcoholic, skull complete.
Type Series. 3 paratypes; MCZ 20144, adult male;
MCZ 20145, juvenile; MCZ 20146, juvenile; all in
alcohol.
Order ARTIODACTYLA Owen, 1848
Family TAYASSUIDAE Palmer, 1897
Genus PEC/\R/ Reichenbach, 1835
Tayassu crusnigrum Bangs, 1 902b
Bull. Mus. Comp. Zool., 39: 20, April.
^Pecari tajacu crusnigrum (Bangs, 1902).
See Hershkovitz (1951: 567).
Holotype. MCZ 10163. Skin and skull. Adult male.
Locality. Panama: Chiriqui, Boquete ( = Ba)o Bo-
quete), 4,000 ft (1,220 m). 13 April 1901.
148 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Collector. W. W. Brown, Jr. Original number 290.
Condition. Skin and skull complete.
Type Series. 2 paratypes; MCZ 10162, adult fe-
male; MCZ 10164, juvenile female.
Comments. P. t. crusnignim was retained as a valid
subspecies by Hall (1981: 1080) under the genus
Dicotijles.
Tayassu torvus Bangs, 1 898k
Proc. Biol. Soc. Washington, 12: 164, 10
August.
= Pecari tajacu torvus (Bangs, 1898). See
comments.
Holotype. B8038. Sldn and skull. Adult male.
Locality. Colombia: Magdalena, Santa Marta. 26
January 1898.
Collector. W. W. Brown, Jr. Original number 50.
Condition. Skin and skull complete.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Ca-
brera (1961: 319) under the genus Tayassu. Use of
the genus Pecari follows Grubb (1993: 380).
Family MONODONTIDAE Gray, 1821
Genus DELPHINAPTERUS Lacepede,
1804
Beluga declivis Cope, 1865
Proc. Acad. Nat. Sci. Philadelphia, 17:
278.
= Delphinapterus leucas (Pallas, 1776).
See Hershkovitz (1966: 111).
Holotype. MCZ 1195. Skull and postcranial skeleton.
Locality. "Arctic Seas" (probably Greenland).
Collector. E. K. Kane.
Condition. Skull partial (right mandibular ramus
missing). Postcranial skeleton complete except for
missing left flipper. Right flipper and tail are un-
cleaned, with tissue largely intact.
Type Series. Holotype only.
Comments. Hershkovitz (1966: 111) erroneously
stated that the holotype of declivis was deposited
in the Academy of Natural Sciences in Philadel-
phia, and the holotype of Beluga concreta Cope,
1865 was in the MCZ. The opposite is in fact true;
Philadelphia holds the type of B. concreta.
Family PHOCOENIDAE Gray, 1825
Genus NEOPHOCAENA Palmer, 1899
Neomeris asiaeorientalis Pilleri and Gihr,
1972
Invest. Cetacea, 4: 126.
= Neophocaena phocaenoides
asiaorientalis (Pilleri and Gihr, 1972). See
van Bree (1973: 17).
Holotype. MCZ 19998 (but see comments). Skull and
postcranial skeleton. Adult male.
Locality. China: Kiangsu (=Jiangsu), Kiangyin, 80
miles (129 km) northwest of Shanghai. 7 April
1922.
Collector F. R. Wulsin.
Condition. Skin and postcrajiial skeleton complete.
Type Series. Holotype only.
Comments. For a discussion of the nomenclature
and synonymy of this form, consult van Bree
(1973). Because Neomeris asiaorientalis Pilleri and
Gihr, 1972 is in fact a replacement name for the
preoccupied name Delpliinus inelas Schlegel, 1841,
the holotype of this new name is the same as that
of Schlegel's name, RMNH 23079.
Family CERVIDAE Goldfuss, 1820
Genus ODOCO//.EL/S Rafinesque, 1832
Cariacus osceola Bangs, 1896b
Proc. Biol. Soc. Washington, 10: 26, 25
February.
= Odocoileus virginianus osceola (Bangs,
1896). See Lydekker (1915: 148).
Holotype. B2394. Skin and skull. Adult female.
Locality. (United States): Florida, Citrus County,
Citronelle. 29 December 1893.
Collector. F. L. Small. Original number 1107.
Condition. Skin and skull complete. Mandible dis-
articulated.
Ttjpe Series. 4 paratypes; B2391, adult male,
B2392, adult male, B2393, adult female, B2395,
juvenile male, all represented by skin and skull.
Comments. O. v. osceola was retained as a valid
subspecies by Hall (1981: 1096) and Smith (1991:
1).
Odocoeleus [sic] virginianus louisianae G.
M. Allen, 1901
Amer. Nat., 35: 449. 28 June.
= Odocoileus virginianus macroura
(Rafinesque, 1817). See Miller and
Kellogg (1955: 804).
Holotype. B9111. Skin and skull. Adult male.
Locality. (United States): Louisiana, Morehouse
Parish, Mer Rouge. 8 November 1898.
Collector B. V. Lilly
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; B9112, B8622, B8623, all
males represented by skin and skull.
Odocoileus americanus borealis Miller,
1900
Bull. New York State Mus. 8: 83, 21
November.
= Odocoilus virginianus borealis Miller,
1900. See Trouessart (1905: 704).
Holotype. B4999. Skin and skufl. Adult male.
Locality. (United States): Maine, (Hancock Coun-
ty), Bucksport. 12 December 1895.
Type Specimens of Recent Mammals • Helaen and McFadden
149
Collector. A. G. Dorr.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. O. v. horealis was retained as a valid
subspecies bv Hall (1981: 1092) and Smith (1991:
1).
Odocoileus virginianus clavium Barbour
and G. M. Allen, 1922
J. Mammal., 3: 73, 9 May.
Holotype. MCZ 19120. Skull and head skin. Adult
male.
Locality. (United States): Florida, (Monroe Coun-
ty), Big Pine Key Winter 1920.
Collector T. Barbour.
Condition. Skin and skaill complete. Mandible dis-
articulated.
Type Series. 2 paratyjDcs, MCZ 18497, sldn and
skull, juvenile male; MCZ 18060, sldn and skull,
juvenile male.
Comments. Retained as a valid subspecies by Hall
(1981: 1093) and Smith (1991: 1).
Genus R/\A/G/FE/=? Hamilton Smith, 1827
Rangifer arcticus caboti G. M. Allen,
1914a
Proc. New England Zool. Club, 4: 104,
24 March.
= Rangifer tarandus caribou (Gmelin,
1788). See Banfield (1962: 70).
Holotype. MCZ 15372. One shed antler. Adult male.
Locality. Canada: northeast coast of Labrador,
about 30 miles (48.3 km) north of Nachvak. 1909.
Collector O. Biyant.
Condition. Single antler; complete.
Type Series. Holotype only.
Rangifer terraenovae Bangs, 1896!
Preliminaiy Description of the
Newfoundland Caribou, Boston, p. 1, 11
November.
= Rangifer tarandus caribou (Gmelin,
1788). See Banfield (1962: 70).
Holotype. B3778. Skull and head sldn. Adult male.
Locality. Canada: Newfoundland, Codroy 8 Sep-
tember 1895
Collector E. Doane.
Condition. Skull and head sldn complete.
Type Series. Bangs remarks that he has "secured a
series of this fine caribou," corresponding to
B3779-B3781 and B5757-B5760.
Comments. J. A. Allen published a description of
Rangifer terraenovae on 21 November 1896, spec-
ifying AMNH 11775, a mounted specimen of a
male adult, as the holotype (1896: 233). Bangs' de-
scription of this taxon pre-dates J. A. Allen's by 10
days and thus has priority.
Family BOVIDAE Gray, 1821
Genus DAIVIALISCUS Sclater and
Thomas, 1894
Damaliscus phillipsi Harper, 1 939
Proc. Biol. Soc. Washington, 50: 90, 5
June.
= Damaliscus pygargus phiillipsi Harper,
1939. See comments.
Holotype. MCZ 35443. Skull and skin. Adult male.
Locality. South Africa: Orange Free State. 23 July
1935.
Collector. P. Andreka. Original number 1958d.
Condition. Skin complete. Skull partial (most of
palate, left maxilla, and left mandibular ramus
missing).
Type Series. Paratype material consists of MCZ
35444, skin and skull of an adult female as well as
the following, which Haij)er examined in the col-
lection of the Academy of Natural Sciences of Phil-
adelphia; "a mounted head, a skull, and a set of
horns purchased in Kimberley, Cape Province; a
mounted head and a set of horns from 'South Af-
rica'; and two skins and skulls from the Zoological
Society of Philadelphia."
Comments. Ansell (1972; 55) used the name Dam-
aliscus dorcas phillipsi. For the use of pygargns
over dorcas, see Rookmaaker (1991; 190).
Order CARNIVORA Bowdich, 1821
Family CANIDAE Fischer de Waldheim,
1817
Genus CANIS Linnaeus, 1758
Canis lupus beothiucus G. M. Allen and
Barbour, 1937
J. Mammal, 18: 230, 14 May.
Holotype. MCZ 351. Skull and postcranial skeleton.
Adult, probably male.
Locality. Canada: Newfoundland. About 1865.
Collector J. M. Nelson.
Condition. Skull and postcranial skeleton com-
plete.
Type Series. 4 paratypes; 348, skull, adult male;
349, skull, adult, probably male; 350, skull, adult
female; MCZ 28726, skin.
Comments. C. I. beothiicus became extinct around
1911. Retained as a vahd subspecies by Hall (1981:
930).
150 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Pachycyon robustus J. A. Allen, 1885
Mem. Mus. Comp. Zool., 10: 4,
December.
= Canis familiaris Linnaeus, 1758. See G.
M. Allen (1920a: 498).
Holotype. MCZ 7091. Postcranial skeleton.
Locality. (United States): Virginia, Lee County, Ely
Cave. Probably 1875.
Collector. N. S. Shaler.
Condition. Skeleton partial (right scapula, right hu-
merus, right femur, right tibia, pelvis).
Type Series. Holotype only.
Comments. This specimen is a domesticated dog of
Native Americans. C. lupus familiaris is the name
now widely used for the domestic dog (Wozencraft
1993: 281). P. robustus is the type species of Pa-
chycyon ]. A. Allen, 1885. Pachycyon is a synonym
oiCanis Linnaeus, 1758, which is commonly over-
looked, for example, in Wozencraft (199.3) and Mc-
Kenna and Bell (1997).
Genus CERDOCYON Hamilton Smith,
1839
Cerdocyon thous germanus G. M. Allen,
1923b
Proc. Biol. Soc. Washington, 36: 55, 28
March.
Holotype. MCZ 19850. Skin and skull. Adult.
Locality. Colombia: high savannali of Bogota, 9,000
ft (2,745 m).
Collector N. Maria. Original number 25.
Condition. Skin and skull complete.
Type Series. 5 paratypes; MCZ 19849, skin and
skull, juvenile; MCZ 20097, sldn and skull, juvenile
male; 3 specimens from the AMNH are also men-
tioned in the description.
Comments. Retained as a valid subspecies by Berta
(1982: 1).
Urocyon aquilus Bangs, 1 898h
Proc. Biol. Soc. Washington, 12: 93, 30
April.
= Cerdocyon thous aquilus (Bangs, 1898).
See Langguth (1969: 178).
Holotype. B8001. Skin and skull. Adult male.
Locality. Colombia: (Magdalena), Santa Marta
Mountains, between 2,000 and 3,000 ft (610-915
m). 10 February 1898.
Collector. W. W. Brown, Jr. Original number 58.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; B8002, skin and skull,
adult female.
Comments. Retained as a valid subspecies by Berta
(1982: 1).
Genus L/ROC/OA/ Baird, 1858
Urocyon cinereoargenteus furvus G. M.
Allen and Barbour, 1923
Bull. Mus. Comp. Zool, 65: 266,
February.
Holotype. MCZ 19774. Skin and skull. Probably fe-
male.
Locality. Panama; Canal Zone, 3 miles (4.8 km)
west of Balboa. April 1922.
Collector T. Barbour and W. S. Brooks.
Condition. Sldn and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Hall
(1981: 943) and Fritzell and Haroldson (1982; 1).
Urocyon cinereoargenteus ocyttious
Bangs, 1899h
Proc. New England Zool. Club, 1: 43, 5
June.
Holotype. B4290. Skin and skull. Adult female.
Locality. (United States): Wisconsin, Grant Coun-
ty, Platteville. 25 Januaiy 1896.
Collector N. E. France.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. Retained as a valid subspecies by Hall
(1981: 943) and Fritzell and Haroldson (1982: 1).
Genus UL/LPES Frisch, 1775
Vulpes deletrix Bangs, 1 898d
Proc. Biol. Soc. Washington, 12: 36, 24
March.
= Vulpes vulpes rubricosa Bangs, 1 898.
See Churcher (1960: 359).
Holotype. B6967. Sldn and skull. Adult female.
Locality. (Canada): Newfoundland, Bay St.
George. 24 April 1897.
Collector E. Doane.
Condition. Sldn and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Vulpes pennsylvanica vafra Bangs, 1 897f
Proc. Biol. Soc. Washington, 11: 53, 16
March.
Name preoccupied by Vulpes vafer
Leidy, 1869.
Type Specimens of Recent Mammals • Helpen and McFadden
151
Vulpes pennsylvanica rubricosa Bangs,
1898a
Science, n. sen, 7: 271, 25 Februaiy.
(Replacement name for Vulpes
pennsi/Ivanica vafra Bangs, 1897)
= Vulpes vulpes rubricosa Bangs, 1898.
See Churcher (1960: 359).
Holotype. B116. Skin and skull. Adult female.
Locality. (Canada): Nova Scotia, Digby. 3 Novem-
ber 1893.
Collector. O. Bangs.
Condition. Skin and skull complete.
Type Series. 4 paratypes; B1991, skin and skull, and
B2001, skull, both adult males; B1992, skin and
sk-ull, and B2002, skull.
Comments. V. v. rubricosa was retained as a valid
subspecies by Hall (1981: 939). Vulpes fulvus nib-
ricatu.s Miller, 1900 (p. 128) was a misspelling and
thus accidental renaming of Vulpes pennsylvanica
rubricosa Bangs, 1898.
Vulpes rubricosa bangsi Merriam, 1900
Proc. Washington Acad. Sci., 2: 667, 28
December 28.
= Vulpes vulpes rubricosa Bangs, 1898.
See Churcher (1960: 359).
Holotype. B8880. Skin and skull. Juvenile female.
Locality. (Canada): Labrador, Lance ( = Lanse) au
Loup. 2 October 1899.
Collector E. Doane.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 1 paratype; B8879, skin and skull,
adult male.
Family URSIDAE Fischer de Waldheim,
1817
Genus L/RSUS Linnaeus, 1758
Ursus (Euarctos) americanus sornborgeri
Bangs, 1898j
Amer. Nat., 32: 500, July.
= Ursus americanus americanus Pallas,
1780. See Bangs (1909: 467).
Holotype. B7411. Skull. Adult, probably female.
Locality. Canada: Labrador, Okkak ( = Okak). Sum-
mer 1897.
Collector J. D. Sornborger, obtained "from the Es-
kimo."
Condition. Skull complete.
Type Series. 2 paratypes; B7412, skull, female;
B7413, skull, female.
Comments. A skull from Hopedale, Labrador
(MCZ 7365), has in the past been erroneously la-
beled as the holotype of sornborgeri; B7411, the
true holoty[3e of sornborgeri, is now correctly la-
beled as such.
Family PROCYONIDAE Gray, 1825
Genus PROCYON Storr, 1780
Procyon gloveralleni Nelson and Goldman,
1930
J. Mammal., 11: 453, 11 November.
= Procyon /otor (Linnaeus, 1758). See
Corbet and Hill (1991: 104).
Holotype. MCZ 18591. Skin and skull. Juvenile male.
Locality. Barbados. 1920.
Collector F. Watts.
Condition. Skin and skull complete
Type Series. Holotype only.
Comments. Considered a valid species by Wozen-
craft (1993: 335) and Nowak (1999: 698) but almost
certainly introduced to Barbados in the 17th cen-
tuiy (Helgen and Wilson, in prep.). The last rac-
coon on Barbados was seen in 1964, and the pop-
ulation is probably extinct.
Procyon lotor elucus Bangs, 1898b
Proc. Boston Soc. Nat. Hist., 28: 219, 15
March.
Holotype. B3502. Skin and skull. Adult male.
Localitij. (United States): Florida, Brevard County,
Oak Lodge, east peninsula opposite Micco. 15 Feb-
ruary 1895.
Collector. O. Bangs.
Condition. Skin complete. Skull partial (condyle,
coronoid, and angular processes of left mandibular
ramus broken). Mandible disarticulated.
Type Series. There is a series of paratypes in the
MCZ.
Comments. Retained as a valid subspecies by Hall
(1981: 968).
Procyon maynardi Bangs, 1898g
Proc. Biol. Soc. Washington, 12: 92, 30
April.
= Procyon /otor(Linneaus, 1758). See
Koopman et al. (1957: 164).
Holotype. B7750. Skin and skull. Juvenile male.
Locality. Bahamas: New Providence Island, Nas-
sau. August 1897.
Collector H. L. Claridge.
Condition. Skin complete. Skull partial (broken
from frontals to occiput). Mandible disarticulated.
Type Series. Holotype only.
Comments. Considered a valid species by Wozen-
craft (1993: 336) and Nowak (1999: 698) but un-
doubtedly a recent introduction to New Providence
Island (see Olson and Pregill, 1982: 5).
152 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Family MUSTELIDAE Fischer de
Waldheim, 1817
Genus LONTRA Gray, 1843
Lutra degener Bangs, 1 898d
Proc. Biol. Soc. Washington, 12: 35, 24
March.
= Lontra canadensis canadensis
(Sclireber, 1776). See van Zyll de Long
(1972: 81).
Holotype. B6965. Sldn and skull. Adult male.
Locality. Canada: Newfoundland, Bay St. George.
23 April 1897.
Collector. E. Doane.
Condition. Skin and skull complete.
Type Series. Paratype material consists of B6966,
skin and skull of an adult female, mentioned by
number in the original description, as well as "two
extra skulls," corresponding to B3755 and B3799,
and "a large series of unsexed otter skulls from
Newfoundland," corresponding to MCZ 494—508.
Lutra hudsonica vaga Bangs, 1 898b
Proc. Boston Soc. Nat. Hist., 28: 224, 15
March.
= Lontra canadensis laxatina F. Cuvier,
1823. See van Zyll de Long (1972: 81).
Holotype. B5749. Skin and skull. Adult male.
Locality. (United States), Florida, Brevard Count)-,
Micco. 17 March 1897.
Collector F. R. Hunter.
Condition. Skin and skull complete.
Type Series. 4 paratypes; B4995, skin and skull,
adult female; B4998, sldn and skull, adult male;
B6092, skin and skull, adult male; B6093, sldn and
skull, adult female.
Genus MARTES Pinel, 1792
Mustela atrata Bangs, 1897b
Amer. Nat., 31: 162, 1 February.
= Martes americana atrata (Bangs, 1897).
See G. M. Allen (1942: 166)
Holotype. B5752. Skin and skull. Adult female.
Locality. Canada: Newfoundland, Bay St. George.
29 September 1896.
Collector E. Doane. Original number 2.
Condition. Sldn and skull complete.
Type Series. 12 paratypes; B5751, skin and skull,
adult female; MCZ 492-93, 509-517, unsexed
skulls.
Comments. M. a. atrata was retained as a valid sub-
species by Hall (1981: 983).
Mustela brumalis Bangs, 1898j
Amer. Nat., 32: 502, July
= Martes americana atrata (Bangs, 1897).
See Clark et al. (1987: 1).
Holotype. B7417. Skull. Adult, probably male.
Locality. Canada: Labrador, Okkak ( = Okak). Sum-
mer 1897.
Collector J. D. Sornborger, obtained "from the Es-
kimo."
Condition. Skull complete.
Type Series. 2 paratypes; B7418, skull; B7419,
skull; both probably male.
Genus MUSTELA Linnaeus, 1758
Mustela cicognanii mortigena Bangs, 1913
Bull. Mus. Comp. ZooL, 54: 511, July
= Mustela erminea rictiardsonii Bonaparte,
1838. See Hall (1951: 110).
Holotype. B3745. Skin and skull. Adult male.
Locality. Canada: Newfoundland, Bay St. George.
27 September 1895.
Collector. E. Doane. Original number 1.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Putorius frenatus neomexicanus Barber
and Cockerell, 1898
Proc. Acad. Nat. Sci. Philadelphia, 1898,
p. 188, May
= Mustela frenata neomexicana (Barber
and Gocl<erell, 1898). See Miller (1912:
100).
Holotype. MCZ 10475. Skin with extra tail, and skull.
Adult male.
Locality. (United States): New Mexico, (Dona Ana
County), Mesilla, shore of Armstrongs' Lake, 3,800
ft (1,159 m). 1 Februaiy 1898.
Collector A. C. Tyson. Original number 58.
Condition. Sldn complete, but tail poorly prepared.
The tail belonging to a discarded topotype is tied
to the holotype as an example. Skull complete.
Type Series. A topotype taken at the same time as
the holotype was partially decomposed and dis-
carded; the tail of this specimen is included with
MCZ 10475. The original description also refers to
"a specimen, without any histoiy, in alcohol ... in
the collection of the New Mexico Agricultural Col-
lege."
Comments. M. f. neomexicana was retained as a
valid subspecies by Hall (1981: 995).
Type Specimens of Recent Mammals • Helpen and McFadden 153
Putorius (Arctogale) longicauda oribasus
Bangs, 1899m
Proc. New England Zool. Club, 1: 81, 27
December.
= Mustela frenata oribasus (Bangs, 1899).
See Hall (1936: 105).
Holotype. B9058. Skin and skull. Adult female.
Locality. Canada: British Cokmibia, source of Ket-
tle River, 7,500 ft (2,288 m). 10 September 1898.
Collector. A. C. Brooks. Original number 1368.
Conclition. Skin and skull complete.
Type Series. Holotype only.
Comments. M. f. oribasus was retained as a valid
subspecies by Hall (1981: 998).
Putorius (Lutreola) lutensis Bangs, 1 898b
Proc. Boston Soc. Nat. Hist., 28: 229, 15
March.
= Musteia vison lutensis (Bangs, 1898).
See Hollister (1913: 474).
Holotype. B7225. Skin and skull. Adult male.
Localitij. (United States), Florida, St. Johns Coun-
ty, salt marsh opposite Matanzas Inlet. 16 February
1897.
Collector. O. Bangs. Original number 7.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. M. v. lutensis was retained as a valid
subspecies by Hall (1981: 1003) and Lariviere
(1999: 1).
Putorius (Arctogale) muricus Bangs, 1 899j
Proc. New England Zool. Club, 1: 71, 31
July.
=Mustela erminea muricus (Bangs, 1899).
See Hall (1945: 77).
Holotype. B9146. Skin and skull. Juvenile male.
Locality. (United States): California, El Dorado
County, Echo, 7,500 ft (2,288 m). 15 July 1897.
Collector W. W. Price and E. M. Nutting. Original
number 266.
Conclition. Skin and skull complete. Mandible dis-
articulated.
Type Series. Holotype only.
Comments. M. e. muricus was retained as a valid
subspecies by Hall (1981: 990) and King (1983: 1).
Putorius noveboracensis notius Bangs,
1899i
Proc. New England Zool. Club, 1: 53, 9
June.
= Mustela frenata noveboracensis
(Emmons, 1840). See Hall (1936: 104).
Holotype. B2678. Skin and skull. Juvenile male.
Locality. (United States): North Carolina, Bun-
combe County, Weavei"ville. 10 July 1892.
Collector. J. S. Cairns. Original number 2214.
Condition. Sldn complete. Skull partial (two frag-
ments only, premaxilla— lachrymal). Mandible dis-
articulated.
Type Series. 2 paratypes; AMNH 1247, adult male;
USNM 32239, adult male.
Putorius occisor Bangs, 18991
Proc. New England Zool. Club, 1: 54, 9
June.
= Mustela frenata occisor {Bangs, 1899).
See Hall (1936: 104).
Holotype. B9102. Skin and skull. Adult male.
Locality. (United States): Maine, Hancock County,
Bucksport, near mouth of Penobscot River. 15 Jan-
uary 1899.
Collector A. G. Dorr.
Condition. Skin complete. Skull slightly damaged
(left zygomatic arch broken).
Type Series. There is a series of paratypes in the
MCZ.
Comments. M. f. occisor was retained as a valid
subspecies by Hall (1981: 997).
Putorius rixosus Bangs, 1896a
Proc. Biol. Soc. Washington, 10: 21, 25
Februaiy.
= Mustela nivalis rixosa (Bangs, 1896).
See Reichstein (1958: 169).
Holotype. B642. Skin and skull. Adult female.
Locality. Canada: Saskatchewan, Osier 15 July
1893.
Collector W. C. Colt. Original number 79/181.
Condition. Skin complete. Skull slightly damaged
(left zygomatic arch broken).
Type Series. Three specimens other than the ho-
lotype are mentioned by number in the original
description; MCZ 5532, USNM 4231, probably fe-
male, and USNM 13904, probably male. All are
unsexed skins.
Comments. M. n. rixosa was retained as a valid sub-
species by Hall (1981: 993) and Sheffield and King
(1994: 1).
Putorius vison energumenos Bangs,
1896c
Proc. Boston Soc. Nat. Hist., 27: 5,
March.
= Mustela vison energumenos (Bangs,
1896). See Miller (1912: 101).
Holotype. B3555. Skin and skull. Adult male.
Locality. Canada, British Columbia, Sumas. 23
September 1895.
Collector A. C. Brooks. Original number 514.
Condition. Skin and skull complete.
154 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Type Series. 1 paratype; B3556, skin and skull, ju-
venile male.
Comments. M. v. energumenos was retained as a
valid subspecies by Hall (1981: 1001) and Lariviere
(1999: 1).
Putorius (Lutreola) vulgivagus Bangs,
1895b
Proc. Boston Soc. Nat. Hist., 26: 539, 31
July.
= Mustela vison vulgivaga (Bangs, 1895).
See Miller (1912: 102).
Holotype. B2751. Skin and skull. Adult male.
Locality. (United States): Louisiana, Plaquemines
Parish, Burbridge. 10 Januaiy 1895.
Collector. F. L. Small. Original number 1439/54.
Condition. Skin and skull complete.
Tijpe Series. 10 paratypes; B2752-B2761, all rep-
resented by sldn and skull, 9 males and 1 female.
Comments. M. v. vulgivaga was retained as a valid
subspecies by Hall (1981: 1004) and Lari\'iere
(1999: 1).
Putorius xanthogenys mundus Bangs,
1899i
Proc. New England Zool. Club, 1: 56, 9
June.
= Mustela frenata munda (Bangs, 1899).
See Hall (1936: 107).
Holotype. B5459. Skin and skull. Adult male.
Locality. (United States): California, Marin Coun-
ty, Point Reyes. 19 June 1896.
Collector C. A. Allen. Original number 931.
Condition. Skin complete. Skull slightly damaged
(left zygomatic arch broken).
Type Series. 1 paratype, B8632 (not B8631, men-
tioned erroneously in the original description), skin
and skull, male.
Comments. M. f. munda was retained as a valid
subspecies by Hall (1981: 995).
Family MEPHITIDAE Bonaparte, 1845
Genus MEPHITIS E. Geoffrey and G.
Cuvier, 1795
Mephistis avia Bangs, 1898c
Proc. Biol. Soc. Washington, 12: 32, 24
March.
=- Mephitis mephitis avia Bangs, 1898. See
Hall (1936: 65).
Holotype. B5747. Skin and skull. Adult male.
Locality. (United States): Illinois, Mason County,
San Jose. 10 March 1897.
Collector. H. H. and C. S. Brimley Original num-
ber 2500.
Condition. Skin and skull complete.
Ti/pe Series. 1 paratype; B5783, skin and skull,
adult male.
Comments. M. m. avia was retained as a valid sub-
species by Hall (1981: 1019) and Wade-Smith and
Verts (1982: 1).
Mephitis mephitica elongata Bangs, 1895b
Proc. Boston Soc. Nat. Hist., 26: 531, 31
July.
= Mephitis mephitis elongata Bangs, 1895.
See A. H. Howell (1921: 39).
Holotype. B3051. Skin and skull. Adult male.
Locality. (United States): Florida, Brevard County,
Micco. 5 March 1895.
Collector O. Bangs.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. Retained as a valid subspecies by Hall
(1981: 1019) and Wade-Smith and Verts (1982: 1).
Mephitis mephitica scrutator Bangs, 1 896i
Proc. Biol. Soc. Washington, 10: 141, 28
December.
= Mephitis mephitis mesomelas
Lichtenstein, 1832. See Hall (1936: 66).
Holotype. B2889. Skin and skull. Adult male.
Locality. (United States): Louisiana, Acadia Parish,
Caitville. 25 May 1895.
Collector F. L. Small. Original number 1842.
Condition. Skin and skull complete. Mandible dis-
articulated.
Ti/pe Series. 1 parats-pe, B2886, skin and skull,
adult female.
Mephitis spissigrada Bangs, 1898c
Proc. Biol. Soc. Washington, 12: 31, 24
March.
= Mephitis mephitis spissigrada Bangs,
1898. See Hall (1936: 67).
Holotype. B3699. Skin and skull. Adult female.
Locality. Canada: British Columbia, Sumas. 30
September 1895.
Collector A. C. Brooks. Original number 518.
Condition. Skin and skull complete. Mandible dis-
articulated.
Type Series. 3 paratypes; B3700, skin and skull,
adult female, and B5548, skin and skull, adult male;
B7435, skull, adult male.
Comments. M. m. spissigrada was retained as a val-
id subspecies by Hall (1981: 1022) and Wade-
Smith and Verts (1982: 1).
Type Specimens of Recent Mammals • Hel^en and McFadden 155
Genus SPILOGALE Gray, 1865
Spilogale ambarvalis Bangs, 1 898b
Proc. Boston Soc. Nat. Hist., 28: 222, 15
March.
= Spilogale putorius ambarvalis Bangs,
1898. See Van Gelder (1953: 255).
Holotype. B3481. Skin and skull. Adult male.
Locality. (United States): Florida, Brevard County,
Oak Lodge, east peninsula opposite Mieco. 30 Jan-
uaiy 1895.
Collector. O. Bangs. Original number 11.
Condition. Skin and skull complete.
Type Series. There is a large series of paratypes in
the MCZ.
Comments. S. p. ambarvalis was retained as a valid
subspecies by Hall (1981: 1014) and Kinlaw (1995:
1).
Family VIVERRIDAE Gray, 1821
Genus EUPLERES Doyere, 1835
Eupleres major Layau6en, 1929
Compt. Rend. Acad. Sci. Paris, 189: 198,
22 July 22.
= Eupleres goudotii major Lavauden,
1929. See Albignac (1973: 23).
Syntypes. MCZ 45691: Skin, skull, and postcranial
skeleton. Subadult female, MCZ 45962: Skin, skull,
and postcranial skeleton. Subadult male.
Locality. Madagascar: (Antsiranana), foot of the
Massif Tsaratanna (=Tsaratanana), Upper Sombi-
rano Valley, above village of Beangona, 1,500 m.
April 1929.
Collector Lavauden.
Condition. MCZ 45691: Skin complete, v^dth bald
spot on dorsum and tail slightly damaged. Skull and
skeleton complete. Mandible disarticulated. Teeth
removed from skull but present. MCZ 45962: Skin
complete. Skull and skeleton complete. Mandible
disarticulated. Teeth removed from skull but pres-
ent.
Type Series. 2 syntypes, described above.
Comments. These are the two specimens from G.
Grandidier's personal collection on which Lavau-
den based his original description of Enpleres ma-
jor Albignac (1973: 23) wrote that these type spec-
imens were "introuvable [nowhere to be found]."
Family HERPESTIDAE Bonaparte, 1845
Genus GALIDICTIS I. Geoffrey, 1839
Galidictis grandidiensis [sic] Wozencraft,
1986
J. Mammal. 67: 561, 8 August.
= Galidictis grandidieri Wozencraft, 1 986.
See Wozencraft (1987: 198).
Holotype. MCZ 45983. Skin, skull, and postcranial
skeleton. Adult.
Locality. Madagascar (no further data available).
The locahty of the paratype, stored in the AMNH,
is "Madagascar, Lac Tsimanampetsotsa, 24°08' S,
43°46' E."
Collector No collection data available. The holo-
type is part of the collection of G. Grandidier.
Condition. Skin prepared flat; incomplete (ven-
trum missing). Skull complete.
Type Series. 1 paraty^^e; AMNH 100478, sldn and
skull, adult male.
Comments. G. grandidieri was considered a valid
species by Wozencraft (1993: 300) and Nowak
(1999: 769).
Family FELIDAE Fischer de Waldheim,
1817
Genus LEPTAILURUS Sevenzoy, 1858
Fells capensis phillipsi G. M. Allen, 1914d
Bull. Mus. Comp. Zool., 58: 337, July.
= Leptallurus serval phillipsi (G. M." Allen,
1914). See comments.
Holotype. MCZ 14908. Skin and skeleton. Adult
male.
Locality. Sudan: Blue Nile, El Garef 10 Januaiy
1913.
Collector J. C. PhiUips.
Condition. Skin and skull complete.
Type Series. Holotype only.
Comments. G. M. Allen (1939: 241) used the name
Felis serval phillipsi. The use of the genus Lep-
tailunis follows Wozencraft (1993: 292). L. s. phil-
lipsi was retained as a valid subspecies by Smithers
(1975: 7).
Genus LVA/XKerr, 1792
Lynx (Cervaria) fasciatus oculeus Bangs,
1899e
Proc. New England Zool. Club, 1: 23, 31
March.
= Lynx rufus callfornlcus Mearns, 1897.
See Grinnell and DLxon (1924: 346).
Holotype. B8633. Skin and skull. Adult male.
Locality. (United States): Cahfomia, Marin Coun-
ty, Nicasio. 11 December 1898.
Collector C. A. Allen. Original number 981.
Condition. Skin and skull complete.
Type Series. 1 paratype; B4789, sldn and skull,
adult male.
Lynx gigas Bangs, 1897e
Proc. Biol. Soc. Washington, 11: 50, 16
March.
= Lynx rufus gigas Bangs, 1897. See
Peterson and Downing (1952: 11).
Holotype. B4951. Skin and skull. Adult male.
Locality. Canada: Nova Scotia, 15 miles (24.1 km)
back of Bear River. 11 December 1895.
156 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Collector. D. R. Ritchie.
Condition. Skin and skull complete.
Type Series. Holotype only.
Comments. L. r gigas was retained as a valid sub-
species by Hall (1981: 1054).
Lynx subsolanus Bangs, 1897e
Proc. Biol. Soc. Washington, 11: 49, 16
March.
= Lynx canadensis subsolanus Bangs,
1897. See ElHot (1901: 296).
Holotype. B1190. Sldn and skull. Adult male.
Locality. Canada: Newfoundland, Codroy. 13 June
1894.
Collector E. Doane.
Condition. Skin and skull complete.
Type Series. 2 paratypes; B5754, skin and skull, ju-
venile female; B3798, skull, adult male.
Comments. L. c. subsolanus was retained as a valid
subspecies by Hall (1981: 1051).
Genus PUMA Jardine, 1834
Felis bangs! Merriam, 1901
Proc. Washington Acad. Sci., 3: 595, 11
December.
= Puma concolor bangs! {Memam, 1901).
See coinments.
Holotype. B8413. Skin and skull. Adult male.
Locality. Colombia: (La Guajira), Dibulla. 8 Oc-
tober 1899.
Collector. W. W. Brown, Jr.
Condition. Skin and skull complete.
Type Series. 3 paratypes; B8147, skin and skull,
adult female, and "two skulls from Peru, in the
American Museum of Natural History."
Comments. Nelson and Goldman (1929: 347) used
the name Felis concolor bangsi. The use of the ge-
nus Puma follows Wozencraft (1993: 296). Re-
tained as a valid subspecies by Currier (1983: 1).
Fells bangsi costaricensis Merriam, 1901
Proc. Washington Acad. Sci., 3: 596, 11
December.
= Puma concolor costaricensis (Merriam,
1901). See comments.
Holotype. MCZ 10118. Skin and skull. Adult female.
Locality. Panama: Chiriqui, Boquete ( = Bajo Bo-
quete), 4,000 ft (1,220 m). 22 April 1901.
Collector W. W. Brown, Jr. Original number 337.
Condition. Skin and skull complete.
Type Series. There is a series of paratypes in the
MCZ.
Comments. Nelson and Goldman (1929: 347) used
the name Felis concolor costaricensis. The use of
the genus Puma follows Wozencraft (1993: 296).
Retained as a valid subspecies by Currier (1983: 1).
Felis coryi Bangs, 1899b
Proc. Biol. Soc. Washington, 13: 15, 31
Januaiy.
= Puma concolor coryi {Bangs, 1899). See
comments.
Holotype. B7742. Sldn and skull. Adult male.
Locality. (United States): Florida, Brevard County,
"wildeniess back of Sebastian". 1 Januaiy 1898
Collector F. R. Hunter.
Condition. Skin and skull complete.
Type Series. 5 paratypes; B5489, adult female,
B5650, adult female, B6992, aduh male, B7743,
adult female, B7744, juvenile female; all repre-
sented by sldn and skull.
Comments. Nelson and Goldman (1929: 347) used
the naine Felis concolor coryi; the use of the genus
Puma follows Wozencraft (1993: 296). Felis coryi
Bangs, 1899 is a replacement name for Felis con-
color floridana Cory, 1896 (1896: 109). Retained as
a valid subspecies by Currier (1983: 1).
Felis improcera Phillips, 1912
Proc. Biol. Soc. Washington, 25: 85, 4
May.
= Puma concolor improcera (Phillips,
1912). See comments.
Holotype. MCZ 12704. Skin and skull. Adult male.
Locality. (Mexico): Lower ( = Baja) California, Cal-
malli. 3 September 1911.
Collector. E. W Funcke. Original number 10.
Condition. Skin and skull complete.
Type Series. Holotype only.
Comments. Nelson and Goldman (1929: 347) used
the name Felis concolor improcera. The use of the
genus Puma follows Wozencraft (1993: 296). Re-
tained as a valid subspecies by Currier (1983: 1).
Order CIMOLESTA McKenna, 1975
Family MANIDAE Gray, 1873
Genus PH/Ar/\G/A/aS Rafinesque, 1821
Phataginus tricuspis mabirae G. M. Allen
and Loveridge, 1942
Bull. Mus. Comp. Zool, 89: 178,
Februaiy.
Holotype. MCZ 39417. Skin, skull, and postcranial
skeleton. Adult male.
Locality. Uganda: (Buganda), Chagwe, Mabira
Forest, Mubango. 12 November 1938.
Collector A. Loveridge.
Condition. Skin and skull complete. Postcranial
skeleton partial (includes atlas, right tibia, and right
fibula).
Type Series. Holotype only.
Type Specimens of Recent Mammals • Helgen and McFadden 157
Comments. Retained as a valid subspecies by
Meester (1972: 2).
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INDEX
Type Specimens of Recent Mammals • Hel^en and McFadden 171
abaconis
Geocapromijs, 132
abbotti
Mijotis, 146
abbreviata
Neotonia, 121
aberti
Sciunis, 99
abietorum
Feromyscus, 125
ablutus
Leggada, 118
acadicus
Microtus, 115
Neosorex, 138
Zapus, 112
aequalis
Aedialops, 140
Scalops, 140
Scalopus, 140
aestuans
GueHinguetus, 106
Sciiirus, 106
Aethalops
aequalis, 140
alecto, 140
Aethoglis
argenteus, 129
/jueij, 129
Aethosciurus
byatti, 106
laetus, 106
africanus
Nycticeius, 147
Afrosoricida, 101
AgOMfi
paca, 130
virgatus, 130
Agoutidae, 130
akka
FuniscUinis, 104
Akodon
apricus, 127
teguina, 127
xerampelinus, 127
alacer
Lepus, 134
Sylvilagus, 134
alashanicus
Spermophilus, 107
albicinctus
Myotis, 146
albigula
Thyroptera, 147
albiventer
Nycticeius, 147
Thyroptera, 147
alboniger
Hylopetes, 105
Fteroinys, 105
alecto
Aethalops, 140
alfari
Microsciunis, 105
alleni
Dasymys, 118
Heteromys, 112
Hylomyscus, 118
Lepus, 133
Liomys, 112
Macrotolagus , 133
Melomys, 118
Rhinolophus, 143
rt/ogfl
Blarina, 137
AZoKflrtfl
luctuosa, 135
palliata, 135
pigra, 135
trabeata, 135
alpinus
Sciuroptenis, 104
altilis
Scotophilus, 147
amazonicus
Nectomys, 121
ambarvalis
Spilogale, 155
americana
Martes, 152
americanus
Euarctos, 151
Lept/s, 133, 134
Odocoileus, 148
Ursus, 151
A77iefn(ia
centurio, 143
minor, 143
ammodijtes
Feromyscus, 125
anastasae
Feromyscus, 124, 125
Scalops, 140
Scalopus, 140
anchietae
Otomys, 120
anchises
Hylopetes, 105
Fteromys, 105
anetianus
Fteropus, 141
angolensis
Mops, 145
Antechinus
mayeri, 100
misim, 100
noso, 100
Antillogale
marcanoi, 137
aorensis
Fteropus, 141
Aplodontia, 103
califomica, 103
Columbiana, 103
rainieri, 103
7^(/a, 103
Aplodontidae, 103
Apodemus
euxinus, 117
mystacinus, 117
apricus
Akodon, 127
Scotinomys, 127
aquaticus
Scalopus, 140
aquilis
Cerdocyon, 150
Craseomys, 113
Urocyon, 150
aquilonius
Fiber, 116
Ondatra, 116
araeum
Flagiodonta, 132
araneus
Sorex, 138
arcticus
Hesperomys, 124
Lepus, 133
Rangifer, 149
ArctogaZe
longicauda, 153
muricus, 153
oribasus, 153
Arcto??ii/.s
avanis, 105
flaviventer, 105
igna-uus, 105
arenarius
Feromyscus, 126
argentatus
Feromyscus, 125
argenteus
Aethoglis, 129
Graphiurus, 129
arJeZ
Fteropus, 141
armatus
Spermophilus, 107
A/tiZjeus
femurvillosum, 144
lituratus, 144
palmarum, 144
Aitiodactyla, 145
Arujco/a
breweri, 114
longipilis, 114
riparia, 114
rufidorsum, 114
terraenovae, 114
Arvicolinae, 112
aslaeorientalis
Neomeris, 148
Neophocaena, 148
Atalapha
brachyotis, 146
Suncus, 139
atrata
Martes, 152
Mustela, 152
atrodorsalis
Thomomys, 111
attioateri
Feromyscus, 124
Geogale, 101
Triaenops, 143
auritus
Flecotus, 147
Eothenomys, 113
Microtus, 113
austerulus
Sigmodon, 127
austini
Fteropus, 141
australis
Cryptogale, 101
Echymipera, 101
Reithrodontomys, 127
austrinus
Geomys, 110
Mannota, 105
Arctomys, 105
Cryptotis, 138
Mephitis, 154
bactrianus
Mus, 121
hadlus
Ictidonujs, 108
Spennophilus, 108
hairdii
Lepus, 133, 134
172 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Mus, 98
Peromyscii.s, 98
haliolus
Peromyscus, 126
baluensis,
Callosciunis, 103
bangsi
Fells, 156
Glaucomys, 104
Perognathus, 112
Puma, 156
Sciuroptenis, 104
Viilpes, 151
bangsii
Lepus, 133
barbouri
Otomys, 121
barrerae
Octoniys, 130
Tympanoctomys, 130
Bathyergidae, 129
hella
Neotoma, 121
belliis
Peromyscus, 124
Beluga
concreta, 148
declivis, 148
beothucus
Cants, 149
bicolor
Crocidura, 137
Hipposideros, 142
Oecomys, 122
Blarina
aloga, 137
brevicauda, 137
compacta, 137
borealis
hasiurus, 146
Odocoileus, 148
Peromyscus, 124
Synaptomys, 116
Tamias, 109
Boromys
torrei, 131
borucae
Sigmodon, 127
Bovidae, 149
brachyotis
Atalapha, 146
Lasiurus, 146
brasiliensis
Sylvilagus, 134
brevicauda
Blarina, 137
Zygodontomys, 128
brevicaudata
Microgale, 102
Arvicola, 114
Microtus, 114
brochus
Syntheosciunis, 108
browni
Microsciunis, 105
Sciurus, 105
bninialis
Mai-tes, 152
Mustela, 152
biifo
Leggada, 118
Mus, 118
burrus
Proechimys, 131
byatti
Aethosciurus, 106
Paraxerus, 106
caboti
Rangifer, 149
cacabatus
Peromyscus, 124
Castor, 109
caecutiens
Sorex, 138
cfi/er
Pedetes, 128
californica
Aplodontia, 103
califoiiticus
Dipodomys, 111
Lynx, 165
Scapanus, 140
callida
Dasyprocta, 130
Callosciunis
baluensis, 103
erythraeus, 103
ferrugineus, 103
haemobaphes, 103
medialis, 103
primus, 103
Caluromyidae, 99
Caluromys
cicur, 99
lanatus, 99
canadensis
Castor 109
Lontra, 152
Lynx, 156
Peromyscus, 125
Canidae, 149
canina
Peropteryx, 142
Cfinzs
beothucus, 149
familiaris, 150
/tipus, 149, 150
Cansumys
canus, 117
canus
Cansumys, 117
capensis
Fells, 165
Pedetes, 128
Pipistrellus , 146
Capromyidae, 131
Capromys
nana, 132
pilorides, 131
relictus, 131
capucinus
Cebus, 136
caraccioli
Vampyrodes, 145
Cariacus
osceola, 148
caribou
Rangifer, 149
carissinia
My Otis, 146
Carnivora, 149
carolinensis
Sciurus, 106
carpenteri
Hijlobates, 136
carroruni
Oryzomys, 123
cascadensis
Lepus, 134
castaneoventris
Sciurus, 103
castanonotus
Sciurus, 99
castanotus
Sciurus, 99
casta nops
Cratogeomys, 111
Pappogeomys, 111
Castor
caecator, 109
canadensis, 109
Castoridae, 109
cavator
Macrogeomys, 110
Orthogeomys, 110
cayennensis
Proechimys, 131
Cebidae, 135
Cebus
capucinus, 136
curtus, 136
celatus
Phenacomys, 116
centu rlo
Ametrida, 143
Cerdocyon
aquilus, 150
germanus, 150
thous, 150
Cervaria
fasclatus, 155
oculeus, 155
Cei-vidae, 148
Cetartiodactyla, 97
Chaerophon
leucostigma, 145
Cheiromys
laniger, 135
madagascariensls, 135
c/ierriei
Zygodontomys, 128
Chilonycteris
pamellii, 143
pusillus, 143
torrei, 143
Chionomys
nivalis, 112
chionopaes
Dicrostonyx, 113
chiriquensis
Guerlinguetus, 106
Sclunis, 106
Chiroptera, 140
Chlorotalpa
tropicalis, 101
chrotorrhinus
Microtus, 114
Chrysochloridae, 101
Chnjsochloris
stidilmanni, 101
tropicalis, 101
chrysogaster
Lenimus, 114
cicognonn
Mustela, 152
cicur
Caluromys, 99
Philander, 99
Cimolesta, 156
cinereoargenteus
Urocyon, 150
cJnereHS
Sorgx, 139
Cingulata, 101
Citellus
obscurus, 107
.siccus, 107
Clavlglls
collaris, 129
soleatus, 129
clavlum
Odocoileus, 149
Clethrionomys
gapperi, 113
proteus, 113
collaris
Claviglis, 129
colondylana
Dasyprocta, 130
colonus
Geomys, 110
coloratus
Type Specimens of Recent Mammals • Hclfien and McFadden 173
On/zoini/s, 123
coliuiibiaiia
Aplodoiifia, 103
coliimbicnnis
Geocapromijs, 132
compacta
Blarina, 137
concoJor
Felis, 156
Puma, 156
concreta
Beluga, 148
condylunts
Mops, 145
confucianus
Niviventer, 119
consobrinus
Sciurofamias, 106
cooper;
Synaptovujs, 116
Tainias, 109
cory;
Fe/;.s, 156
Puma, 156
Conjnorhinus
phijllotis, 146
casta ricensis
Felis, 156
Pinna, 156
cowani
Microgale, 102
Craseomijs
aquilus, 113
crassus
Phenacornys, 116
Cratogeomys
castanops. 111
rubellus. 111
crgper
ReUhrodontomys , 127
Cricetinae, 117
Cricetomylnae, 117
cricefulus
Saccostomus, 117
crinitus
Peromyscus, 125
Crocidura
bicolor, 137
fuscomurina, 137
geata, 138
hildegardeae, 138
maurisca, 138
phaios, 138
tephragaster, 137
cnisiiignim
Pecan, 147
Tayassu, 147
Cryptogale
a u straits, 101
Cryptomys
hottentotus, 129
occlusus, 129
ivhijtei, 129
Cryptotis
avia, 138
tJioinasi, 138
cubanus
Geocaproinys, 132
Solenodon, 137
cumbedandius
Geoim/s, 110
c;/ppra
Ochotona, 133
ctirttis
CeZpjf.s, 136
Hipposideros, 142
Damaliscus
pJiiUipsi, 149
pygargus, 149
daphaenodon
Sorex, 139
darlingtoni
Eptesicus, 145
PipistreUus, 145
Dasogale
fontoynonti, 103
Dasyinys
alleni, 118
inconitus, 118
Dasypodidae, 101
Dasyprocta
callida. 130
colombiana, 130
leporina, 130
noblei, 130
nuchalis, 130
punctata, 130
Dasyproctidae, 130
Dasypus
hoplites, 101
novemcinctus, 101
Dasyuiidae, 100
Das)airomoi"phia, 100
Daubentonia
n ladagasca riens is , 135
Daubentoniidae, 135
davidanus
Sciurotamias, 106
decaryi
Microgale, 102
decUvis
Beluga, 148
degener
Lutra, 152
delectonim
Praomys, 119
deletrix
Vidpes, 150
Delphinapterus
leucas, 148
Delphinus
nielas, 148
demidovii
Galago, 135
Dendromurinae, 117
desmarestianus
Heteromys, 111
Oryzomys, 123
diadeina
Propithecus, 135
Dicrostonyx
chionopaes, 113
exsid, 113
groenlandicus, 113
torquatus, 113
Didelphidae, 100
Didelphis
marsupialis, 100
particeps, 100
pigra, 100
virginiana, 100
Didelphimorphia, 99
Diplomys
labilis, 131
Dipodidae, 112
Dipodomys
californicus. 111
ordii. 111
pallidulus. 111
palmeri. 111
Dipodops
ordii. 111
palmeri. 111
dispar
Sorex, 139
distincta
Neotoma, 122
dorcfls
Damaliscus, 149
dorsalis
Tamias, 109
dorsatum
Erethizon, 130
Dremomys
flavior, 104
pemyi, 104
se?iex, 104
droidtardi
Microgale, 102
douglasii
Tamiasciunis, 109
dupreanum
Eidolon, 98
Echimyldae, 131
Echyniipera
australis, 101
kalabu, 100
rufescens, 101
Eidolon
dupreanum, 98
sakalava, 98
eZegans
Spennophilus, 108
elongata
Mephitis, 164
Proct/on, 151
Emballonuridae, 142
energuinenos
Mustela, 153
Putorius, 153
Microtus, 115
Eonycteris
glandifera, 140
spelaea, 140
Eothenomys
aurora, 113
eua, 113
melanogaster, 113
mucronatus, 113
Epimys
zappeyi, 119
Eptesicus
darlingtoni, 145
pliasma, 146
pumilus, 145
erem/cus
Hesperomys, 98
Peromyscus, 98, 126
Erethizon
dorsatum, 130
picinus, 130
Erethizontidae, 130
erigens
Hipposideros, 142
Erioryzomys
monochromos, 128
Mustela, 152, 153
Erophylla
sezekomi, 144
syops, 144
erythraeus
Callosciurus, 103
Euarctos
americanus, 151
sornhorgeri, 151
Eulipotyphla, 97
Eu pie res
goudotii, 155
major, 155
Eutamias
minimus, 109
neglectus, 109
ewxmus
Apod emus, 117
eufl
Eothenomys, 113
Evotomys
proteus, 113
exsputus
Sigmodon, 127
174 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
exsul
Dicrostonyx, 113
extinius
Sciurus, 106
familiaris
Cants, 150
fasciatus
Cervaria, 155
Lynx, 155
fatuus
Synaptomys, 116
Felidae, 155
Fells
bangsi, 156
capensis, 155
concolor, 156
coryi, 156
costaricensis, 156
floridana, 156
improcera, 156
philUpsi, 155
serval, 155
femurvillosu m
Artibeus, 144
ferrugineus
Callosciunis, 103
feroidus
Sigmodon, 128
Fiber
aquilonius, 116
obscunis, 116
rivalicius, 116
zibethicus, 116
fischeri
Haplonycteris, 141
flavicans
Oecomys, 122
Oryzomys, 122
flavidus
Isthniomys, 121
Megadontomys, 121
flavior
Dremomys, 104
flavipes
Tat era, 117
flaviventer
Arctomys, 105
flaviventris
Marmot a, 105
floridana
Felis, 156
Neotoma, 122
floridanus
Geomys, 110
Sylvilagus, 134
fontigenus
Microtus, 115
fontoynonti
Dasogale, 103
frenata
Mustela, 152, 153, 154
frenatus
Putorius, 152
fidvescens
Oligoryzomys, 122
fulviventer
Mannosa, 99
fumatus
Myoniys, 119
Praomys, 119
Fimisciurus
akka, 104
pyrrhopiis, 104
victoriae, 104
furcuhis
Triaenops, 143
furvus
Sigmodon, 128
Urocyon, 150
fuscipes
Neotoma, 98
fuscomurina
Crocidura, 137
fusus
Peromyscus, 126
Galago
demidovii, 135
orimis, 135
Galagoides
orinus, 135
Galagonidae, 135
Galidictis
grandidiensis, 155
grandidieri, 155
Clethrionomys, 113
geata
Crocidura, 138
Myosorex, 138
Geocapromys
abaconis, 132
colutnbianus, 132
cubanus, 132
ingrahami, 132
irrectus, 132
GeogaZe
aurita, 101
orientalis, 101
Geomyidae, 110
Ggoinys
austriniis, 110
colonus, 110
cumberlandius, 110
floridanus, 110
go#, 110
pinetis, 110
fwz.a, 110
Gerbillinae, 117
gerhillus
Leggada, 119
germanus
Cerdocyon, 150
gibbsi
NeurotricJius, 140
giganteiis
Pteropus, 141
gtgas
Lynx, 155
glaber
Heterocephalus, 129
glandifera
Eonycteris, 140
Glaucomys
bangsi, 104
lascivus, 104
makkovikensis, 104
querceti, 105
sabrinus, 104
uoZans, 104, 105
Glossophaga
longirostris, 144
gloveralleni
Procyon, 151
Sorex, 138
goffi
Geomys, 110
gorgonag
Proechimys, 131
gorilla
Gorilla, 136
Troglodytes, 136
Gorilla
gorilla, 136
gossypinus
Hesperomys, 123
Peromyscus, 123, 124,
125
goudotii
Eupleres, 155
Grammonujs
macmillani, 118
granatensis
Sciurus, 106, 107
grandidiensis
Galidictis, 155
grandidieri
Galidictis, 155
grandis
Orthogeonujs, 111
Graphiurus
argenteus, 129
griseus, 129
/lue^i 129
lorraineus, 129
microtis, 129
murinus, 129
schwabi, 129
surdiis, 129
griseus
Graphiunis, 129
griswoldi
Tana, 134
Guerlinmietus
aestuans, 106
chiriquensis, 106
guyannensis
Proechimys, 131
gymnicus
Sciurus, 109
Tamiasciunis, 109
haemobaphes
Callosciunis, 103
Sciurus, 103
Haplonycteris
fischeri, 141
hardyi
Zapus, 112
haiyia
Harpiocephalus, 146
Hai~piocephalus
harpia, 146
nifuhis, 146
harringtoni
TateriUus, 117
helleri
Platyrrhinus, 144
Herpestidae, 155
hesperia
Lonchophylla, 144
Hesperomys
arcticus, 124
eremicus, 98
gossypinus, 123
leucopus, 124
nebrascensis, 124
sonoriensis, 124
Heterocephalus
glaber 129
stygius, 129
Heteromyidae, 111
Heteromys
alleni, 112
desmarestianus. 111
repens. 111
hildegardeae
Crocidura, 138
Hipposideros
bicolor, 142
curtus, 142
erigens, 142
turpis, 143
hirsutiis
Sigmodon, 128
hispid a
Suillomeles, 100
hispidus
Sigmodon, 127, 128
Hominidae, 138
hoplites
Dasypus, 101
hottentotus
Cryptomys, 129
hudsonica
Lutra, 152
Type Specimens of Recent Mammals • Helpen and McFadden 175
luidsonlciis
Sciunis, 109
Tainiasciurus, 109
Zapus, 112
htieti
Aedioglis, 129
Graphiurus, 129
huinidis
Reifli rodoiifoinys, 127
hyacindiits
Neiirotriclius, 140
hijleae
Proechimijs, 131
Hylobates
carpenteri, 136
lai; 136
Hylobatidae, 136
Hijlomyscus
alleni, 118
simus, 118
Hijlopetes
alboniger, 105
anchises, 105
oriniis, 105
phaijrei, 105
hypoxanthus
Oenomys, 119
Hypudaeus
nivicola, 112
Ictidomys
badius, 108
trideceinllneatus, 108
Idionycteris
phyllotis, 146
ignaua
Marmot a, 105
jgnfluxs
Arctomys, 105
igni/er
Rhinolophus, 143
illectiis
Oecotm/s, 122
Oitjzomys, 122
impiger
Reldirodontoinys, 127
improcera
Fells, 156
Puma, 156
incitatus
Lepus, 134
Sylvilagus, 134
Tapeti, 134
Dasymys, 118
Indridae, 135
ingrahami
Geocapromys, 132
innultus
Mictomys, 116
Synaptomys, 116
insulanus
Peromyscus, 125
insularis
Pteropus, 98
Geocapromys, 132
irroratus
Liomys, 112
Isdimomys
flavidus, 121
kalabit
Echymipera, 100
T«f<?ra, 117
kivtiensis
Thamnomys, 120
kodiacensis
Spennopliilus, 99
Microtus, 115
Sorex, 138
labilis
Diplomys, 131
Loncheres, 131
lacustris
Otomxjs, 120
ladas
Zapus, 112
laetus
Aethosciunis, 106
Lagomoi'plia, 133
lambertoni
Nesomys, 120
lanatus
Cahiromys, 99
laniger
Cheiromys, 135
Pteropus, 98
lanigera
Pteropus, 98
Hylobates, 136
lascivus
Glaucomys, 104
Sciuroptenis, 104
Lasiurus
borealis, 146
brachyotis, 146
latimanus
Scapanus, 140
laxatina
Lontra, 142
lecontii
Reithrodontomys, 127
Leggada
ablutus, 118
fou/o, 118
gerhtllus, 119
Lemr7ius
chrysogaster, 114
paulus, 114
sibiricus, 114
lepida
Neotoma, 121
Leporidae, 143
leporina
Dasyprocta, 130
Leptailunis
phillipsi, 155
serval, 155
LepM5
alacer, 134
alleni, 133
americanus, 133, 134
arcticus, 133
Z?air<ii, 133, 134
bangsii, 133
cascadensis, 134
incitatus, 134
palitans, 133
paludicola, 134
stnithopus, 133
sylvaticus, 134
transltionalis, 134
Delphinaptenis, 148
leucogaster
Scotophilus, 147
leucopus
Hesperomys, 124
Peromyscus, 126
leucostigma
Chaerophon, 145
Mops, 145
levipes
Melomys, 118
hiomys
alleni, 112
irroratus, 112
Lipotyphla, 136
lituratus
Artibeus, 144
Loncheres
labilis, 131
Lonchophylla
hesperia, 144
longicauda
Arctogale, 153
Putorius, 153
longicaudata
Microgale, 102
longimembis
Perognathus, 112
longipilis
Arvicola, 114
longirostris
Glossophaga, 144
Lonfra
canadensis, 152
laxatina, 152
Sciurus, 109
Tamiasciurus, 109
lorraineus
Graphiurus, 129
Zotor
Procyon, 151
louisianae
Odocoileus, 148
lucifugus
My Otis, 146
luctuosa
Alouatta, 135
ludovicianus
Sciurus, 106
lupus
Canis, 149, 150
lutensis
Lutreola, 153
Mustela, 153
Putorius, 153
degener, 151
hudsonica, 151
uago, 151
Lutreola
lutensis, 153
vulgivagus, 154
Lynx
calif ornicus, 155
canadensis, 155, 156
fasciatus, 155
gigas, 155
oculeus, 155
nifus, 155
subsolanus, 156
mabirae
Phataglnus, 156
macmillani
Grammomys, 118
Macrogeomys
cavator, 110
pansa, 110
macropygmaeus
Sorex, 138
macrotis
Nycteris, 142
Peropteryx, 142
Macrotolagus
alleni, 133
palitans, 133
macrou ra
Odocoileus, 148
macnirus
Sorex, 139
madagascariensis
Cheiromys, 135
Daubentonia, 135
Nycteris, 142
Rousettus, 141
major
Eupleres, 155
Vampyrodes, 145
makkovikensis
176 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Glaucomys, 104
Sciuroptenis, 104
maniculatus
Peromyscus, 98, 124,
125
Manidae, 156
marcanoi
Antillogale, 137
Solenodon, 137
Marmosa
fulviventer, 99
mitts, 99
robinsoni, 99, 100
Marmosidae, 99
Marmota
avara, 105
flaviventris, 105
ignava, 105
monax, 105
inarsupialis
Didelphis, 100
Martes
americana, 152
atrata, 152
bnimalis, 152
maurisca
Crocidura, 138
mayeri
Antechinus, 100
rnaynardi
Procyon, 151
meamsz
Saccostomus, 117
niedialis
Callosciunis, 103
niedioximii.s
Mictomys, 116
Synaptomys, 116
Megadontomys
flavidus, 121
megaphyllus
Rhinolophus, 143
melanogaster
Eothenomys, 113
melanops
Taterillus, 117
melanotis
Oryzomys, 123
melanotus
Praomys, 119
meZas
Delphinus, 148
Myoictis, 100
Melornys
alleni, 118
levipes, 118
mollis, 118
moncktoni, 118
ruhex, 118
stevensi, 118
mephitica
Mephitis, 154
Mephitidae, 154
mephitis
Mephitis, 154
Mephitis
avia, 154
elongata, 154
mephitica, 154
mephitis, 154
mesomelas, 154
scrutator, 154
spissigrada, 154
Mesocap romys
nanus, 132
mesomelas
Mephitis, 154
Neotoma, 122
mexicanus
Peromyscus, 124
Microgale
brevicaudata, 102
cowani, 102
decanji, 102
droiihardi, 102
longicaiidata, 102
parvula, 102
principtda, 102
prolixicaudata, 102
pu/Zfl, 102
Microsciurus
alfan, 105
browni, 105
microtis
Graphiuriis, 129
Microtus
acadtcus, 115
aurora, 113
breweri, 114
chrotorrhinus, 1 14
enixus, 115
fontigenus, 115
koreni, 115
mucronatus, 113
oeconomus, 115
pennsylvanicus, 114,
115
provectus, 115
ravus, 114
shattucki, 115
terraenovae, 114
Nesophontes, 136
Mictomys
innuitus, 116
medioximus, 116
minimus
Tamias, 99, 109
Ametrida, 143
Suncus, 139
minusculus
Scapanus, 140
mjsczx
Sorex, 139
misim
Antechinus, 100
mitis
Marmosa, 99
mollipilosus
Tamiasciunis, 109
mollis
Melonujs, 118
Stenomys, 120
Molossidae, 145
monax
Marmota, 105
moncktoni
Melomys, 118
monochromos
Erioryzomys, 128
Oryzomys, 128
Thomasomys, 128
Monodontidae, 148
monoensis
Pteropus, 141
Mops
angolensis, 145
condylunis, 145
leucostigma, 145
orientis, 145
Mormoopidae, 143
mortigena
Mustela, 152
morulus
Sciurus, 107
mucronatus
Microtus, 113
munda
Mustela, 154
mundus
Putorius, 154
muricola
Myotis, 146
initnctis
Arctogale, 153
Mustela, 153
Putorius, 153
Muridae, 112
Murinae, 117
murinus
Graphiuriis, 129
Pseudohydromys, 120
Mms
bactrianus, 119
bairdii, 98
Z;r//o, 118
niusculus, 119
tantillus, 119
tenellus, 119
musculiis
Mus, 119
Mustela
atrata, 152
bnimalis, 152
cicognanii, 152
energiimenos, 153
erminea, 152, 153
frenata, 152, 153, 154
lutensis, 153
mortigena, 152
munda, 154
muricus, 153
neomexicana, 152
nivalis, 153
noveboracensis, 153
occisor, 153
oribasus, 153
richardsonii, 152
rixosa, 153
vison, 153, 154
vulgivaga, 154
Mustelidae, 152
Myoictis
nielas, 100
wallacei, 100
wavicus, 100
Myomys
funmtus, 119
Myopus
schisticolor, 115
thayeri, 115
Mi/osorex
geata, 138
Myofjs
abbotti, 146
albicinctus, 146
carissima, 146
lucifugus, 146
muricola, 146
nugax, 146
sodalis, 146
Myoxidae, 129
mystacinus
Apodemus, 117
nana
Capromys, 132
Nycteris, 142
nanus
Mesocapromys, 132
nrtso
Antechinus, 100
Rhynchiscus, 142
natator
Oryzomi/s. 123
naiyws
Oryzomi/s, 122
nebrascensts
Hesperomys, 124
Peromyscus, 124
Nectomys
aniazonicus, 121
squamipes, 121
Type Specimens of Recent Mammals • Helgen and McFadden 111
neglectus
Tainias, 109
Neomeris
asiaeorientalis, 148
neomexicana
Musfelfi, 152
neoinexica)}its
Putoriiis, 152
Neophocaena
asiaorientalis, 148
phocaenoides, 148
Neosoi-ex
acadicus, 138
palttstris, 138
Neotonia
abhreviata, 121
beUa, 121
distincta, 122
floridana, 122
fuscipes, 98
lepida, 121
mexicana, 122
riibida, 122
nesaeiis
Scitinis, 107
Nesomylnae, 120
Nesomys
lambertoni, 120
Nesophontes
micriis, 136
Nesophontidae, 136
Neil rot richits
gibbsi, 140
Jiyaciiidihiiis, 140
Sciiiriis, 106
nigrictihis
Peromysciis, 125
StenoDiys, 120
nitellinits
Nyctoinys, 122
nivalis
Chionomys, 112
Mustela, 153
nivicola
Hypudaeiis, 112
Niviventer
confuciainis, 119
noblei
Dasyprocta, 130
Pipistrelliis, 147
Putorius, 153
Scabrifer, 147
noveboracensis
Mustela, 153
Putorius, 153
novemcinctus
Dasypus, 101
nuchalis
Dasyprocta, 130
nugax
My Otis, 146
Steatomys, 117
Nycteridae, 142
Nycteris
macrotis, 142
madagascariensis, 142
nana, 142
revoili, 97
tristis, 142
Nycticeiiis
africanus, 147
albiventer, 147
schlieffeni, 147
Nyctomys
nitellinus, 122
sumichrasti, 122
ohsciinis
Citellus, 107
FAer, 115
Ondatra, 115
obsoletus
Spennophilus, 99
occidentalis
Paramicrogale, 102
occisor
Mustela, 153
Putorius, 153
occlusus
Cryptonu/s, 129
Ochotona
cupppes, 133
princeps, 133
saxatilis, 133
Ochotonidae, 133
ochraceiis
Thamnomys, 118
Octodontidae, 130
Octomys
barrerae, 130
Cervaria, 155
Lynx, 155
ocythoiis
Urocyon, 150
Odocoileus
americanus, 148
borealis, 148
clavium, 149
louisianae, 146
macroura, 146
osceola, 146
virginianus, 146, 147
bicolor, 122
flavicans, 122
illectus, 122
trabeatus, 122
oecono»?i;<s
Microtus, 115
Oenomys
hypoxanthus, 119
talangae, 119
Oligoryzornys
fulvescens, 122
vegetus, 123
Ondatra
ac/uilonius, 116
obscunis, 116
rivalicius, 116
zibethictis, 116
orariiis
Sciunis, 109
Zapus, 112
ordii
Dipodomys, 111
Dipodops, 111
oreas
Peromyscus, 126
oribasus
Arctogale, 153
Mustela, 153
Putorius, 153
orientalis
Geogale, 101
orientis
Mops, 145
onn«.S'
Gala go, 135
Galagoides, 135
Hylopetes, 105
Pteromys, 105
Orthogeoniys
cavator, 110
grandis. 111
pansa, 110
p/((fo. 111
Oryzoinys
carrorum, 123
coloratus, 123
devius, 123
flavicans, 122
fulvescens, 122
illectus, 122
nielanotis, 123
monochronios, 128
natator, 123
navus, 122
palustris, 123
rostratus, 123
vegetus, 123
osceoZa
Cariacus, 148
Odocoileus, 146
Otomops
papuensis, 145
Otomyinae, 120
anchietae, 120
barboiiri, 121
lacustris, 120
typus, 121
uzungwensis, 121
Praoinys, 119
Agouti, 130
Pachycyon
robustus, 150
paitana
Tupaia, 134
palitans
Lepus, 133
Macrotolagus, 133
palliata
Alouatta, 135
pallidulus
Dipodomys, 111
pallidus
Spennophilus, 99
Tainias, 99
pahnarius
Peromyscus, 125
pabnarum
Artibeus, 144
pahneri
Dipodomys, 111
Dipodops, 111
paludicola
Lepus, 134
St/lvilagus, 134
palustris
Neosorex, 138
Onjzomys, 123
Sorex, 138
Sylvilagus, 134
pansa
Macrogeomys, 1 10
Ortliogeonujs, 110
Pappogeomys
castanops. 111
ntbellus. 111
papuensis
Otomops, 145
Paracaproniys, 132
Paramicrogale
occidentalis, 102
Paraxenis
byatti, 106
vexillarius, 106
paniellii
Chilonycteris, 143
Pteronotus, 143
parryi
Spennophilus, 99
particeps
Didelphis, 100
parvula
Microgale, 102
paulus
Leninuis, 114
178 Bulletin Museum of Comparative Zoology, Vol. 157, No. 2
Pecari
ci-usnigrum, 147
tajacu, 147, 148
torvus, 148
Pedetes
cafer, 128
capensis, 128
taborae, 128
Pedetidae, 128
pennsylvanica
Vulpes, 150, 151
pennsylvanicus
Microtus, 114, 115
Peramelia, 100
pernyi
Dremomys, 104
Perognathus
hangsi, 112
longiinembris, 112
Peromyscus
abietoruin, 125
ammodytes, 125
anastasae, 124, 125
arenarius, 126
argentatus, 125
attwateri, 124
bairdii, 98
baliolus, 126
foeZZtis, 124
borealis, 124
cacabatiis, 124
canadensis, 125
crinitus, 125
eremicus, 98, 126
fusus, 126
gossypinus, 123, 124,
125
insidanus, 125
leucopus, 126
maniculatus, 98, 124,
125
mexicaniis, 124
nebrascensis, 124
nigriculus, 125
oreas, 126
pahnarius, 125
phasma, 126
polionotus, 126
rhoadsi, 126
saturatus, 126
scitulus, 125
subgrisetis, 126
texanus, 126
Peropteryx
canina, 142
macrotis, 142
phaea, 142
Peroiyctidae, 100
perneri
Propithecus, 135
personatus
Sorex, 139
phaea
Peropteryx, 142
phaios
Crocidura, 138
phasma
Eptesicus, 147
Peromyscus, 126
Pipistrellus, 147
Phataginus
niabirae, 156
tricuspis, 156
phayrei
Hylopetes, 105
Pterornys, 105
Phenacomys
celatus, 116
crassus, 116
ungava, 116
Philander
ciciir, 99
philippinensis
Rhinolophus, 143
phillipsi
Damaliscus, 149
Fe/is, 155
Leptatlunis, 155
phocaenoides
Neophocaena, 148
Phocoenidae, 148
Phyllostomidae, 143
phyllotis
Corynorhinus, 146
Idionycteris, 146
picnii/s
Erethizon, 130
pigra
Alouatta, 135
Didelphis, 100
piloroides
Capromys, 131
pinetis
Geomys, 110
Pipistrellus
capensis, 147
notius, 147
phasma, 147
rendalli, 147
Plagiodonta
araeum, 132
Platyrrhinus
helleri, 144
umbratus, 144
Plecotus
aiiritus, 147
sacrimontis, 147
pluto
Orthogeomys, 111
poeyanus
Solenodon, 137
polionotus
Peromyscus, 126
Praomys
delectorum, 119
fumatus, 119
melanotus, 119
oweni, 119
tuHbergi, 119
pratensis
Steatomys, 117
Primates, 135
primus
Callosciimts, 103
pnuceps
Ochotona, 133
principula
Micro gale, 102
prisons
Rhynchiscus, 142
Procyon
elucus, 151
gloveralleni, 151
Zofor, 151
maynardi, 151
Procyonidae, 151
Proechimys
burnis, 131
cayennensis, 131
gorgonae, 131
guyannensis, 131
hyleae, 131
semispinosus, 131
p rolixicaudata
Microgale, 102
Propithecus
diadema, 135
perrieri, 135
proteus
Clethrionomys, 113
Evotomys, 113
provectus
Microtus, 115
Pseudohydromys
murinus, 120
Pferojnys
alboniger, 105
anchises, 105
orinus, 105
phayrei, 105
Pteronotus
pamellii, 143
pusillus, 143
quadridens, 143
Pteropodidae, 140
Pteropus
anetianus, 141
aorensis, 141
rtrie/, 141
austini, 141
giganteus, 141
insidaris, 98
laniger, 98
lanigera, 98
monoensis, 141
rayneri, 141
woodfordi, 141
pulla
Microgale, 102
Pu??ifl
hangsi, 156
concolor, 156
coryi, 156
costaricensis, 156
improcera, 156
punctata
Dasyprocta, 130
pusillus
Chilonycteris, 143
Pteronotus, 143
putorius
Spilogale, 155
Putorius
energumenos, 153
frenatus, 152
longicauda, 153
lutensis, 153
mundus, 154
muricus, 153
neomexicanus, 153
notius, 153
noveboracensis, 153
occisor, 153
oribasus, 153
rixosus, 153
vison, 153
vidgivagus, 154
xanthogenys, 154
pj/gargMS
Damaliscus, 149
pyrrhopus
Funisciunis, 104
quadridens
Pteronotus, 143
qiiadrivittatus
Tamias, 99, 109
querceti
Glaucomys, 105
Sciuropterus, 105
ramieri
Aplodontia, 103
Rangifer
arcticus, 149
caboti, 149
caribou, 149
tarandus, 149
terraenovae, 149
rat>u.s
Microtus, 114
rayneri
Pteropus, 141
Reithrodontomys
australis, 127
creper, 127
Type Specimens of Recent Mammals • Helaen and McFadden 179
hinutills, 127
itnpiger, 127
lecontii, 127
sumichrasti, 127
viilcaniiis, 127
relicttis
Capromys, 131
rendalli
Pipistrelhis, 147
repens
Heteroimjs, 111
Nijcteris, 97
Rhinolophidae, 142
Rliinolopliiis
alleni, 143
ig;nifer, 143
megaphyllus, 143
philippinensis, 143
rhoadsi
Peroinijscus, 126
Rhynchiscus
naso, 142
priscus, 142
RJn/nchonycteris
naso, 142
richardsonii
Mustela, 152
riparia
Arvicola, 114
rivalicius
Fiber, 116
Ondatra, 116
rixosrt
Mustela, 153
nxosrts
Piitorius, 153
robinsoni
Mannosa, 99, 100
roboratus
Sorex, 139
robustiis
PacJu/cyon, 150
Rodentia," 103
ro stratus
Oryzomys, 123
Rousettus
madagasca riens is , 141
nibellus
Cratogeomys, 111
Pappogeoniys, 111
nibex
Melomys, 118
rubida
Neotoma, 122
rubricatus
Vulpes, 151
nibricosa
Vulpes, 150, 151
Aplodontia, 103
nifescens
Echymipera, 101
rufidorsunx
Arvicola, 114
rufulus
Haiyioceplialus, 145
rufus
Lynx. 155
sabrinus
Glaucoinys, 104
Sciuroptenis, 104
Saccostomus
cricetulus, 117
meamsi, 117
sacrimontis
Plecotus, 147
sakalava
Eidolon, 98
saltuensis
Sciunis, 107
sanctaeniartae
Sigmodon, 128
sanguinidens
Sorex, 139
saturatus
Peroniyscus, 126
saxatilis
Ochotona, 133
Scabrifer
notius, 147
Scalops
aereus, 140
anastasae, 140
texanus, 140
Scalopus
aereus, 140
anastasae, 140
aquaticus, 140
Scandentia, 134
Scapanus
califomicus, 140
latirnanus, 140
minusculus, 140
schisticolor
Myopus, 115
schliejfeni
Nycticeius, 147
Schwab i
Graphiurus, 129
scitulus
Peroniyscus, 125
Sciuridae, 103
Sciuroptenis
alpinus, 104
bangsi, 104
lascivus, 104
niakkovikensis, 104
querceti, 105
sabrinus, 104
.siZiis, 104
ooZons, 104, 105
Sciurotaniias
consobrinus, 106
davidianus, 106
thayeri, 106
Scit<n<s
aberti, 99
aestuans, 106
browni, 105
carolinensis, 106
castaneoventris, 103
castanonotus, 99
castanotus, 99
chiriquensis, 106
extimus, 106
granatensis, 106, 107
gymnicus, 109
haemobaphes, 103
hudsonicus, 109
loquax, 109
ludovicianus, 106
mondus, 107
nesaeus, 107
niger, 106
orarius, 109
saltuensis, 107
variabilis, 107
vicinus, 106
vulpinus, 106
ijucatanensis, 106
Scotinomys
apricus, 127
teguina, 127
xeranipelinus, 127
Scotophilus
altilis, 147
leucogaster, 147
scrutator
Mephitis, 154
seniispinosus
Proechimys, 131
senex
Drenionujs, 104
Zygodontomys, 128
serval
Felis, 155
Leptailunis, 155
Setifer
setosus, 103
setosus
Setifer, 103
Erophylla, 144
shattucki
Microtus, 115
sibiricus
Lemrnus, 114
siccus
Citellus, 107
Sigmodon
austendus, 127
borucae, 127
exsputus, 127
fervidus, 128
furvus, 128
hirsutus, 128
hispidus, 127, 128
sanctaeniartae, 128
spadicipygus, 128
Sigmodoiitinae, 121
siZus
Sciuroptenis, 104
SiHUiS
Hylomyscus, 118
sodalis
Myotis, 146
soleatus
Claviglis, 129
Solenodon
cubanus, 137
marcanoi, 137
poeyanus, 137
Solenodontidae, 137
sonoriensis
Hesperomys, 124
Sorex
araneus, 138
caecutiens, 138
cinereus, 139
daphaenodon, 139
dispar, 139
gloveralleni, 138
koreni, 138
inacropygniaeus, 138
niaci~urus, 139
miscix, 139
palustris, 138
personatus, 139
roboratus, 139
sanguinidens, 139
tundrensis, 138
ultinuis, 138
oir, 139
Soricidae, 137
sornborgeri
Euarctos, 151
L/rs((s, 151
soro/-
Trtfera, 117
spadicipygus
Sigmodon, 128
spelaea
Eonycteris, 140
Spennophilus
alashanicus, 107
arniatus, 107
badius, 108
elegans, 108
kodiacensis, 99
obsoletus, 99
pallidus, 99
parryi, 99
180 Bulletin Museum of Comparative Zoology, Vol. 157, No.
spilosoma, 99
texensis, 108
tridecemlineatus, 99,
108
Spilogale
ainbarvalis, 155
putorius, 155
spissigrada
Mephitis, 154
squarnipes
Nectomys, 121
Steatomys
nyasae, 117
pratensis, 117
Stenoinys
mollis, 120
niobe, 120
stevensi, 120
verecundus, 120
Melomys, 118
Stenoinijs, 120
striatus
Tamias, 107
stnithopus
Lepiis, 133
stuhhnanni
Chnjsochloris, 101
stygius
Heterocephalus, 129
subgriseus
Peromyscus, 126
suhsolanus
Lynx, 156
Suillomeles
hispida, 100
sumichrasti
Nyctomys, 122
Reithrodontomys, 127
Suncus
ater, 139
minor, 139
varilla, 139
Graphiunis, 129
sylvaticus
Lepus, 134
Sylvilagus
alacer, 134
brasiliensis, 134
floridanus, 134
incitatus, 134
paludicola, 134
palustris, 134
transitionalis , 134
Synaptomys
boreal is, 116
cooperi, 116
fatuus, 116
innuitus, 116
medioxitnus, 116
Syntheosciwins
broclius, 108
syops
Erophylla, 144
taborae
Pedetes, 128
tajacu
Pecari, 147, 148
talangae
Oenomys, 119
Talpidae, 140
Tamias
borealis, 109
cooperi, 108
dorsalis, 108
minim.us, 99, 109
neglectus, 109
pallidus, 99
quadrivittatus, 99
striatus, 109
townsendii, 108
venustus, 109
Tamiasciunis
douglasii, 109
gym,nicus, 109
hudsonicus, 109
loquax, 109
mollipilosus, 109
Tanfl
grisiooldi, 134
tona, 134
fana
Tana, 134
Tupaia, 134
tantilliis
Mas, 119
incitatus, 134
ta rand us
Rangife7', 149
Tafera
flavipes, 117
kempi, 117
soror, 117
valida, 117
Taterillus
liarringtoni. 117
melanops, 117
Tayassu
crusnignim, 147
torvus, 148
Tayassuidae, 147
teguina
Akodon, 127
Scotinomys, 127
tenellus
Mus, 119
Tenrecidae, 101
tephragaster
Crocidura, 137
terraenovae
A)~vicola, 114
Microtus, 114
Rangifer, 149
texanus
Peromyscus, 126
Scalops, 140
fexe'n.sz.s
Spernwpliilus, 108
Thamnomys
kivuensis, 120
ochraceus, 118
venustus, 120
thayeri
My opus, 115
Sciurotamias, 106
thoniasi
Crypt Otis, 138
Thomasomys
nionochromos, 128
Thoniomys
atrodorsalis. 111
unibrinus. 111
Cerdocyon, 150
Thyroptera
albigula, 147
alhiventer, 147
tricolor, 147
Thyropteridae, 147
torquatus
Dicrostonyx, 113
torrei
Boromys, 131
Chilonycteris, 143
toruw5
Pecari, 148
Tayassu, 148
townsendii
Tamias, 109
trabeatiis
Alouatta, 135
Oecomys, 122
transitionalis
Lepus, 134
Sylvilagus, 134
Triaenops
aurita, 143
furculus, 143
fricoZor
Thyroptera, 147
tricuspis
Phataginus, 156
tridecemlineatus
Spennophilus, 99, 108
trinotatus
Zapus, 112
Nycteris, 142
Troglodi/tes
gorilla, 136
tropicalis
Chlorotalpa, 101
Chnjsochloris, 101
tullbergi
Praomys, 119
tundrensis
Sorex, 138
Tupaia
paitana, 134
tona, 134
Tupaiidae, 134
turpis
Hipposideros, 143
Geo my s, 110
Tympanoctomys
barrerae, 130
typus
Otomys, 121
ultimus
Sorex, 138
umbratus
Platyrrhinus, 144
Vampyrops, 144
umbrinus
Thomonu/s, 111
ungava
Phenacomys, 116
Urocyon
aquilus, 150
cinereoargenteus, 150
furvus, 150
ocythous, 150
Ursidae, 151
americanus, 151
somborgeri, 151
uzungwensis
Otomys, 121
vafra
Vulpes, 150, 151
uaga
L«fra, 152
valida
Tatera, 117
Vampijrodes
caraccioli, 145
major, 145
Vampyrops
umbratus, 144
zarhinus, 144
variabilis
Sciurus, 107
varilla
Suncus, 139
uege"f!/.s
Oligoryzonujs, 123
Oryzomys, 123
venustus
Tamias, 109
Thamnomys, 120
verecundus
Type Specimens of Recent Mammals • Helpen and McFadden
181
Stenoinys, 120
Vespertilionidae, 145
vexillariiis
Paraxerus, 106
vicinus
Sciunis, 106
victoriae
Fitnisciiinis, 104
vir
Sorex, 139
virgatus
Agouti, 130
virginiana
Didelphis, 100
virginianus
Odocoileus, 148, 149
vison
Mnstela, 153, 154
Putoriiis, 153
Viverridae, 155
volans
Glaucomys, 104, 105
Sciuroptenis, 104, 105
vidcanitis
Reithrodoutomi/s, 127
vidgivaga
Mustela, 154
vtdgivagus
Lutreola, 154
Putorius, 154
vidpes
Vtdpes, 150, 151
Vidpes
bnngsi, 151
deletrix, 150
pennsylvanica, 150, 151
ruhricatus, 151
nibricosa, 150, 151
ufl/ra, 150, 151
vulpes, 150, 151
vulpinus
Sciun.is, 106
ivallacei
Myoictis, 100
wavicus
Myoictis, 100
whijtei
Cryptomys, 129
woodfordi
Pteropus, 141
xanthogenys
Putorius, 154
xerampelinus
Akodon, 127
Scotinoniys, 127
yucatanensis
Sciurus, 106
zappeyi
Epirnys, 119
Zapus
acadicus, 112
luirdyi, 112
hudsonicus, 112
ladas, 112
orarius, 112
trinotatus, 112
zarhinus
Vampyrops, 144
zibethicus
Fiber, 116
Ondatra, 116
Zygodontomys
brevicauda, 128
cheniei, 128
seorsus, 128
(US ISSN 0027-4100)
OF THE
seum
A New Species of Lizard Related to
Stenocercus caducus (Cope) (Squamata:
Iguanidae) From Peru and Bolivia, With a
Key to the "Ophryoessoides Group"
JOHN E. CADLE
IJBRARY
OCT 2 3 2002
HARVARD
HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSEI IS, U.S.A.
VOLUME 157, NUMBERS
14 November 2001
(US ISSN 0027-4100)
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SPECIAL PUBLICATIONS.
1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and
Evolution of Crustacea. 192 pp.
2. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere-
dinidea (Mollusca: Bivalvia). 265 pp.
3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino-
derms. 284 pp.
4. Eaton, R. J., 1974. A Flora of Concord from Thoreau's Time to the
Present Day. 236 pp.
5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology
and Evolutionary Biology: Essays in Honor of Ernest E. Williams.
725 pp.
6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp.
Other Publications.
Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine.
Reprinted 1964.
Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of
Insects. (Bulletin of the M. C. Z, Vol. 108.) Reprinted 1971.
Creighton, W. S., 1950. The Ants of North America. Reprinted 1966.
Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In-
ternational Symposium on Natural Mammalian Hibernation. (Bulletin
of the M. C. Z, VoL 124.)
Orinthological Gazetteers of the Neotropics (1975-).
Peter's Check-list of Birds of the World, vols. 1-16.
Proceedings of the New England Zoological Club 1899-1947. (Complete
sets only.)
Proceedings of the Boston Society of Natural History.
Price list and catalog of MCZ publications may be obtained from Publica-
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This publication has been printed on acid-free pennanent paper stock.
©The President and Fellows of Harvard College 2001.
A NEW SPECIES OF LIZARD RELATED TO STENOCERCUS
CADUCUS (COPE) (SQUAMATA: IGUANIDAE) FROM PERU AND
BOLIVIA, WITH A KEY TO THE "OPHRYOESSOIDES GROUP "
JOHN E. CADLE1
CONTENTS
Abstract -- 183
Resumen 183
Introduction 184
Materials and Methods 184
Description of a New Species of Stenocersus 187
Stenocercus prionotus new species _ 187
Diagnosis and Comparisons 191
Description „ 193
Distribution Patterns in Stenocercus prionotus
197
Patterns of Sympatry and Geographic Variation
in Stenocercus prionotus, and the Need for
Additional Fieldwork 200
Natural Histoiy of Stenocercus prionotus 205
Comparison of Stenocercus prionotus with S.
caducus 206
Distributions of Stenocercus prionotus and S.
caducus in Eastern Bolivia 210
Is the Distribution of Stenocercus aculeatus
disjunct? 212
Key to Species of the " Ophryoessoides group"
of Stenocercus 212
Acknowledgments 215
Appendix: Specimens Examined 216
Literature Cited 218
Abstract. A new species of iguanid lizard, Steno-
cercus prionotus, is described from eastern Peiii and
Bolivia (known range froin San Martin Department,
Peru, to northern La Paz and El Beni departments,
Bolivia). Most localities are in the Andean foothills
and immediately adjacent lowlands. Stenocercus
prionotus is similar to several other species of Sten-
ocercus with large posterior head scales, an enlarged
row of supraoculars, and keeled ventral scales. These
similar species are referred to as the "Ophryoessoides
group" without implying that it is a monophyletic as-
semblage. Based on their common possession of a
' Department of Herpetology, Chicago Zoological
Society, 3300 Golf Road, Brookfield, Illinois 60513.
Research Associate, Department of Herpetology,
Museum of Comparative Zoology.
unique scaly flap concealing a portion of the posthu-
meral mite pocket, the new species is apparently
closely related to S. caducus (Cope), which is known
from central Bolivia south to northern Argentina and
Paraguay. Stenocercus prionotus is distinguished from
S. caducus by having a more prominent vertebral
crest and a pattern of alternating light and dark bars
on the throat (rarely obseived in S. caducus, which
usually has light throat spots). These two species also
occupy different physiographic regions (western Am-
azonian rainforest for S. prionotus; chaco for S. cad-
ucus).
Populations of S. prionotus from northern Peru
have a higher vertebral crest tlian those in southern
Peru and Bolivia. Northern populations are also
broadly sympatric with two other species of the
"Ophryoessoides group," S. aculeatus and S. fimbria-
tus. However, in southern Peru S. prionotus is not
known to be sympatric with other species of that
group. I postulate that the higher vertebral crest in
northern populations of S. prionotus functions as a
species recognition signal in the multispecies assem-
blages. A key to species of the "Ophryoessoides
group" is provided and distributions of the species in
Peru and Bolivia are summarized.
Resumen. Se describe una nueva especie de lagar-
tija iguanida, Stenocercus prionotus, del Peru Orien-
tal y de Boli\aa. Se conoce la nueva especie desde el
departamento de San Martin, Peni, hasta el norte de
los departamentos La Paz y El Beni en Bolivia. La
mayoria de las localidades se encuentran en las estri-
baciones andinas y adyacentes tierras bajas. Stenocer-
cus prionotus es similar a varias otras especies de
Stenocercus con grandes escamas sobre el posterior
de la cabeza, una fila amplia de supraoculares, y es-
camas ventrales quilladas. Se refiere estas especies
como el "grupo Ophryoessoides," sin implicar su
monophyletismo. Basada en su posesion de un lobulo
escamoso unico que oculta una porcion del bolsillo
antehumeral, se considera la nueva especie cercana-
mente relacionada a Stenocercus caducus (Cope), que
se conoce desde Bolivia central hasta el norte de la
Bull. Mus. Comp. ZooL, 157(3): 183-222, November, 2001 183
184 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
Argentina y del Paraguay. Stenocercus prionotus se
distingue de S. caduciis al tener una cresta vertebral
mas prominente y un patron de alternativas barras
claras y oscuras sobre la garganta (un patron rara-
mente observado en S. caducus, que usualmente ti-
ene manchas claras sobre la garganta).
Las poblaciones de Stenocercus prionotus del norte
del Peni tienen una cresta vertebral mas alta que la
cresta en poblaciones del sur de Peru y Bolivia. Las
poblaciones del norte de Peru tambien son amplia-
mente simpatricas con dos otras especies del "gnipo
Ophryoessoicles," S. aculeatus y S. finibriatus. Sin em-
bargo, en el sur del Peni y Bolivia no se conocen lo-
calidades donde se encuentra S. prionotus simpatrica
con otras especies del giaipo. Postulo que la cresta ver-
tebral mas alto en las poblaciones nortefias de S. prion-
otus functiona como un serial para reconocimiento de
especies en comunidades donde hay varias especies
simpatricas. Se provee una clave para las especies del
"grupo Ophnjoessoides" y se resumen las distribu-
ciones de las especies Peruanas y Bolivianas.
INTRODUCTION
Stenocercus sensu Frost (1992), includ-
ing the nominal genera Ophryoessoicles and
Proctotretus, is a moderately diverse assem-
blage of South American iguanid lizards
(sensu Macey et al., 1997; Schulte et al.,
1998) with about 50 species currently rec-
ognized. Most of the species are in the An-
des and adjacent lowlands of Colombia, Ec-
uador, and Pei"u, but a few are primarily
Amazonian or have distributions in the
physiographically diverse terrain south of
the Amazon basin. Although new species of
Stenocercus continue to be discovered in
the field, others have been known from old
collections and are only now being de-
scribed (e.g., Cadle, 1991, 1998; Avila-Pi-
res, 1995). In this category is a new species
from the lowlands of eastern Peru and Bo-
livia that has been referred erroneously to
the names aculeatus O'Shaughnessy (1879)
or caducus Cope (1862) in previous litera-
ture, and associated with the genera Lei-
ocephalus or Ophnjoessoides before the
current understanding of these names
came into use (see Etheridge, 1966; Frost,
1992). Rodriguez and Cadle (1990) left the
new species nameless in a checklist pend-
ing resolution of its status. The new species
is apparently closely related to Stenocercus
caducus (Cope) and is described herein.
MATERIALS AND METHODS
Frost (1992; see especially footnote 5)
and Cadle (1991) discussed reasons for re-
ferring new species such as the one de-
scribed here to Stenocercus Dumeril and
Bibron sensu lato (including Ophnjoesso-
ides Dumeril and Proctotretus Dumeril
and Bibron). Externally, the new species is
similar to those species that Fritts (1974)
placed in Ophnjoessoides, that is, those
species with keeled ventral scales, large
posterior head scales (usually including
well-differentiated interparietal, parietals,
postparietals, and occipitals), and one
moderately to greatly enlarged supraocular
row. In addition to the new species de-
scribed here, species included in the
''Ophnjoessoides group" are aculeatus
O'Shaughnessy, 1879; caducus Cope,
1862; dunierilii Steindachner, 1867; ery-
throgaster Hallowell, 1856; finibriatus Avi-
la-Pires, 1995; huancahamhae Cadle,
1991; iridescens Giinther, 1859; liniitaris
Cadle, 1998; scapidaris Boulenger, 1901;
tricristatus Dumeril, 1851; and two un-
described species noted later in this paper
under Key to Species of the "Ophryoesso-
ides group" o/ Stenocercus. I use the term
''Ophnjoessoides group" as a convenience
to refer to this group of phenotypically
similar species without implication as to its
status as a monophyletic or nonmonophy-
letic assemblage within Stenocercus. m
General descriptive protocols follow Ca-
dle (1991), who defined terminology of the
scales, neck folds, and mite pockets used
herein, based in part on Frost (1992). Bi-
lateral scale counts (e.g., subdigitals) were
made only on one side (the left, unless it
was damaged), except for the holotype, for
which both left and right counts were re-
corded (1, r). A summary of selected scu-
tellational and qualitative characters for
the new species and similar species from
eastern Peru and Bolivia is presented in
Table 1.
All measurements are in millimeters.
The abbreviation SVL refers to the head-
body length, from snout to vent. The con-
New Species of Stenocercus from Peru and Bolivia • Cadle
185
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186 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
Figure 1. Distribution of species of Stenocercus empfnasized in tfiis paper (western South America, Ecuador to Paraguay and
northern Argentina). Open symbols for S. fimbhatus are literature records from Avila-Pires (1995); locality for S. caducus in
northern Argentina is the southernmost locality in Argentina reported by Cei (1993). Otherwise, all records are based on spec-
imens examined. Numbered localities are documented or suspected cases of sympatry referred to in the text and noted in the
Appendix: 1, Pampa Hermosa; 2, Pampa Seca, Rio Mixiollo; 3, Tingo Maria; 4, Manu National Park. Upper left quadrant outlined
with dotted line is the area shown in greater detail in Figure 2.
figurations of neck folds and mite pockets
vaiy considerably among species of Sten-
ocercus and are useful in distinguishing
species. The most important qualitative
characteristics of these features used here-
in are the following, which are discussed
more fully by Cadle (1991):
Neck and Body Folds and Crests. In
contrast to many species of Stenocercus,
neck folds are usually absent or weakly de-
veloped in the "Opliryoessoides group."
When present, they are better character-
ized as crests rather than folds because
they are usually indicated by strongly
keeled rows of scales instead of actual
folds of skin. The position of such crests
corresponds to the position of folds seen
in other species of Stenocercus, but only
two are commonly seen in the "Ophryoes-
soides group": an antehumeral crest, which
New Species of Stenocercus from Peru and Bolivia • Cadle
187
is a more or less vertical row of strongly
keeled scales immediately anterior to the
forelimb insertion, and usually highlighted
with white scales; and a supra- auricular
crest, a strongly keeled horizontal row of
scales extending from the posterior tem-
poral region to the shoulder region. All
species of the ^'Ophnjoessoides group"
have a distinct vertebral crest formed by
the strongly keeled, and often projecting,
scales of the vertebral row. In addition,
some of these species have a more or less
prominent dorsolateral crest formed by a
row of strongly keeled scales separating
the flanks from the dorsum proper. The
dorsolateral crest varies in length. In some
species it is exceptionally long, extending
from the proximal portion of the tail to the
neck region, where it is confluent with the
supra-auricular crest. In other species it is
present only anteriorly or posteriorly.
Posthiimeral (Axillary) and Postfemoral
Mite Pockets. Type 1 — pocket absent; no
skin modification. Type 2 — rudimentaiy
pocket manifested by skin modification,
such as bare skin, a series of wrinkles, or
a shallow depression lined with scales dif-
ferent from surrounding body scales. Type
3 — similar to Type 2, but with an over-
hanging fold of skin or a thickened border.
Type 4 — a deep pocket, usually with a
broad circular opening, whose depth is
greater than half the diameter of its open-
ing. Type 5 — a deep pocket with a narrow,
slit-like opening and a depth greater than
half the diameter of its opening. In two
species discussed herein the posthumeral
pocket is partially concealed by a scaly flap
of skin, which I term a posthumeral or ax-
illaiy flap. This structure is described more
fully later.
Angulate temporal scales are distinctly
enlarged, keeled scales posterior to, and in
line with, the superciliaiy scales. When
present, they form a distinct border be-
tween the posterior head scales and the
lateral temporal scales, and they are mor-
phologically distinguishable from these se-
ries (Cadle, 1991:^6-7; see Fig. 4). Angu-
late temporal scales are equivalent to su-
pratemporals as used in some literature
(e.g., Avila-Pires, 1995). In several species
of Stenocercus the angulate temporals are
not only keeled but they bear a projecting
bladelike vane from the keels; in such cas-
es I refer to the scales as "projecting."
Coordinates for localities were obtained
from the ornithological gazetteers of the
Neotropics (Stephens and Traylor, 1983;
Paynter, 1989, 1992, 1993, 1997), and
from Lamas (1976), Morales and Mc-
Diarmid (1996), Schulenberg and Awbrey
(1997b), and Peruvian department maps
produced by the Instituto Geografico Na-
cional, Lima. I also consulted the on-line
versions of the Peru and Bolivia gazetteers
of the U.S. Board on Geographic Names
at the GEOnet® Names Sei-ver: http://
164.214.2.59/gns/html/index.html. Brack-
eted data in localities are inferences from
these or other cited sources. Distributions
of the new species and others emphasized
in this paper are given in Figures 1 and 2.
Institutional abbreviations are given at the
beginning of the Appendix.
DESCRIPTION OF A NEW SPECIES OF
STENOCERCUS
Stenocercus prionotus^
new species
Figures 3-7, Figure 12; Table 1
Liocephalus cadiicus (Cope, 1862): Boulenger (1898),
specimen from "Barraca, Rio Madidi" [Boli\da]
(BMNH 98.6.9.4) and probably two other northern
BoHvian locahties discussed in the text.
Ophnjoessoldes caducus (Cope, 1862): Fugler (1989:
63), specimens from San Marcos Ranch, El Beni
Department, Boli\da, including ROM 12815.
Ophryoessoides acideatus (O'Shaughnessy, 1879):
Fugler (1983, 1986, 1989), specimens from Tumi
Chucua, El Beni Department, Bolix-ia (USNM par-
atypes).
Ophryoessoides sp.: Rodriguez and Cadle (1990),
specimen from Cocha Cashu, Manu National Park,
Madre de Dios Department, Peru (MCZ 150243).
- Stenocercus pnonotus was recognized as new by
R. Etheridge, P. E. Vanzolini, and E. E. Williams
many years ago. Vanzolini and Williams applied the
unpublished name Stenocercus dorsatus to labels of
many specimens in various collections.
188 Bulletin Museum of Comparative Zoologij, Vol. 157, No. 3
7511^0°
New Species of Stenocercus from Peru and Bolivia • Cadle 189
Figure 3. Dorsal and ventral views of the holotype of Stenocercus prionotus (USNM 193683). Approximately xO.87.
Figure 2. Northern Peru and Ecuador (see Fig. 1) showing distributions of species in the "Ophryoessoides group." Numbered
localities are documented or suspected cases of sympatry referred to in the text and the Appendix (see Fig. 1 for names). Open
circle at 06°S, 77°W is the type locality for Stenocercus aculeatus (Moyabamba, San Martin Department). All other localities are
based on specimens examined; see Cadle (1991, 1998) for S. iridescens, S. limitaris, and S. huancabambae. The known
distributions of all species are indicated by the localities plotted; however, the distribution of S. /r/descens continues farther north
in western Ecuador than the area covered by the map.
190
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
Holotype (Figs. 3—5). United States
National Museum of Natural History
(USNM) 193683 (field number WCS
2421). PERU: Depto. Huanuco: Jardin
Botanico de la Universidad Agraria de la
Selva, Tingo Maria, vicinity of Rio Hual-
laga, 670 m elevation [09°18'S, 75°59'W].
Adult male collected 29 June 1966 by
Wade C. Sherbrooke.
Faratypes from the Vicinity of the Type
Locality. PERU: Depto. Huanuco: Ca. 2
mi. by trail W. Tingo Maria, west bank of
Rio Huallaga in vicinity of confluence with
Rio Monzon, 670 m elevation (9 October
1966, W. C. Sherbrooke), USNM 193685.
Universidad Agraria de la Selva, Tingo
Maria, Rio Huallaga, 670 m elevation (18
August 1967, unknown collector for W. C.
Sherbrooke), USNM 193686. Vicinity of
Cueva de las Lechuzas, ca. 3 mi. SW Tingo
Maria, Rio Monzon, ca. 700 m elevation
(17 April 1968, Vito Yaringano for W. C.
Sherbrooke), USNM 193687. Picuriacu,
ca. 2 mi. NW Tingo Maria, Rio Huallaga
(20 April 1968, W. C. Sherbrooke), USNM
193688.
Other Faratypes. PERU: Depto.
Huanuco: Buena Vista, Valley of the
Chimchao [ = Rio Chinchao] [approximate-
ly 9°31'S, 75°52'W] (1-15 September
1923, E[dmund] Heller), FMNH 5582-
83. Hacienda Pampayacu [09°33'S,
75°54'W] (17 July-16 August 1936, Dr.
Snonge), MCZ 43758-59, 43761-62.
Rio Llullapichis, 4—5 km upstream from
Rio Pachitea, 200 m elevation [09°37'S,
74°55'W] (Januaiy 1969, Hans W. Koep-
cke), KU 179058. [Depto. Loreto]:
E [astern] Peru, Pampa Hermosa, near
mouth of Rio Cushabatay, Rio Ucayali Val-
ley 500 ft. [152 m] [07°12'S, 75°17'W]
(date unknown, H[ai-vey] Bassler), AMNH
56760-64. Depto. Madre de Dios: Co-
cha Cashu Biological Station, Manu Na-
tional Park [11°51'S, 71°19'W] (July 1975,
John W. Fitzpatrick), MCZ 150243. Ex-
plorer's Inn, Tambopata Reserve, ca. 30
km (straight line) SSW Puerto Maldonado,
280 m [12°50'S, 69°17'W] (2 September-
7 October 1983, native collectors), USNM
247468-69, 247680; (23 May 1986, Vic-
tor R. Morales), USNM 269022. Depto.
Puno: Prov. Sandia: Tambopata, San Juan
[del Oro], 1520 m or 5000 ft. [14°12'S,
69°08'W]^ (22 November-20 December
1950, Hilda H. Heller), FMNH 64788-
92, 64794-811. [Depto. San Martin]:
Juanjui [07°11'S, 76°45'W] (collector and
date' unknown), MCZ 121233. Tarapoto,
370 m [06°30'S, 76°25'W] (25 July 1984,
Rainer Schulte), KU 212629.
ROLIVIA: Depto. Reni: Provincia Va-
cadiez, Tumi Chucua [176 m; 11°08'S,
66°10'W] (23 October-18 November
1981, Charles M. Fugler), USNM
280246—51. Puerto Cruzeiro, San Marcos
Ranch at confluence of Rios Isiboro and
Ichoa [15°17'S, 65°45'W] (10 February
1977, J. Lovisek), ROM 12815. [Depto.
La Paz]: Barraca, Rio Madidi [12°35'S,
67°02'W] (1893, Luigi Balzan), RMNH
98.6.9.4.
Distribution (Fig. 1). Stenocercus prion-
otiis is known from the lowlands and An-
dean foothills of eastern Peru and adjacent
Bolivia (San Martin department, Peru,
south to the Rio Beni in northern Bolivia).
The known elevational range is 176—1,520
m, but the highest recorded elevation is
more than twice the elevation of the next
lower one. Most localities in Peru are ad-
jacent to or near the Andean foothills, and
several localities in La Paz and El Beni
departments, Bolivia, are unconfirmed. No
specimens are known from a broad geo-
graphic hiatus between central and south-
ern Peru (Fig. 1). See Distribution Fat-
terns in Stenocercus prionotus for further
discussion.
Etymology. The epithet prionotus is de-
^ Hilda Hellers notes on this collection in the Karl
P. Schmidt archives of the FMNH describe San Juan
as the site of an agricultural station on "the left side
of die Rio Tambopata at 5000 feet." With some hes-
itancy I identify Heller's locality as the town kno\\'n
as San Juan del Oro, which is on the left bank of the
Rio Tambopata at approximately the elevation given
by Heller. I have located only two other places named
San Juan in Puno Department, but neither is on the
Rio Tambopata.
New Species of Stenocercus from Peru and Bolivia • Cadle
191
rived from the Greek adjective prionotos
meaning jagged or serrate. The name re-
fers to the strongly serrate vertebral crest
of Stenocercus prionotus, which is the
most obvious character distinguishing this
species from its apparent closest relative,
S. caduciis.
Data on the Holotype. Adult male, hem-
ipenes partially everted. SVL, 83 mm. Tail
length, 201 mm. Total length, 284 mm.
Tail/total length, 0.71. Vertebral scales be-
tween the occipital and the posterior mar-
gin of the hind limb, 31. Midbody scales,
42. Gular scales between the ears, 16. In-
ternasals, 6. Subdigital scales on fourth fin-
gers and toes, respectively, 18—18, 25—25.
Color pattern well presei-ved: top of head
brown with narrow dark brown interorbital
bar extending laterally onto supraoculars;
dorsum brown with narrow blackish chev-
rons middorsally (1 on neck, 1 above fore-
limbs, 2 others anterior to midbody; pos-
terior chevrons poorly defined); dark
brown scapular blotch bordered anteriorly
by white antehumeral stripe; ill-defined
grayish dorsolateral streaks between ear
and anterior body; throat grayish with
poorly defined oblique light grayish
stripes; venter brown without distinct pat-
tern.
Definition. A species of Stenocercus
characterized by the following features: (1)
Dorsal head scales subimbricate and
strongly keeled to multicarinate; temporals
keeled, imbricate or subimbricate. (2) Pos-
terior head scales larger than anterior
ones, with distinct interparietal, a pair of
parietals, a pair of postparietals, and a
large median occipital (often surrounded
by several small irregular scales). (3) In-
ternasals usually 7, but pattern irregular
and may be 5 or 6. (4) One row of supra-
oculars distinctly enlarged. (5) One canthal
on each side between the superciliaries
and the lateralmost internasal. (6) A pair
of strongly keeled angulate temporals in
line on each side (rarely, 3 angulate tem-
porals are present), each with a low pro-
jecting blade; partially or completely sep-
arated from enlarged posterior head scales
by a single row of small scales. (7) Anterior
and posterior gular scales strongly keeled.
(8) Parietal eye distinct. (9) Neck folds ab-
sent; a vertical, strongly keeled row of
scales in the antehumeral region and oc-
casionally a much less distinct raised series
in the supra-auricular region. (10) Dorsal
and ventral body scales imbricate, mucro-
nate, strongly keeled; dorsal scales at mid-
body 36—48. (11) Vertebral row continu-
ous, bearing a strongly projecting serrate
crest in adults; a dorsolateral crest present
on posterior body and the base of the tail.
(12) Deep posthumeral pocket (Type 4)
partially concealed by a scaly posthumeral
flap originating on its anteroventral bor-
der; postfemoral pocket absent (Type 1).
(13) Scales of posterior thigh imbricate,
keeled. (14) Tail strongly compressed in
adults, anteriorly with low vertebral and
dorsolateral crests continuous with those
of the body. (15) Dorsal coloration of
males in preservative (Figs. 3, 6) brown
with or without distinct chevrons; a dis-
tinct white vertical antehumeral stripe ex-
tending ventrally to the proximal ventral
surface of forelimb; a large dark scapular
blotch; in well-presei"ved specimens the
throat bears oblique alternating dark and
light stripes (see Description); females
similar but pattern elements often more
subdued.
DIAGNOSIS AND COMPARISONS
In having enlarged posterior head
scales, an enlarged row of supraoculars,
and strongly keeled ventral scales, Steno-
cercus prionotus is like other species in the
"Ophryoessoides group" of Stenocercus.
These are the species most likely to be
confused with S. prionotus. Five other de-
scribed species of the ''Ophryoessoides
group" occur in eastern Peru or Bolivia: S.
aculeatus (O'Shaughnessey, 1879); S. cad-
ucus (Cope, 1862); S. finibriatus Avila-Pi-
res, 1995; S. huancabanibae Cadle, 1991;
and S. scapularis (Boulenger, 1901). An
undescribed species occurs in the Rio
Maraiion valley of eastern Peru (see Key
to Species of the "Ophryoessoides Group"
192 Bulletin Museum of Comparative Zoologtj, Vol. 157, No. 3
of Stenocercus). Stenocercus prionotus
and S. caducus are unique among known
species of Stenocercus (perhaps unique
within iguanids) in having deep posthu-
meral mite pockets (Type 4) that are par-
tially concealed anteroventrally by a scaly
flap, which may be termed a posthumeral
or axillary flap (Fig. 5). Stenocercus prion-
otus and S. caducus are compared in
greater detail below, but S. pmonotus is
distinguished from S. caducus (character-
istics in parentheses) by: (1) a strongly pro-
jecting, serrate vertebral crest (low and
scarcely projecting); (2) 2 (usually) or 3 en-
larged, strongly keeled, projecting angu-
late temporal scales on each side (scales
not greatly enlarged, less projecting); and
(3) a gular pattern consisting, when evi-
dent, of oblique alternating dark and light
lines or bars, or oblique light lines on a
dark ground color (usually light spots on a
darker ground color, unicolor, or [rarely] a
pattern similar to that of S. prionotus).
Readily determined characters distin-
guishing Stenocercus prionotus from the
other four species of the "Ophryoessoides
group" known from eastern PeiTi and Bo-
livia include the extent of keeling on dorsal
head and body scales, relative develop-
ment of the postfemoral pockets, and the
number of midbody scale rows (Table 1
and key presented later herein). Stenocer-
cus fimbriatus and S. aculeatus are known
to be sympatric with S. prionotus at several
localities in eastern Peru. In addition to
having a posthumeral flap (absent in S.
fimbriatus and S. acideatus), S. prionotus
is distinguished from S. fimbriatus (char-
acteristics in parentheses; see Avila-Pires,
1995) in having strongly keeled dorsal
scales in adults (smooth or weakly keeled),
a dorsolateral crest prominent only on the
posterior body (prominent anteriorly and
continuous with antehumeral and supra-
auricular folds or crests), and in lacking
"fimbriate" scales on the posterior distal
portion of the thigh (present). Stenocercus
prionotus is distinguished from S. aculea-
tus (characteristics in parendieses) in lack-
ing a postfemoral pocket (moderate to
deep); in having strongly keeled, often
multicarinate, head scales (smooth or
weakly striated in adults, wrinkled in ju-
veniles); 5—7 internasals (4—5); and only
moderately enlarged supraoculars, usually
5—6 supraoculars across the widest part of
the orbit (greatly enlarged, usually 4 across
the orbit).
Stenocercus prionotus differs from S.
scapularis (characteristics in parentheses)
in lacking squarish or rectangular project-
ing superciliaiy scales (present) and in
having fewer than 50 midbody dorsal scale
rows (59—70 rows). Stenocercus prionotus
differs from S. huancabambae (characters
in parentheses) in lacking a postfemoral
pocket (deep, Type 5) and in having prom-
inent dorsolateral crests on the posterior
body (weak, restricted to anterior body
when present).
Two species of Stenocercus from west-
ern Ecuador and Peru, S. iridescens and
S. limitaris, have enlarged posterior head
scales and supraoculars. In contrast to
Stenocercus prionotus, S. iridescens has
smooth head plates, 2 canthals, a poorly
developed posthumeral pocket (Type 1 or
2), and lacks keeled angulate temporals
and dorsolateral crests on the body (see
Cadle, 1991, fig. 10). Stenocercus limitaris
has a deep postfemoral pocket (Type 5), 2
canthals, a single strongly keeled (but non-
projecting) angulate temporal, and lacks
dorsolateral crests.
Other non-Peruvian species of the
"Ophryoe.ssoides group" can be distin-
guished from Stenocercus prionotus by
features in the key presented later and
other superficial characters, such as the
presence of 2 canthals and 4 internasals
(usually) in S. erythrogaster (1 and 5—7 in
S. prionotus), and enlarged, projecting py-
ramidal or conical postsuperciliary scales
in S. dumerilii and S. tricristatus (Avila-
Pires, 1995).
Other species of Stenocercus are distin-
guished from S. prionotus by a combina-
tion of features such as smaller head
plates, smooth ventrals, and absence of
dorsolateral crests. Most species of Steno-
New Species of Stenocercus from Peru and Bolivia • Cadle
193
cercus except the "Ophnjoessoides group"
have smooth or weakly keeled head plates.
Other characters, such as the number of
dorsal scale rows, morphology of the pos-
thumeral and postfemoral mite pockets,
extent of sexual dimoi*phisin, and degree
of differentiation of the vertebral scale row
and crest also aid in distinguishing the spe-
cies (see descriptions and discussion in
Fritts, 1974; Cadle, 1991, 1998).
Apart from Stenocercus fimbriatus, S.
scapularis, and S. acideatus, two other spe-
cies of Stenocercus are known from local-
ities close to or synipatric with known pop-
ulations of S. prionotus: S. crassicaudatus
and S. roseiventris. These species are dis-
tinguished from S. prionotus by lacking
prominent serrate vertebral crests (low
crests may be present), having smooth
ventral scales, and having prominently spi-
nose tails with the spines arranged in dis-
tinct whorls.
DESCRIPTION
Head (Fig. 4). Dorsal head scales sub-
imbricate (a tendency to be more juxta-
posed posteriorly); strongly keeled to mul-
ticarinate or wrinkled. Rostral in contact
with first supralabial, first lorilabials, and a
series of postrostrals. Usually 7 elongate,
strongly keeled internasals between the
nasals dorsally; however, the anterior dor-
sal head scales are very irregular and oc-
casionally only 5 or 6 internasals are pre-
sent. One canthal scale between the an-
terior superciliary and the lateralmost in-
ternasal, separated from the nasals by tiny
postnasals. Canthus very strongly angled.
Nostril in posterior portion of an elongate
nasal scale, which may contact the rostral
scale anteriorly or be separated from it by
small postrostrals. Four or 5 strongly over-
lapping, elongate anterior superciliaries
followed by 2 or 3 shorter posterior su-
perciliaries slightly overlapping in the re-
verse direction (but more or less in a
straight line). One supraocular row inod-
erately enlarged, 2 mediocentral scales
much larger than the others. Five or 6
scales across the supraocular area at its
widest part. Interparietal distinct and elon-
gate, diamond-shaped or pentagonal (apex
posteriorly). Parietal eye visible. A pair of
parietals in contact behind the inteqDari-
etal, flanked posterolaterally by a postpar-
ietal on each side. Postparietals separated
medially by a single median transversely
elongate occipital; occasionally 1 or 2 small
scales are intercalated at the juncture of
the parietal, postparietal, and/or occipital
(e.g.. Fig. 4).
Lateral temporal scales strongly keeled,
imbricate to subimbricate; separated from
posterior dorsal head scales on each side
by 2 (occasionally 3) elongate, strongly
keeled angulate temporal scales bearing a
low projecting vane. Keels of adjacent an-
gulate teinporals aligned. Posterior angu-
late temporals separated from postparie-
tals by 2 or 3 small scales in a longitudinal
row. Anterior angulate temporals may con-
tact postparietal and one other larger pos-
terior head scale or be separated from
them by small scales. Anterior border of
ear weakly denticulated; posterior border
rounded, bordered with keeled imbricate
scales.
Anterior and posterior gulars strongly
keeled. Mental smooth, in contact with
first pair of postmentals and first pair of
infralabials. Enlarged postmentals 3 or 4
on each side, only the first pair in contact
medially.
Neck and Body. Dorsal and lateral
scales of neck and body imbricate, mucro-
nate, strongly keeled. Vertebral row pro-
duced into a prominent projecting serrate
crest in adults of both sexes that is contin-
uous from the nuchal region to the base
of the tail, gradually disappearing on the
anterior Va to V2 of tail. Dorsolateral crest
(a raised, strongly keeled row of scales) on
posterior % of body, continuing onto base
of tail. The dorsolateral crest occasionally
appears very indistinctly farther anteriorly
on the body, but only on the posterior
body does it shaiply delimit the dorsolat-
eral (paradorsal) scales from the flank
scales. Three rows of scales between dor-
solateral and vertebral crests at anterior
194 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
Figure 4. Head scales of the holotype of Stenocercus prionotus (USNM 193683) in dorsal and lateral views. Scale bars = 3
mm. To facilitate coordination with the text, the following scales are indicated on one side (interparietal and occipital are median
scales); A, angulate temporal; I, interparietal; O, occipital; P, parietal; PP, postparietal.
New Species of Stenocercus from Peru and Bolivia • Caclle
195
edge of dorsolateral crest (i.e., just anterior
to the pelvic region), 2 rows posteriorly
(dorsal to hindlimbs); scales between the
crests strongly imbricate and keeled only
on posterior part of scale. Flank scales
mostly fully keeled (sometimes only the
posterior part of each scale), imbricate,
mucronate, slightly smaller than dorsolat-
eral scales. Ventral body scales mucronate,
strongly keeled (keels running the length
of the scales). Ventrals approximately the
same size as the dorsolateral scales, larger
than flank scales.
Neck Folds. Distinct neck folds absent.
Poorly developed antehumeral crest pre-
sent.
Tail. Tail strongly compressed, anteriorly
bearing low projecting vertebral and dor-
solateral crests continuous with those of
the body. Dorsal scales moderately keeled,
ventral scales strongly keeled.
Limbs. Dorsal and ventral scales of fore-
limbs, hindlimbs, and posterior thigh
strongly keeled, unicarinate, mucronate;
some scales of the shank larger than any
thigh scales. Supradigitals and subdigitals
unicarinate. Palmar scales strongly multi-
carinate. Plantar scales strongly unicari-
nate.
Posfhiimeral and Postfemoral Mite
Pockets. Posthumeral mite pocket a deep
cavity (Type 4) with a prominent axillaiy
flap concealing the anteroventral aspect
(Fig. 5). Postfemoral pocket absent (Type
The flap associated with the posthumer-
al pocket projects from the anteroventral
and ventral edges of the pocket. Antero-
dorsally, a similar but much smaller flap is
present in some specimens (e.g.. Fig. 5).
The posthumeral flap consists of a fleshy
ridge covered anteriorly and posteriorly
(or externally and internally when the flap
is lying flat against the body) by keeled im-
bricate scales. Externally, usually 3 or 4
larger scales cover the flap ventrally and a
series of much smaller scales is present
dorsally. One or 2 of the larger scales are
sometimes highlighted with white. When
appressed against the body (i.e., against
Figure 5. Posthumeral (axillary) flap of Stenocercus priono-
tus (USNM 193683, holotype). Anterior to the right. The broad
oval on the right is the deflected forelimb and the posthumeral
mite pocket is the heavily stippled cavity deep to the flap. The
posterior border of the axillary flap is marked by the scales
with heavily outlined posterior borders and it extends anteriorly
to the ventral part of the forelimb. A smaller dorsal flap is also
present in this specimen (small patch of heavily outlined scales
on anterodorsal edge of the pocket; see text). Scale bar = 1
mm.
the opening of the posthumeral pocket)
the flap conceals approximately the ventral
Vs to Vi of the vertical dimension of the
pocket. The flap is equally prominent in
adults of both sexes and is proportionally
as well developed in subadults (including
hatchlings) as in adults.
Size and Propoi~tions. Largest inale
(USNM 193683) 89 mm SVL, 323 mm to-
tal length (sample size of males with SVL
>70 mm = 13). Largest female (KU
212629) 93 mm SVL, 329 mm total length
(sample size of females with SVL >70 mm
= 16). Tail relatively long, 69-74% of total
length in adults (67-71% in juveniles).
Coloration and Pattern of Adult Males
in Life. The following color descriptions
are paraphrased from the field notes of
Wade C. Sherbrooke. USNM 193683 (ho-
lotype):
Lower half of side between limbs is lavender-
brown. This color extends from both sides across
the belly approximately Vs of the way on each side,
leaving a tan-brown central strip down the belly.
General base color of the body is brown, darkest
196 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
Figure 6. Lateral view of Stenocercus prionotus from northern Peru (USNM 193685; male, snout-vent length 76 mm). Note
the high vertebral crest and the following pattern elements: dark subocular bar, dark blotches on the dorsal and lateral surfaces
of the neck, pale antehumeral bar, indistinct pale dorsolateral stripe, and indistinct oblique bars on the trunk.
near the hind legs and tan just in from of the fore-
legs. A distinct white line runs dorsally from the
top of each foreleg to three quarters of the way to
the dorsal crest; it runs through a large black patch
just above the forelimb. Head markings consist of
a dark brown line i-unning between the eyes on die
top of die head; this continues through the eye to
broaden slightly at the rear portion of the jaw. The
gular area is streaked by several light cream lines.
Very slight lavender tinge to all of body behind
forelimbs. [Sketch in notes shows black middorsal
patches that don't extend to flanks].
USNM 193685 (WCS 2543):
This specimen closely resembles [USNM 193683]
in color, with one exception. There are several
green spots on . . . the right dorsal surface between
the limbs and on the dorsal portions of the tail
behind the hind limbs and the dorsal tibio-fibula
portion of the right hind leg.
Coloration of Adult Females in Life. Un-
known.
Coloration in Preservative (Figs. 3, 6, 7).
Specimens of Stenocercus prionotus vary
greatly in coloration due mainly to varia-
tion in initial preservation and the length
of time in presei-vative. Poorly preseived
specimens may be more or less uniform
brown all over, although obscure pattern
eleiTients are usually present. Well pre-
served specimens are brown dorsally with
darker brown to blackish chevrons mid-
dorsally. One chevron dorsal to each pair
of limbs and one on the neck are usually
evident, and these are darker than others
that iTiay be present. Three to five mid-
dorsal chevrons are between the limbs.
Dark spots or an additional chevron are
often present on the dorsal neck and usu-
ally on the base of the tail. The light an-
tehumeral/humeral line is universally pre-
sent and evidence of the dark shoulder
patch is usually present (often veiy prom-
inent). Flanks usually unicolor and some-
what darker than the dorsum between the
dorsolateral crests; however, soixie speci-
mens (e.g., KU 179058, USNM 280246)
have distinct dirty white vertical bars or
chevrons on the flanks (five between the
limbs), and such bars appear occasionally,
but more obscurely in other specimens
(see Fig. 6). The dorsolateral crest is often
highlighted for a variable length with a dis-
tinct or indistinct light line, giving the im-
pression of a light dorsolateral stripe.
Forelimbs more or less unicolor brown or
with obscure pattern; hindlimbs brown
with darker brown bands. Dark subocular
bar distinct. Top of head often with an ob-
scure or distinct dark brown interocular
bar. Oblique bars on throat (Fig. 7) often
visible but throat may be unicolor or have
an obscure pattern. Venter of most speci-
mens unicolor, dirty white, gray, or beige;
however, some specimens (e.g., USNM
280246) have a series of irregular longi-
tudinal dark brown streaks.
Scale Counts and Qualitative Features
(Table 1). Stenocercus prionotus has rela-
tively low midbody, vertebral, and gular
scale counts. The scales are relatively large
and strongly keeled over most of the body.
New Species of Stenocercus from Peru and Bolivia • Cadle
197
Figure 7. Gular patterns in Stenocercus prionotus. Top, typ-
ical throat pattern (oblique view) consisting of light and dark
stripes that extend medially to the midline (USNM 193687).
Bottom, specimen in which the ground color is lighter and
therefore the contrasting pale stripes are less distinct (USNM
193685).
Sexual Dimorphism. Stenocercus prion-
otus does not exhibit strong sexual dimor-
phism. Males and females attain approxi-
mately the same size and have the same
general pattern, but whether the colora-
tion in life reported above for adult males
pertains to females as well is unknown.
The vertebral crest is only slightly more
developed in males than in females of the
same population but this character shows
strong clinal variation (northern popula-
tions with higher crests; further discussed
below). Other characters that sometimes
vary between the sexes in Stenocercus
show little variation in S. priontotus. Nei-
ther standard meristic counts (Table 1) nor
the relative development of the posthu-
meral and postfemoral pockets (Types 4
and 1, respectively, in adults of both sexes
and in subadults) show obvious sexual di-
morphism.
DISTRIBUTION PATTERNS IN
STENOCERCUS PRIONOTUS
Absence of Stenocercus prionotus from
Lowland Localities in Eastern Fern. Sten-
ocercus prionotus is widespread in the
lowlands along the Andean front from
northern Peru to northern Bolivia (Fig. 1).
However, all Peruvian localities are close
to the Andean foothills and south of the
broad extension of the Cordillera Oriental
separating the great bend of the Rio Mar-
aflon from upper reaches of the Rio Hual-
laga (Fig. 2). The absence of specimens in
comprehensive collections from the Iqui-
tos region (Dixon and Soini, 1986), Balta
(Ucayali Department; specimens at
LSUMNS and University of Arizona), and
Cuzco Amazonico (Madre de Dios De-
partment; Duellman and Salas, 1991) sug-
gest that S. prionotus may be absent from
the lowlands distant from the Andean foot-
hills, at least in Peru. Similarly, collections
from northern Loreto Department (Duell-
man and Mendelson, 1995) and northern
Amazonas Department (J. E. Cadle and R.
W. McDiarmid, unpublished data from the
Rio Cenepa and Rio Santiago) suggest that
S. prionotus does not occur north of the
Rio Marafion.
However, these sites have been sampled
unevenly. For example, species accumula-
tion curves for lizards at Cuzco Amazonico
(Duellman and Koechlin, 1991) reached
an asymptote after about 15 person-weeks
of effort, whereas only 5 person-weeks
were expended in northern Loreto and the
species accumulation curve for the total
herpetofauna showed no asymptote
(Duellman and Mendelson, 1995; data not
presented for lizards only). Quantitative
data are not available for the other sites,
but large collections are available for the
Iquitos region, including more than 1,000
198 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
Table 2. Patterns of presumed sympatry of Stenocercus in eastern Peru. Only species of the
"Ophryoessoides group" and the superficially similar species, S. roseiventris, are listed. 1 Locali-
ties ARE listed roughly NORTH TO SOUTH; FOR PRECISE LOCALITIES SEE THE APPENDIX AND LOCALITIES
FOR THE TYPE SERIES OF S. PRIONOTUS..
Iquitos San Martin/ Pampa
Region VV. Loreto- Hermosa
Rio
Mishollo
Tingo
Maria
Balta
Cuzco
Amazonico
Explorer's
Inn
S. prionotus
X
X
X
X
S. aculeatus
X
X
S. fimbriatus
X
X
X
X
X
S. roseiventris
X
X
^ The only other species of Stenocercus known from primarily the lowlands and lower Andean foothills in
this region is S. crassicaiidatus, which lacks a projecting vertebral crest and has a very spiny tail.
^ The distributions of S. prionotus and S. aculeatus overlap both altitudinally and latitudinally in northern
Peru but they have not been taken together at the same locality. See Figure 2 for localities.
^ In Manu National Park, S. fimbriattis and S. roseiventris are sympatric at Pakdtza, whereas S. prionotus is
known only from Cocha Cashu.
lizards obtained by Dixon and Soini (1986)
and additional collections froin the region
made by Harvey Bassler and deposited in
the AMNH. Similarly large collections re-
sulted from the efforts of Cadle and
McDiarmid in Amazonas (specimens in
the MVZ and USNM). Thus, barring ar-
tifacts introduced by the difficulties of col-
lecting cryptic rainforest lizards, S. prion-
otus seeiTis to be absent froixi these sites.
Documenting and explaining patterns of
absence is always difficult, but the appar-
ent absence of Stenocercus prionotus froin
the lowlands distant from the Andes in
Peru is not due to failure to collect Sten-
ocercus at these localities because at least
one other species of Stenocercus is known
from each (Table 2). However, no lowland
locality is known in which inore than two
species of the "Ophryoessoides group" of
Stenocercus are syinpatric. Of interest in
this connection is that S. prionotus has not
been taken in the upper reaches of the Rio
Perene and its tributaries (Junin Depart-
ment), although it is known from north
and south of that region. Numerous spec-
iinens of Stenocercus of at least five spe-
cies (S. hoettgeri, S. crassicaudatus, S. for-
mosus, S. scapularis, and S. variabilis'^)
have been collected along an elevational
^ Based on examination of two of the three synty-
pes of Stenocercus variabilis Boulenger (BMNH
1946.8.11.89, 1946.8.11.91) I concur with Fritts
(1974: 66) that Boulenger (1901: 553) erred in as-
cribing these specimens to "Palca, Bolixda" rather
than Palca (Junin Department), Peru. One specimen
I collected near Palca, Peru (MCZ 178166) is nearly
identical with BMNH 1946.8.11.91 in scale counts,
pattern, and qualitative characters (both of these
specimens differ considerably from BMNH
1946.8.11.89 in color pattern). However, the attri-
bution of the specimens to Peru is not without some
equivocation. The types were collected by P. O. Si-
mons (see Cadle, 1998: footnote 6), who collected in
the vicinity of Palca, Peru, in March and April, 1900,
although that specific locality is not listed in his pub-
lished itinerary (Chubb, 1919). However, Simons's
itineraiy places him at "Palca, 18 miles E of La Paz"
[Bolivia] on 9 November 1900 (Chubb, 1919: 5). Si-
mons's field tags attached to the syntypes record sim-
ply "Palca 3000 m," which is close to the elevation of
Palca, Peru (2,740 m), but not that in Bolivia (4,600
m). Adding to the confusion are entries in the
BMNH registries for the syntypes, concerning which
Colin J. McCarthy provided the following comments
via e-mail:
Boulenger originally wrote "Palca Peru 3000m" but
later struck through Peru and wrote "Bolivia."
There are two sheets of notes (presumably from
Simons) stuck in at this page of the register about
the localities in the batch. With regard to Palca he
has written "Palca, just S. of La Paz, Bolivia." I
assume it was that information that caused Boulen-
ger to alter his original entry!
Thus, the confusion may have originated with the
note that Simons provided subsequent to cataloguing
of the collection at the BMNH. In any case, no spec-
imens resembling S. variabilis are definitely known
from Bolivia and the only species of the genus defi-
nitely known to occur above 4,000 m is Stenocercus
chrysopt/gus from northern Peru.
New Species of Stenocercus from Peru and Bolivia • Cadle
199
transect along this well-traveled route
(Fritts, 1974; Cadle, 1991 and unpublished
data). Thus, analysis of circumstantial dis-
tributional data suggests that the distribu-
tion of S. prionotus in the lowlands of east-
ern Peru may be influenced by the num-
ber of sympatric species of Stenocercus. At
all localities from which S. prionotus has
been taken, only one other species of Sten-
ocercus is known (Table 2). These patterns
of sympatry are discussed in the next sec-
tion with reference to patterns of geo-
graphic variation in S. prionotus.
In contrast, the apparent restriction of
Stenocercus prionotus to the Andean foot-
hills and immediately adjacent lowlands in
Peru does not seem related to the distri-
bution of any major habitat type or phys-
iographic region. Most of the extensively
sampled localities (e.g., Iquitos and Cuzco
Amazonico) include a variety of lowland
habitats characteristic of western Amazon-
ia. Stenocercus prionotus is known from
both floodplain forests (Cocha Cashu) and
more upland forests on river terraces in
southeastern Peru (Explorer's Inn); Foster
(1990) and Dallmeier et al. (1996) de-
scribed these floristic communities in
southeastern Peru. Thus, the apparent ab-
sence of S. prionotus at the localities dis-
cussed above is not due to some simple
relation to local habitat availability. For ex-
ample, it is unclear why S. prionotus was
not obtained at Pakitza (Morales and
McDiarmid, 1996), even though it occurs
at nearby Cocha Cashu. On a broader geo-
graphic scale, the restriction of S. priono-
tus to lowlands and foothills adjacent to
the Andes may be related to the present
or historical influence of the Andes on the
climate and vegetation (rainfall, tempera-
ture, and major soil types) of neighboring
regions.
A curious and unexplained hiatus in the
distribution of Stenocercus prionotus oc-
curs between the vicinity of Tingo Maria—
Rio Llullapichis (Huanuco Department)
and Cocha Cashu in Manu National Park
(Madre de Dios Department), a gap of
some 600 km that includes the entire up-
per reaches of the Rio Ucayali-Urubam-
ba— Ene system. Additionally, populations
north and south of this gap differ in some
qualitative and quantitative characters (see
Patterns of Sympatry and Geographic Var-
iation in Stenocercus prionotus, and the
Need for Additional Fieldivork). Scattered
collections (e.g., maps in Fritts, 1974; Avi-
la-Pires, 1995), but no comprehensive her-
petofaunal surveys or collections, are avail-
able from this vast region. Thus, whether
the geographic hiatus is real or a sampling
artifact cannot be discerned. Efforts to re-
solve this issue need to be made.
Unconfirmed Bolivian Localities for
Stenocercus prionotus. Because of the
previous confusion of Stenocercus priono-
tus wath S. caducus, specimens perhaps re-
ferable to S. prionotus from several local-
ities in Bolivia are unconfirmed. Boulen-
ger (1898) reported specimens of "Lioce-
phalus caducus. Cope" in the Museo
Civico Storia Naturale Giacomo Doria in
Genoa collected by Luigi Balzan from four
localities (Balzan, 1931). I have not at-
tempted to verify the existence of these
specimens. However, one of Balzan s spec-
imens from "Barraca, Rio Madidi" was ex-
changed to the BMNH (now BMNH
98.6.94), and is confirmed as S. prionotus.
The "Leiocephalus caducus" specimens
from the other three Balzan localities are
outside the known distribution of S. cad-
tictis but are close to other known localities
for S. prionotus in northern Bolivia (Fig.
1). I suspect these are S. prionotus based
on geographic location. The localities are,
as listed by Boulenger (1898) (see Fig. 1),
(1) "Coroico and Chulumani, Prov. Yun-
gas, 1,600 metres alt." [La Paz Depart-
ment; Coroico, 1,725 m, 16°10'S, 67°44'W;
Chulumani, 1,905 m, 16°24'S, 67°31'W].
(2) "Reyes, right bank of Rio Beni" [El
Beni Department, 232 m; 14°19'S,
67°23'W]. (3) "Missiones [sic] Mosetenes"
[approximately 15°31'S, 67°25'W].5 These
^ The Moseten Indians inhabited upper reaches of
the Rio Beni and its tributaries in the Andean foot-
hills of the present department of La Paz (Metraux,
200
Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
localities would not be unusual for S.
prionotus, although the first two localities
are the highest elevations recorded for the
species (the species occurs at 1500 m in
nearby Puno Department, Peru). All are
in the upper reaches of the Rio Beni,
whereas the two confirmed Bolivian local-
ities for S. prionotus are farther north in
the same drainage.
Burt and Burt (1931: 273) listed two
specimens of ''Leiocephalus [ = Stenocer-
cus] scapularis" (AMNf the
area; females with enlarged eggs were
found in early November (Fugler, 1986:
table 4).
Hilda H. Heller provided the foUov^dng
Figure 10. Diagrammatic representation of geographic and size-related variation in the height of the vertebral crest in Steno-
cercus prionotus and S. caducus. Drawings were made with a camera lucida to emphasize the form and height of the vertebral
crest. Sketches are drawn to an approximately uniform interval between the external ear opening (EE) and the white antehumeral
stripe (AS). For each specimen the geographic location and the SVL are given (all specimens are adult males): (a) MCZ 43759,
(b) FMNH 64799, (c) USNM 280250, (d) CM 970. Note especially the differences between the northern specimen of S. prionotus
(a) compared to southern ones (b and c), especially given the size differences among these; and the differences between size-
matched specimens of S. prionotus and S. caducus (b and d).
206 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
notes on the San Juan [del Oro] locality
from which she obtained a series of Sten-
ocercus prionotus in the early 1950s (K. P.
Schmidt archives, FMNH):
Steep forest with deep undergrowth. Steep fields.
Rainfall probably somewhat greater than at Pampa
Grande,^ due to its colder climate and steep ex-
posure; I have no figures. Brushy second growth
may be burned in fairly wide patches in August,
and [the] resort is frequently made to burning, in-
choating a moderately dry winter period.
As of the mid-1980s very little undisturbed
forest was left in the vicinity of San Juan
del Oro (personal observations). Although
Heller provided detailed notes on some of
the snakes and frogs from her collection,
she makes no specific comments about the
lizards.
Most observations suggest that Steno-
cercus prionotus prefers open habitats,
such as areas of human disturbance and
light gaps within forests (e.g., created by
trails), rather than deep rainforests. Alter-
natively, the observations may simply in-
dicate the ease of observation and capture
in more open habitats. A combination of
cryptic coloration and escape behavior
(rapid flight followed by immobility) pos-
sibly makes S. prionotus very difficult to
observe in closed-canopy rainforest, as re-
ported for the similar species, S. fimbria-
tus (Dixon and Soini, 1986; Avila-Pires,
1995) and S. caducus (Scrocchi et al.,
1985).^ However, we currently lack obser-
vations to support these statements for S.
prionotus.
COMPARISON OF STENOCERCUS
PRIONOTUS WITH S. CADUCUS
A scaly flap associated with the posthu-
meral pockets is a unique and unquestion-
ably derived character shared by Stenocer-
cus prionotus and S. caducus, which sug-
gests that these are sister species (Figs. 5,
11).^" I am unaware of a similar structure
in any other lizards. Some individual vari-
ation in the precise form and size of the
flap occurs in both species but it seems
extremely iinprobable that these structures
are not homologous in the two species.
Moreoever, Stenocercus prionotus and
S. caducus are similar in standard meristic
characters (Table 1; Figs. 8, 9), and the
siinilarity among Bolivian populations of
both species has caused confusion about
the identity of particular populations (see
citations in the synonymy of S. prionotus) .
Geographic, ontogenetic, or individual var-
iation of some characters within both spe-
cies, especially the height of the vertebral
crest and the number of midbody scale
rows (Fig. 8), further clouds the distinc-
tions between them. Differentiating the
northern populations of S. prionotus from
S. caducus is unequivocal and facile based
solely on the size of the vertebral crest and
on associated meristic counts. However,
specimens of S. prionotus from southern
Peru and northern Bolivia are more diffi-
cult to distinguish from S. caducus. For
example, animals from populations of S.
prionotus in southern Peru and Bolivia
have less prominent vertebral crests
(hence, higher vertebral scale counts), and
generally lower numbers of midbody scale
rows, than do specimens from northern
populations (Figs. 8, 9); in these respects
they are more similar to S. caducus. Nev-
ertheless, even accounting for these diffi-
culties, a combination of three qualitative
characters is sufficient to distinguish S.
prionotus from S. caducus, and the species
^ I have been unable to localize Pampa Grande.
® Cei (1993) claimed that Stenocercus caducus was
arboreal, but Scrocchi et al. (1985) reported the be-
havior of this species in more detail and stated that
it was terrestrial.
'" In my comparisons I have emphasized Bolivian
specimens referred to Stenocercus caducus, whereas
the type locality is "Paraguay." I have not fully con-
vinced myself that specimens referred to this species
from Bolivia, Paraguay, and Argentina are, in fact, all
the same taxon. Considerable variation exists in some
aspects of coloration and scale characters in diese
specimens. However, my concept of S. caducus cor-
responds to that used in current literature (e.g., Gal-
lardo, 1959; Scrocchi et al, 1985; Cei, 1993). Only a
thorough study of S. caducus across its range will re-
solve this issue.
New Species of Stenocercus from Peru and Bolivia • Codle 207
may differ in patterns of sexual size di-
morphism.^^
The Form of the Vertebral Crest. De-
spite geographic variation in the promi-
nence of the vertebral crest, males and fe-
males of Stenocercus prionotiis have a dis-
tinctly projecting serrate vertebral crest ex-
tending from the nuchal region to the
anterior portion of the tail (Figs. 6, 10, 12).
The scales of the crest are strongly trian-
gular in lateral view, are flaplike (i.e., they
bend easily), project vertically from the
dorsum, and are strongly differentiated
from the adjacent dorsal scales. Although
the crest is somewhat less developed in fe-
males, it is prominent in both sexes. Spec-
imens from northern Bolivia and southern
Peru have a substantially lower crest than
specimens from central and northern Peru
(Fig. 10). Nonetheless, the form and pro-
jection angle of the crest scales is the same
as in the northern populations.
In contrast, the scales of the vertebral
crest in Stenocercus caducus are only mod-
erately differentiated from adjacent dorsal
scales in being more strongly keeled and
mucronate. The crest in S. caducus is only
slightly projecting in males (Fig. 10) and
even less so in females (Fig. 12); the crest
is mainly apparent on the neck and ante-
rior body. In S. caducus, the scales of the
crest are stiff and prismatic, and the main
axis of projection is posterior rather than
vertical, as in S. prionotiis.
Crest height in Stenocercus varies posi-
tively with size and thus it is critical to
compare similar-sized specimens when
documenting differences among popula-
tion samples or species. This realization
has been critical to differentiating Steno-
cercus prionotiis from S. caducus in south-
" I am uncertain how Fugler (1983, 1986, 1989)
distinguished Boh\dan specimens he referred to Sten-
ocercus aculeatus and S. caducus. In 1983 and 1986
he referred specimens from Tumi Chucua (Beni, Bo-
livia) to S. aculeatus. In 1989 he Usted these again,
along with ROM specimens from San Marcos Ranch
(Beni, Bolivia) identified as S. caducus. Fugler spec-
imens from these localities that I have examined are
all S. prionotus (see list of paratypes).
Figure 1 1 . Axillary region of Stenocercus caducus showing
the posthumeral flap (MCZ 34215). Anterior to the left. The
posthumeral flap comprises the heavily outlined scales pos-
teroventral to the forelimb. The opening of the posthumeral
pocket is the heavily stippled area deep to the flap. Top, Pos-
thumeral flap in its normal orientation covering the anterov-
entral portion of the pocket. Bottom, the flap deflected ven-
trally, with its posterior scales viewed from their tips. Approx-
imately X8.5.
ern Peru and Bolivia. Large adult males of
S. prionotus from southern populations
are scarce in collections. For example, al-
though only 18 specimens of S. prionotus
are available from northern Peru, one half
of these are males with SVL >60 mm. In
208 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
contrast, twice as many specimens each of
S. prionotus and S. caducus are available
from southern Peru and Bolivia. Yet, only
one third of the available specimens of ei-
ther species from these areas are males
>60 mm SVL, and no males of S. caducus
were >72 mm SVL.
Figure 10 shows differences in the
height of the vertebral crest in a series of
males of Stenocercus prionotus from
southern Peru and Bolivia compared with
similar-sized specimens of S. prionotus
from northern Peru and with S. caducus
(the largest males of S. caducus studied
were 72 mm SVL; see also Fig. 6). The
trend toward lower crests in S. prionotus
from the southern part of its range is evi-
dent, as is the difference between S. cad-
ucus and S. prionotus. A comparison of all
specimens suggests that the difference in
the height of the vertebral crest between
S. prionotus and S. caducus males begins
to be apparent by approximately 65 mm
SVL and becomes pronounced at around
70-75 mm SVL. No males of S. caducus
>72 mm SVL were among the specimens
examined, despite the availability of a large
number of specimens froin southern Bo-
livia, including a series of 31 specimens
(10 males >60 inm SVL) from the vicinity
of Santa Cruz. A siniilar contrast in crest
height appears in females of the two spe-
cies (Fig. 12).
Angulate Temporal Scales. Stenocercus
prionotus has two (occasionally three) very
strongly keeled, projecting angulate tein-
poral scales on each side (Fig. 4). These
are much larger than adjacent scales on
the head and they are partially or com-
pletely separated from the large posterior
head scales (parietals, postparietals, and
occipital) by one row of small keeled scales
(occasionally partially doubled). Stenocer-
cus caducus usually has two (occasionally
three) angulate temporals that are smaller
and less projecting than those in S. prion-
otus. In S. caducus the angulate temporals
may or may not be larger than adjacent
posterior head scales and they are not
Figure 12. Size-matched females of Stenocercus prionotus
and S. caducus from Bolivia. Top, S. prionotus (USNM
269022, snout-vent length [SVL] 91 mm). Bottom, S. caducus
(UTA 38046, SVL 93 mm). Note the subtle difference in crest
height between the two specimens and their otherwise similar
patterns.
strongly differentiated from other posteri-
or head scales.
Color Pattern of the Gular Region.
Many specimens of Stenocercus prionotus
have a regular pattern of alternating diag-
onal light and dark stripes on the throat.
These usually converge closely toward the
midline (Fig. 7) and are most easily visu-
alized in preseived specimens submerged
in alcohol. This pattern consists of a dark
stripe beginning at a point on the lower
labials in line with, but broader than, the
subocular dark bar. The stripe projects
posteromedially, gradually fading and
blending with the ventral ground color on
the neck anterior to the pectoral region.
The dark stripe is bordered on either side
by a distinct pale stripe. Anteriorly, this se-
ries is preceded by another dark and an-
other pale stripe. The dark stripes are usu-
ally approximately twice as wide as the
pale ones, although not always (e.g., the
dark stripes are only slightly wider than
the pale ones in MCZ 150243). In life the
pattern may manifest itself as a series of
pale stripes on a darker background (e.g.,
the "gular area streaked by several light
New Species of Stenocercus from Peru and Bolivia • Cadle
209
cream colored lines" in the life colors of
the holotype).
The gular region appears uniform in
many preserved specimens of Stenocercus
prionotus, but I suspect this is a preser-
vation artifact. Occasional specimens have
pale spots in the pectoral region, and oth-
ers are essentially unicolor and without ap-
parent pattern (again, probably a preser-
vation artifact).
On the other hand, the throat pattern of
Stenocercus caducus is highly variable and
irregular. When a distinctive pattern is
present, it most often consists of light
spots rather than alternating stripes (Fig.
13). Cope (1862) described the holotype
of S. caducus from Paraguay as having a
dark throat that was "light varied" (i.e.,
variegated, or spotted), and some speci-
mens I examined have this pattern (Fig.
13). None of several color descriptions for
Argentinian specimens of S. caducus men-
tion stripes or spots on the throat. Scrocchi
et al. (1985) described living examples as
having pale spots in parallel transverse
rows in the pectoral region or with pale
spots on the abdomen, but did not com-
ment on the throat pattern; Gallardo
(1959) described the ventral coloration as
"pale olive with some scattered pale spots;
throat darker"; and Cei (1993) described
the venter as "dark brownish with series of
rounded pale spots, sometimes anastomos-
ing along the length of a median line." Al-
though no authors mention alternating
light and dark stripes on the throat in S.
caducus, UTA 38046 does have this pat-
tern (Fig. 13). But in this specimen the
stripes are confined to the lateral edges of
the throat (i.e., do not closely approach the
midline as in S. prionotus). Apart from the
throat pattern, the coloration of S. prion-
otus and S. caducus seems to be veiy sim-
ilar judging from descriptions of S. cadu-
cus in the literature (Gallardo, 1959;
Scrocchi et al, 1985; Cei, 1993).
A Possible Dijference in Sexual Size Di-
morphism. Data presented in Table 1 sug-
gests another contrast between Stenocer-
cus prionotus and S. caducus: S. prionotus
Figure 13. Gular patterns in Stenocercus caducus. Top, typ-
ical throat pattern consisting of light spots on a dark back-
ground (BMNH 1927.8.1.163). Bottom, variant pattern consist-
ing of stripes confined to the lateral portion of the throat (UTA
38046). Connpare to Figure 7.
210 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
is not dimorphic in the maximum sizes at-
tained by males and females, whereas fe-
males of S. caducus apparently attain
about 20 mm greater SVL than males.
However, estimating maximum size is sub-
ject to considerable sampling error so this
distinction should be considered to be only
provisional. Nonetheless, males of S.
prionotus with SVL >80 mm are known
from the northern (USNM 193685) and
southern (USNM 280250, BMNH
98.6.9.4) portions of its range, even though
the three largest males from the largest
population sample (23 specimens in
FMNH from Puno Department, Peru)
had an SVL of 73 mm (this sample includ-
ed six adult females with an SVL of 78-89
mm).
In contrast, a sample of 39 Stenocercus
caducus from Bolivia included 12 adult
males, none of which had an SVL >72
mm; in the total sample of S. caducus iJSJ
= 43) 17 females had an SVL >80 mm
(range 80-93 mm).^- Thus, unless a sys-
tematic collecting bias against males exists,
the different pattern of sexual size dimor-
phism provides another character distin-
guishing S. prionotus and S. caducus. Data
presented in Table 1 suggests that other
species of the " Ophrijoessoides group"
may be size dimoi*phic (S. fimbriatus, S.
hiiancabambae, and ?S. scapularis) or not
(S. aculeatus), and either males (S. huan-
cahambae) or females (S. caducus and S.
fimbriatus) may attain a larger body size.
DISTRIBUTIONS OF STENOCERCUS
PRIONOTUS AND S. CADUCUS IN
EASTERN BOLIVIA
The ranges of Stenocercus prionotus
and S. caducus approach one another in
eastern Bolivia, but do not overlap. ^^ Cur-
rently, the two closest documented locali-
ties are, for S. prionotus, near the junction
of the Rio Madidi and the Rio Beni
(BMNH 98.6.9.4), and for S. caducus,
many specimens from the vicinity of Santa
Cruz de la Sierra (see above discussion for
S. prionotus and Appendix; Fig. 1). Sten-
ocercus caducus is also known from the
Bolivia— Brazil border in the region of the
Serrania de Huanchaca in northern Santa
Cruz Department, Bolivia. ^^
I am aware of no specimens of Steno-
cercus referable to either S. caducus or S.
prionotus between the Rio Beni valley and
roughly a line connecting Santa Cnaz and
the vicinity of Serrania de Huanchaca.
Southeast of the Rio Beni, the central part
of the Beni basin (the Llanos de Mojos) is
characterized by flooded savanna grass-
lands, palm savannas, swamps, and other
habitats that are inundated for significant
portions of the year; terra firme forests are
restricted to somew^hat elevated levees
along rivers (Clapperton, 1993: 196; Han-
agarth, 1993). Stenocercus prionotus or S.
caducus seem unlikely to occur in this area
except possibly in these galleiy forests, al-
though Fugler (1986) reported S. priono-
tus (as Ophryoessoides aculeatus) in sea-
sonally flooded forest during the dry sea-
son.
Stenocercus prionotus and S. caducus
probably are segregated by habitat in Bo-
livia and their distributions may not over-
lap. Stenocercus prionotus is associated
^- These sizes are somewhat larger than those pre-
viously reported (81 mm; Gallardo, 1959; Scrocchi et
al, 1985; Marcus, 1986). Cei (1993) stated that S.
caducus reaches only 75 mm SVL in Argentina. Sexes
were not given for any individual or sexed specimens
in these reports.
'"^ All references to "Ophryoessoides aculeatus" in
Bolivia (e.g., Fugler, 1983, 1986) that I verified have
referred to Stenocercus prionotus. However, given
the general confusion of species in this complex,
some records not traced will have to be checked to
rule out the possibility that they do not refer to S.
caducus or perhaps some other species of the
"Ophryoessoides group," such as S. fimbriatus or S.
scapidaris (see Distribution Patterns in Stenocercus
prionotus).
i-i See the Appendix, UTA 38048. Michael Hawey
(personal communication) recently obtained speci-
mens of Stenocercus caducus at El Refugio, a lowland
locality at the southern end of the Serrania de Huan-
chaca (14°44'S. 61°01'W).
New Species of Stenocercus from Peru and Bolivia • Cadle
211
with upper Amazonian and lower montane
rainforests with annual rainfall greater
than 2,000 mm in both Peru and Bolivia.
On the other hand, confirmed localities of
S. caducus are within the physiographic
domain broadly referred to as chaco, in-
cluding a mixture of diy forests, palm sa-
vannas, galleiy forests, deciduous forests,
and ecotonal areas (Scrocchi et al., 1985;
Marcus, 1986; Cei, 1993). Short (1975)
and Parker et al. (1993) described the di-
versity of chaco habitats. Gallardo (1979:
table 12.1) listed S. caducus as a species
"basically restricted to the chaco." Average
annual rainfall in this area is less than
1,000 mm. Stenocercus caclucus is known
from Parque Nacional Noel Kempff Mer-
cado and vicinity in Bolivia (see footnote
14; Haivey, 1998). This area is character-
ized by a complex mixture of habitat types,
including deciduous forests and cerrado
enclaves, and with an annual rainfall of
1,400-1,500 mm (Killeen, 1998). Hai-vey
(1998) encountered S. caducus at granitic
outcrops covered by semideciduous forests
and more open habitats. The herpetofauna
of this site is a mixture of species that are
typical of Amazonian and of chaco envi-
ronments (Hai^vey, 1998; personal obser-
vations).
The range of Stenocercus caducus ex-
tends outside the strictly defined chaco re-
gion (see Short, 1975, and Clapperton
1993, for discussion) on the southeastern
edge of its range east of the Rio Paraguay
and in the Andean foothills of southern
Bolivia and northern Argentina (Fig. 1).
Harvey (1997) reported S. caducus from
"subtropical wet forests" (1,150—2,050 m
elevation) in southern Bolivia. He charac-
terized S. caducus as a "Chacoan species
that invade[s] the Andean foothills . . . [in-
cluding] those distributed within the Gran
Chaco or that occur in diy forests sur-
rounding the Gran Chaco" (Hai-vey, 1997:
35). The montane wet forests (yungas) of
this area are restricted to ridges high
enough for cloud formation during much
of the year (generally > 1,500 m elevation),
and they are surrounded by deciduous dry
forest (Schulenberg et al., 1997). The cli-
mate of this area is generally dry and it
receives only about 1,200 mm of rainfall
per year (Hoist, 1997).
The transition between the wet rainfo-
rests of Peru and northern Bolivia (range
of Stenocercus prionotus) and the chaco
habitats (range of S. caducus) occurs in a
very broad ecotone consisting of savannas,
evergreen shrublands, and gallery forests
of the Beni basin and Rio Mamore drain-
age, from which no specimens of either S.
prionotus or S. caducus have been report-
ed. The piedmont forests of the Andes be-
tween the known ranges of S. prionotus
and S. caducus, which are wetter than ad-
jacent lowland forests because of the mod-
erating effect of the Andes, provide one
potential route for contact or overlap of
their ranges.
The eastern distributional limits of Sten-
ocercus caducus along the Bolivia— Brazil
frontier are not well understood. I am un-
aware of verified records from Brazil, al-
though the species does occur close to the
Brazilian border in the vicinity of the Ser-
rania de Huanchaca in Parque Nacional
Noel Kempff Mercado. Some references
to "Stenocercus caducus" from western
Brazil (e.g., Mato Grosso State; Cope,
1887; Boulenger, 1903) likely refer instead
to an undescribed species veiy similar to
S. caducus (P. E. Vanzolini and E. E. Wil-
liams, personal communication; personal
observations). However, the ranges of S.
caducus and the undescribed species in
eastern Bolivia-Paraguay and western Bra-
zil are not well defined; the two species
may be separated by the seasonally inun-
dated savannas of the pantanal. Addition-
ally, few specimens of S. caducus appar-
ently exist from the chaco of northwestern
Paraguay, although Aquino et al. (1996) re-
ported specimens from Parque Nacional
Defensores del Chaco (approximately
20°30'S, 60°20'W), as well as other Para-
guayan localities in more mesic regions
east of the Rio Paraguay.
212 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
IS THE DISTRIBUTION OF
STENOCERCUS ACULEATUS
DISJUNCT?
In the process of diagnosing Stenocercus
prionotus I reviewed the characters and
distribution of S. aculeatus. In addition to
variation in some characters of uncertain
significance, some aspects of the distribu-
tion of S. aculeatus seem pecuHar (Fig. 2).
First, the distribution of S. aculeatus ap-
pears to be disjunct between northern
Peru and eastern Ecuador. Although the
type locality is in northern Peru (Moya-
bamba, San Martin Department), most
specimens are from eastern Ecuador (Fig.
2). The two areas from which specimens
are known (Fig. 2) are separated by a
broad geographic gap through which
courses the main tributary of the upper
Amazon, the Rio Marafion. Neither S. acu-
leatus nor any other species of Stenocercus
was obtained during hei"petofaunal surveys
in northern Loreto Department, Peru
(Duellman and Mendelson, 1995), north-
ern Amazonas Department, Peru (Rio Ce-
nepa and Rio Santiago; J. E. Cadle and R.
W. McDiarmid, unpublished data), or dur-
ing a rapid biological assessment of the
Cordillera del Condor region of southeast-
ern Ecuador and northern Peru (Schulen-
berg and Awbrey, 1997a). Stenocercus acu-
leatus is known from many localities in ad-
jacent regions of Ecuador.
Second, all Ecuadorian localities for
Stenocercus aculeatus are in the drainages
of the Rio Pastaza and the Rio Curaray.
No specimens are known from the Rio
Napo drainage just to the north, even
though no recognized physiographic or
faunal break seems to separate the Rio
Napo drainage from the Rio Curaray
drainage. However, all localities that have
been sampled comprehensively from the
Rio Napo are on the left (northern) bank
of the river (e.g., Duellman, 1978; Vitt and
De la Torre, 1996; unpublished list from a
large collection from the Jatun Sacha Bi-
ological Station assembled and under
study by Gregoiy Vigle). The absence of
S. aculeatus from Santa Cecilia (Duell-
man, 1978) is probably real rather than
sampling error, given the intensity of col-
lecting over several years at the site. Thus,
S. aculeatus possibly does occur on the
right (south) bank of the Rio Napo and
will be recorded once large collections are
made there.
The apparent geographic disjunction of
Stenocercus aculeatus between northern
Peru and eastern Ecuador may correspond
to some character differences among sam-
ples that should be studied more thor-
oughly (Cadle, unpublished data). For ex-
ample, Peruvian specimens of S. aculeatus
have veiy deep postfemoral pockets (Type
5) in both sexes, whereas the postfemoral
pockets are more weakly developed in
specimens from Ecuador (Type 2 or 3 in
both sexes). Ecuadorian specimens also
appear to have more scales in the vertebral
row and fewer subdigital scales on the
fourth toe than do Peruvian specimens. All
of these impressions are based on small
sample sizes (Appendix).
The significance of diese differences is
unclear without a more detailed study of
variation among populations of Stenocer-
cus aculeatus. However, one possibility is
that two or more species are represented
in specimens currently referred to S. acu-
leatus, in which case the distributions of
individual taxa may be not be contiguous.
This is analogous to the previous confusion
of S. finihriatus and S. prionotus with S.
aculeatus. Taxonomic recognition of S.
finihriatus and S. prionotus has concomi-
tantly reduced the geographic distribution
understood for S. aculeatus. Consequently,
a more comprehensive systematic analysis
of S. aculeatus with special reference to a
comparison of Ecuadorian and Peruvian
populations is warranted. If two species
are recognized, the name Liocephalus an-
gulifer Werner (1901) is available for the^
Ecuadorian populations.
Key to Species of the "Ophryoessoides
group" of Stenocercus
Because of the general confusion about
the species considered herein (e.g., see the
New Species of Stenocercus from Peru and Bolivia • Codle 213
synonymy of Stenocercus prionohis) , I pro-
vide the following key as a guide for iden-
tifications. The key will work for those spe-
cies of Stenocercus in Peru or Bolivia with
keeled ventral scales, enlarged posterior
head scales, and one row of moderately to
greatly enlarged supraoculars COphryoes-
soides group" as used herein). I have also
included the three other currently recog-
nized species having these characteristics,
S. enjthrogaster (Hallowell), S. dumerilii
(Steindachner), and S. tricristatus (Du-
meril), although these are not known from
Peru or Bolivia and are unlikely to occur
there. Character and distributional data in
the key for S. dunierilii and S. tricristatus
follow Avila-Pires (1995).
I also include in the key an undescribed
species with keeled ventrals and enlarged
head plates and supraoculars from Ama-
zonas Department, Peru, but I am un-
aware of other undescribed species of the
"Ophrijoessoides group" from Peru or Bo-
livia. However, an undescribed species
similar to Stenocercus caducus (but lacking
a posthumeral flap) is known from western
Brazil (Mato Grosso) and is not included
in the key. Additional study of S. iridescens
from the Pacfic lowlands of Peru and Ec-
uador is needed (Cadle, 1998: footnote 4)
and, as indicated above, a thorough mod-
ern study of variation in S. aculeatus (Am-
azonian Ecuador and Peru) is also war-
ranted. Other undescribed species may re-
side within either of these named taxa.
The key Mdll permit identification of all Pe-
ruvian and Bolivian taxa previously con-
fused with S. aculeatus (e.g., DLxon and
Soini, 1975, 1986 [S. Jimbriatus]; Fugler,
1983, 1986, 1989 [S. prionotus]) and S. ir-
idescens (e.g., S. huancahambae and S.
limitaris; see Cadle, 1991, 1998). The key
also should work for Ecuadorian species,
with the caveat that I have paid less atten-
tion to Ecuadorian Stenocercus except as
necessary in conjunction with work on Pe-
ruvian species. Of the species covered,
only S. acideatus, S. iridescens, and S. lim-
itaris are definitely known from Ecuador.
The key should be viewed as a means of
identifying a set of phenotypically similar,
but not necessarily closely related, species
within Stenocercus in the broad sense. All
other species of Stenocercus in Peru and
Bolivia have smooth (or at most only very
weakly keeled) ventrals and more frag-
mented supraoculars and head plates; see
Fritts (1974), Frost (1992), and Cadle
(1991, 1998) for discussion and illustra-
tions. Many of these species also have
granular scales on the body or posterior
surface of the thigh, neither of which is
present in species covered by the key. The
keys and discussions in Fritts (1974) and
Cadle (1991, 1998) are useful for identi-
fying these other species.
The key assumes familiarity with char-
acters of the mite pockets, head scales, and
neck folds and crests outlined in Cadle
(1991) (see also Materials and Methods).
In most cases I have used characters that
show minimal sexual dimoqDhism so that
specimens of either sex can be identified;
exceptions are noted. It is useful to keep
in mind that, in most species of Stenocer-
cus, scales of juveniles are more promi-
nently keeled than in adults, even when
the corresponding scales of adults, such as
head scales and dorsal body scales, are
smooth. Instances of possible confusion in
the key are indicated. The extent of de-
velopment of posthumeral and postfemor-
al mite pockets varies according to sex and
size in many species of Stenocercus, al-
though such variation seems less extensive
in this set of species than in many others;
I have indicated the range of variation in-
cluding juveniles and adults of both sexes
in the key. Summaiy geographic distribu-
tions are given for each species as a rough
guide to known occurrences. However,
these should be used cautiously as ancil-
laiy information in identifying specimens
because distributions of species are some-
times poorly circumscribed. For greatest
utilit}^ the key should be used in conjunc-
tion with illustrations herein and in Cadle
(1991, 1998) and Avila-Pires (1995).
214 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
1. Canthal and supraciliaries forming a pro-
nounced crest that ends in an enlarged,
erect, postsupraciliaiy that may be dis-
tinctly pointed or blunt. Posthumeral and
postfemoral mite pockets absent (Type 1
in both instances) 2
Cantlials and supraciliaries not forming a
pronounced crest; no enlarged, erect post-
supraciliaiy. Posthumeral pocket absent
(Type 1) to deep (Type 4). Postfemoral
pocket absent (Type 1) to deep (Type 5)
3
2. Enlarged postsupraciliary distinctly pointed.
Two enlarged scales above ear opening.
Tibia approximately equal to thigh length
Stenocercus dumerilii (Steindachner)
(northeastern Para, Brazil)
Enlarged postsupraciliary blunt. No enlarged
scales above ear opening; tibia distinctly
shorter than thigh
Stenocercus tricristatus (Dumeril)
(known only from the holotype, probably
from the state of Minas Gerais, Brazil)
3. Superciliary scales projecting laterally shelf-
like above the orbit in adults, rectangular
in dorsal view.'^' Midbody dorsal scale rows
more than 55 (59-70). Postfemoral pocket
well developed (Type 3 or 5)
Stenocercus scapularis (Boulenger)
(intermediate elevations on the Andean
slopes of central and southern Peru; known
elevations greater than 1,000 m)
Superciliaiy scales not projecting laterally,
the anterior ones elongate, strongly over-
lapping. Midbody dorsal scale rows fewer
than 55 (30—53). Postfemoral pocket vari-
able (Type 1, 2, 3, or 5) 4
4. Posthumeral mite pocket deep (TjqDe 4) and
with an associated scaly flap extending
from its anteroventral border and partally
concealing it 5
Posthumeral mite pocket absent to deep
(Type 1, 2, 3, or 4) but without an asso-
ciated flap 6
5. Vertebral crest strongly projecting in both
sexes, serrate, extending from the nuchal
region to the proximal portion of the tail;
its individual scales triangular, flaplike.
Throat often with alternating oblique dark
and light stripes. Two enlarged, strongly
keeled and projecting angulate temporal
'' The superciliaries in juveniles of Stenocercus sca-
pularis have a more typical shape. The extent of
shelf-like projection and change to a more rectan-
gular shape seem positively correlated with body size
and thus develop with age. The number of dorsal
scale rows and the development of the postfemoral
pocket are useful clues for subadults.
scales on each side
Stenocercus prionotiis Cadle
(rainforested lowlands and Andean foothills
of eastern Peru and northern Bolivia)
Vertebral crest low, nonprojecting in both
sexes (slightly higher in males), evident
mainly on neck and anterior body; its in-
dividual scales prismatic and lying more or
less flat. Throat pattern variable, but usu-
ally consisting of light spots on a darker
background when evident. Angulate tem-
poral scales keeled, but not greatly en-
larged, and nonprojecting
Stenocercus caducus (Cope)
(deciduous woodlands and ecotonal areas of
southern Bolivia, northern Argentina, and
the chaco of Paraguay)
6. A fringe of enlarged fimbriate scales on the
distal posterodorsal surface of thigh. Sev-
eral longitudinally oblique rows of large,
strongly keeled scales on shank.''' Scales
between vertebral and dorsolateral crests
smooth or occasionally veiy weakly keeled.
.-. Stenocercus finibriatus Avila-Pires
(lowlands of eastern Peru and central western
Amazonian Brazil)
No fringe of fimbriate scales on thigh or
strongly keeled oblique scales on shank.
Scales between vertebral and dorsolateral
crests moderately to strongly keeled at
least posteriorly; dorsolateral crest may be
weakly developed, but dorsal scales still
strongly keeled 7
7. Posthumeral mite pocket variable (Type 1, 2,
3, or 4). Postfemoral mite pocket variable
(Type 1, 2, 3, or 5). Head scales smooth
or keeled. Angulate temporal scales
keeled, may be projecting and bladelike.
Internasals usually 4 or more (occasionally
3, never 2), often irregular in pattern and
shape. 8
Posthumeral mite pocket absent or weakly
developed (Type 1 or 2). Postfemoral mite
pocket absent (Type 1). Head scales
smooth. Angulate temporal scales smooth;
none bladelike and projecting. Two polyg-
onal internasals in contact on the midline,
each broader laterally than medially.
Stenocercus iridescens (Giinther)
(Pacific lowlands and intermediate elevations
of western Ecuador and northwestern
Peru)
8. One to 3 strongly keeled, but nonprojecting,
angulate temporal scales in line with the
superciliary row between the lateral tem-
"^ The fimbriate scales form a projecting fringe on
the distal portion of the thigh. Both the fimbriate
scales and the oblique scales on the shank are rela-
tively more prominent in juveniles than adults.
New Species of Stenocercus from Peru and Bolivia • Cadle 215
porals and the posterior dorsal head scales.
Two subequal can thai scales on each side.
Head scales keeled, at least posteriorly 9
Two projecting bladelike angulate temporals
in line with superciliary row. A single can-
thai on each side (rarely, 2 are present but
in that case 1 is much larger than the oth-
er). Head scales smooth or keeled. 11
9. Posthumeral pocket moderately developed
in males (Type 2 or 3), absent in females
(Type 1). Postfemoral pocket absent in fe-
males (Type 1), moderate to deep in males
(Type 3 or 5). Anterior gular scales weakly
to strongly keeled.
Stenocercus erythrogaster (Hallowell)
(northern Colombia)
Posthumeral and postfemoral pockets deep
in both sexes (Types 4 and 5, respective-
ly).'' Anterior gular scales smooth to weak-
ly keeled. 10
10. Inteiparietal indistinct, parietal eye not visi-
ble. Three occipitals. Dark subocular bar
absent. Three angulate temporals separat-
ed from large posterior head scales by a
row of tiny scales Stenocercus new species
(known from a single specimen [Appendix]
from the inter-Andean valley of the Rio
Maranon near Balsas, Amazonas Depart-
ment, Peru)
Inteiparietal distinct, parietal eye visible.
Two occipitals. Dark subocular bar pre-
sent. One angulate temporal much larger
than others and in contact with at least 1
other enlarged posterior head scale.
Stenocercus liniitaris Cadle
(intermediate elevations [600—2,200 m] of the
Andes on the Pacific versant of southwest-
ern Ecuador and northwestern Peru)
11. Head scales smooth to slightly wrinkled in
adults; weakly keeled, wrinkled, or rugose
in juveniles. Prominent dorsolateral crest
on body from neck to base of tail and con-
tinuous with both supra-auricular crest
and antehumeral crest. Postfemoral pock-
et moderate to deep (Type 2, 3, or 5).
Stenocercus aculeatus (O'Shaughnessy)
(rainforested lowlands and intermediate ele-
vations of northern Peru adjacent to the
Andes and in eastern Ecuador)
Head scales strongly keeled or multicarinate
in juveniles and adults. Dorsolateral crest,
when present, weak and restricted to neck
and anterior body. Postfemoral pocket
deep (Type 5).
Stenocercus huancahanibae Cadle
'' An undescribed species in the ne.xt couplet of the
key is known only from a single adult male. The dis-
tributions of species in couplets 9 and 10 should be
used as ancillaiy data for identification.
(diy inter-Andean valleys of the upper Rio
Maraiion in Cajamarca and west central
Amazonas departments, northern Peru)
ACKNOWLEDGMENTS
Loans and other assistance were facili-
tated by Linda Ford, Barrel Frost, and
Charles W. Myers (AMNH); E. Nicholas
Arnold and Colin J. McCarthy (BMNH);
John Wiens (CM); Cassy Redhed, Alan
Resetar, and Harold Voris (FMNH); Wil-
liam E. Duellman, Christopher J. Raxwor-
thy, and John E. Simmons (KU); Frank
Burbrink and Douglas Rossman
(LSUMNS); Ross MacCulloch and Robert
W. Murphy (ROM); Roy W. McDiarmid,
Steven W. Gotte, W. Ronald Heyer, and
Robert R Reynolds (USNM); and Jona-
than Campbell and Michael B. Harvey
(UTA). Victor Morales permitted me to ex-
amine a specimen of Stenocercus fimbria-
tus in his care. I am grateful to Wade C.
Sherbrooke for providing copies of his
field notes and other information on spec-
imens he collected. I owe a great debt to
the late Ernest E. Williams, who was ex-
tremely generous with discussion, notes, il-
lustrations, and encouragement. Williams,
Paulo E. Vanzolini, and Richard Etheridge
long ago distinguished Stenocercus prion-
otus and two other species I described (S.
huancabainbae and S. liniitaris) but kindly
let my work on the group unfold with their
gracious consent. Vanzolini supplied a
copy of a portion of Balzan (1931), helped
interpret Balzan's localities, and pointed
out Metraux's work to me. I am indebted
to several people for the special efforts
they made in tracking down information
about particular collections: Bruce Patter-
son (FMNH) supplied information on the
collections of Colin Sanborn and Hilda
Heller from Puno Department, Peru; Alan
Resetar and Cassy Redhed (FMNH) dug
into the Schmidt archives and found ad-
ditional information about Hellers collec-
tion; Charles W. Myers (AMNH) did the
same for Haivey Basslers journeys in
northern Peru and provided the base map
used to prepare Figure 1; Robert S. Voss
216 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
checked AMNH sources for information
on Keays's Peruvian localities; and Colin J.
McCarthy (BMNH) clarified the confusion
about P. O. Simons's "Palca" locality, if not
the locality itself. Robin Andrews, Tom
Jenssen, and A. Stanley Rand discussed as-
pects of geographic variation and the uses
of vertebral crests in S. prionotus with me.
Laszlo Meszoly drew Figures 4, 5, and 11.
For comments on the manuscript I thank
Richard Etheridge and Charles W. Myers.
The research was supported in part by a
faculty grant from the School of Arts and
Sciences of Harvard University; publica-
tion costs were supported by the Colles
Fund of the MCZ.
APPENDIX: SPECIMENS EXAMINED
Institutional abbreviations are as fol-
lows:
AMNH American Museum of Natural
History, New York
ANSP Academy of Natural Sciences
of Philadelphia
BMNH The Natural History Muse-
um, London
CM Carnegie Museum of Natural
History, Pittsburgh, Pennsyl-
vania
FMNH The Field Museum, Chicago
KU Natural History Museum,
University of Kansas, Law-
rence
LSUMNS Louisiana State University
Museum of Natural Science,
Baton Rouge
MCZ Museum of Comparative Zo-
ology, Harvard University,
Cambridge
ROM Royal Ontario Museum, To-
ronto
USNM National Museum of Natural
History, Washington, D.C.
UTA University of Texas at Arling-
ton
Bracketed information was inferred
from sources listed in the Materials and
Methods. For specimens of other species
of the "Ophryoessoides group" examined
(S. huancabamhae, S. iridescens, and S.
limitaris), see Cadle (1991, 1998). Bold-
face numbered localities 1—4 are known or
suspected areas of sympatry between the
species listed and Stenocercus prionotus.
They correspond to numbered localities in
Figures 1 and 2 and in the text discussion.
Stenocercus aculeatus
ECUADOR: Prov. Morona-Santiago: Chiguaza
[ca. 1,000 m; 01°59'S, 77°58'W] (USNM 200882-84).
[?Prov. Napo/Tungurahua]: Llanganates area'**
(FMNH 2.3527). Prov. Pastaza: Rio Pastaza, Abita-
gua [01°23'S, 78°05'W] (FMNH 2.5803-05, 26892,
28011, 28057 [ = 17 specimens]). Rio Pastaza, Alpay-
acu [01°28'S, 78°07'W] (FMNH 3926-27; MCZ
8081). Canelos [01°35'S, 77°45'W; 530 m] (MCZ
38530). Montalvo, Rio Robonaza [314 m; 02°04'S,
76°58'W] (USNM 200892). [?Prov. Pastaza]: Ranos,
Mera Trail [? = between Raiios and Mera'^; approx-
imately 01°30'S, 78°10'W] (FMNH 28012).
PERU: [Depto. La Libertad]: E Peru, Pampa
Seca, Rio MLxiolla [ = Rio Mishollo] Valley, Upper
Huallaga region, 4300 ft [2, 1,311 m; approximately
08°16'S, 76°58'W]2« (AMNH 57085). [Depto. Lor-
'' I have been unable to localize this. The Cordil-
lera de los Llanganates is a high range (to >4,500 m)
in the Cordillera Oriental north of the Rio Pastaza
(Paynter, 199.3). The locality may refer to lower ele-
vations in this range.
"^ Rafios is a famous collecting locality at the foot
of Volcan Tunguraliua at 1,820 m. That would be an
altitudinal record for Stenocercus aculeatus. I inter-
pret the locality as stated in the FMNH catalogues
as being on the trail between Raiios and Mera, which
is at 1,160 m. See Rrown (1941) and Chapman (1926)
for discussion.
-" Hai"vey Rassler collected Stenocercus for the
AMNH at two localities on the Rio Mixiolla ( = Rio
Mishollo): Pampa Seca and La Pinita (see Stenocer-
cus finibriatus), as listed in AMNH catalogues. The
Rio Mishollo originates in southeastern La Libertad
Department, flows eastward, and joins the Rio Hual-
laga in southwestern San Martin Department. The
elevations given for these localities, 1,067 m and
1,311 m, indicate that they lie in the narrow stretch
of the valley that straddles the boundary between La
Libertad and San Martin departments (departmental
maps produced by the Instituto Geografico Nacional,
Lima). I identify these localities as Pampaseca and
Piiiita, respectively, in extreme southeastern La Lib-
ertad Department, as indexed by Stiglich (1922).
Roth localities are in Ongon District and the coor-
dinates given are those for the town of Ongon. Stig-
lich (1922) states that Piiiita is a small village on the
Quebrada Pedernal, a left tributary of the Rfo Mish-
ollo. Apart from indicating that Pampaseca is a farm.
New Species of Stenocercus from Peru and Bolivia • Cadle 217
ETo]: NE Peru, Front Range between Moyabamba
and Cahuapanas. 3000 ft [915 m; approximately
05°37'S, 77°00'W] (AMNH 57083). Northeastern
Peru: Icuta on Balsapuerto-Moyabamba trail, 3500 ft
[1,067 m; 05°58'S, 76°40'W; given as "Icuto" or "len-
to Cuesta" by Lamas, 1976] (AMNH 56413).
Stenocercus caducus
BOLIVIA: No specific locality: BMNH
1946.8.29.76 (holotype of Leiocephahis bolivianus
Boulenger); CM 4583-84. Depto. Chuquisaca: Sud
Cinti, trail from Rinconada Bufete to El Palmar
[1,170-2,000 m; approximately 20°50'S, 64°21'W]
(UTA 39102). Depto. Santa Cruz: Buena Vista, ca.
500 m [17°27'S, 63°40'W] (MCZ 20625-26, 29023;
FMNH 16165, 21486, 21511; BMNH 1927.8.1.163-
164; CM 4527, 4550-51, 4558, 4587-88, 4605, 4607,
4616, 4626, 4634-36, 4641). Las Yuntas [ = Las Jun-
tas;-' 18°38'S, 63°08'W] (CM 970). Provincia Chiqui-
tos, Santiago (Serrania and nearby), 700-750 m
[18°19'S, 59°34'W] (FMNH 195983). [Provincial
Chiquitos, Canton El Cerro, Finca Dos Milanos,
17°27'30"S, 62°20'00"W (UTA 38046). Provincia Sara,
eastern Bolivia, 600 m [17°27'S, 63°40'W] (BMNH
1907.10.31.7-8). Provincia Sara, Santa Cruz de la Si-
erra [17°48'S, 63°10'W] (CM 966, 969, 13018). Prov-
incia Sara, Rio Surutu W of Buena Vista [17°24'S,
63°51'W] (CM 4590). Provincia Sara, Rio Colorado
[17°38'S, 63°54'W] (CM 4598). [Provincia] Velasco,
Inselbergs near Florida [14°38'S, 6ri5'W] (UTA
38048). [Depto. Takija]: Villa Monies [21°15'S,
63°30'W] (MCZ 28634). Misidn San Francisco
[21°15'S, 63°30'W] (BMNH 98.7.7.5; specimen col-
lected by Alfredo Borelli, whose San Francisco = Vil-
la Monies fide Paynter, 1992).
PARAGUAY: [Depto. Caaguazu]: Pastoreo [ap-
proximately 25°23'S, 55°52'W] (MCZ 34214-15).
[Depto. Central]: Asuncion [25°16'S, 57°40'W]
(FMNH 9496). Colonia Nueva Italia [25°37'S,
57°30'W] (FMNH 42281).
Stenocercus erythrogaster
COLOMBIA: [Depto. Magdalena]: Rio Frio
[30-450 m; 10°55'N, 74°10'W] (MCZ 29707). Santa
Malta Mountains [approximately 10°50'N, 73°40'W]
(MCZ 11303). Rio Toribio, Hacienda "Papare," sec-
ond river on road from Cienaga to Santa Marta
[11°03'N, 74°14'W] (FMNH 165153). [Depto. San-
tander]: San Gil [1,095 m; 06°33'N, 73°08'W]
(ANSP 24136, MCZ 36877).
Stiglich (1922) gives no further information about its
location.
-' The specimen was collected in November or De-
cember 1913 by Jose Steinbach, who collected at a
locality known as "Las Juntas" during that same pe-
riod (Paynter, 1992). The two locahties are assumed
to be the same. The variant spelling "Yuntas" does
not appear in any sources consulted.
Stenocercus fimbriatus
PERU: No specific locality: (FMNH 56070).
Depto. Huanuco: ca. 35 km NE Tingo Maria, Hcda.
Santa Elena, ca. 1000 m [approximately 08°57'S,
76°02'W] (LSUMNS 26966-67). Approximately Vi
mile E Universidad Agraria de La Selva, Tingo Maria,
vicinity of Rio Huallaga [3, 09°18'S, 75°59'W],
USNM 193684. [Depto. La Lihertad]: E Peru, La
Pinita, Rio Mixiolla, tributaiy of upper [Rio] Hualla-
ga, 3500 ft [1,067 m; approximately 08°16'S, 76°58'W;
see footnote 20] (AMNH 56797-98). [Depto. Lor-
ETO]: E Peru, Contamana, Ucayali River valley [134
m; 07°15'S, 74°54'W] (AMNH 56803). E Peru, E of
Contamana on trail to Contaya, 700 ft [213 m; ap-
proximately 07°15'S, 74°54'W] (AMNH 56781-82). E
Peru, Pampa Hermosa, mouth of Rio Cushabatay,
500 ft [152 m] [1, 07°12'S, 75°17'W] (AMNH 56788,
56790-92, 56794-96, 56801-02). Mishana, Rio Na-
nay, Estacion Biologica Cauicebus, 150 m [03°53'S,
73°27'W] (USNM "222377). Mishuana [ = Mishana;
150 m, 03°53'S, 73°27'W] (KU 212628). Depto. Ma-
DREDE Dios: Pakitza Station [Rio Manu], Manu Na-
tional Park [4, 11°56'S, 71°17'W] (Victor R. Morales
18235). Depto. Ucayali: Rio Curanja, Balta, ap-
proximately 300 m [approximately 10°08'S, 71°13'W]
(LSUMNS 17519, 25402-04, 26720-23). Alto [Rio]
Purris, Alto [Rfo] Curanja, Igarape Champuiaco
[9°34'S, 70°36'W] (MCZ 61226)" Peru/Brazil frontier,
Utoquinia Region, 1000 ft. [305 m; approximately
08°00'S, 74°00'W]22 (AMNH 56789, 56799-800).
Stenocercus scapulari^^
PERU: No specific locality: (FMNH 56444).
[Depto. Junin]: Chanchamayo, 1200 m [approxi-
mately 11°03'S, 75°47'W] (FMNH 40608-11). Pere-
ne, 1200 m [10°58'S, 75°13'W] (MCZ 49580-81).
Tarma, Chanchamayo, 1300 m [11°25'S, 75°42'W]
(FMNH 45522). [Depto. Puno]: Sagrario, Rio Qui-
tun [approximately 1,020 m; 13°55'S, 69°41'W]
-- The region referred to is north to northeast of
Pucallpa. The variant spellings Utoquinia, Utoquinea,
and Uroquinea are in the literature and are applied
to a right-bank tributary of the Rio Ucayali, a village
on the Rio Ucayali, and an airstrip on the Rio Uto-
quinia near the Brazilian border. The entire region is
less than 500 m in elevation except for a small raised
area near the Brazilian border that attains nearly 800
m and that is apparently the source of the Rio Uto-
quinia.
^^ The occurrence of Stenocercus scapularis at Rur-
renabaque. El Beni Department, Bolivia, as reported
for two specimens in the AMNH (Burt and Burt,
1931: 273) is apparently based on a misidentification.
These specimens are probably either S. prionotus
(most likely) or S. caducus (see Distribution Patterns
in Stenocercus prionotus).
218 Bulletin Museum of Comparative Zoology, Vol. 157, No. 3
(FMNH 40408). "Camp 4" [between Santo Domingo
and La Pampa; approximately 13°44'S, 69°37'W]-^
(FMNH 40409). Juliaca, Lake Aracona, 16,600 ft.
[shipping point only; correct locality is on the right
laank of the Rio Inambari, 1,830 m, 13°30'S,
70°00'W]^'^ (AMNH 1701).
-^ According to the field catalogue in the FMNH
Mammal Di\dsion the collector, Colin Sanborn, was
in Santo Domingo on 20 October 1941 and in La
Pampa on 23 October (see also notes in Stephens and
Traylor, 1983). The specimen FMNH 40409 was col-
lected 21 October, and thus "Camp 4" is assumed to
be between these points.
-''The specimen was collected in 1900 by H. H.
Keays, who collected many mammals and other ver-
tebrates in southern Peru, primarily for the American
Museum of Natural History. It is clear that most of
the specimens labeled with the locality "Juliaca" (a
town on the Peruvian altiplano near Lake Titicaca)
actually came from farther north in the Rio Inambari
valley. Allen (1900: 219; 1901: 41) provides the fol-
lowing information:
The Museum has recently received two small col-
lections of mammals made by Mr. H. H. Keays, at
Juliaca, in southeatern Peru, a little to the west-
ward of Lake Titicaca. Mr Keays writes: "Our
camp is situated in the loop of the Inambaiy River.
The countiy is very broken, with deep narrow can-
ons, and is covered with a dense undergrowth oi
shrubs and vines, with here or there a palmetto or
a cedar rising above the surrounding vegetation."
He gives the" altitude as 6000 feet [1,830 m], and
the position as latitude 13°30' S., longitude 70° W.
... it is necessary to correct a misleading statement
in my former paper in respect to the locality where
the . . . collections were made. Mr. Keays's post-
office address was Juliaca, and through lack of ex-
plicit information, it was inferred that the Inca
Mines, where he collected, were in the immediate
vicinity of Juhaca . . . the Inca Mines are situated
about 200 miles northeast of JuHaca, on the east
side of the Andes, on the Inambaiy River, a trib-
utaiy of the Amazon, and at a much lower altitude
than Juliaca. The altitude and geographical position
were correctly given in the former paper, but in
place of Juliaca, . . . read Inca Mines.
Keays's information quoted by Allen places the local-
ity on the right bank of the Rio Inambari in the foot-
hills of an outlying Andean spur separating the Rio
Inambari from upper tributaries of the Rio Tambo-
pata. I have not located a Lake Aracona and suspect
that this is an error for Lake Aricoma, a high Andean
lake on tlie route between Juliaca and the location of
Keays's camp. However, it is not at all clear why this
name is associated with the locality. No notes or cor-
respondence of Keays are in the AMNH mammal
department archives for further clarification (R. S.
Stenocercus sp.
PERU: Depto. Amazonas: 17 km ENE Balsas
[06°49'S, 7S°00'W] (ROM 16458).
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OF THE
seum
(US ISSN 0027-4100)
On the Subfamily Xylophagainae
(Family Pholadidae, Bivalvia, Mollusca)
RUTH D.TURNER
MCZ
'.IBRARY
JAN 2 7 2003
HARVARD
UNIVERSITY
HARVARD university
CAMBRIDGE, MASSACHUSETTS, U.S.A.
VOLUME 157, NUMBER 4
31 October 2002
(US ISSN 0027-4100)
PUBLICATIONS ISSUED
OR DISTRIBUTED BY THE
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Breviora 1952-
bulletin 1863-
Memoirs 1865-1938
JoHNSONiA, Department of Mollusks, 1941-1974
Occasional Papers on Mollusks, 1945-
SPECIAL PUBLICATIONS.
1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and
Evolution of Crustacea. 192 pp.
2. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere-
dinidea (Mollusca: Bivalvia). 265 pp.
3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino-
derms. 284 pp.
4. Eaton, R. J., 1974. A Flora of Concord from Thoreau's Time to the
Present Day. 236 pp.
5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology
and Evolutionary Biology: Essays in Honor of Ernest E. Williams.
725 pp.
6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp.
Other Publications.
Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine.
Reprinted 1964.
Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of
Insects. {Bulletin of the M. C. Z, Vol. 108.) Reprinted 1971.
Creighton, W. S., 1950. The Ants of North America. Reprinted 1966.
Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In-
ternational Symposium on Natural Mammalian Hibernation. (Bulletin
of the M. C. Z, Vol 124.)
Orinthological Gazetteers of the Neotropics (1975-).
Peter's Check-list of Birds of the World, vols. 1-16.
Proceedings of the New England Zoological Club 1899-1947. (Complete
sets only.)
Proceedings of the Boston Society of Natural Histoiy.
Price list and catalog of MCZ publications may be obtained from Publica-
tions Office, Museum of Comparative Zoology, Harvard University, Cambridge,
Massachusetts 02138, U.S.A.
This publication has been printed on acid-free pennanent paper stock.
© The President and Fellows of Harvard College 2002.
ON THE SUBFAMILY XYLOPHAGAINAE
(FAMILY PHOLADIDAE, BIVALVIA, MOLLUSCA)
RUTH D. TURNERS
CONTENTS
Editorial Note 223
Abstract 223
Introduction 224
Species Groups in the Genus Xylophaga 225
Variation 227
Preliminaiy Report on the Results of
Experiments on the Ecology of Deep-
Sea Wood Borers and the Role of Wood
in the Deep Sea 234
Systematic Account 236
Genus Xylophaga Turton 1822 236
Xylophaga concava Knudsen 236
Xylophaga gerda Turner new species 237
Xylophaga grevei Knudsen 238
Xylophaga clenchi Turner and Culliney ... 239
Xylophaga supplicata Tald and Habe 241
Xylophaga whoi Turner new species 242
Xylophaga profunda Turner new species 243
Xylophaga ahyssonuu Dall 245
Xylophaga dupUcata Knudsen 247
Xylophaga muraokai Turner new species 247
Xylophaga atlantica Richards 249
Xylophaga washingtona Bartsch 250
Xylophaga rikuzenica Tald and Habe 252
Xylophaga depalmai Turner new species 253
Xylophaga guineensis Knudsen 256
Xylophaga niexicana Dall 257
Xylophaga tipperi Turner new species 259
Xylophaga hayeri Turner new species 260
Xylophaga japonica Tald and Habe 261
Genus Xylopholas Turner 1972 262
Xylopholas altenai Turner 263
Genus Xyloredo Turner 1972 264
Xyloredo nooi Turner 265
Xyloredo ingolfia Turner 266
Xyloredo naceli Turner 267
Acknowledgment 268
Literature Cited 268
' Department of Mollusks, Museum of Compara-
tive Zoology, Harvard University, Cambridge, Mas-
sachusetts 02138.
Editorial Note. Professor Emerita Ruth Dixon
Turner died on 30 April 2000 and was for the last
several months of her life severely disabled; in fact,
her active work as a researcher was considerably fore-
shortened by medical problems beginning in about
1995. Among her Nachgelassene Werke was an im-
portant manuscript on the systematics of the deep-
sea pholadid bivalve genera Xylophaga, Xyloredo, and
Xylopholas, a manuscript that she had been preparing
for a number of years and one that had the active
support of the U.S. Department of Defense's, then.
Office of Naval Research (ONR). Professor Turner
was unquestionably a leading world authority on
these taxa and had posted this document, in its pre-
liminary draft form, on a Web site; after her retire-
ment and the beginning of the illnesses that plagued
her, the manuscript was removed from the Web site
witli tlie mtent of readying it for formal publication. Two
outside authorities. Dr. Jorgen Knudsen of the Zool-
ogisk Museum, K0benhavns Universitet, K0benhavn,
Denmark, and Dr. K. Elaine Hoagland, then at the
Association for Systematic Collections, Washington,
D.C., were solicited to make criticisms, and these,
along with my own, were incoi-porated into a more
advanced revision of the text prepared by Ms. Helene
Ferranti, a long time coworker and associate of Pro-
fessor Turner. Ms. Ferranti agreed to revise this
Nachlass in accordance with the comments of the re-
viewers and to update its content and organization.
Having collaborated with Professor Turner on the
subject of deep-sea bivalves, Ms. Ferranti is credited
herein as the person responsible for the final com-
pletion and revision of this valuable text. The new
species described, for which specimens are available
for study in the Museum of Comparative Zoology
(MCZ), and the taxonomic suggestions incorporated
into the text are to be credited to Professor Turner.
Kenneth J. Boss
Editor
Abstract. The provisional grouping of the species
of the bivalve genus Xylophaga suggested by Turner
and Culliney is further elaborated, with 37 species
assigned to six groups depending on characteristics of
Bull. Mus. Comp. ZooL, 157(4): 223-307, October, 2002 223
224 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
the mesoplax, siphons, muscle scars, and method of
reproduction. Three cases of variation in species of
Xylophaga are discussed: variation in response to dif-
ferent substrates, variation that possibly is genetic,
and variation in a normal growth series. Some obser-
vations are made regarding the ecology of deep-sea
wood borers based on experiments carried out with
wood panels; these support the hypothesis that wood
contributes to the growth and diversity of deep-sea
organisms and that the Xylophagainae contribute sig-
nificandy to the food chain by converting wood to a
usable form. A section on systematics considers 19
species of Xylophaga, of which 7 are new, as well as
the monospecific genus Xylopholas and three species
of Xyloredo. Detailed descriptions are given of new
species.
INTRODUCTION
The Xylophagainae, a subfamily of the
Pholadidae, is composed of the genera Xy-
lophaga Turton, 1822, Xyloredo Turner,
1972, and Xylopholas Turner, 1972. Species
in the genus Xijlophaga range in depth
from just below low tide (X dorsalis Tur-
ton) to depths of 7,000 m (X grevei Knud-
son), whereas species in Xyloredo and Xy-
lopholas are restricted to the deep sea
(depths of 239 to more than 2,000 m).
Species in Xyloredo range in depth from
1,737 to more than 2,000 m (X nooi Turn-
er, 1972) and species in Xylopholas range
from 239 to 366 m (X altenai Turner,
1972), with one lot dredged in 2,550 m off
Port Victoria, Sao Tome, Gulf of Guinea.
However, the Sao Tome specimens were
boring in coconut shells and may not have
been living at that depth.
The Xylophagainae are marine, cosmo-
politan, and range from moderate to abys-
sal depths. All of the Xylophagainae, so far
as known, are wood borers, and all are
sublittoral. Only in high latitudes do they
compete with shipworms (Teredinidae) in
cold boreal waters. So far as known, the
Xylophagainae do not occur intertidally, or
in floating wood. Wood containing speci-
mens of the Xylophagainae usually has
been obtained by dredging. Occasionally,
specimens may be obtained from water-
logged wood that has been brought up and
thrown ashore during a storm after being
on the bottom for some time.
Until recently, the Xylophagainae were
considered to be deep-sea organisms of lit:
tie or no economic importance. They were
rare curiosities, of interest mostly in their"
role of recycling wood on the continental
shelf and slope and in the abyss, largely
beyond the depth range of the teredinids.
The Xylophagainae often were referred to
as shipworms and because of the ephem-
eral, patchy distribution of wood in the
deep sea were thought to have little im-
pact on ecological processes. With the ex-
tension of human activities into the deep
sea for fishing (especially trap fishing for
lobsters and crabs), as well as for archae-
ology, mining, monitoring currents, and
other activities, these borers are now gen-
erally considered to be pests.
Species in the genus Xylophaga (Xylo =
wood, phaga = eating) are restricted to
wood, woody plants, and structures made
of wood found in the deep sea. In com-
mon with the teredinids (shipworms), they
have symbiotic bacteria in their gills (Wa-
terbuiy et al., 1983). These bacteria are be-
lieved to have the ability to digest cellulose
and probably to fix nitrogen. Collaborative
work with Dr. Waterbury, microbiologist at
Woods Hole Oceanographic Institute, was
unable to culture cellulose-digesting bac-
teria from the gills of X atlantica, but ev-
idence was found of cellulose enzymes in
the gill tissue.
The three genera of Xylophagainae may
be briefly characterized as follows:
Genus Xylophaga Turton. Siphons rela-
tively short, of equal length or with the
excurrent siphon truncated, and often
capable of retraction between the
valves. Burrow seldom more than five
times the length of the valves and often
with a chimney of fecal pellets lining the ,
posterior end of the burrow. |
Genus Xylopholas Turner. Shell typical
but with the animal extended and with
lateral plates on the siphons.
Genus Xyloredo Turner. Shell typical but
animal elongated and producing a tere- j
dinidlike burrow that is lined with a cal-
Xylophagainae • Turner
225
careous tube marked with distinct
growth rings and margined anteriorly
with a periostracal band.
The Xylophagainae are often confused
with teredinids, but the gills and digestive
and reproductive organs in the Xylopha-
gainae do not extend posteriorly beyond
the valves. In addition, the Xylophagainae
do not have pallets to close the entrance
to their burrows or apophyses for the at-
tachment of the foot muscles. In common
with the teredinids, but unlike species in
the pholadid genera Martesia and Ligno-
pholas, the only other genera of wood-bor-
ing pholadids, the Xylophagainae have a
large wood-storing cecum and probably
utilize the wood in which they bore as
food. For details of anatomy, see Purchon
(1941) for Xijlophaga dorsalis Turton, and
Turner (1955) for X atlantica.
SPECIES GROUPS IN THE
GENUS XYLOPHAGA
Genus Xijlophaga Turton. Xylophaga Turton, 1822,
Conchylia Insularum Britanicarum, p. 253 (type
species, Teredo dorsalis Turton, 1819).
Species in this genus are characterized
by teredolike shells that lack apophyses
and have a divided mesoplax that is vari-
able in shape and size. A chrondrophore
and internal ligament are present. The si-
phons are variable, united for part or all of
their length, with the excurrent siphon of-
ten truncated. The visceral mass and gills
do not extend beyond the valves posteri-
orly. The wood-storing cecum is large.
In his discussion of the taxonomy of Xy-
lophaga, Knudsen (1961) believed that the
use of subgenera was not feasible and
would only lead to the creation of a large
number of monotypic subgenera that
would be of limited value. This is still par-
tially true but new species described in
this report and the additional material now
available concerning other species have
made possible a provisional grouping of
the species, as suggested by Turner and
Culliney (1971). See also Hoagland and
Turner (1981) and Hoagland (1983). This
grouping is helpful when discussing rela-
tionships and geographic distribution. The
characters used for grouping the species
are those mainly of the mesoplax and si-
phons in conjunction with the muscle scars
and methods of reproduction (see Text-
Fig. 1). The mesoplax is a transverse plate,
usually wider than long, that straddles the
valves at the umbos and partially or com-
pletely covers the posterior end of the an-
terior adductor muscle. The mesoplax may
be composed of one or two parts. The im-
portant character of the mesoplax is the
presence or absence of a ventral portion
and tubes; the more detailed characters,
such as the presence of lobes, seem to be
of specific value only. Siphonal characters
include the relative length of the two si-
phons, the presence or absence of cirri at
the apertures, and the type of siphonal
folds, which may or may not have lappets
or fringes.
Not all characters are known for all spe-
cies and a few species seem to be transi-
tional between groups. There is no ques-
tion that more material is needed before
definite statements can be made concern-
ing the formal use of subgenera. However,
the grouping of species as presented here
does offer an opportunity to speculate on
the possible origin and evolution of the ge-
nus and to focus attention on the types of
information that should be considered in
future studies. Comparative anatomical
and molecular studies are greatly needed
but it probably will be some time before
these can be completed because well-pre-
served specimens of deep-sea Xylophaga
are rare and difficult to obtain.
If we consider species with simple si-
phons of equal length and a mesoplax of
two simple flat to slightly curved plates to
be the basic type, it is possible to group
the species in what appears to be a devel-
opmental series of six groups. This list
does not include all nominal Xylophaga.
The groups may be characterized as fol-
lows:
226 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
siphonal retractor
pedal retractor
posterior adductor
umbonal reflection
anterior adductor
ventral adductor
1 bona! -ventral ridg
Text-Figure 1 . Nomenclature of parts of Xylophaga. (1) Internal view of left valve showing relative position of muscle scars. (2)
External view of left valve. (3) Dorsal view of animal with siphons retracted. (4) Lateral view of entire animal with siphons
extended.
Group 1. Mesoplax composed of two
simple flat or slightly cuived plates lo-
cated posterior to the anterior adductor
muscle and standing erect. Siphons of
equal length or with the excurrent si-
phon slightly shorter, cirri on one or
both siphons present or absent. Group
1 includes X. erecta Knudsen, X. con-
cava Knudsen, and X. gerda Turner n.
sp.
Group 2. Mesoplax composed of two
plates that may be curved, flat, sculp-
tured, or smooth, set at an acute angle
to each other, lack dorsal tubes and a
ventral portion, but cover the anterior
adductor muscle dorsally. Siphons of the
same length or with the excurrent si-
phon slightly shorter and with large cirri
on the sides of the excurrent siphon and
small cirri at the incurrent siphon.
Group 2 includes X. grevei Knudsen, X.
wolffi Knudsen, X. hadalis Knudsen, X.
galatheae Knudsen, X. murraiji Knud-
sen, X. africana Knudsen, X. pananien-
.sis Knudsen, and X. clenchi Turner and
Culliney.
Group 3. Mesoplax composed of two
nearly flat plates set at an acute angle to
each other forming an inverted V, with
tubes extending from the posterior dor-
sal surface or longitudinally folded with
anterior lobes or pores. Mesoplax lack-
ing a ventral portion and set in a tentlike
fashion over the anterior adductor mus-
cle. Siphons nearly the same length and I
usually with small cirri on both open-
ings. Group 3 includes X. supplicata
Taki and Habe, X. lohata Knudsen, X.
tuhidata Knudsen, X. bniuni Knudsen,
X. obtusata Knudsen, X. whoi Turner n.
sp., and X. profunda Turner n. sp.
Group 4. Mesoplax composed of two
plates that have a small to large ventral
portion, the dorsal portion being
smooth, folded, or lobed. Siphons of the
same length or with the excurrent si-
phon slightly shorter and with cirri or
papillae at one or both openings. Group
4 includes X. abyssorum Dall, X. dupli-
cata Knudsen, X muraokai Turner n.
sp., X. foliata Knudsen, and X. atlantica
Richards.
Xylophagainae • Turner
227
Group 5. Mesoplax composed of two
plates that are more or less triangular in
outline, with a ventral portion ranging
froin very narrow to more than one half
the width of the dorsal portion. The ex-
current siphon may vary in length from
one half to three quarters that of the
incurrent siphon and have cirri, or it
may be truncated just posterior to the
valves and have dorsal lobes or folds ex-
tending from the truncation along the
dorsal surface of the incurrent siphon
for part or all of its length. Group 5 in-
cludes X. washingtona Bartsch, X ri-
kuzenica Taki and Habe, X. aurita
Knudsen, X. turnerae Knudsen, and X.
praestans E. A. Smith.
Group 6. Mesoplax composed of two
more or less ear-shaped plates some-
what coiled posteriorly. Excurrent si-
phon truncated near the posterior end
of the valves, continuing as lateral lobes
extending from the truncation along the
dorsal surface of the incurrent siphon.
These lobes may vary in width but are
always fringed. Group 6 includes X. dor-
salis Turton, X. depahnai Turner n. sp.,
X. guineensis Knudsen, X. mexicana
Dall, X. tipperi Turner n. sp., X. baijeri
Turner n. sp., X. globosa Sowerby, X.ja-
ponica Taki and Habe, and X. indica
Smith.
Most species are known only from the type
series and these have all been studied by
the author except X. indica, and the spe-
cies described by Taki and Habe. Howev-
er, paratype specimens received through
the kindness of Dr. Habe are in the col-
lection of the MCZ. They include X. ja-
ponica, X. rikuzenica, X. teramachi, and X.
supplicata, although unfortunately all lack
the mesoplax except the last. Two species,
X. teramachi Taki and Habe (Taki and
Habe, 1950) and X. tomlini Prashad (Pras-
had, 1932); are known only from the valves
and remain unassigned. A map showing
the distribution of species of Xylophaga is
provided in Text-Figure 2.
Nineteen of the 37 species oi Xylophaga
listed in the groups above as well as the
monospecific genus Xylopholas and the
three species of Xyloredo are considered
in the section on systematics. Some spe-
cies are discussed more fully than others
but the distinctive characters have been
given for all. For example, Xylophaga mex-
icana Dall and Xijlophaga abyssorum Dall
are fully described because these names
were based on valves only and were vir-
tually nomina dubia. By matching the
valves of the holotypes with complete
specimens, it has been possible to fix the
names of these species. If additional char-
acters or records are given for a well-de-
scribed species, a reference is made to the
original description. Detailed descriptions
are given for new species.
VARIATION
Knudsen (1961) aptly stated that very
little was known about variation in species
of Xylophaga. Unfortunately, large series
of any one species seldom have been avail-
able for study because inaterial usually is
obtained from small pieces of wood or oth-
er plant material that has been dredged or
occasionally thrown ashore as driftwood.
Of the 30 species listed by Knudsen and
the 7 species described as new in this pa-
per, 24 are known from fewer than 10
specimens; only 7 species are known from
series of more than 100 specimens. Most
are known from only one or two localities
and often all specimens are froin a single
piece of wood, and may all be of the same
set, that is, have settled at the same time.
Consequently, it is not surprising that all
specimens in any one lot are quite similar.
Only since the beginning of deep-sea test-
ing and the use of the submersible DSV
Alvin to place experimental wood islands
at great depths has it been possible to ob-
tain sufficient material to study intraspe-
cific variation in this subfamily.
The earliest work of this sort was done
by the U.S. Naval Civil Engineering Lab-
oratory (USNCEL) and the Navy Ocean-
ographic Office (NOO). Three cases of
variation based on this material are re-
228 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Text-Figure 2. Distribution map of species of Xylophaga.
ported here. The first case involves varia-
tion in response to different substrata, the
second case exliibits variation that possibly
is genetic because the substrate and all
other parameters were as nearly uniforin
as possible, and the third case illustrates
variation in a normal growth series.
Variation Due to Different Substrates.
Variation in Xylophaga mashingtona in re-
sponse to the substrate can be deinonstrat-
ed with material from the USNCEL tests.
A series of 10 panels of different types of
wood were attached to a submersible test
unit (STU) that was submerged from April
1965 to May 1966 off San Miguel Island,
California (34°06'N, 120°42'W) at 2,370 ft
(730 m) (see Table 1 and Text- Figs. 3-9).
The 2 X 6 X 0.5-in (50.8 X 152.4 X 12.7-
min) wood panels were all attached to the
same rack on the STU so that they would
be resting just above the mudline. Con-
sequently, all factors affecting the borers
were as nearly identical as possible except
the substrate (i.e., the species of wood) on i
which the borer larvae settled and into j
which they would bore. Text-Figures 3—8
illustrate typical specimens from each of
the wood panels; Text-Figure 9 shows
specimens from a phenolic laminated rod.
It is interesting to note that the dorsal
plates in all specimens are remarkably uni-
form, varying only slightly in length/width
proportions. Even the specimens taken
from the phenolic laminated rod could be
identified by the dorsal plates.
The general shape of the valves with the
high posterior slope also remained rather
constant except in the extremely steno-
morphic (stunted) specimens from Afam-
beaii and the phenolic laminated rod. It is
difficult to explain the proportionate size
of the larval valves on specimens boring
into harder materials except that these
speciinens had not greatly increased in di-
Xylophagainae • Turner
229
Table 1. Variation of Xylophaga washingtona burrows in different types of wood.
Wood
No.
Burrow
Burrow
Specimens
Examined
Lens^th
Diameter
(mm)
(mm)
Remarks
Cedar
Ash
Maple
Pine
Oak
Fir
Redwood
Greenheart
Afambeou
Antidesma pulvinatum
125
21.0
5.0
75
11.0
5.5
35
15.0
5.0
±50
15.0
6.0
50 25.0 4.5 many dead, often three or four specimens in one
enlarged cavity where burrows ran together.
Heavily attacked, particularly at one end.
many dead, burrows running together, panel
heavily attacked, particularly at one end.
well distributed, with a little more concentration
around the hole at one end.
clustered at one end, newly settled to adult.
evenly distributed, shells yellowish-green from
wood.
46 12.0 4.5 many newly settled, often cut into burrows of oth-
er specimens.
specimens stained dark red brown by wood.
concentrated around edges.
concentrated at one end, all very small, many in
umbo stage veliger, 15 small depressions with-
out animals.
150 0.10 0.05 many newly settled, just beginning metamorpho-
46
14.0
4.5
75
2.75
1.75
19
0.75
0.05
Cedar
5 mm
Pine
5 mm
I 1
1 mm
1 mm
Text-Figure 3. Typical specimens of Xylophaga washingtona collected from submerged cedar (top) and pine (bottom).
230 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Ash
5 mm
Text-Figure 4. Typical specimens of Xylophaga washingtona collected from submerged ash (top) and maple (bottom).
Oak
5 mm
Fir
5 mm
Text-Figure 5. Typical specimens of Xylophaga washingtona collected from submerged oak (top) and fir (bottom).
Xyloph AGAIN AE • Turner 231
Redwood
Greenheart
Text-Figure 6. Typical specimens of Xylophaga washingtona collected from submerged redwood (top) and greenheart (bottom).
Afambeau
0.1 mm
Text-Figure 7. Typical specimens of Xylophaga washingtona collected from a submerged panel of Afambeau.
232 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Antidesma pulvinatum
Text-Figure 8. Typical specimens of Xylophaga washingtona collected from a submerged panel of Antidesma pulvinatum.
ameter as they bored so that the lai"val
valves were not inturned with the growth
of the umbos. Evidently little or no attri-
tion had occurred at the umbonal area or
the embryonic valves would have been
worn away.
The most variable characters in this se-
ries are the size of the valves and the num-
ber of denticulated ridges on the anterior
slope in relation to the length of the valves.
Specimens from cedar, pine, ash, maple,
oak, and fir (Text-Figs. 3-5) all were drawn
to the same scale, as shown by the 5-miTi
scale bar, and are arranged in order of de-
creasing size. The nuiTiber of denticulated
ridges on the anterior slope of these spec-
iiTiens varies from 12 to 20. Specimens
froin redwood, greenheart, Afambeau, and
Antidesma pulvinatum (Text-Figs. 6—8)
"were drawii to their own scales and, al-
though these speciinens are much siTialler,
they have as many or more ridges on the
anterior slope. The specimen removed
from the phenolic laminated rod (Text-Fig.
9) has 44 denticulated ridges. Correlated
with the increased hardness of the sub-
strate and the additional denticulated ridg-
es is a proportional increase in the size of
the posterior adductor muscle and its scar.
This suggests the enlargement of the pos-
terior adductor inuscle in response to in-
creased activity of boring. The general
shape and sculpturing of the inuscle scar
was similar in all specimens. No noticeable
variation was found in the siphons, except
size, regardless of the substrate. However,
specimens that were able to bore deeply
into the wood usually formed a chimney
composed of compacted fecal inaterial lin-
ing the posterior end of the burrow.
Two periods of settlement apparently
occurred on some of the panels, because
specimens of two age groups could be
found. However, it is impossible to say
whether the specimens removed from the
Antidesma were of the second set or if
Xylophagainae • Turner
233
1 mm
Phenolic Laminated Rod
Text-Figure 9. Typical specimens of Xylophaga washingtona collected from a submerged phenolic rod.
they had simply been unable to increase
in size because of the hardness, chemical
composition, or both of the wood. Because
the specimens apparently were alive at the
time the wood was reinoved from the wa-
ter and because only a few rows of dentic-
ulated ridges were present, the inference
was made that these specimens probably
belonged to a second set. Certainly the lar-
val shells shown in Text- Figure 8 must be
from a second set. The number of speci-
mens examined and the maximum length
and diameter of the burrows for each type
of wood are given in Table 1 .
Variation in the Mesoplax o/ Xylophaga
depalmai Turner n. sp. Approximately 300
specimens of X. depalmai n. sp. were ob-
tained from tests conducted by the NOO
about 2—3 miles east of Fort Lauderdale,
Florida (26°04'N, 80°04'W) in depths
from 100 to 500 ft (30.5 to 152.5 m) (see
Table 14 under the description of X de-
palmai n. sp. for information giving panel
numbers, depth, and dates of exposure).
In this species, the general shape of the
valves, the siphons, and the muscle scars
show little variation but the mesoplax is
extremely variable. The mesoplax is typi-
cally bilaterally symmetrical, ear-shaped,
longer than wide, with the two halves
coiled inward at the posterior end, and
with a long medial line where the two
halves meet (Plate 24, Figs. 16, 17). In nu-
merous specimens, the mesoplax was not
bilaterally symmetrical, but one half was
considerably shorter than the other and of-
ten appeared malformed (Plate 24, Figs.
11—15). In several specimens, the ventral
surface of the two halves of the mesoplax
was fused by the periostracal covering, al-
though the dorsal surface still appeared di-
vided. In other specimens, the mesoplax
was elongated, the coiled posterior ends,
instead of curling inward toward each oth-
er, remained nearly straight or curled
slightly outward, with the ventral surface
being completely fused (Plate 24, Figs. 1-
5). In two specimens, the two halves of the
mesoplax had completely fused dorsally
and ventrally, although the lines of fusion
remained clearly visible. The tapered pos-
terior end of this cornucopialike mesoplax
coiled slightly ventral and to the left.
Such variation is in marked contrast to
the uniformity seen in the mesoplax of X.
washingtona Bartsch. It is impossible to
say whether this variation is genetic or eco-
logic but we are able to say that all types
of the mesoplax of X. depalmai were found
in a single panel retrieved from a depth of
approximately 300 ft (91.44 m). Variation
in the mesoplax is a factor that must be
234 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
taken into consideration when evaluating
species in this genus.
Variation Exhibited in Growth Series.
In most species of Xylophaga, the dorsal
plate is quite simple and the mesoplax of
the young is similar to that of the adult, as
seen for X. washingtona. Dons (1929a,b)
described briefly and illustrated a similar
situation in X praestans Smith and X. dor-
salis Turton.
Some species have more elaborate dor-
sal plates and for some of these (i.e., X.
muraokai Turner n. sp., X. hayeri Turner
n. sp., and X. abyssorum Dall), it is pos-
sible to build up what appears to be
growth series. All begin with a simple pos-
terior covering to the anterior adductor
muscle that is difficult to differentiate
among the species in the young stage. As
calcification of the periostrascal covering
progresses, the adult form gradually ap-
pears. In the three species mentioned, the
characters of the valves and the siphons of
the young agree with those of the adult
specimens. Therefore, although living ma-
terial has not been available and develop-
mental studies have not been possible, it
seems reasonable to consider these as
growth series. Xylophaga muraokai (Plate
19, Fig. 3) is relatively simple, with the
dorsal portion becoming thickened and
joining laterally with the basal portion
while the ventral flanges enlarge. In X.
hayeri (Plate 31, Fig. 3), the broad trans-
verse dorsal portion becomes very con-
spicuous and in X. abyssorum (Plate 15,
Figs. 4, 5), the dorsal incurving of the lat-
eral arms produce an elaborately lobed
mesoplax. Nothing is known of the allo-
metric changes that take place during
growth of the peculiar dorsal plates of X.
tiibulata Knudsen, X. bruuni Knudsen, X.
obtusata Knudsen, and X. whoi Turner n.
sp.
Detailed studies of these growth series
will have to await improved techniques for
handling living material under deep-sea
conditions, and the ability to rear deep-sea
species in the laboratoiy. In the meantime,
it seems best to consider these forms as
members of growth series rather than dif-
ferent species, particularly because the se-
ries in each case was built up from mate-
rial taken from a single piece of wood.
PRELIMINARY REPORT ON THE
RESULTS OF EXPERIMENTS ON THE
ECOLOGY OF DEEP-SEA WOOD
BORERS AND THE ROLE OF WOOD IN
THE DEEP SEA
The results of the first exposures of
wood panels at the Woods Hole Oceano-
graphic Institution (WHOI)-AZuin per-
manent bottom station south of Woods
Hole (39°46'N, 70°41'W, in 1,830 m) were
reported in 1973 (Turner, 1973). At that
time, the Xylophagainae were postulated !
to be the most important deep-sea organ-
isms involved in converting woody plant
material to an available food source for
other organisms.
Pursuing this theory and to continue the
studies of the ecology and life history of
the Xylophagainae, wood panels were ex-
posed at the Alvin permanent station in
the Tongue of the Ocean, Bahama Islands,
on 19-22 January 1974 {Alvin dives 492,
493, 494, and 495) at a depth of 2,032 m.
The first of these panels was picked up on
7 March 1974, frozen immediately, and re-
turned to Woods Hole where it was ex-
amined. Newly settled larvae and meta-
moi^phosing Xylophaga with one to two
rows of denticulated ridges were removed
from the panel. The specimens were ap-
proximately 300 |JLm in length and the
greatest penetration was about twice the
depth of the shell. The debris rings sur-
rounding the burrows were much coarser
than those made by teredinids. The distri-
bution of the entrance holes was some-
what patchy and varied from 5 to 20 cm^.
The specimens were too young to identify
because none of the dorsal plates had been
formed but examination with scanning
electron microscopy showed a well-devel-
oped distinctive sculpture on the larval
shell.
This first panel from the Tongue of the
Ocean established that Xylophaga were
XYLOPHAGAINAE • Turner
235
just beginning to settle on the wood a max-
imum of 48 days after it was implanted in
the bottom and that settlement of larvae
could occur in early March, at least at this
site.
Three more panels were removed from
the Tongue of the Ocean station (Tower
1 — west arm) on 19 April 1975 during Al-
vin dive 552. I was an observer on this dive
and as we approached the panels I noticed
an increase in the number of shrimp and
galatheid crabs. The panels had numerous
crabs crawling all over them. Some of the
crabs had crawled under the plastic mesh
bags covering the panels and had grown so
large they could not escape. (Note: After
the near loss of the panels at the northern
station because of the heavy attack of bor-
ers, the decision was made to put the pan-
els in plastic mesh bags so that the pieces
could be retrieved if they began to disin-
tegrate.) The specimens inside the bag
were carried to the surface v^dth the pan-
els. When the panel was disturbed, the
specimens on the outside of the mesh fell
off. The largest of the 12 crabs was 43 mm
in length. The diamond-shaped opening of
the mesh was 5 X 10 mm. The smallest
crab ineasured 8 mm in length; others
measured 40, 33, 32, 30, and 24 mm. It is
obvious that the crabs were finding suffi-
cient food either in or on the wood to grow
at a fairly rapid rate.
The first young crabs to find the wood
may have fed on the newly settled Xijlo-
phaga laiA^ae and this might explain the
patchy distribution of the borers in the
panels. However, the larger crabs would
not have stayed on the wood unless there
was something for them to eat. The crabs
had to be under 10 mm in leng-th to gret
under the mesh and if the larvae were not
settling until early March it would be at
least early May before the borers had
grown sufficiently to be a good food source
for the crabs. Therefore, I think we can
postulate that the largest crab measured
grew at least 33 mm in a period of 10
months.
Examination of the panels showed a
rather heavy attack of three species of Xy-
lophagainae. These included Xyloredo
nooi Turner and two Xylophaga species,
Xylophaga clenchi Turner and Culliney
and X profunda Turner n. sp. The X nooi
were typical with valves that reached 5
mm in length and burrows that were 18—
22 mm in length. The calcareous lining of
the burrow of the largest specimen was 13
mm long and 2.5 mm in diameter at its
anterior end. The smaller species of Xy-
lophaga, X. clenchi Turner and Culliney,
also had been obtained previously from
wood exposed in the Tongue of the Ocean
by John DePalma of the NOO. This is a
fairly small species. The valves were 8—10
mm in length and several of the specimens
were canying lai"vae on the umbonal area
of the valves. The laivae measured 0.2 mm
in length. The large species o{ Xylophaga,
X. profunda Turner n. sp., had not been
seen before. The valves were 14 mm in
length, and one specimen measured 40
mm to the tip of the siphons. The burrows
were 45—50 mm in length. Both X clenchi
and X profunda lined the posterior end of
their burrows with consolidated fecal pel-
lets and the burrows of all dead specimens
contained one or more specimens of cap-
itellid worms that were feeding on the pel-
lets as well as the remains of the Xylopha-
ga. Often the spaces between the valves of
the borers were filled with the smaller fe-
cal pellets of the worms. Breaking these
balls of pellets apart, I always found one
or two capitellid worms. In the Xylophaga
burrows and on the surface of the wood,
I also found two other polychaete worms.
One belonged to the family Chiysopetali-
dae and the other to a family of polynoid
wonns.
A preliminaiy examination of the stom-
ach contents of a broken specimen of a
galatheid showed that the crab had ingest-
ed some fine chips of wood because iden-
tifiable cells remained in the material.
Consequently, we can postulate that the
crabs were feeding on the Xylophaga,
probably dead ones. The tissues of the Xy-
lophaga are so soft that they are unrec-
236 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
ognizable in such a preliminary examina-
tion. The crab's stomach also contained se-
tae of the chrysopetalid worms, a small
nematode, and some sponge spicules. The
chrysopetalids probably were feeding on
the capitellid worms.
I think five points are worthy of notice:
1) Xijlophaga in the Tongue of the Ocean
as well as at the northern Alvin station are
growing much faster than one would ex-
pect. 2) The lai-vae of X profunda n. sp.
were settling in early March. 3) Xijlophaga
clenchi broods its young and was carrying
young in mid-April. 4) Probably the crabs
and worms were also growing faster than
normal for the deep sea. 5) A food chain
based on wood and Xijlophaga was being
developed.
This lends support to my hypothesis that
wood is an important source of enrich-
ment in the deep sea, that it contributes
to both the diversity of organisms in a giv-
en area and to their rate of growth, and
that the Xylophagainae are the most im-
portant organisms involved in converting
the wood to a usable form. To my knowl-
edge, this is the first documented food
chain for invertebrates in the deep sea. On
the basis of these simple experiments, it
now seems conceivable that the slow
growth rates usually attributed to deep-sea
animals may be due to lack of food, at least
for epifaunal forms, rather than being an
inherent characteristic of the species in-
volved.
SYSTEMATIC ACCOUNT
GENUS XYLOPHAGA TURTON 1822
Xytophaga concava Knudsen
Plate 1
Xijlophaga concava Knudsen, 1961, Galathea Report,
5: 167-169, figs. 4, 5 {Galathea, station 726, Gulf
of Panama [5°49'N, 78°52'W] in 3,270-3,672 m).
Holotype, Zoological Museum, University of Co-
penhagen; paratyjae, MCZ 235796.
Distinctive Characters. Posterior slope
of valves concave when viewed dorsally
Mesoplax composed of two rather wide,
erect, curved plates that extend above the
umbos. Siphons nearly the saine length,
excurrent slightly shorter with a few large
cirri, incurrent siphon with many small cir-
ri (Plate 1, Figs. 2, 3). Chimney apparently
lacking, not mentioned by Knudsen and
not found with the single specimen re-
ported here.
Remarks. At the time Knudsen de-
scribed this species he had 4 specimens
from Galathea station 726 and 25 speci-
mens from Galathea station 739. Both of
these stations were given as in Gulf of Pan-
ama. However, station 726 is about 95
miles west of the Gulf of Tibuga, Golom-
bia, whereas station 739 is about 90 miles
west of Ensenada Guayabo, Panama. A
single specimen of X. concava was taken
by the RA^ Pillshury at station 526. This
locality is about midway between the two
Galathea stations.
The PiUsbury specimen agrees closely
with the description and figures given by
Knudsen (1961) except that the incurrent
siphon has a double row of about 25 small
cirri around the aperture and the excur-
rent siphon has 6 large cirri (Plate 1, figs.
2, 3). Concerning the siphons, Knudsen
stated that "both openings are at the distal
end, close together, and around theiu 15-
16 small cirri are present." In Knudsen's
illustration, the distal ends of the siphons
appear to be contracted; neither the two
openings nor the cirri are apparent. The
contracted condition of Knudsen's speci-
mens probably accounts for the differenc-
es noted here.
Xijlophaga concava is closely related to
Xijlophaga gerda Turner n. sp. but differs ;
in the size of the mesoplax, the type of
siphonal openings, and the chimney. (See
Remarks under x. gerda.) Xijlophaga con-
vava also is related to X. erecta Knudsen
(1961) from the Sulu Sea. Knudsen (1961) i
reported that no cirri were visible on the!
siphons of X. erecta, that the posterior ad-
ductor scar was much broader in X. con-
cava, and that the posterior slope of the
valve viewed dorsally was convex in X. er-
ecta rather than concave. Knudsen's de-
scription of X. erecta was based on 20
Xylophagainae • Turner
237
specimens and unfortunately no further
records have been obtained.
Range. From off Ensenada Guayabo,
Panama, south to off the Gulf of Tibuga,
Colombia, in depths from about 915 to
3,670 m.
Specimens Examined. COLOMBIA: Galathea, sta-
tion 739, Gulf of Panama, about 90 miles W of En-
senada Guayabo (7°22'N, 79°32'W) in 915-975 m
(dried specimens); Pillsbunj, station 526, about 110
miles W of Cabo Marzo (6°53'N, 79°27'W) in 3,193-
3,211 m; Galathea, station 726, Gulf of Panama
(5°49'N, 7S°52'W) in 3,270-3,670 m.
Xylophaga gerda^ Turner new species
Plates 2, 3
Holofijpe. MCZ 328378. Paratypes,
MCZ 316741, 316742.
Type Locality. Gerda, station 499, about
3 miles off Southwest Point, Great Baha-
ma Island, Bahama Islands (26°37'N,
78°56'W) in 155 fathoms (283.96 m).
Distinctive Characters. Posterior slope
of valves concave when viewed dorsally.
Mesoplax composed of two narrow, erect
cui-ved plates at the posterior end of the
anterior adductor muscle. Siphons of
equal length, with a periostracal sheath
and four or five cirri surrounding the ap-
ertures. Chimney composed of fecal ma-
terial agglutinized to a periostracal base
(Plate 2, Fig. 7).
Description. Shell globose, fragile,
reaching 3.0 mm in length and 2.8 mm in
height, umbos inflated. Pedal angle 110—
115°. Anterior slope with up to 45 rather
evenly spaced denticulated ridges. Umbo-
nal— ventral sulcus narrow and only slightly
depressed. Disc and posterior slope sculp-
tured with fine, incised growth lines. Pos-
terior slope high, flaring, and somewhat
ear-shaped.
Inner surface of valves smooth and glis-
tening. Umbonal— ventral ridge low and in-
distinct except near the wide, low ventral
condyle. Chondrophore and internal liga-
Table 2. Measurements of Xylophaga gerda.
Length
(mm)
lIHiihl
(min)
Location
1.1
1.0
Gerda, station 266
1,5
1.4
Gerda, station 266
1.8
2.0
Gerda, station 266
2.5
3.0
2.3
2.5
paratype
holot)/pe
3.0
2.8
Pillsbunj, station 328
3.3
3.0
Pillsbunj, station 944
3.8
3.5
Pillsbunj, station 944
- Named for RA^ Gerda, Rosenstiel School of Ma-
rine and Atmospheric Sciences, University of Miami,
Miami, Florida, whose station 499 is tiie type locality.
ment small. Posterior adductor muscle
scar large with faint transverse impres-
sions, which are best seen externally on an
entire specimen. Disc separated from pos-
terior slope by a shelflike ridge (Plate 3,
figs. 2—5). Pedal and siphonal retractor
scars not visible.
Mesoplax composed of two erect, nar-
row, curved, slightly calcified plates, locat-
ed just posterior to the anterior adductor
muscle and not extending above the um-
bos.
Siphons long, probably not capable of
retraction between the valves, of equal
length, united nearly to the tip, with a thin
periostracal sheath. Siphonal apertures of
about equal size, each with four or five
comparatively large cirri, which appear as
a common ring of cirri when the siphons
are retracted. A browii periostracal cylin-
der containing fecal material may extend
nearly one half the length of the excurrent
siphon (Plate 2, Figs. 1-4). Chimney built
in sections, composed of fine fecal material
agglutinized on a periostracal lining with
"leaves" of periostracum extending to the
outer surface (Plate 2, Figs. 6, 7). Arrange-
ment of the gills and labial palps typical
for the genus, foot large but not muscular,
cecum veiy large and showing through the
foot (Plate 2, Fig. 5). Pedal and siphonal
retractor muscles weak, their arrangement
typical for the genus.
Measurements. See Table 2.
Remarks. On the basis of the shell and
the mesoplax, this species is closely related
to X concava Knudsen from the Gulf of
Panama. It differs in being much smaller
238 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
(none of the 19 specimens reached 4 mm
in length, whereas Knudsen gives 8.6 mm
for X concava) and in having a narrow me-
soplax that does not extend above the um-
bos. In addition, the excurrent and incur-
rent siphons of X. gerda are the same
length, are separate at the tip, and the ap-
ertures of both siphons have four or five
relatively large cirri. In X. concava, the si-
phons are joined for their entire length
and the excurrent siphon is slightly shorter.
The type of chimney produced by X gerda
is unlike any other known to date in this
genus.
Unfortunately, all the specimens of X
gerda are small, extremely fragile, and
rather poorly preserved. Consequently, it
has been impossible to do anatomical work
beyond that mentioned in the description.
One specimen from which the valves were
removed appeared to have an accessory
genital organ similar to that described by
Purchon (1941).
Range. Probably throughout the Carib-
bean in depths from about 283 to 2,072 m.
Specimens Examined. BAHAMA ISLANDS; Ger-
da, station 499, about 3 miles off Southwest Point,
Great Bahama Island (26°37'N, 78°56'W) in 155 fath-
oms (283 m). UNITED STATES, FLORIDA: Gerda,
station 266, off Fowey Rocks, Florida Keys (25°39'N,
79°58'W) in 185-187 fathoms (338-342 m). LESSER
ANTILLES: Pillsbunj, station 944, 45 miles N of
Port Louis, Guadeloupe Island (I6°32,2'N,
61°36.8'W) in 364-421 m. PANAMA: Pillsbunj, sta-
tion 328, about 25 miles N of Punta San Bias, Gulf
of San Bias (9°55.8'N, 78°59.8'W) in 2,069-2,072 m.
Xylophaga grevei Knudsen
Plate 4
Xylophaga grevei Knudsen, 1961, Galathea Report,
5: 176, figs. 16-18 (Galathea, station 495, Banda
Trench, south of Geram [5°26'S, 130°58'E] in
7,250-7,290 m). Holotype, Zoological Museum,
University of Copenhagen.
Distinctive Characters. Mesoplax com-
posed of two triangular plates that are flat
dorsally, in contact the length of their iTie-
dian edge, bent downward on their outer
edge to meet the umbonal reflection, and
lack a ventral portion. Posterior adductor
muscle scar with oblique radiating impres-
Table 3. Measurements of Xylophaga grevei.
Length
(mm)
Height
(mm)
1,9 1.6 Galathea. station 444
12.5 11.0 holotype
sions. Siphons nearly the same length, the
aperture of the excurrent siphon much
smaller in diameter than the incurrent si-
phon and with about 6 cirri; incurrent si-
phon with about 35 small cirri. Young car-
ried on the uinbonal area of the adult.
Measurements. See Table 3.
Remarks. Through the kindness of Jor-
gen Knudsen, it was possible to borrow the
preserved dredged wood from the Zoolog-
ical Museum, University of Copenhagen.
In a small piece taken by the Galathea at
station 444, I found several additional
speciiTiens of X. grevei. They are much i
smaller than that figured by Knudsen, the
anterior slope is much narrower, and the
denticulated ridges are more widely
spaced. However, they appear to be young
but sexually mature specimens of that spe-
cies. From one to five rather large young
were attached posterior to the umbos on
the dorsal surface of the parent shells.
Xylophaga wolffi Knudsen, based on i
only two specimens, also was fro in Gala-
thea, station 444. It has valves, muscle
scars, and siphons similar to those of X.
grevei. The outstanding difference be-
tween these species is the flat plates of the
mesoplax of X. wolffi, which are set in
tentlike fashion at an acute angle to each
other. It has not been possible to compare
the type of X. wolffi with this new material
from Galathea, station 444, but it appears
that these two forms (i.e., X. grevei and X.
wolffi) may be equivalent to the condition
found in X. clenchi Turner and Culliney,
where occasional specimens have a bent,
flat-topped mesoplax (Plate 4, Figs. 5, 6).
Further collecting may show X. grevei and
X. wolffi to be forms of a single species.
The denticles on the ventral edge of the
mesoplax of X. wolffi described by Knud-
Xylophagainae • Turner
239
sen may be an age factor but more mate-
rial is needed to prove this.
Range. Mindanao Sea south to the Ban-
da Trench, Banda Sea in depths from
about 545 to 7,290 m.
Specimens Examined. PHILIPPINE ISLANDS:
Galathea, station 444, Sulu Sea, W of Basilan Island
(7°54'N, 121°30'E) in 5,050 m.
Xylophaga clench P Turner and Culliney
Plates 5-8
Xylophaga clenclii Turner and Culliney, 1971, Amer-
ican Malacological Union Annual Report for 1970,
p. 66 (U.S. NOO test site. Tongue of the Ocean,
about 4 miles off northeastern tip of Andros Island,
Bahama Islands [24°54'N, 77°49'W] in 1,737 m).
Holotype, MCZ 316743; paratypes, MCZ 316744,
316745.
Distinctive Characters. Mesoplax small,
composed of triangular, nearly flat plates
lacking a basal portion; the two plates usu-
ally meeting at an acute angle in frontal
view. Burrow lined with a chimney of
coarse, loosely consolidated fecal pellets.
Young held on posterior dorsal surface of
the adult. Excurrent siphon shorter than
incurrent siphon and with two large papil-
lae on either side.
Description. Shell globose, reaching 14
mm in length and 13.5 mm in height, thin,
fragile, with a thin, light brown periostrac-
um that is thickened along the dorsal and
posterior margin of the valves. Umbos
prominent and strongly incurved. Pedal
gape angle about 108°. Beaked portion of
the anterior slope sculptured with numer-
ous denticulated ridges that are widely
spaced in the young, becoming increasing-
ly compacted toward the ventral margin in
older specimens. Specimens 5 mm in
length have up to 25 ridges. Posterior por-
tion of the anterior slope rather narrow.
Umbonal— ventral sulcus moderately to
deeply impressed becoming shallower
with age and bounded posteriorly by a
broad, low, rounded ridge. Disc and pos-
^ Named for Dr. William J. Clench, Curator of
Mollusks, 1926—1966, Museum of Comparative Zo-
ology, Harvard University.
terior slope sculptured with uniform, rath-
er pronounced growth ridges that are par-
ticularly prominent in young specimens.
Inner surface of valves smooth and glis-
tening. Umbonal-ventral ridge prominent,
distinctly segmented, and with a large ven-
tral condyle. Chondrophore and internal
ligament well developed. Posterior adduc-
tor muscle scar kidney-shaped in outline
and with irregular transverse, often anas-
tomosing impressions that become more
numerous with age. Pedal retractor scar
more or less oval in outline and located
just anterior to the embayment in the pos-
terior adductor scar. Siphonal retractor
scars small, not impressed, located just
posterior to the umbonal-ventral ridge
about midway between the umbo and the
ventral condyle.
Mesoplax small, composed of two flat to
slightly arched triangular plates lacking a
ventral portion, set in a tentlike fashion
and held in place by the periostracum and
the anterior adductor muscle. Plates usu-
ally meet dorsally at 90° (Plate 5, Fig. 3),
occasionally at an obtuse angle. They are
rarely bent longitudinally (Plate 8, Fig. 3).
Outer surface of plates sculptured with
ridges paralleling the anterior margin and
lining up with the denticulated ridges of
the valves w^here they come in contact.
Siphons short, extending only about one
third the length of the valves. Excurrent
siphon about one half the diameter of and
slightly shorter than the incurrent siphon,
with a well-developed sphincter muscle
surrounding the aperture and two large
papillae on each side (Plate 7, Fig. 3). In-
current siphon has minute cirri surround-
ing the aperture and a second set of larger
cirri anterior to them within the siphon
(Plate 6, Fig. 4).
Posterior end of burrow lined with
coarse, loosely compacted fecal pellets.
Young held by byssus threads on the pos-
terior dorsal surface of the adult shell
(Plate 7, Figs. 1, 2).
Measurements. See Table 4.
Remarks. This species is most closely re-
lated to X africana Knudsen from the
240 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Table 4. Measurements of Xylophaga clenchi.
Length
Height
(mm)
(mm)
Locatioii
2.3
2.1
Ingolf, Station 67
3.2
2.0
Atlantis II, station 124
3.5
3.5
holotype
4.0
3.5
Tongue of the Ocean
5.0
4.8
Tongue of the Ocean
10.0
9.5
Tongue of the Ocean
10.0
10.0
Pillsbury, station 104
14.0
13.5
Pillshury, station 394
Gulf of Guinea, West Africa (1°42'N,
7°51'E in 2,550 m), but the mesoplax dif-
fers in having a longer median line where
the two plates meet, a weaker sculpture,
and in lacking the rounded nodule on the
ventral surface. Unfortunately, the valves
of the two known specimens of X. africana
were too fragixientaiy to allow comparisons
on this basis. The siphons of these two
species are similar but X clenchi differs in
having only two rather than three large cir-
ri on either side of the excurrent siphon
and in having two rings of small cirri bor-
dering the aperture of the incurrent si-
phon. The incurrent siphon of X. africana
lacks cirri, according to Knudsen (1961).
Two other species, X wolffi Knudsen from
the Sulu Sea and X murrayi Knudsen
from off Zanzibar, also have similar dorsal
plates. The former differs from X. clenchi
in having denticles on the basal margin of
the mesoplax with corresponding denticles
on the umbonal reflection. In addition, on
each side of the excurrent siphon of X.
wolffi there are "7 finger-like tentacles on
a common base and somewhat larger ten-
tacle dorsally." The siphons of X. murrayi
differ in having a circle of about 35 "short
tentacles" surrounding both openings.
Knudsen (1961) compared X. wolffi with
X. supplicata Taki and Habe but exami-
nation of paratypes of X. supplicata re-
ceived from Habe showed that species to
have tubules on the mesoplax. (See also
under X. supplicata Taki and Habe.)
Xylophaga clenchi Turner and Culliney
also appears to be closely related to X.
panamensis Knudsen but the latter differs
in having the plates of the mesoplax
sinooth. The posterior adductor muscle
scar of X. panamensis is oval and irregu-
larly lobed anteriorly, the weak umbonal-
ventral ridge is bounded posteriorly by a
groove, and the condyles are lacking. The
siphons of X. panam,ensis are unknown. \
The variation in the inesoplax of X. clen-
chi is interesting and presents some prob-
lems. In most specimens, the plates are flat
or nearly so and none has a ventral portion
or cavity of any kind. The angle at w^hich
the plates meet dorsally is typically acute;
however, specimens from shallower waters
(i.e., less than 900 m) occasionally have
plates that meet at an obtuse angle. Two
of the five specimens from Atlantis II, sta-
tion 124, have the dorsal plate bent lon-
gitudinally at a right angle (Plate 8, Fig.
3), whereas others are typical. Because the
shell characteristics of these specimens fit
within the range of variation of the typical
forms, they are considered to be ecologic
variants. Unfortunately, none of the spec-
imens from shallower water has extended
siphons, so comparisons cannot be made
on that basis.
Although the holotype of X. clenchi is
small, it was selected because it was the
most nearly perfect and the only specimen
for which dorsal plates, young, siphons,
and chimney all were present. The large
specimen from IW Pillsbury, station 394,
and the two from Pillsbury, station 104,
were dead when collected. At least one
specimen from all other localities was alive
at the time of collection. The specimens
froiTi the Tongue of the Ocean were from
test panels submerged by the NOO from
4 April 1962 to 17 February 1965. For a
description of the test site see DePalma
(1969). One specimen taken by the IW
Atlantis II at station 119 and three speci-
mens taken at station 131 were from epi-
benthic sled hauls in which no wood or
plant material was found. The specimens
were alive and apparently normal although
they were badly crushed, for they were mi-
nute and veiy delicate. This raises the
question as to whether or not at least some
Xylophagainae • Turner
241
species of Xylophaga can survive in a firm
muddy bottom if wood is not available. So
far as known, these are the only living
specimens of Xylophaga not taken from
wood or other plant material, except those
that were boring in plastic at the navy test
site off California and possibly those spec-
imens of Xylophaga foliata Knudsen that
came from a station in Macassar Strait
from which it was reported that no wood
was taken. The specimens from off Iceland
were removed from wood dredged by the
Ingolf Expedition in 1896. All specimens
were small, the largest being 2.5 mm in
length. Many had from 1 to 10 large young
attached to the valves in the umbonal area
(Plate 7, Fig. 2). This record extends the
range of the species far to the north but it
has been impossible to find any characters
to distinguish these specimens from those
occurring to the south.
The difference in the size of the young
attached to the dorsal surface of the spec-
imens figured in Plate 5, Figures 1 and 2,
and Plate 7, Figures 1 and 2, reflects the
age of the lan^ae, with those in the latter
plate having well-developed umbos and
appearing fully mature. It is interesting to
note the position of the larvae on the two
adults and to speculate that perhaps the
larvae move gradually toward the umbos
as they mature.
Range. From off Iceland south to Ve-
nezuela in depths ranging from 35 to
4,862 m.
Specimens Examined. ICELAND: Ingolf, station
67, S of Eyrabakki (61°30'N, 22°30'W) in 1,836 m.
UNITED STATES, NEW JERSEY: Albatross, station
2550, about 160 miles E of Barnegat Bay (39°44'N,
70°30'W) in 1,977 m. VIRGINIA: Albatross, station
2731, off Cape Henry (36°45'N, 74°28'W) in 1,428
m. NORTH CAROLINA: Atlantis II, station 131,
about 420 miles E of Currituck Sound (36°28.9'N,
67°58.2'W) in 2,178 ni; off Cape Hatteras (35°44'N,
75°15'W) in 35 m. GEORGIA: Pillsburij, station 104,
about 80 miles SE of Brunswdck (3r00'N, 79°50'W)
in 247 m. BAHAMA ISLANDS: Tongue of the
Ocean, about 4 miles off NE tip of Andros Island
(24°54'N, 77°49'W) in 1,737 m; Tongue of die Ocean
(24°53.2'N, 77°40.2'W) in 2,066 m. BERMUDA: At-
lantis II, station 124, about 750 miles E of Cape
Charles (due N of Bermuda) (37°26'N, 63°59'W) in
Table 5. Measurements of
Xylophaga supplicata.
Length Height
(mm) (mm)
8.5
8.8
parat>pe MCZ 194820
4,862 m; Atlantis II, station 119, just S of Bermuda
(32°15'N, 64°32'W) in 2,095-2,223 m. PANAMA:
Pillsbunj, station 328, about 25 miles off Punta San
Bias, Gulfo San Bias (9°55.8'N, 78°59.8'W) in 420-
640 m. VENEZUELA: Pillsbunj, station 719, about
100 miles N of Pertigalete Bay (11°35'N, 64°35.4'W)
in 770-890 m.
Xylophaga supplicata Taki and Habe
Plate 9
Metaxi/lophaga supplicata Taki and Habe, 1950, Il-
lustrated Catalogue of Japanese Shells No. 7, p. 47,
text-figs. 1, 2 (Tosa Bay, Shikoku, Japan, in 100
fathoms). Holotype, T Habe Collection; paratype,
MCZ 194820; Knudsen 1961, Galathea Report, 5:
188.
Distinctive Characters. Mesoplax com-
posed of two flat triangular plates set at a
sharp angle to each other, dorsal margin
more than one half the total length, and
with minute tubules at the posterior end.
The tubules sit deep within the cavity
formed by the umbos and cannot be seen
when the shell is viewed anteriorly. Chon-
drophore of left valve with a large tooth
(Plate 9, Figs. 5, 6).
Measurements. See Table 5.
Remarks. Taki and Habe (1950) de-
scribed the dorsal plate of X supplicata
simply as a "small triangular protoplax
[ = mesoplax]" that is attached vertically. A
paratype received from Dr. Habe has small
tubes on the dorsal posterior end of the
mesoj^lax.
Xylophaga bniuni Knudsen, 1961, from
the Mindanao Sea is very close to if not
synonymous with X supplicata. Unfortu-
nately, Taki and Habe did not mention the
tubes in their original description and
Knudsen did not see the types. Also, un-
fortunately, Knudsen had only a single
specimen; hence, it will be necessaiy to
obtain more material before a definite
statement can be made concerning the sta-
242 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
tus of X bniuni. (See also Remarks under
Xylophaga ivhoi Turner n. sp.)
Specimens Examined. JAPAN: Tosa Bay, Shikoku,
in 100 fathoms (183 m). "
Xylophaga whoh Turner new species
Plates 10, 11
Holotijpe. MCZ 275015.
Type Locality. R/V Atlantis, station
3471, off Cardenas, Matanzas Province,
Cuba (23°21'N, 80°56'W) in 500 fathoms
(914 m).
Distinctive Characters. Mesoplax com-
posed of two flat triangular plates with a
short median line, set at a shai-p angle to
each other and with a large hollow tube
extending outward from the posterior dor-
sal surface of each plate.
Description. Shell globose, reaching 7
mm in length and 6.2 mm in height, thin,
fragile, and with a veiy thin, light tan per-
iostracum. Pedal gape angle about 95°.
Beaked portion of the anterior slope sculp-
tured with numerous low denticulated
ridges; the posterior portion with fine in-
distinct ridges. Umbonal-ventral sulcus
only slightly impressed and irregularly
sculptured. Sculpture on the disc and pos-
terior slope consisting of pronounced
growth ridges. Umbonal reflection narrow,
closely impressed near the umbos, free
and vertical anteriorly.
Inner surface of valves smooth and shin-
ing. Umbonal-ventral ridge becoming ev-
ident only near the ventral margin but rap-
idly increasing in size to a large ventral
condyle. Chondrophore and internal liga-
ment well developed; chondrophore of the
left valve with a small tooth on the poste-
rior upper margin. Posterior adductor
muscle scar large, elongate, oval in outline,
irregularly sculptured, set high on the pos-
terior slope and well in from the posterior
margin. Pedal retractor scar rather large,
elongate, somewhat irregular, and located
Table 6. Measurements of Xylophaga whoi.
Leiiytli
(mil.)
Height
(mm)
Location
2.5
2.3
Gerda, station 266
3.1
3.0
Gerda, station 266
6.9
7.0
6.2
6.8
holotype
Pillsbiii-y, station 394
7.2
7.0
Pillsbury, station 944
9.5
12.5
13.8
14.9
9.0
11.5
13.1
14.0
Pillsbury, station 944
Pillsbury, station 944
Pillsbury, station 944
Pillsbury, station 944
just anterior to the posterior adductor scar.
Siphonal retractor scar not impressed.
Mesoplax composed of two triangular
plates that are set at an acute angle to each
other dorsally and have two large, hollow,
tubular projections extending from the
posterior dorsal margin (Plate 10, Figs. 3—
10). Length of dorsal margin less than one
half the length of the plate, outer surface
sculptured with distinct ridges paralleling
the anterior margin; inner surface smooth,
with a small internal opening into the tube
(Plate 10, Figs. 11, 12).
Siphons short, of about equal length,
and probably not capable of extending
more than one half the length of the shell.
Aperture of incurrent siphon large and ap-
parently lacking cirri, that of the excurrent
siphon small and with a few fine cirri on
each side ventrally.
Measiirenients. See Table 6.
Remarks. Knudsen (1961) described
three species of Xylophaga with tubulate
mesoplaxes, and studies for this report
have shown that X supplicata Taki and
Habe also is tubulate. Three of these spe-
cies are from the western Pacific (X. tuh-
ulata Knudsen from Macassar Strait, X.
briiiini Knudsen from the Mindanao Sea,
and X. supplicata Taki and Habe from Ja-
pan), and one from the eastern Pacific (X.
obtusata Knudsen from the Gulf of Pana-
Xijlophaga whoi Turner n. sp. is the
ma)
' An acronym for Woods Hole Oceanographic In-
stitution, whose research vessel Atlantis collected the
holotype.
first tubulate species reported from the At-
lantic. It is most closely related to X. bni-
uni Knudsen from the Mindanao Sea but
differs in that the mesoplax has a propor-
Xylophagainae • Turner
243
tionately shorter dorsal margin and larger
tubes. In addition, the posterior adductor
muscle scar is set well in from the poste-
rior margin and the umbonal-ventral ridge
is evident only near the ventral margin of
tlie shell but increases rapidly to a large
condyle. Xtjlophaga whoi Turner n. sp. dif-
fers from X. obtusata Knudsen in the po-
sition of the posterior adductor scar, the
shape of the mesoplax, the larger tubes on
the mesoplax, the narrower anterior slope,
and smaller umbonal reflection. The third
tubulate species described by Knudsen, X.
tubulata froni Macassar Strait, differs from
X. whoi in having very large tubules that
extend to the anterior margin of the me-
soplax and in having the plates bent lon-
gitudinally at a right angle so they are flat
dorsally rather than meeting at an acute
angle. Examination of a paratype specimen
of X. supplicata Taki and Habe, 1950, from
Japan shows that this species has minute
tubes on the mesoplax similiar to those in
X. briiiini and it may be synonymous with
that species. (See also under X supplicata
Taki and Habe.)
The specimen of X. whoi Turner n. sp.
taken by the RA^ Pillsbiinj from off Punta
Piedras, Colombia, had 13 young attached
to the umbonal area (Plate 11, Fig. 2).
Among the other species in this group
(Group 3), Knudsen (1961) reported that
the type of X obtusata had two young and
the type of X. bruuni had four young on
the dorsal surface of the adult shell,
whereas X. tubulata had five young at the
ventral base of the siphons in a depression
of the mantle tissue. The young at the base
of the siphons of X. tubulata were very
small, about one half the size of those on
X. bixiuni. They possibly had only recently
been extruded from the excurrent siphon
of the adult and had not yet crawled to the
dorsal surface of the adult. It has not been
proven that the juveniles clinging to the
mantle and shells of adult Xylophaga are
definitely the young of the specimen to
which they are attached, and this has been
questioned by some workers. However,
until the species concerned are cultured in
the laboratory, it is the only assumption
that can be made safely. The only way the
young of another specimen or species
could get into the burrow would be via the
incurrent siphon. For the veliger lai'vae of
another specimen to get to the umbonal
area, they either would have to pass
through the gills into the epibranchial
chamber and then out the excurrent si-
phon, or go through the digestive tract to
the excurrent siphon. Neither of these al-
ternatives seems likely. It is, of course, pos-
sible that the young of X. tubulata com-
plete their development at the base of the
siphons of the parent.
Range. From off southern Florida to
Colombia in depths from about 336 to
910 m.
Specimens Examined. UNITED STATES, ELOR-
IDA; Gerda, station 266, off Eowey Rocks (25°39'N,
79°58'W) in 185-187 fathoms (338-345 m). CUBA:
Atlantis, station 3471, off Cardenas, Matanzas Prov-
ince (23°21'N, 80°56'W) in 500 fathoms (914 m).
LESSER ANTILLES: PiUsbunj, station 944, 45 miles
N of Port Louis, Guadeloupe Island (16°32.2'N,
61°36.8'W) in 364-421 m. COLOMBIA: PiUsbunj,
station 394, off Punta Piedras (9°28'N, 76°26'W) in
230-350 fathoms (419-640 m).
Xylophaga profunda Turner new species
Plates 12, 13
Holotijpe. MCZ 316751.
Type Locality. Tongue of the Ocean, off
NE tip of Andros Island, Bahama Islands
(25°54'N, 77°49'W) in 1,722 m. From test
panel submerged from 26 July 1962 to 17
February 1965.
Distinctive Characters. Valves with a
well-impressed umbonal-ventral sulcus,
bounded posteriorly by a low broadly
rounded ridge. Posterior slope sculptured
with narrow, concentric grooves. Mesoplax
of two triangular plates, lacking a basal
portion, set at an acute angle to each other,
folded longitudinally with the anterior
margin bent inward and with a small pore
in each anteriorly. Umbonal-ventral ridge
on inner surface of valves narrow, high,
and strongly segiTiented.
Description. Shell globose, reaching
about 11 mm in length, thin, fragile, and
244 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Table 7. Measurements of
Xylophaga profunda.
Lengtli Height
(mm) (mm)
5.9 5.6 holotype
Tongue of the Ocean, off NE tip of
8.0 7.9 Andros Island, Bahama Islands
Tongue of the Ocean, off NE tip of
9.5 8.8 Andros Island, Bahama Islands
Tongue of the Ocean, off NE tip of
10.0 9.8 Andros Island, Bahama Islands
Tongue of the Ocean, off NE tip of
10.9 10.8 Andros Island, Bahama Islands
with a relatively heavy, light brown perios-
tracum. Pedal gape angle about 95°.
Beaked portion of anterior slope sculp-
tured with numerous strong, denticulated
ridges that are more closely spaced toward
the ventral margin in adults. Umbonal—
ventral sulcus narrow, deep, bounded pos-
teriorly by a low, broad, rounded ridge.
Posterior slope sculptured with widely
spaced, incised grooves. Umbonal i^eflec-
tion narrow, high, and strongly segmented.
Incised grooves of posterior slope ex-
pressed internally as faint ridges just pos-
terior to the umbonal— ventral ridge gi\^ng
a "back bone and ribs effect." Posterior ad-
ductor muscle scar broadly oval, set high
on the posterior slope, and lightly marked
with irregularly anastomosing depressions.
Pedal retractor scar lightly impressed,
dumbell-shaped, and located just anterior
to the midportion of the posterior adduc-
tor. Siphonal retractor scar large, elongate,
lightly impressed, and located about mid-
way between the posterior adductor and
the umbonal— ventral ridge. Chondrophore
large, that of the left valve with two small
teeth on the posterior edge.
Mesoplax composed of two triangular
plates lacking a basal portion, set at an
acute angle to each other, folded longitu-
dinally with the anterior margin bent in-
ward and with a small pore in each ante-
riorly at the outer edge of the fold.
Measurements. See Table 7.
Remarks. This species is superficially
similar to X abijssonini Dall but differs in
having a low, broad, rounded ridge poste-
rior to the umbonal— ventral sulcus, in lack-
ing the deep groove posterior to the um-
bonal— ventral ridge on the inner surface,
in having a pronounced sculpture on the
posterior slope, and in being larger. In ad-
dition, the inesoplax of X. profunda lacks
a ventral portion, has only a single pore in
each plate, does not produce tubes, and
appears narrow when viewed anteriorly.
Xylophaga profunda Turner n. sp. is
probably most closely related to X lohata
Knudsen from the Sulu Sea. Xijlophaga
profunda differs in having a mesoplax that
is much longer than wide, whereas in X
lobata the structure is wider than long.
Both species have a strongly segmented
umbonal— ventral ridge and a similarly
marked posterior adductor scar, although
in X. profunda the markings are more ex-
tensive and elaborate. Knudsen (1961)
compared his X. galatheae from the Tas-
man Sea with X lobata, noting differences
in the mesoplax and muscle scars. Unfor-
tunately, he had only a single complete
specimen of X. galatheae, so he could not
determine range of variation. Xylophaga
galatheae possibly is a young X. lobata but
more material from inteivening areas is
needed to ascertain this.
On the basis of the general shape and
attachment of the dorsal plates, X. profun-
da, along with X. lobata and X. galatheae,
appear to belong in species Group 3, al-
though X. profunda is not closely related
to any species in the group and is placed
here tentatively. The major differences in-
clude the sculpturing of the posterior ad-
ductor muscle scar, the presence of ante-
rior pores, and the lack of posterior tubes
on the mesoplax. The chondrophore of the
left valve of X. profunda has two small
teeth, a character that relates it to X. ivhoi
and X. supplicata, both of which have a
single large tooth. Unfortunately, Knudsen
did not mention the presence of teeth in
the three tubulate species he described.
Consequently, it cannot be stated definite-
ly that this is characteristic of the group.
The siphons of X. profunda are short.
Xylophagainae • Turner 245
probably of the same length and capable
of complete retraction within the valves.
Unfortunately, none of the specimens was
sufficiently well preserved for anatomical
work. Four specimens carried young on
the umbonal area, the smallest had 5 and
the largest had 75. The young have pro-
nounced umbos, distinct concentric sculp-
ture, and measure about 0.30 mm in
length.
Xylophaga profunda Turner n. sp. is
known from five specimens taken from
one panel submerged in the Tongue of the
Ocean at 25°54'N, 77°49'W, and from 14
specimens taken from an asbestos-backed
panel submerged at 24°53.2'N, 77°40.2'W.
Specimens Examined. BAHAMA ISLANDS:
Tongue of the Ocean, off NE tip of Andros Island
(25°54'N, 77°49'W) in 1,722 ni; Tongue of the Ocean,
Tower 3 (24°53.2'N, 77°40.2'W) in 2,066 m.
Xylophaga abyssorum Dall
Plates 14-16
Xylophaga abyssorum Dall, 1886, Bulletin Museum
of Comparative Zoology, 12: 317, pi. 9, fig. 7, 7a
{Blake, station 215, off St. Lucia, Lesser Antilles
[13°51'N, 61°03'W] in 226 fathoms). Holotype,
MCZ 8135; Turner, 1955, Johnsonia, 3(34): 156, pi.
93.
Distinctive Characters. Valves with a
proiTiinent ridge just posterior to the um-
bonal— ventral sulcus and with a slightly to
strongly concave profile posterior to the
ridge when viewed dorsally Umbonal—
ventral ridge on the inner surface devel-
oped only near the ventral condyle and
bounded posteriorly by a deep groove.
Mesoplax composed of two more or less
triangular plates having a ventral portion
and a variously lobed dorsal portion; lobes
varying with age and coalescing to form
pores or tubes (Plate 15, Figs. 4, 5).
Description. Shell globose but appear-
ing constricted posteriorly when viewed
dorsally, reaching 5.5 mm in length, thin,
fragile, and with a thin, light brown per-
iostracum. Pedal gape angle about 95°.
Beaked portion of the anterior slope sculp-
tured with fine, denticulated ridges, widely
spaced in young specimens but compacted
toward the ventral margin in older speci-
mens. Umbonal— ventral sulcus narrow,
rather shallow, and with a narrow promi-
nent ridge just posterior to it. Profile of
valves concave posterior to the ridge when
viewed dorsally. Posterior slope sculptured
with faint concentric growth lines, umbo-
nal reflection erect, the ventral margin of
the mesoplax meeting it and attached by a
periostracal fold. Umbos inflated, particu-
larly in young specimens.
Inner surface of valves white and glazed.
Muscle scars well marked. Posterior ad-
ductor scars somewhat pear-shaped, set
well in from the posterior margin, and
marked with irregular, elongate depres-
sions extending inward from the posterior
margin. Anterior adductor scar covering
most of the umbonal reflection. Pedal re-
tractor scar nearly circular and located just
anterior to the widest position of the pos-
terior adductor scar. Siphonal retractor
scar small, elongate, lightly impressed, and
located anterior and ventral to the pedal
retractor. Umbonal— ventral ridge not well
developed except near the ventral condyle
but bounded posteriorly by a deep groove
that is a reflection of the external ridge.
Chondrophore well developed, internal
ligament strong.
Mesoplax variable, with a well-devel-
oped, more or less triangular basal portion,
occasionally with lateral notches in young
speciniens. Dorsal portion developing and
vaiying with age; in young specimens, con-
sisting of lobes extending anteriorly from
the posterior ends of the plates (Plate 14,
Fig. 2) followed by lobed lateral folds
(Plate 14, Figs. 3-5, 7), which grow and
eventually coalesce to form pores or short
tubes (Plate 15, Figs. 4, 5, and Plate 16,
Figs. 4, 5). Aperture of the tubes in living
specimens covered by a periostracal mem-
brane.
Siphons short, about equal in length,
and capable of complete retraction within
the valves. Diameter of the incurrent si-
phonal aperture about twice that of the ex-
current siphon. The margins of both ap-
pear to have cirri.
246 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Table 8. Measurements of
Xylophaga abyssorum.
Length Height
(mm) (mm)
3.0 2.6 Pillsbury, station 944
3.0 2.9 Tongue of the Ocean, off NE tip of
Andros Island, Bahama Islands
3.3 3.0 Pillsbunj, station 944
3.9 3.2 Pillsbunj, station 944
4.0 3.5 holotype
4.0 3.6 Gerda, station 266
5.4 4.5 PiUsbun/, station 944
Measurements. See Table 8.
Retnarks. Dall based his description of
X abyssorum on two isolated valves of a
dead specimen, with the main distinguish-
ing character being the pronounced ridge
posterior to the umbonal— ventral sulcus.
All specimens that have been assigned to
this species since that time have been so
named on the basis of this ridge. Among
the specimens taken from boards sub-
merged in the Tongue of the Ocean off
Andros Island, Bahama Islands, by the
U.S. NOO (DePalma, 1969), was a single
specimen the valves of which coincided
with those of the holotype. Two specimens
were taken from wood dredged off Gua-
deloupe Island, Lesser Antilles. All of
these specimens have a pronounced ridge,
concave posterior slope, and lobed meso-
plax. With this new material, it has been
possible to redescribe X abyssonnn giving
the characters of the mesoplax and a range
of variation. Plate 14, Figure 6, illustrates
the dorsal view of the right valve of the
holotype of X. abyssorum; Figure 3 illus-
trates the dorsal view of the right valve of
the specimen from the Tongue of the
Ocean. These two valves so closely resem-
ble each other that, except for a slight dif-
ference in size, they could be from the
same specimen. Variations in the valves
and dorsal plates of specimens from Flor-
ida and Guadeloupe Island are illustrated
on Plates 14-16. Plate 15, Figure 5, illus-
trates the position in which the young are
carried on the adult. The largest number
of young on any one specimen was five;
the average measurement of the young
was 0.3 mm in length.
Xylophaga abijssorum has a small shell
but is a distinctive species, particularly in
the adult stage. It is probably most closely
related to X. lobata Knudsen from the
Sulu Sea but differs in having a sharp ridge
posterior to the umbonal— ventral sulcus
and a far more elaborately lobed mesoplax
in the adult. From X. profunda Turner n.
sp., the only other species with lobes on
the dorsal portion of the mesoplax, X
abyssorum differs in having broad dorsal
plates with two or more pores or tubes in
each. In addition, X. profunda is a much
larger species, the umbonal— ventral ridge
on the outer surface of the valves is broad
and rounded, and the deep groove on the
inner surface is lacking. The posterior ad-
ductor muscle scars of X. profunda are
lightly and irregularly marked, whereas the
inuscle scars of X. abyssorum are well
marked. In the young stage, the ixiesoplax
of X abyssorum is similar to that of X. baij-
eri Turner n. sp. and X. profunda, but the
valves of these species do not have a pro-
nounced ridge. The shape of the valves of
X. abyssorum somewhat resembles those
of X. japonica Taki and Habe but the me-
soplax of that species is ear-shaped and not
lobed. (See under X. japonica.)
Range. Based on the valves of dead
specimens lacking a mesoplax, the range of
this species extends from off Atlantic City,
New Jersey, south to St. Lucia, Lesser An-
tilles (Turner, 1955: 157). Living speci-
mens are known only from Florida, the
Bahamas, and Guadeloupe Island, Lesser
Antilles, in depths ranging from 342 to
1,722 m.
Specimens Examined. UNITED STATES, FLOR-
IDA: Gerda, station 266, off Fowey Rocks (25°38'N,
79°58'W) in 185-187 fathoms (338.3-342 m). BA-
HAMA ISLANDS: Tongue of the Ocean, about 4
miles off NE tip of Andros Island (24°54'N, 77°47'W)
in 1,722 m. LESSER ANTILLES: Pillsbunj, station
944, 4.5 miles N of Port Louis, Guadeloupe Island
(16°32.2'N, 61°36.8'W) in 360-420 m.
XylOPHAGAINAE • Turner
247
Table 9. Measurements of
Xylophaga duplicata.
Length
(mm)
Height
(mm)
3.7 3.5 holoty].ie
5.2 5.0 Gulf of Tehuantepec, Me.\ico
5.5 5.0 Gulf of Tehuantepec, Mexico
Xylophaga duplicata Knudsen
Plate 17
Xylophaga duplicata Knudsen, 1961, Galathea Re-
port, 5: 175, figs. 14, 15 {Galathea, station 745,
Gulf of Panama [7°15'N, 79°25'W] in 915 m). Ho-
lotype. Zoological Museum, University of Copen-
hagen.
Distinctive Characters. Plates of the
mesoplax elongate oval, with a large ven-
tral portion, somewhat inflated, diverging
anteriorly, and standing off from the sur-
face of the valves. Anterior adductor mus-
cle extending into the cavity of the meso-
plax. Anterior slope scLilptured with nu-
merous exceedingly fine, closely set den-
ticulated ridges. Umbonal reflection
narrow and erect. Umbonal— ventral sulcus
narrow, slightly impressed, and bounded
posteriorly by a faint, rounded ridge. Pos-
terior adductor muscle scar oval and
smooth. Umbonal— ventral ridge narrow,
high, and segmented; ventral condyle
small. Siphons of equal length, united ex-
cept at the tip, the posterior three-fourths
covered with a brown periostracal sheath;
siphonal openings (apertures set on two
short tubes) each with six to eight cirri.
Measurements. See Table 9.
Remarks. At the time Knudsen de-
scribed this species he had only two spec-
imens from the Gulf of Panama. Two ad-
ditional specimens received from D. Shas-
ky were taken by the San Juan Expedition
at station N-12 from a sunken log dredged
in 60 fathoms (109 m) in the Grflf of Te-
huantepec, Mexico (15°08'N, 93°28'W).
Although the specimens were in rather
poor condition, the valves and siphons
agree with those described by Knudsen.
The mesoplax stands off from the surface
of the shell and, as stated by Knudsen, is
double, that is, has a basal portion, and the
anterior muscle extends into the cavity of
the mesoplax. The siphons are the same
length, combined in a common sheath ex-
cept at the tip, and are covered with a thin,
brown periostracum. The white spots
mentioned by Knudsen are not evident,
but the few cirri surrounding the siphonal
openings are similar.
Range. This record extends the geo-
graphic range of the species about 500
miles to the north. The depth range is
from 109 to 915 m.
Xylophaga muraokai^ Turner new species
Plates 18, 19
Holotype. MCZ 316746; paratypes,
MCZ 316747, 316748, 316749, 316750.
Type Locality. U.S. Naval Civil Engi-
neering Laboratory Test Site I, about 81
miles SW of Port Hueneme, California,
or about 25 miles S of San Miguel Is-
land, Santa Barbara Islands (33°44'N,
120°45'W) in 5,640 ft (1,720 m). The ho-
lotype was taken from a panel exposed on
STU 1-2 from October 1963 to October
1965.
Distinctive Characters. Plates of the
mesoplax wedge-shaped, the basal portion
large and the small dorsal portion covering
only the posterior part of the muscle. Si-
phons smooth, of rmequal length, the ex-
current slightly shorter and with 8-10
proininent cirri.
Description. Shell globose, valves reach-
ing 14 mm in length and 13 mm in height;
the width of apposed valves about 12 mm.
Valves thin but strong, white, with a thin
clear, transparent to bright yellow perios-
tracum. Angle of the pedal gape about
115°. Anterior slope sctilptured with nar-
row, finely denticulated ridges that are
widely spaced during early growth, becom-
ing more closely spaced toward the ventral
margin in older specimens. Holotype with
17 ridges on the anterior slope. Umbonal—
ventral sulcus rather narrow, only slightly
' Named for James Muraoka, Biologist, U.S. Naval
Engineering Laboratory, Port Hueneme, California.
248 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
depressed and sculptured widi fine growth
lines. Disc and posterior slope sculptured
with growth lines only.
Inner surface of valves smooth and shin-
ing. Unibonal— ventral ridge narrow and
marked with distinct transverse ridges that
appear knobby in some specimens. Chon-
drophore and internal ligament well de-
veloped. Posterior adductor muscle scar
large, elliptical, covering most of the pos-
terior slope, and marked with irregular,
faint depressions that generally cross the
short axis of the scar. Pedal retractor mus-
cle scar broadly oval in outline and located
just anterior to and about midway on the
posterior adductor scar. In some speci-
mens, the two scars appear to be adjacent.
Siphonal retractor muscle scars not clearly
defined but located just posterior to the
umbonal— ventral ridge about midway dor-
soventrally. Ventral adductor muscle scar
long, narrow, and traversing the ventral
end of the umbonal— ventral ridge. Ante-
rior adductor muscle scar covering the
umbonal reflection and the ventral flange
of the mesoplax.
Mesoplax small and divided. The ventral
portion is a large, cuived, wedge-shaped
shield, fitting closely against the surface of
the valves. Posteriorly, the mesoplax curves
upw^ard and foi"Avard covering the posterior
portion of the anterior adductor muscle
(Plate 18, Figs. 3-6).
Siphons smooth, united, and capable of
extending over three times the length of
the valves. Excurrent siphon only slightly
shorter than the incurrent siphon. Aper-
ture of the incurrent siphon bordered by
18—22 long, slender cirri that, in preserved
specimens, curl inward and are not easily
seen. At the base of these are numerous
small cirri and, anterior to them on the
inner wall of the siphon, 8—10 broad,
branched cirri. Distal margin of excurrent
siphon with two broad, short cirri on each
side adjacent to the incurrent siphon, and
8—10 long, slender cirri, the longest locat-
ed middorsally, with those on either side
becoming progressively shorter.
The anterior adductor muscle inserts
Table 10. Measurements of
XYLOPHAGA MURAOKAl.
Length
Height
(mm)
(mm)
Location
6.5
6.2
paratype
8.5
8.0
paratype
11.8
12.0
paratype
12.0
11.5
paratype
12.0
11.9
paratype
12.8
12.8
holotype
14.0
13.0
paratype
both on the anterior reflection and on the i
basal portion of the mesoplax. Siphonal re-
tractor muscles straplike, extending ante-
riorly from the base of the siphons along
the midportion and over the outer surface
of the visceral mass to insert on the valves
just posterior to the umbonal— ventral
ridge. Ventral adductor muscle, formed as
a thickened area of the fused mantle mar-
gin, inserts over and to each side of the
ventral condyles. Pedal retractor muscles
pass through the visceral mass and insert
just anterior to the posterior adductor
muscle.
Measurements. See Table 10. I
Remarks. The young of X. muraokai
Turner n. sp. superficially resemble those
of X concava Knudsen, found in the Gulf
of Panama, but differ in having a mesoplax
with a large ventral portion, in having the
excurrent siphon slightly shorter than the
incurrent siphon, and in having a few large
cirri on the excurrent siphon. From X. du-
plicata Knudsen, also from the Gulf of
Panama, X. muraokai differs in lacking the
brown periostracal sheath covering the si-
phons, in having an elongate rather than
nearly circular posterior adductor scar, and
in having the dorsal portion of the meso-
plax reduced, covering only the posterior
portion of the anterior adductor muscle.
As with all Xylophaga known to date,
specimens of X. muraokai show consider-
able variation in size and scLilpture de-
pending on the type of wood in which they
are boring. A specimen boring in green-
heart with valves only 6 mm in length had
45 closely spaced denticulated ridges on
Xylophagainae • Turner
249
the anterior slope, whereas a specimen of
the same size in white pine had only 7
ridges. Although it is impossible to tell the
exact age of these specimens, we know
that the greenheart panel was exposed for
35 months and the white pine panels were
exposed for 24 months. Other specimens
in a white pine panel (exposed on the
same rack for the same length of time as
the greenheart panel) reached 12 mm in
length and had only 24 ridges.
Large numbers of X. muraokai were
taken from panels submerged for 2 years
at 5,640 ft (1,720 m). The openings of the
burrows averaged about 1 mm in diameter
and increased rapidly. When specimens
were uncrowded, the anterior end of bur-
rows that had reached 15 mm in length
averaged 12 mm in diameter; burrows 55
mm long averaged about 15 mm in diam-
eter anteriorly. The tunnels often ran to-
gether and frequently two to six specimens
occupied a large irregular cavity. Appar-
ently, several specimens occasionally used
a single opening to the surface.
Xylophaga muraokai Turner n. sp. is
known only from the USNCEL Test Site
I. It has never been taken from panels sub-
merged at Test Site II, which is just north
of San Miguel Island in about 2,370 ft (722
m). Differences in temperature, dissolved
oxygen content of the water, and hydro-
static pressure between the two sites are
probably the responsible factors. It is im-
possible to say whether or not they are all
of equal importance.
Xylophaga washingtona Bartsch, a spe-
cies of wide geographic and depth range,
has been taken from wood exposed at Test
Site I. It is the only species found at Test
Site II, where it is abundant. For a com-
parison of the test sites, see Table 11. For
a description of the test sites, see Muraoka
(1964, 1965, 1966a, 1966b, 1966c, 1967).
Range. Known only from USNCEL Test
Site I.
Specimens Examined. UNITED STATES, CALI-
FORNIA: USNCEL Test Site I (33°44'N, 120°45'W)
about 81 nautical miles SW of Port Hueneme (about
25 miles S of San Miguel Island, Santa Barbara Is-
Table II. Comparison of the U.S. Naval Civil
Engineering Laboratory test sites (average
values taken from muraoka, 1967).
Locality
Deptli
(fi)
TeinpeiE
ture
(°C)
Dissolved Hydrostatic
Ox\'gen Pressure
(ml/L) (psi)
Test Site I
Test Site II
5,640
2,370
2.5
5.0
1.26
.30
2,482
1,043
lands); STU I-l at 5,300 ft (1615.4 m) e.xposed March
1962-February 1965; STU 1-2 at 5,640 ft (1,720 m)
exposed October 1963-October 1965; STU 1-4 at
6,800 ft (2,072.6 m) exposed June 1964-7 July 1965;
STU 1-5 at 6,000 ft (1,828.8 m) exposed 25 August
1965-12 Febi-uaiy 1966.
Xylophaga atlantica Richards
Plate 20
Xylophaga atlantica Richards, 1942, Nautilus, 56: 68,
pi. 6, fig. 4 (east coast of the United States). Ho-
lotype. Academy of Natural Sciences Philadelphia
178741; Turner 1955, Johnsonia, 3(34): 152-154,
pi. 91, figs. 1—6 (type locality. Mount Desert Island,
Maine). (See Turner, 1955: 153.) [Not Turner,
1954, Johnsonia, 3(33): 5-6, pi. 4 = X. clenchi
Turner and Culliney; see below.]
Distinctive Characters. Mesoplax small,
anterior to and between the umbos, com-
posed of two triangular plates that are in
contact the length of the dorsal inargin
and meet at a broadly obtuse angle. Ven-
tral jDortion of the mesoplax narrow, form-
ing a small posterior cavity into which the
posterior end of the anterior adductor
muscle extends. Umbonal— ventral sulcus
shallow with a median threadlike groove.
Posterior adductor muscle scar elongate
and irregularly marked. Excrn-rent siphon
slightly shorter than the incurrent, the ex-
current aperture with 15—20 large papillae,
incurrent aperture with a double row of
nrmierous minute papillae (Plate 20, Fig.
3).
Remarks. Between 16 June 1964 and 16
July 16 1965, the U.S. NOO submerged
test panels off Mark Island, Penobscot
Bay, Maine, in water 200 ft in depth. The
panels were set in a vertical array at depths
of 50, 100, 150, and 195 ft. Dissection of
these panels showed that the bottoin one
had 85 specimens of X atlantica, the one
250 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
at 150 ft had six specimens, and the one
at 100 ft had only one specimen. This sug-
gests that, hke X. washingtona Bartsch, X
muraokai Turner n. sp., X. depalmai Turn-
er n. sp., and probably all Xijlophaga, the
larvae of this species do not rise very high
in the water column and that the greatest
decrease in attack occurs within a few feet
of the bottom. Probably a panel touching
or partially submerged in the substrate at
this locality would have been heavily at-
tacked. These tests also showed that the
valves of X. atlantica may reach a length
of at least 10.5 mm in a year. Several spec-
imens removed from wood dredged off
Ipswich, Massachusetts, in about 80 m av-
eraged 14 mm in length. The maximum
length for X. atlantica appears to be
15 mm.
The Ipswich specimens were well pre-
served and allow a more detailed descrip-
tion of the siphons than that given in the
paper by Turner (1955) cited above. The
siphons can be extended 1.5—2 times the
length of the valves. The excurrent siphon
is slightly shorter than the incurrent si-
phon and is combined with it for most of
its length. The excurrent aperture is sur-
rounded by 15—20 relatively large papillae
that on the dorsal surface appear to be
grouped in two lobes. The incurrent si-
phon is surrounded by an outer rim of
small papillae and an inner one of slightly
larger, stouter papillae (Plate 20, Figs. 1,
2).
Xylophaga atlantica is oviparous and
does not brood its young. The reproduc-
tion and lai'val development of X. atlantica
are fully discussed by Culliney and Turner
(1976); they detail inethods of laboratory
culture and illustrate the various lai"val and
growth stages. A reexamination of the
specimen reported by Turner (1954: 5—6,
pi. 4) has shown it to be X. clenchi Turner
and Culliney, a species that usually is
found with young attached to the parent.
The adult of X. clenchi differs from that of
X. atlantica in having a mesoplax that lacks
a ventral portion, in the shape and type of
marking on the posterior adductor muscle
scar, in having a broad, rounded ridge pos-
terior to the umbonal— ventral sulcus, and
in the arrangement of the papillae on the
siphons.
On the basis of the material then avail-
able, I formerly considered this species
most closely related to X. washingtona
Bartsch (Turner, 1956). It is now evident
that these species belong to two different
species groups. In X. atlantica, the excur-
rent siphon is only slightly shorter than the
incurrent siphon and cirri surround the
aperture of both siphons (Plate 20, Figs.
1—3). In addition, the posterior adductor
scar is irregularly marked, whereas that of
X. washingtona has regular herringbone
markings.
Range. From Newfoundland south to
Cape Heniy, Virginia, in depths ranging
from about 15 to 1,242 m.
Specimens Examined (New Records Since Turner,
1955). CANADA, NEWFOUNDLAND: Vi miles off
Argentia in 18.3 m (test panel). NOVA SCOTIA:
North Bay, Cape Breton Island (46°20'N, 61°50'W)
in 128 m. UNITED STATES, MAINE: off Mark Is-
land, Penobscot Bay, in 15.2 m, 30.5 m, 45.7 m, and
59.5 m (test panels). MASSACHUSETTS: 15 miles
off Ipswich in 73 m; NE of Cape Cod Light off Truro
in 40 m; E of Nantucket Island (41°23'N, 68°46'W).
Xylophaga washingtona Bartsch
Plate 21
Xylophaga washingtona Bartsch, 1921, Proceedings
of the Biological Society of Washington, 34: 32 (San
Juan Island, Washington). Holotype, U.S. National
Museum (USNM) 334478; Turner, 1955, Johnson-
ia, 3(34): 154, pi. 92; Turner, 1956, Nautilus, 70:
10; Tipper, 1968, Ecological Aspects of Two Wood-
Boring Molluscs from the Continental Terrace Off
Oregon, Thesis, Department of Oceanography,
Oregon State University, pp. 8-13, 64-118.
Distinctive Characters. Mesoplax small,
anterior to and between the umbos, com-
posed of two triangular plates that are in
contact for the length of the dorsal margin
and are set at a moderately obtuse angle
to each other. Ventral portion of the me-
soplax usually greater than one half the
width of the dorsal portion and keeled.
Umbonal-ventral sulcus broad and shal-
low. Posterior adductor muscle scar elon-
gate oval in outline with regular herring-
Xylophagainae • Turner 251
Table 12.
Penetration into wood of Xylophaga washingtona at various depths
and exposure times.
LocalitN'
Exposure
3 miles off Oceanside, California
STU* II (1 and 2), California
STU* I (1-1), California
19 miles off Depoe Bay, Oregon
25 miles off Depoe Bay, Oregon
40 miles off Depoe Bay, Oregon
109
4 months
Turner, 1956
722
6.5, 13.4 months
Muraoka, 1965,
1967
,615-2,066
4, 13, 25, and 35
Muraoka, 1964,
1966a,
months
1966b, 1966c
200
38 and 72 days
Tipper, 1968
500
2 months
Tipper, 1968
1,000
2 months
Tipper, 1968
STU, submersible test unit.
bone markings. Proximal end of the coixi-
bined siphons usually having a thin perios-
tracal sheath. Excurrent siphon one third
to one half the length of the incurrent si-
phon, truncated, and ha\4ng a narrow
ridge extending from each side for a short
distance along the dorsal surface of the in-
current siphon. Excurrent siphonal open-
ing small, located at the end of a short
tube extending between the lateral ridges,
and apparently lacking papillae (Plate 21,
Figs. 2, 3). Incurrent siphonal opening
margined by 15 inwardly extending papil-
lae (Plate 21, Fig. 4).
Remarks. In recent years a great deal
has been added to our knowledge of the
distribution of X. ivashingtona, and inter-
esting obseivations have been made con-
cerning its ecology and variation. The
many specimens taken from the STU pan-
els exposed by the USNCEL off San Mi-
guel Island, California, have shown that a
great deal of variation exists in the shape
and sculpture of the valves as well as the
size of the posterior adductor muscle in
response to the hardness of the substrate
in which they are boring. This is discussed
in the section on Variation. From these
tests we have also learned that lai"val and
adult X ivashingtona can tolerate a wide
range in dissolved oxygen concentration.
They were the only borers taken from the
STU II test site where the concentration
was 0.30 ml/L. The temperature at this site
was 5.0° C (Muraoka, 1965). Based on
adults dissected from new wood exposed
at various sites with known dates and
depth of submergence, it has been possi-
ble to determine the depth at which the
larvae can successfully penetrate. These
data are summarized in Table 12.
A single large specimen was taken in
18.3 m from wood that had been removed
from old street cars and used to make
Hermosa Reef, V2 mile west of Hennosa
Beach, Santa Monica Bay, California. This
is the shallowest record known for the set-
tlement of lai-vae. The apparent scarcity of
specimens suggests that this is the upper
limit of the depth range. Living specimens
have been taken from a fir log dredged
from 73 m off Vancouver Island, British
Colombia. Although no testing has been
done in this area, it is likely that, in these
colder waters, X. washingtona could occur
at shallower depths.
Muraoka (1966c) found that "wood
specimens which were exposed near the
sediment were damaged considerably
more severely than those specimens which
were exposed about 3 feet above the sed-
iment." This, he said, "indicates that the
deep sea borers are very active in large
numbers immediately above the sediment
layer and that their numbers tend to de-
crease in seawater as the distance from the
sediment layer increases." Tipper (1968)
showed that the settlement of lai-vae of X.
ivashingtona off Depoe Bay, Oregon, was
densest in proximity to the sea— sediment
interface and that the drop in borer pen-
etration usually occurred within the first 6
cm upward from the interface. He also
showed that, from the initial penetration
252 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
very close to the sea— sediment interface,
the attack progressed upward with increas-
ing time of exposure. Analyses of these
data suggest that 1) the free-swimming lar-
vae of X. washingtona probably do not rise
more than 10-25 ft above the sea floor; 2)
as the attack increases, competition occurs
for space; and 3) the second crop of larvae,
prevented from settling on the lower levels
because of the activity of the siphons of
specimens already in the wood, are kept
swimming and eventually settle on surfac-
es higher up in the water column.
Tipper (1968) also pointed out that the
depth of penetration decreased with in-
creased density of the wood. Cedar, pine,
fir, and oak panels exposed for 50 days at
a depth of 200 m showed an average pen-
etration of 3.8 mm in cedar, 3 mm in pine,
1.5 mm in fir, and 0.5 mm in oak. De-
crease in depth of penetration also is re-
lated to increase in depth of exposure.
This, he postulated, may be correlated
with lower temperatures resulting in slow-
er growth rates or with the increased den-
sity of the wood from compression at great
depth.
These obseivations agree with those ob-
tained from the tests off southern Califor-
nia (see section on Variation in the Intro-
duction). From the STU samples we found
that the softer the wood (except balsa), the
deeper the Xylophaga penetrated and the
greater was the development of the chim-
ney of fecal pellets. In a cedar panel ex-
posed on STU 1-5 for 6 months, two sets
of X washingtona were found; the earlier
set had tunnels averaging 10—12 mm in
length, and the second set had tunnels av-
eraging 5 mm in length. Another panel on
STU I-l exposed for 35 months had spec-
imens with tunnels reaching 35 mm in
length. These tunnels averaged 10 mm in
diameter at the anterior end; a chimney of
rather coarse fecal pellets lined the pos-
terior 15 mm of the burrow. Considering
the greater depth of the STU panels and
realizing that the rate of growth decreases
with age, these growth rates agree with
those reported by Tipper (1968).
Based on data gathered to date from
test panels, it would appear that X. wash-
ingtona probably breeds throughout the
year and that the entire larval life is spent
in the sea. Young specimens have never
been found attached to the shells of the
hundreds of adults examined.
Range. From Vancouver Island, British
Columbia, south to the Santa Barbara Is-
lands, California, in depths ranging from
20 to 2,000 m.
Specimens Examined (New Records Since Turner,
1955). CANADA, BRITISH COLOMBIA: Satellite
Channel, between Saltspring and Vancouver Islands
in 93 m (C. Carl). UNITED STATES, OREGON:
about 30 miles W of Seaside (46°00'N, 124°48'W) in I
464.5 m; about 50 miles W of Tillamookhead
(45°54'N, 125°09'W) in 1,554.4 m; about 40 miles W
of Silver Point (45°52'N, 124°54'W) in 822.9 m (all
Oregon State University); 19 miles W of Depoe Bay
(44°52'N, 124°54'W) in 200 m; 25 miles W of Depoe
Bay (44°52'N, 124°36'W) in 500 m; 40 miles W of
Depoe Bay (44°52'N, 125°01'W) in 1,000 m (all R.
Tipper). CALIFORNIA: Hermosa Reef, Vi mile off '
Hermosa Beach in 18.3 m (J. Fitch); 3 miles off :
Camp Pendleton pier, Oceanside, in 100 m (F. Snod- .
grass); USNCEL STU II (1 and 2), 75 miles W of:
Port Hueneme or about 5 mi NW of San Miguel I.,
Santa Barbara Islands (34°06'N, 120°42'W) in 722 ni;
USNCEL STU I (1, 4, and 5), 81 miles SW of Port
Hueneme or 25 miles SW of San Miguel Island, San-
ta Barbara Islands (33°44'N, 120°45'W) in 1,524-
2,066 m (both J. Muraoka); Allan Hancock station
1372—41, about % mile E of Empire Landing, Santa
Catalina Island, Santa Barbara Islands (.3.3°25'50"N,
118°24'50'^V) in 84 m (Allan Hancock Foundation);
2 miles off Eel Point, San Clemente Island, Santa
Barbara Islands in 72.2 m (F. Snodgrass).
Xylophaga rikuzenica Taki and Habe
Plate 22
Xylophaga rikuzenica Taki and Habe, 1945, Japanese
Journal of Malacology (Venus), 14: 112 (off Riku-
zen, Honshu, Japan). Holotype, T Habe Collec-
tion; paratype, MCZ 194821. Neoxylophaga riku-
zenica (Taki and Habe) 1950, Illustrated Catalogue
of Japanese Shells, no. 7, p. 46, text-figs. 4, 5.
Distinctive Characters. Mesoplax com-
posed of two inflated triangular plates with
a large basal portion and ventral keel;
plates in contact both dorsally and ven-
trally for the length of the medial margin
(Plate 22, Figs. 5-7). Posterior adductor
muscle scar elongate oval, tapering dorsal-
Xylophagainae • Turner
253
ly, and with regular herringbone markings.
Valves and dorsal plates covered with a
heavy, brown periostracum. Umbonal-
ventral sulcus wide and deep (Plate 22,
Figs. 2, 4).
Remarks. Taki and Habe (1945) briefly
described this species in Japanese but in-
cluded no figures. In 1950, they created
the genus Neoxylophaga with rikuzenica as
the type species, mentioning the small, tri-
angular mesoplax and thick, brown perios-
tracum as generic characters. Turner
(1956) related X rikuzenica to X. wash-
ingtona Bartsch on the basis of the dorsal
plates. A paratype specimen, received
through the kindness of Dr. Habe, shows
that the posterior adductor muscle scar is
similar to that of X washingtona Bartsch,
X praestans Smith, and X. aiirita Knud-
sen, the other species in this group with
simple, unfringed lappets extending from
the truncation of the excurrent siphon. Xy-
lophaga rikuzenica is closely related to but
differs from X washingtona mainly in hav-
ing a heavy, brown periostracum covering
the valves and mesoplax as well as in hav-
ing a much wider, deeper umbonal— ventral
sulcus. Unfortunately, the siphons of X ri-
kuzenica are unknown so comparisons
cannot be made on that basis. However,
all species with regular herringbone mark-
ings on the posterior adductor scar known
to date have similar siphons so X. rikuzen-
ica can be assumed to belong to the same
group. No Xylophaga have been taken
north of Vancouver, British Columbia, but
continuous collecting across the north Pa-
cific may show that X. rikuzenica merges
with X. washingtona. Until this is done,
the two species are being maintained be-
cause they are recognizable and may not
merge.
Range. Known only from off Rikuzen
and Toyama Bay, Honshu, Japan, in depths
ranging from 100 to 700 fathoms (183 to
1,270 m) (Taki and Habe, 1950).
Specimens Examined. JAPAN: off Rikuzen, Hon-
shu in 183 m.
Xylophaga depalmai*^ Turner new species
Plates 23, 24
Holotype. MCZ 316735.
Type Locality. 3.2 miles off Fort Lau-
derdale, Florida (26°04'N, 80°04'W) in
152.4 m (500 ft) in a test panel.
Distinctive Characters. Shell globose,
anterior slope with fine, closely set, den-
ticulated ridges; umbonal— ventral sulcus
shallow and bounded posteriorly by an in-
distinct ridge; posterior slope smooth.
Umbonal— ventral ridge narrow and high.
Muscle scars barely impressed and
smooth. Mesoplax composed of two inflat-
ed, elongate plates that are coiled poste-
riorly, vary in shape, and are occasionally
fused. Excurrent siphon truncate; lappets
on dorsal surface of incurrent siphon with
exceedingly fine serrations.
Description. Shell globose, valves reach-
ing 9.8 mm in length and 9 mm in height,
thin, fragile, and with a thin transparent
periostracum. Pedal gape angle about 95-
105°. Anterior slope sculptured with ex-
ceedingly fine, closely set, denticulated
ridges; specimens 8 mm long had 40—66
ridges. Posterior portion of the anterior
slope rather wide and the ridges extending
to the umbonal— ventral sulcus more
coarsely denticulate. Umbonal— ventral sul-
cus barely impressed, nearly smooth, and
bounded posteriorly by a fine, indistinct
ridge. Disc and posterior slope faintly
marked with growth lines. Posterior slope
with a low, smoothly cui"ved dorsal margin
that is not reflected.
Inner surface of valves smooth and glis-
tening. Umbonal-ventral ridge narrow,
high, and not enlarging at the ventral mar-
gin to form a knoblike condyle. Posterior
adductor muscle scar barely impressed,
rather small, oval, smooth, and set high on
the posterior slope. Pedal retractor scar
dumbbell-shaped or as two adjacent cir-
'^ Named for Jolin DePalma, U.S. Naval Oceano-
grapliic Office, wlio was responsible for the tests off
Fort Lauderdale, Florida; in the Tongue of the
Ocean, Bahama Islands; and in Penobscot Bay,
Maine.
254 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Table 13. Measurements
OF
Xylophaga depalmai.
Length
Height
(mm)
(mm)
Locati(jn
6.0
5.5
off Fort Lauderdale,
Florida
7.6
7.0
off Fort Lauderdale,
Florida
8.0
7.5
holotype
8.2
8.2
off Fort Lauderdale,
Florida
8.5
7.0
off Fort Lauderdale,
Florida
9.8
9.0
off Fort Lauderdale,
Florida
cles just anterior to the posterior adductor
scar. Ventral adductor scar large, elongate,
irregLilarly oval, located posterior to the
umbonal— ventral ridge and paralleling the
ventral margin of the valve. Siphonal re-
tractor scar small, broadly oval, and locat-
ed just posterior to the umbonal-ventral
ridge at the level of the pedal retractor
scar. Anterior adductor scar covering the
umbonal reflection and the lower surface
of the mesoplax. Chondrophore and inter-
nal ligament well developed.
Mesoplax composed of two inflated,
posteriorly coiled plates that may be asym-
metrical, have a well-developed ventral
portion, are longer than wide, with the
medial margin of the plates parallel and
occasionally with the two parts fused
(Plate 24, Figs. 3-17).
Burrow length of specimens with valves
4—8 mm in length varying from 1.5 to 4
times the length of the valves. Posterior
end of the burrows usually with a smooth,
firmly packed chimney composed of fine
wood fragments.
Siphons capable of complete retraction
within the valves. Excurrent siphon trun-
cated just posterior to the valves and lack-
ing cirri. Finely serrated lappets extend
from the truncation along either side of
the dorsal sru'face of the incurrent siphon.
Incurrent siphon with fine cirri surround-
ing the aperture. Base of the siphons cov-
ered by a fine periostracal sheath that ex-
tends along the sides of the siphons and
contains fine, irregular, glasslike granules
(Plate 23, Figs. 1, 4, 5).
Measurements. See Table 13.
Remarks. This species is most closely re-
lated to X. guineensis Knudsen from the
Gulf of Guinea, West Africa. Xylophaga
depalmai differs in being larger, having an
elongate mesoplax, the posterior end of
which fits between the umbos, and in hav-
ing more coarsely serrated lappets on the
siphons. (See also Remarks under X gui-
neensis.) From X. tipperi Turner n. sp., to
which it is also related, X. depalmai differs
in having an elongate inflated mesoplax
that fits between the umbos rather than a
broad flattened one that covers them, in
having finely serrated lappets on the si-
phons, in having the granules on the si-
phons only in the periostracum rather than
being embedded in the tissue, and in pro-
ducing a chimney. From X. mexicana Dall,l
which it also somewhat resembles, X. de-
palmai differs in the shape of the meso-
plax, in having narrower, more finely
fringed lappets on the siphons, an incon-
spicuous ridge posterior to the umbonal-
ventral sulcus, and an exceedingly thin
periostracum. In addition, the granular in-
clusions on the siphons of X mexicana are
coarse, chalky, white, and arranged in a,
single line extending nearly the length of
the siphons.
Variation exhibited in the mesoplax of X.
depalmai is unusual and unexpected, for
in most species of Xylophaga the mesoplax
is remarkably uniform vmless the specimen
is injured or stenomoiphic. The extreme
forms (Plate 24, Figs. 11-15) are suffi-
ciently different to be considered of spe-
cific value in this genus. However, in the
more than 300 specimens obtained from
the test site off Fort Lauderdale, connect-
ing forms exist between them. The char-
acters of the shell, the siphon, and soft'
parts are similar in all specimens. Conse-
quently, because the large majority are
typical (Plate 24, Figs. 16, 17), the ex-.
tremes are considered to be variants.
Xylophaga depalmai Tm^ner n. sp. was
the most common species occm-ing in the
Fort Lauderdale test. It ranged from 1 to
3.2 miles off shore and occurred in boards
placed near the bottom in depths from 100
Xyloph AGAIN AE • Turner
255
Table 14. Data for all of the specimens of Xylophaga depalmai n. sp. found in the collecting
panels off fort lauderdale, florida.
Distance
From
Bottom
Dentil ol
No.
Shore
Deptli
Panel
Date
ol
Dati
of
.Sj5eeimens
Examined
Panel no.
(miles)
in Feet (i
i) in Feet (m)
Subniei
genee
Rem
jval
2 panels without
2.5
300
on bottom
Sept. 1961
June 1962
±250
numbers
(91.44
W44
1.8
100
on bottom
Jan. 1964
Oct. 1964
1
(30.48
H50
2.3
200
100
Jan. 1964
Oct. 1964
2
(60.96
(30.48)
H56
2.3
200
on bottom
Jan. 1964
Sept. 1964
5
(60.96
G62
2.6
300
200
Jan. 1964
Sept. 1964
6
(91.44
(60.96)
G68
2.6
300
on bottom
Jan. 1964
Sept. 1964
31
(91.44
J59
2.3
200
on bottom
24 Jan.
1964
4 Jan. 1965
3
(60.96
492
3.2
500
(152.4
200
(60.96)
20 Oct.
1965
13 Oct.
1965
3
493
3.2
500
(152.4
200
(60.96)
20 Oct.
1965
13 Oct.
1966
10
497
3.2
500
(152.4
300
(91.44)
20 Oct.
1965
13 Oct.
1966
5
498
3.2
500
(152.4
300
(91.44)
20 Oct.
1965
13 Oct.
1966
7
500
3.2
500
(152.4
400
(121.92)
20 Oct.
1965
13 Oct.
1966
3
501
3.2
500
(152.4
400
(121.92)
20 Oct.
1965
13 Oct.
1966
2
502
3.2
500
(152.4
400
(121.92)
20 Oct.
1965
13 Oct.
1966
3
504
3.2
500
(152.4
on bottom
20 Oct.
1965
13 Oct.
1966
8
505
3.2
500
(152.4
495
20 Oct.
1965
13 Oct.
1966
6
to 500 ft (30.5 to 152.4 m). In addition, a
few specimens were found in boards 100,
200, and even 300 ft off the bottom, which
means that, in this species at least, the lar-
vae range well up in the water column. As
can be seen from Table 14, panels on the
bottom in 300 ft (91.4 m) were the most
heavily attacked. This may reflect cyclic
variation in population density because the
panels were exposed at different times, or
that, because of the configuration of the
bottom and the currents, inore wood was
available on the bottom in this vicinity,
supporting a native population and a ready
source of larvae. However, this also may
indicate that this is the optimum depth for
this species.
According to DePalma (1969), the salin-
ity at the test site was uniformly high and
the temperature of the water ranged from
24 to 30° C at the surface, from 20 to 25°
C at 100 m, and from 7 to 9.5° C at 153
in. The northward-flowing Florida Current
averaged 1.5 knots.
Range. From Florida north to Massa-
chusetts in depths ranging from about 30
to 174 m.
Specimens Examined. UNITED STATES, FLOR-
IDA: off Fort Lauderdale (26°04'N, 80°04'W). See
Table 14. Additional records include: FLORIDA: 160
256 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
miles W of Tampa, OTEC station 3089 in 126 ni; and
OTEC station 3086 in 96 m. RHODE ISLAND:
From rosewood panels tied to lobster pot set at
39°57'N, 69°19'W in 45.7 m (Knutton, from Al Ea-
gles ship Reliance). MASSACHUSETTS: Delaware
II, station 369 (42°37'N, 66°27'W) from wood
dredged from 174 m.
Xylophaga guineensis Knudsen
Plates 25, 26
Xylopliaga guineensis Knudsen, 1961, Galathea Re-
port, 5: 195-196, fig. 38 (Galathea, Station 52, off
West Africa, 01°42'N, 07°51'E in 2,550 m). Holo-
type. Zoological Museum, University of Copenha-
gen.
Distinctive Characters. Valves globose,
with exceedingly fine, closely spaced, den-
ticulated ridges; umbonal— ventral sulcus
barely impressed, bounded by a weak an-
terior ridge and a well-marked, narrow
posterior ridge. Mesoplax composed of
two inflated cornucopialike plates.
Description. Valves globose, reaching
2.2 mm in length, thin, and fragile. Pedal
gape angle about 90°. Anterior slope sculp-
tured with exceedingly fine, evenly spaced,
denticulated ridges, there being about 45
in a specimen 2 inm long. Umbonal— ven-
tral sulcus only slightly impressed, bound-
ed anteriorly by a low, inconspicuous,
rounded ridge and posteriorly by a narrow,
well-defined ridge, occasionally almost
bladelike. Disc and posterior slope
smooth.
Inner surface of valves smooth and glis-
tening. Muscle scars barely visible. Poste-
rior adductor scar smooth, lobed anteriorly
with the small pedal retractor just anterior
to it in the upper embayment. Umbonal—
ventral ridge and condyle prominent, with
a parallel groove posterior to the ridge.
This groove is the internal expression of
the ridge bounding the sulcus on the outer
surface.
Mesoplax composed of two smooth, in-
flated, cornucopialike plates, the horns of
which curl toward each other (Plate 25,
Figs. 3, 6, 8, 9); the two halves usually
equal and mirror images but occasionally
unequal and varying in size and shape.
Siphons about equal to the length of the
Table 15. Measurements of
Xylophaga guineensis.
Length
Height
(mm)
(mm)
Location
0.9
Pillsbury, station 260
1.3
1.2
Pillsbunj, station 260
1.3
1.5
paratype (estimated from Knudsen,
1961, fig. 38b)
1.5
1.3
Atlantiqiie Sud, station 33
2.2
2.2
Atlantique Sud, station 33
shell, capable of complete retraction with-
in the valves. Excurrent siphon truncated,
about one third the length of the incurrent
siphon, with narrow and minutely serrated
lappets extending from the truncation
along the dorsal surface of the incurrent ,
siphon. Base of siphons covered by a filmy'
periostracal sheath with fine, irregular,
whitish granules imbedded in it along the
sides of the siphons (Plate 25, Figs. 1, 2).
Measurements. See Table 15.
Remarks. When Knudsen described X
guineensis he had only "very fragmented
shells of 5—6 individuals removed froin a
piece of wood; not a single complete shell I
present." One specimen had a complete
mesoplax although only the umbonal area '■
of the valves was present. The species was
diagnosed on the basis of the ixiesoplaxi
and the large sculptured, bright yellow i
prodissoconch.
In the course of examining wood
dredged from off the western coast of Af-
rica, I obtained four dead specimens, one
with a complete mesoplax {Atlantique Sud,
station 33), and seven small living speci-
inens, two complete {Pillsbury, station
260), all of which appear to be this species.
The two plates of the mesoplax of these
specimens are mirror images of each other
(Plate 26, Figs. 2-4). As shown under X.
depalmai (Plate 24, Figs. 11, 12), the me-
soplax is not always bilaterally syinmetrical.
Therefore, these new specimens from off
the western coast of Africa are being con-
sidered X. guineensis and a description of
the species based on entire specimens is
given.
Xyloph AGAIN AE • Turner
257
Although stenomorphs are commonly
found in Xijlophaga (see section on Vari-
ation), the specimens dissected from the
wood dredged by the Atlantique Sud Ex-
pedition and the Fillsbunj did not appear
to be stunted or malformed in any way
They were not crowded and the wood was
not particularly hard. Consequently, on the
basis of the large number of evenly spaced
ridges on the anterior slope, the well-de-
veloped inesoplax, the strength of the
ridge bounding the umbonal— ventral sul-
cus posteriorly, and the small size of the
specimens, I consider this to be a small
species.
All the major characters of the siphons
mentioned in the description could be
seen with a dissection scope at 250 X.
However, the fringe on the lappets could
only be detected under a compound mi-
croscope at 430 X. It was impossible to de-
termine whether or not the fringe extend-
ed the entire length of the lappet on the
two specimens that had extended siphons.
The siphons appear similar to those of X
depalmai.
Xylophaga guineensis is most closely re-
lated to X. depalmai taken from waters off
Florida and the Bahamas. Xijlophaga gui-
neensis differs in being smaller; in having
a smooth mesoplax composed of inflated,
tubular, cornucopialike plates; in having
more numerous and closely spaced, den-
ticulated ridges on the anterior slope when
comparing specimens of equal size; and in
having a larger, more prominent, strongly
sculptured prodissoconch. (See also Re-
marks under X. depalmai. )
The specimens that Knudsen described
were taken from wood dredged in 2,550 m
but the three lots of new material were
from 147, 145, and 46 m. Although X. gui-
neensis may extend into deep water, it is
probably a fairly shallow water species,
and this would agree with the known
depth range of X depalmai, the species to
which it is most closely related. The fact
that Knudsen's specimens were all dead
and fragmented further substantiates the
possibility that the wood was carried into
deeper water after it was invaded by X.
guineensis. The continental shelf in this
area is narrow and such a movement of
wood on the bottom could easily take
place.
Range. Gulf of Guinea, West Africa, in
depths ranging from 46 to 147 m (living
material) and in 2,550 m (dead).
Specimens Examined. CAMEROONS: Pilhhitrtj,
station 260, off Santa Isabel Island (3°45'N, 9°05'E)
in 46 m. GABON: Atlantique Sud, station 146, about
46 miles NNE of Port Gentil (0°0.3'S, 9°07'E) in 147
m; Atlantique Sud, station 33, 35 miles W of Ambri-
zette (7°16'S, 12°17'E) in 145 m.
Xylophaga mexicana Dal I
Plates 27, 28
Xylophaga mexicana Dall, 1908, Bulletin Museum of
Comparative Zoology, 43(6): 425 {Albatross, station
3422, off Acapulco, Mexico [16°47'N, 99°59'W] in
141 fathoms [257.9 m]). Holotype USNM 122947;
Turner, 1955, Johnsonia, 3(34): 150, pi. 90.
Distinctive Characters. Umbonal— ven-
tral sulcus bounded posteriorly by a nar-
row, shaij) ridge. Mesoplax ear-shaped, in-
flated, the two halves not meeting medially
except at the posterior end. Excurrent si-
phon truncated; lappets extending along
the dorsal surface of the incurrent siphon
finely fringed. Siphons with a single row of
opaque white granules imbedded along
the sides. Posterior quarter of the incur-
rent siphon papillose.
Description. Shell globose, valves reach-
ing 10.5 mm in length and 9.5 mm in
height, thin, fragile, and with a thin, light
straw-colored periostracum on the poste-
rior slope. Pedal gape angle about 110°.
Anterior slope with exceedingly fine, close-
ly set, denticulated ridges. Umbonal-ven-
tral sulcus rather narrow, moderately deep,
and bounded posteriorly by a narrow,
sharp ridge. Disc and posterior slope
sculptured with fine growth lines. Poste-
rior margin broadly rounded. Posterior
slope becoming proportionately more
elongate with increased size and age, as
indicated in the measurements. Umbonal
reflection free and recui-ved anteriorly, ap-
258 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
pressed posteriorly, and covering the an-
terior portion of the umbonal area.
Inner surface of the valves white,
smooth, and glazed. Umbonal-ventral
ridge narrow, slightly segmented, and
broadening only slightly toward the ventral
margin to form the condyle. Posterior ad-
ductor muscle scar well impressed,
smooth, broadly oval in outline, set high
on the posterior slope and at the posterior
margin of the valve. Pedal retractor scars
nearly circular, double, and adjacent to the
embayment on the anterior margin of the
posterior adductor scar. Ventral adductor
scar large, irregularly oval in outline, and
well posterior to the umbonal-ventral
ridge. Siphonal retractor scar small, bi-
lobed, and located posterior to the um-
bonal-ventral ridge at a level correspond-
ing to the pedal retractor scars. Anterior
adductor scar covering the umbonal re-
flection. Chondrophore and internal liga-
ment well developed.
Mesoplax somewhat variable but usually
slightly longer than wide, with the medial
margin of the two halves diverging. Plates
ear-shaped, inflated, coiled posteriorly, and
covering the umbos.
Siphons capable of complete retraction
within the valves and of extension to at
least 1.5 times the length of the valves. Ex-
current siphon truncated, about one third
the length of the incurrent siphon, and
lacking cirri. Lappets extending from the
truncation of the excurrent siphon finely
fringed. Posterior portion of the incurrent
siphon papillose; inner edge of siphonal
opening margined with numerous minute
cirri. Siphons with thin periostracal sheath
posteriorly and with a single row of white,
opaque, irregular granules embedded
along the sides.
Measurements. See Table 16.
Remarks. Xylophaga mexicana Dall was
described on the basis of two right valves
that were in rather poor condition and
lacked a mesoplax. Consequently, it has
been impossible up to now to relate it to
other species. Valves of living specimens
taken off California agree with those of the
Table 16. Measurements of
Xylophaga mexicana.
Length
Height
(mm)
(mm)
Locatioi 1
4.8
4.5
Malibu Reef, California
5.0
4.2
holotype
7.5
7.3
Malibu Reef, California
8.0
7.3
Santa Monica Reef, California
9.0
9.5
Hermosa Reef, California
10.5
9.0
Santa Monica Reef, California
holotype of X. mexicana. Dall's specimen
was from much deeper water (141 fath-
oms) than the Californian series (about 19
fathoms) but it could have been advecti-
tious at that depth. Xylophaga mexicana
Dall could, of course, be considered a no-
men duhium but this would necessitate de-
scribing the Californian specimens. Be-
cause the valves of these specimens resem-
ble the holotype of X mexicana sufficiently
well to carry the name, it seems best to
establish that taxon firmly on the basis of
this new material.
The size of the beaks, the smooth mus-
cle scars, and the narrow, rather deep um-
bonal-ventral sulcus with a pronounced
ridge at its posterior margin relate X mex-
icana to Xylophaga glohosa Sowerby from
Chile (Turner, 1955). Xylophaga mexicana
differs from X. globosa in having a much
finer fringe on the lappets, in having a se-
ries of irregular, opaque granules along the
sides of the siphons, and in having the pos-
terior portion of the incurrent siphon pa-
pillose. In addition, the mesoplax of X.
mexicana is proportionately much smaller
than that of X. globosa, and the two plates
diverge medially. In his original descrip-
tion, Dall (1908) did not relate X. mexi-
cana to any other species in the genus.
One year later (Dafl, 1909), he gave the
range of X. globosa Sowerby as from Pan-
ama to Chile but cited no specific locali-
ties. The Panamanian specimens on which
he based this range may well have been X.
mexicana. I have not seen a specimen of
X. glohosa from north of Chiloe Island,
Chile. The valves of X. cluplicata some-
Xylophagainae • Turner
259
what resemble those of X rnexicana; how-
ever, the siphons of these two species
place them in different groups. (See also
Remarks under X. tipperi Turner n. sp.
and X japonica, the species to which X.
rnexicana is most closely related.)
Through the kindness of John Fitch and
Charles Turner of the California State
Fisheries Laboratory, Terminal Island,
California, I received 55 specimens of X.
rnexicana taken from test panels exposed
on the "replication reef experiments" con-
ducted by that laboratory. Old streetcars,
automobile bodies, concrete shelters, and
rocks were dumped offshore in several lo-
calities to sei've as settling areas for marine
organisms and to make the area more at-
tractive to fish. Within a short time, the
test panels and wooden portions of the
streetcars were attacked by teredinids and
Xijlophaga. Because all the specimens had
been dissected from the wood, it is im-
possible to say whether or not they pro-
duced a chimney
Range. Living specimens known only
from Santa Monica Bay, California, in
about 35 m.
Specimens Examined. MEXICO: Bocochibampo,
Sinaloa (dead). UNITED STATES, CALIFORNIA:
All from "replication reefs" off Hermosa, Malibu, Re-
dondo, and Santa Monica (about 33°50'N, 118°30'W)
in about 35 m.
Xylophaga tipperi Turnev new species
Plate 29
Holotijpe. MCZ 316736; paratype, MCZ
316737.
Type Locality. 3.2 miles off Fort Lau-
derdale, Florida (26°04'N, 80°04'W) in
152.4 m (500 ft) in a U.S. NOO test panel,
submerged from October 1965 to October
1966.
Distinctive Characters. Umbonal— ven-
tral sulcus bounded by low rounded ridg-
es. Mesoplax ear-shaped, slightly longer
than wide, compressed, and with a sharp
peripheral keel. Excurrent siphon truncat-
ed; lappets on dorsal surface of incurrent
siphon with a coarse fringe. Siphons with
a single row of minute glasslike plaques
embedded along the side and with knobby
pustules at the posterior end.
Description. Shell globose, reaching 9
miTi in length and 8.5 mm in height, thin,
fragile, with prominent umbos and a dull,
light brown periostracum. Pedal gape an-
gle about 110°. Anterior slope sculptured
with numerous close-set, denticulated
ridges, there being 25 on the holotype.
Umbonal— ventral sulcus slightly de-
pressed, bounded by a threadlike rounded
ridge anteriorly and a somewhat heavier
one posteriorly. Posterior slope low and
sculptured with distinct growth lines. Um-
bonal reflection free for most of its length,
the ventral edge of the mesoplax fitting be-
neath it anteriorly.
Inner surface of the valves smooth and
glistening. Umbonal— ventral ridge very
prominent, nearly sinooth, and not greatly
enlarged ventrally at the condyle. Chon-
drophore and internal ligament well de-
veloped. Posterior adductor muscle scar
oval, tapering dorsally, only slightly im-
pressed, and smooth. Pedal retractor scar
kidney-shaped and located adjacent to the
anterior margin of the posterior adductor
scar, about midway dorsoventrally. Siphon-
al retractor scar lightly impressed, located
just posterior to the umbonal— ventral ridge
at the level of the ventral margin of the
beak. Ventral adductor scar usually not vis-
ible but located near the ventral margin
posterior to the umbonal— ventral ridge.
Mesoplax large, ear-shaped, covering
the umbos, longer than wide, compressed
dorsoventrally, with a shaq? peripheral
keel, and a coiled early portion.
Siphons united, excurrent siphon trun-
cated, about one third the length of the
incurrent siphon. Lappets extending from
the truncation along the dorsal surface of
the incurrent siphon with a coarse fringe.
Incurrent siphon papillose posteriorly,
with a single row of glasslike plaques im-
bedded along the side at the juncture of
the lappets and the excurrent siphon and
with numerous small, broad papillae sur-
rounding the inner rim of the aperture.
260 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Table 17. Measurements of Xylophaga tipperi.
Lens;tli
Height
(iTini)
(mm)
Location
2.0
2.0
off Fort Lauderdale,
Florida
4.6
4.1
off Fort Lauderdale,
Florida
7.5
7.0
off Fort Lauderdale,
Florida
8.2
7.5
off Fort Lauderdale,
Florida
9.0
holotype
Burrows reaching 20 mm in length. Fe-
cal pellets not formed into a chimney.
Measurements. See Table 17.
Remarks. This species is closely related
to X. mexicana Dall from California but
differs in having a compressed, sharply
keeled mesoplax; in having glasslike rather
than white, chalky plaques imbedded in
the siphons; and in having much more
coarsely fringed lappets on the siphons.
Xylophaga tipperi differs from Xylophaga
dorsalis Turton in having a definitely
sculptured mesoplax, an excurrent siphon
one third to one half the length of the in-
current (in X. dorsalis it is truncated at the
posterior end of the valves), and in having
glasslike plaques imbedded in the walls of
the siphons. In addition, on the basis of
the limited material now available, X. tip-
peri apparently does not build a chimney.
Xylophaga tipperi differs from X. depahnai
Turner n. sp., also from off Fort Lauder-
dale, in having a compressed, sharply
keeled mesoplax, a much coarser fringe on
the lappets, and in having glasslike plaques
imbedded in a single line along the sides
of the siphons. In addition, the posterior
end of the incru-rent siphon of X. tipperi
is papillose. Xylophaga hayeri Turner n.
sp., which also is found in Florida and the
West Indies, differs from X. tipperi and all
other species in Group 6 in having a
broad, horizontal mesoplax (Plate 31, Fig.
3).
Only eight specimens of X tipperi Turn-
er n. sp. were obtained. They were all
from panel 505 and were collected along
with X. depahnai Turner n. sp. The spec-
imens, even the smallest, were consistent
in having a broad, compressed mesoplax as
well as in the characteristics of the si-
phons. For data on the testing site, see in-
formation under X. depahnai.
Range. Known only from off Fort Lau-
derdale, Florida, in 152.4 m.
Specimens Examined. UNITED STATES, FLOR-
IDA: 3.2 miles off Fort Lauderdale (26°04'N,
80°04'W) in 152.4 m (500 ft) in a test panel.
Xylophaga bayerr Turner new species
Plates 30, 31
Holotype. MCZ 316738; paratype, MCZ
316739.
Type Locality. U.S. NOO test site, about
3.2 miles off Fort Lauderdale, Florida
(26°04'N, 80°04'W) in 152.4 m (500 ft).
From test panels submerged froixi October
1965 to October 1966.
Distinctive Characters. Posterior margin
of the umbonal-ventral sulcus bounded by
a pronounced, sharp ridge. Mesoplax of
adult much broader than long, extending
laterally as wings and with a sharp periph-
eral margin. Excurrent siphon truncated,
about one third the length of the incurrent
siphon. Lappets extending along the dorsal
surface of incurrent siphon finely fringed.
Description. Shell globose, reaching 8
mm in length, thin, fragile, with prominent
umbos and a heavy, golden-brown perios-
tracum covering the shell and the meso-
plax. Pedal gape angle about 120°. Beaked
portion of anterior slope sculptured with
numerous, closely set, denticulated ridges,
there being 32 on the holotype. Posterior
portion of the anterior slope narrow, the
ridges very closely packed. Umbonal re-
flection broad, closely appressed over the
umbos, free anteriorly, and with a funnel-
like pit beneath. Umbonal-ventral sulcus
moderately impressed, sculptured with
growth lines, and bounded posteriorly by
a pronounced, shaip ridge. Posterior slope
rather low and sculptured with well-
marked growth lines.
" Named for Frederick M. Bayer, Rosenstiel School
of Marine and Atmospheric Sciences, University of
Miami, Miami, Florida, who kindly loaned material
collected by the research vessels Pillsbunj and Gerda.
XylophaGAINAE • Turner
261
Table 18. Measurements of Xylophaga bayeri.
Length
Height
{mm)
(mm)
Location
8.0
7.5
holotyj:
)e
4.2
4.0
off Mona Island, Puerto Rico
5.4
4.9
Gercla,
station 266
6.0
5.2
Gerda.
station 266
6.0
5.8
Gerda,
station 266
6.2
6.0
Gerda,
station 266
6.8
6.0
Gerda,
station 266
8.0
S.2
Gerda,
station 266
Inner surface of valves smooth and glis-
tening. Umbonal— ventral ridge very prom-
inent and distinctly segmented but not
greatly enlarged ventrally at the condyle.
Chondrophore and internal ligament well
developed. Posterior adductor muscle scar
elongate oval in outline, tapering dorsally,
and smooth. Pedal retractor scar broadly
oval and contiguous with the anterior mar-
gin of the posterior adductor. Ventral ad-
ductor scar large, located adjacent to the
groove margining the umbonal— ventral
ridge posteriorly. Siphonal retractor scar
lightly impressed and located just posterior
to the umbonal— ventral ridge on a level
with the ventral margin of the beak. An-
terior adductor scar well marked and cov-
ering most of the umbonal reflection.
Mesoplax in young specimens com-
posed of a large ventral plate with a small,
triangular dorsal portion (Plate 31, Fig. 5).
Dorsal plate of the adult is much wider
than long, with a sharp perphery (Plate 31,
Figs. 2, 3) and pronounced concentric
sculpture.
Siphons capable of complete retraction
within the valves and of extension proba-
bly not more than the length of the valves.
Excurrent siphon truncated, about one
third length of the incurrent siphon. Lap-
pets extending from the truncation along
the dorsal surface of the incurrent siphon,
finely fringed. Periostracal sheath, if pres-
ent, extremely thin and lacking calcareous
or glasslike inclusions.
Measurements. See Table 18.
Remarks. Xylophaga bayeri Turner n.
sp. is a distinctive species apparently not
closely related to any other species in this
genus. Its smooth muscle scars, ear-shaped
mesoplax, truncated excurrent siphon, and
fringed lappets place it in Group 6 and the
lack of granular inclusions in the perios-
tracal sheath of the siphons places it more
closely to X. globosa and X dorsalis than
the others in the group. The mesoplax of
X. bayeri is basically like that of X. dorsalis
but is greatly extended laterally.
Only two specimens were obtained from
the panels placed off Fort Lauderdale,
Florida, and unfortunately these were not
sufficiently well preserved for detailed an-
atomical work. The 14 specimens taken
from wood dredged off Fowey Rocks were
in situ, but all were dead and the soft parts
had disintegrated, although the mesoplax
was in place. All had typical valves but
some had unusually wide dorsal plates
(Plate 31, Figs. 2, 3). Four complete spec-
imens and about 25 dead and disarticulat-
ed specimens were extracted from the
wood dredged off Mona Island. These
were remarkably uniform in the characters
of the shell and dorsal plates.
Range. From off the coast of Florida
probably throughout the Caribbean, in
depths ranging from 150 to 365 m.
Specimens Examined. UNITED STATES, FLOR-
IDA: 3.2 miles off Fort Lauderdale (26°04'N,
80°04'W) in 152.4 m (500 ft) in test panel; Ge7'da,
station 266, off Fowey Rocks (25°39'N, 79°58'W) in
about 340 ni. PUERTO RICO: Johnson-Smithsonian
Expedition, station 37, midway between Mona and
Desecho islands (18°11'55"N, 67°42'50'W) in about
365 m.
Xylophaga japonica Taki and Habe
Plate 32
Xylophaga japonica Taki and Habe, 1950, Illustrated
Catalogue of Japanese Shells, No. 7, p. 45, text-
figs. 6, 7 (Tosa Bay, Shikoku, Japan, in about 100
fathoms). Holotype, T. Habe Collection; paratype,
MCZ 194822.
Distinctive Characters. Posterior slope
elongate in adult specimens. Umbonal—
ventral sulcus well impressed, bounded by
an inconspicuous ridge anteriorly and a
high narrow ridge posteriorly. Posterior
adductor muscle scar smooth. Mesoplax
262 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Table 19. Measurements of
Xylophaga japonic a.
Lengtli Height
(mm) (mm)
13.2 10.8 holotype (from Tald and
Raise, 1950)
5.8 5.0 paratype Tosa Bay, Shikoku,
Japan
3.8 3.0 off Clara Island, South Burma
2.0 1.8 off Java
large, covering the upper one half of the
anterior area, extending over the umbos,
ear-shaped, moderately inflated, with a
narrowly rounded peripheiy and distinct
concentric sculpture (Plate 32, Figs. 2-5).
Siphons capable of extending two to three
times the length of the shell. Excurrent si-
phon about one third the length of the in-
current, lappets extending along the dorsal
surface of the incurrent siphon finely
fringed, opaque white granules imbedded
along the sides of the siphons. Chimney of
coarse fecal pellets lining the burrow.
Measurements. See Table 19.
Remarks. At the time Taki and Habe
(1950) described this species, they did not
have the soft parts and therefore could not
describe the siphons. However, it is clear
from their description and figures that the
species belongs to the dorsalis group
(Group 6). Taki and Habe related their
species to X indica Smith, but noted its
more elongate posterior area. It further
differs from X indica in its broad meso-
plax for, according to Knudsen (1961), the
mesoplax of X. indica is "oblong pear-
shaped" and his figure [of it] shows it to
be nearly three times as long as wide. Xy-
lophaga indica is probably more closely re-
lated to X. guineensis and X. depalmai n.
sp., but because the siphons are unknown,
this cannot be stated definitely.
Xylophaga japonica appears to be most
closely related to X. mexicana Dall but dif-
fers in having a larger, more highly sculp-
tured mesoplax that extends over the um-
bos; in having the posterior slope more
elongate; and in having the ridge bounding
the umbonal— ventral sulcus inore pro-
nounced.
Unfortunately, the material from off
Java and South Burma noted in the re-
cords below was poorly presei'ved, but it
was possible to ascertain the basic char-
acters of the siphons. The specimens from
Java had been kept in alcohol since they
were collected by Mortensen in 1929. The
specimens from South Burma, dredged by
the Anton Bniun and taken from a small
piece of wood and the husk of a nut, had
been allowed to dry.
Range. From Tosa Bay, Japan, south and
west to Burma and Java, in depths ranging
from 183 to 384 m."
Specimens Examined. JAPAN: Tosa Bay, Shikoku, p
in 183 m. JAVA: Danish Java-South African Expedi-
tion, station 10, off SE tip of Java (08°36'S, 114°34'E)
in 300 m. BURMA: Anton Bniun, station 23, about
77 miles W of Clara Island, South Burma (10°30'N,
96°35'E) in 3S4 m. I
GENUS XYLOPHOLAS TURNER 1972
XylopJiolas Turner, 1972, Basteria, 36(2): 97-99.
Tijpe Species. Xylopholas altenai Turner,
original designation. I
Distinctive Characters. Valves and me- f
soplax typical for the genus Xylophaga. \
Animal long, not capable of retraction
within the valves, with a periostracal si-
phonal sheath posterior to the valves and |
a pair of lateral, chitonlike, siphonal plates
at the posterior end. Siphons short, ex- i
tending between the plates, the siphonal
retractor muscles inserted on their inner |
surface. Gills and visceral mass contained
between the valves as in typical Xylophaga. j
Wood-storing cecum large. i
Remarks. This genus differs from Xylo-
phaga in having extended excurrent and
incurrent canals contained in a common j
sheath with siphonal plates at the posterior
extremity. These plates probably arose in-
dependently but may be homologous with i
the siphonoplax of other pholads (i.e., Pho- j
ladidea), which was carried posteriorly as i
the animal elongated. However, the si- I
phonal retractor muscles of these species
insert on the valves rather than on the si-
Xylophagainae • Turner
263
phonoplax. The siphonal plates of Xylo-
pholas probably function as do the pallets
of the Teredinidae and the siphonoplax of
other pholadids to close the end of the
burrow. Embiyological studies are needed
to prove the affinities of the plates.
From Xijloredo, Xijlopholas differs in
having siphonal plates, a periostracal
sheath on the animal posterior to the
valves, and in not lining the burrow with
a chitonlike or calcareous tube.
Range. To date the genus is known only
from the type species found off the Lower
Florida Keys in the western Atlantic and
in the Gulf of Guinea in the eastern At-
lantic in depths from 239 to 366 m. (Two
specimens were dredged in 2,550 m but
these may have been advectitious.)
Xylopholas a/tena/ Turner 1972
Plates 33, 34
Xylopholas altouii Turner, 1972, Basteria, 36(2): 99-
103, figs. 1-12 (Gerda, station 66, about 13 miles
SE of Fowey Rocks, Florida [25°25'N, 79°59'] in
366 m). Holotype, MCZ 279315.
Distinctive Characters. Aniinal elon-
gate, not capable of retraction within the
valves, and with lateral, paddlelike siphon-
al plates at the posterior end. Shell similar
to that of Xijlopliaga. Mesoplax composed
of two flat, elongate plates that are held in
place by the periostracum extending be-
tween the beaks dorsally. Posterior adduc-
tor muscle scar large and with transverse
forking impressions. Young carried on the
ventral surface just posterior to the valves.
Description. Shell globose, reaching 2.5
mm in length, thin, fragile, and with a rel-
atively heavy, golden-brown periostracum
covering the valves and mesoplax. Pedal
gape angle about 90°. Beaked portion of
the anterior slope recurved dorsally and
sculptured with numerous strong, dentic-
ulated ridges. Umbonal reflection narrow.
Umbonal— ventral sulcus narrow and only
slightly impressed. Disc and posterior
slope sculptured with fine growth lines
only.
Inner surface of valves smooth and glis-
tening. Umbonal-ventral ridge low, indis-
tinct except near the ventral margin,
slightly segmented, and with a small ven-
tral condyle. Chondrophore and internal
ligament well developed. Posterior adduc-
tor muscle scar large, covering most of the
posterior slope, elongate oval in outline
and marked with transverse, forking im-
pressions. Pedal retractor scar irregularly
and broadly oval and located about mid-
way on the anterior margin of the poste-
rior adductor scar. Siphonal retractor mus-
cles inserted on the siphonal plates and
collar.
Mesoplax small, not filling the gape be-
tween the beaks, composed of two flat,
elongate, subrectangular plates, somewhat
pointed posteriorly, sculptured with fine
transverse ridges, covering the dorsal sur-
face of the anterior adductor muscle and
held in place by the periostracum.
Animal long, with a periostracal sheath
covering the portion posterior to the valves
and with a pair of lateral, paddle-shaped,
chitonlike plates at the posterior end
(Plate 33, Fig. 4). The siphons extend be-
tween the plates, and the siphonal retrac-
tor muscles insert on the inner surface of
them. Siphons separate, excurrent siphon
longer than the incurrent siphon, the ap-
ertures of both with fine cirri.
Gills and visceral mass contained entire-
ly between the valves, the portion of the
animal extending beyond the valves com-
posed of a dorsal excurrent and a ventral
incurrent canal combined in a common
muscular and periostracal sheath, with a
chitinous collar and two lateral, paddle-
shaped plates posteriorly.
Measurements. See Table 20.
Remarks. Isolated valves of this species
would be difficult if not impossible to dis-
tinguish from several species of Xylopha-
ga; however, its reduced, flat mesoplax,
elongate soft parts, and siphonal plates
readily distinguish it from all other species
in the Xylophagainae. Nothing is known of
the biology of the species except that it has
a large wood-storing cecum and, therefore,
probably utilizes wood as food. The young
are held within the burrow to the late ve-
264 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Table 20. Measurements of
Xylopholas altenai.
Length
(mm)
Height
1.8
1.8
holotype
1.0
1.0
Atlantique Sud, station 147
1.5
1.4
Galathea, station 52
1.2
1.1
Gerda, station 66
1.9
1.8
Gerda, station 266
2.0
2.1
Gerda, station 266
2.0
2.5
Gerda, station 266
2.5
2.5
Gerda, station 266
liger stage when the foot has developed.
They are attached to the ventral surface of
the animal just posterior to the valves. The
number of attached young ranged from
two to eight and they averaged 0.28 mm
in length. The hinge plate of the young is
well developed, with two teeth and a sock-
et in the right valve and a corresponding
single tooth and two sockets in the left
valve (Plate 33, Figs. 1, 2).
About 60 specimens were taken at the
three stations off Florida and 10 speci-
mens were taken from the three stations
in the Gulf of Guinea. The shells of many
of the specimens were in poor condition
and the valves had largely dissolved, pos-
sibly a result of the wood being veiy acid.
However, the characteristic siphonal plates
readily identified the species. The two
specimens taken from a coconut husk
dredged off Sao Tome in 2,550 m may
have been advectitious; this species may
not typically occur at that depth. Neither
specimen carried young. The shelf in this
area is very narrow and steep so that plant
material could easily be carried into deep
water. All other records are from depths of
239-366 m.
Range. Known only from off the Florida
Keys in the western Atlantic and the Gulf
of Guinea in the eastern Atlantic in depths
ranging from 239 to 2,550 m.
Specimens Examined. Western Atlantic: UNITED
STATES, FLORIDA: Gerda, station 266, off Fowey
Rocks, about 16 miles SE of Miami (25°39'N,
79°58'W) in 340 m; Gerda, station 66, off Turtle
Reef, about 13 miles SE of Fowey Rocks (25°25'N,
79°59'W) in 366 ni; Gerda, station 220, about 30
miles S of Alligator Reef (24°25'N, 80°33.5'W) in 311
m. Eastern Atlantic: GABON: Galathea, station 52,
off Port Victoria, Sao Tome Island (1°42'N, 7°51'E)
in 2,550 m (in coconut shell); Atlantique Sud, station
147, about 45 miles N Port Gentil, Cape Lopez (0°S,
8°58'E) in 250 m; Atlantique Sud, station 154, about
35 miles NE of Port Gentil (0°15'S, 8°47'E) in 239 m.
GENUS XYLOREDO TURNER 1972
Xyloredo Turner, 1972, Breviora, 397: 3 (type species,
Xyloredo nooi Turner, original designation); Turner,
1973, Science, 180: 1377-1379.
Distinctive Characters. Species in this
genus are characterized by having typical
Xylophaga shells, which lack apophyses
and have a mesoplax composed of two flat
plates, and by making a long teredolike
burrow. The posterior two thirds of the
burrow has a thin calcareous lining,
marked with distinct growth lines and cov-
ered with periostracum that extends to the
calcareous portion anterior as a band. The
part of the animal extending beyond the
valves into the calcareous tube is covered
by a golden-brown periostracal sheath that
is continuous anteriorly with the covering
of the valves and posteriorly with the per-
iostracum of the tube. A fold of the mantle
is attached to the growing end of the tube
where both periostracum and calcium are
added. In young specimens the tube may
be composed entirely of periostracum.
Posterior to the valves, the combined in-
current and excurrent canals extend the
length of the tube and are attached lightly
to it at the base of the short separate si-
phons. Two dorsolateral ridges within the
incurrent canal appear to be ciliated and
possibly aid in water transport.
Remarks. Members of this genus differ
from Xylophaga and Xylopholas in making
a burrow that may reach more than 30
times the length of the shell and is lined
with a calcareous tube. Several species of j
Xylophaga make a burrow more than five 1
times the length of the shells and form a |
chimney composed of mucous-cemented [
fecal pellets at the posterior end of the |
burrows. These are not homologous with |
the calcareous tubes of Xyloredo but rath
Xylophagainae • Turner 265
er with the chimney of rock-boring pho-
lads, as in Parapholas Conrad (Turner,
1955: 123).
The discovery of this teredohke genus
in the Pholadidae requires a reexamination
of the fossil teredinids, especially those re-
corded as having ringed tubes. On the ba-
sis of our present knowledge, it may be
impossible to distinguish Xyloredo from
teredinids in fossilized wood. However, if
tubes are present, a microscopic analysis
of their structure may aid in distinguishing
between them because teredinid tubes are
amorphous, whereas tubes of Xyloredo
have a definite structure with growth rings
and periostracum. Certainly Xyloredo
should be considered when examining
drilled wood thought to have come from a
deep water fossilized deposit.
Although Xyloredo superficially resem-
ble the Teredinidae, this is entirely con-
vergent and does not in any way indicate
relationship, nor does it suggest the evo-
lution of the Teredinidae from the Xylo-
phagainae. The latter lack apophyses and
pallets, and have a mesoplax. In addition,
none of the visceral mass or gills of the
Xylophagainae extend beyond the valves
posteriorly as they do in the Teredinidae.
Range. To date three species of Xylo-
redo are known, two in the western Atlan-
tic and the other in the eastern Pacific. All
occur at depths greater than 1,500 m.
Xyloredo noo/ Turner
Plate 35
Xyloredo nooi Turner, 1972, Breviora, 397: 5—7, pis.
1, 2 (Tongue of the Ocean, about 4 miles off north-
east tip of Andros Island, Bahama Islands [25°.54'N,
77°49'W] in 1,737 m). Holotype, MCZ 279631;
paratypes from the same and other panels exposed
at the same locality, MCZ 279632, 279633, 279634,
and 279635, and the Zoological Museum, Univer-
sity of Copenhagen; Turner, 1973, Science, 180:
1377-1379.
Distinctive Characters. Burrow long,
teredinidlike, lined with a thin calcareous
tube marked with growth rings and cov-
ered with periostracum. Shell similar to
Xylophaga, anterior slope narrow, umbo-
nal— ventral sulcus lightly impressed, pos-
terior slope high and reflected dorsally
Posterior adductor muscle scar subellipti-
cal, set high on the posterior slope, and
divided into two distinct areas. Disc sep-
arated from the posterior slope by a
groove on the inner surface of the valves.
Mesoplax small, the two flat triangular
plates composed almost entirely of perios-
tracuiTi. Periostracal sheath between the
valves and the tube smooth.
Description. Shell globose, valves reach-
ing 10 mm in length and 10.5 mm in
height, thin, fragile; umbos prominent.
Periostracum relatively thick, golden-
brown, glistening, and covering entire
valve. Pedal gape angle about 110°. Ante-
rior slope sculptured with numerous den-
ticulated ridges, there being 24 on the ho-
lotype. UiTibonal— ventral sulcus narrow,
slightly impressed, and sculptured with
fine, irregular growth lines. Posterior slope
high, reflected near the dorsal margin, and
sculptured with fine growth lines. Umbo-
nal reflection rather wide, thin, adhering
to the valves in the umbonal area, free an-
teriorly.
Inner surface of valves smooth and glis-
tening. Umbonal— ventral ridge narrow,
high, and segmented. Chondrophore and
internal ligament prominent. Disc separat-
ed from the posterior slope by a pro-
nounced narrow groove extending from
the umbo to the posterior ventral margin.
Posterior adductor muscle scar large, ellip-
tical, and divided into two areas, the upper
part marked with irregular impressions,
the lower with regular chevron-shaped im-
pressions. Anterior adductor scar covering
most of the umbonal reflection. Pedal re-
tractor scars elongate, the ixiuscles insert-
ing in the groove separating the disc from
the posterior slope.
Mesoplax small, flat, the two broadly tri-
angular plates composed almost entirely of
periostracum and located anterior to the
umbos.
Burrow long, teredinidlike, and lined
with a thin calcareous tube that is sculp-
tured with distinct growth rings and cov-
ered with periostracum that extends as a
266 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Table 21. Measurements of Xyloredonooi.
Length
Hei2;ht
(mm)
(mm)
Location
5.0
5.1
paratype
6.7
6.9
paratype
7.2
7.8
paratype
9.5
9.8
paratype
9.5
10.0
holotype
border at the anterior end. Tube in veiy
young specimens composed entirely of
periostracum. Between the valves and the
anterior end of the tube the animal is cov-
ered with a smooth periostracal sheath,
which is continuous anteriorly with the
covering of the valves and posteriorly with
the tube. Siphons short, separate, and ap-
parently with a few small cirri.
Measurements. See Table 21.
Remarks. Xyloredo nooi is related to
both Xyloredo naceli Turner from the east-
ern Pacific and X ingolfia Turner from the
North Atlantic. Xyloredo nooi differs from
them in having a much thinner burrow lin-
ing, a high, reflected posterior slope on the
valves, and a proportionately smaller, more
highly placed and divided posterior adduc-
tor muscle scar. In addition, the periostra-
cal sheath extending between the valves
and the calcarious tube is smooth.
The larvae of X. nooi apparently do not
rise very high in the water column, inas-
much as a panel 25 ft off the bottom
showed no trace of them, whereas the
panel directly beneath it on the bottom
was riddled. On the basis of the prodis-
soconch still visible on some of the speci-
mens, it would appear that the lai'vae are
similar to those ofXylophaga. No evidence
was found in the three riddled panels ex-
amined that the young were brooded or
held in the tubes.
The burrows resemble those of teredi-
nids and intertwine with each other but
basically follow the grain of the wood.
They may reach 200 mm in length and 15
mm in diameter at the anterior end. The
largest tube with intact shells and animal
remaining was 145 mm long; the valves
were 9.5 mm in length and 10 mm in
height. The panels in the Tongue of the
Ocean were submerged for 34 months and
inasmuch as the larger burrows were emp-
ty or contained only fragments of shells,
the length of life may be about 2—2.5
years. The calcareous lining of a burrow
122 mm long extends to about 30 nam
from the anterior end, thus leaving room
for boring activities and changing the di-
rection of the burrow.
Range. Knowii only from the type lo-
cality.
Specimens Examined. BAHAMA ISLANDS:
Tongue of the Ocean, about 4 miles off NE tip of
Andros Island (24°54'N, 77°49'W) in 1,737 m.
Xyloredo ingolfia Turner
Plates 36, 37
Xyloredo ingolfia Turner, 1972, Breviora, 397: 7-9,
pis. 3-5 (from wood dredged by the Ingolf Expe-
dition at station 67, south of Eyrabakld, Iceland
[61°30'N, 22°30'W] in 975 fathoms [1,783 m]). Ho-
lotype, MCZ 279636; paratypes, MCZ 279637, and
the Zoological Museum, University of Copenha-
gen; Turner, 1973 Science 180: 1377-1379.
Description. Shell globose, valves reach-
ing 2.5 mm in length and 2.0 mm in
height, thin, fragile, with prominent um-
bos; thin, glistening, almost colorless per-
iostracum covering disc and posterior
slope. Beaked portion of anterior slope
wide, extending more than one half dis-
tance to ventral margin; sculptured with i
close-set and very finely denticulated ridg-
es. Posterior portion of anterior slope
about two thirds width of beak, sculptured !
with close-set ridges that extend to very
slightly impressed umbonal— ventral sulcus.
Disc sculptured with well-marked growth
lines. Posterior slope small, low, and not
clearly demarcated on outer surface of
valve. Umbonal reflection thick, narrow,
short, and free except at posterior end.
Inner surface of valves smooth, slightly
shiny to chalky (perhaps owing to long '
preservation). Umbonal-ventral ridge wide,
flattened, often varying in width, irregularly
segmented, and not enlarged at ventral
condyle. Chondrophore and internal liga-
Xyloph AGAIN AE • Turner
267
Table 22. Measurements of
Xyloredo ingolfia.
Range. Known only from the type loeal-
Lrimtll
Heidit
(mill)
(mm)
Location
2.5
2.3
holotype
2.5
2.0
paratype
2.1
2.0
paratype
2.0
1.9
paratype
1.5
1.4
paratype
1.5
1.2
paratype
ment large. Disc not clearly separated from
posterior slope. Posterior adductor muscle
scar large, slightly raised, elliptical, extend-
ing nearly to ventral margin, with irregular,
transverse impressions. Anterior adductor
muscle scar covering umbonal reflection.
Siphonal retractor muscle scars not im-
pressed. Pedal retractor muscle scar small,
elongate to oval, and located just anterior
to posterior adductor muscle scar. Meso-
plax of two veiy small, narrow, subrectan-
gular, flat, calcified plates lying on dorsal
surface of anterior adductor muscle.
Burrow 10—15 times length of valves; cal-
careous tubular lining three fourths length
of bruTow. Tube relatively heavy, marked
with rmiform, close-set, raised rings, and
covered with light tan periostracum that ex-
tends anteriorly as border. Portion of ani-
mal between valves and tube covered by
tliin, irregrflarly ridged periostracal sheath.
Siphons short; incurrent siphon slightly lon-
ger than excurrent siphon. Protoconch
large, medium golden-brown, and sculp-
tured with fine, concentric ridges.
Measurements. See Table 22.
Remarks. This species differes from Xy-
loredo naceli in having a less well-devel-
oped posterior slope, a shallow, indistinct
umbonal— ventral groove, a flattened um-
bonal— ventral ridge, and in having the
valves longer than high. It differs from X.
nooi in having valves longer than high, in
having a low, rounded posterior slope, in
lacking the distinct groove on the inner sur-
face separating the disc from the posterior
slope, and in having the plates of the me-
soplax sub rectangular and well calcified.
ity.
Specimens Examined. ICELAND: Ingolf Expedi-
tion, station 67, S of Eyrabakld (61°30'N, 22°30'W)
in 1,783 m.
Xyloredo nace// Turner
Plate 38
Xyloredo naceli Turner, 1972, Breviora, 397: 9—11, pi.
6, figs. 1-5 (USNCEL STU 1-4 about 30 miles S
of San Miguel Island, off Port Hueneme, Santa
Barbara Islands, California [33°46'N, 120°45'W] in
6,800 ft [2,072.6 m] from panels submerged from
June 1964 to July 1965). Holotype, MCZ 279638;
paratope, 279639.
Distinctive Characters. Burrow teredi-
nidlike, lined with a thin calcereous tube
marked with growth rings and covered
with periostracum. Shell similar to Xijlo-
phaga, with a narrow, slightly impressed
umbonal— ventral sulcus. Posterior adduc-
tor muscle scar elliptical, almost complete-
ly covering the posterior slope and uni-
formly marked with transverse impres-
sions. Mesoplax small, the two flat, trian-
gular plates composed largely of
periostracimi. Periostracal sheath covering
the animal between the valves and the
tube papillose.
Description. Shell globose, reaching 1.5
mm in length, thin, fragile, white, with a
thin, pale yellow periostracum. Pedal gape
angle about 100°. Anterior slope sculp-
tured with 8—12 widely spaced, pro-
nounced, denticulated ridges. Umbonal—
ventral sulcus narrow and only slightly im-
pressed. Discs and posterior slope sculp-
tured with fine growth lines only.
Inner surface of valves smooth and glis-
tening. UiTibonal— ventral ridge narrow,
high, and indistinctly segmented. Chon-
drophore and internal ligament well de-
veloped. Posterior adductor muscle scar
elliptical, extending from the dorsal nearly
to the ventral margin of the posterior slope
and regularly marked with transverse im-
pressions. Pedal retractor scar not im-
pressed. Ventral adductor only lightly im-
pressed and located just posterior to the
268 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Table 23. Measurements of Xyloredo naceli.
Length
(mm)
1.1
1.0
1.2
1.5
1.2
1.0
1.3
1..5
holotype
paratype
paratype
paratype
umbonal— ventral ridge. Siphonal retractor
muscles inserting on the tube.
Mesoplax small, thin, set between and
anterior to the umbos, the two triangular
plates composed almost entirely of perios-
tracum. Burrow of largest specimen about
six times the length of the shell. Calcare-
ous tube lining the burrow thin, distinctly
marked with growth rings, and covered
with periostracum that extends as a border
anteriorly. Aperture of the burrow small,
round, the white lining visible within. The
portion of the animal between the valves
and the calcareous tube covered with a pa-
pillose periostracal sheath that is continous
with the periostracum covering the valves
anteriorly and the periostracal border of
the tube posteriorly. Siphons short, of
equal length, and apparently lacking cirri.
Measurements. See Table 23.
Remarks. This species is most closely re-
lated to X nooi Turner from the western
Atlantic. Only eight specimens of X. naceli
were found and they were all very small,
but these appear to be sufficiently distinct
to consider them members of a separate
species. Xyloredo naceli differs from X
nooi in having a heavier tube that is cal-
careous, even in very yotmg specimens;
and a posterior adductor muscle scar that
is proportionately larger and not divided
into two areas. In addition, it lacks the pro-
nounced internal groove separating the
disc from the posterior slope, and the per-
iostracal sheath covering the animal ante-
rior to the tube is papillose.
Nothing is known of the biology of the
species except that at the sites where they
were collected the temperature of the wa-
ter was 2.1° C, the salinity was 34.52%o,
the dissolved oxygen content was 1.26
ml/L, the pH was 7.84, and the hydrostatic
pressure 3000 psi (Mru-aoka, 1966b).
The embryonic valves still visible on
some of the specimens suggest that the
mature larvae are similar to those in Xy-
lophaga. Xylophaga niuraokai was the
most common borer in the panels from
which the X. naceli were taken.
Range. Known only from the type lo-
cality.
Specimens Examined. UNITED STATES, CALI-
FORNIA: USNCEL STU 1-4 S of San Miguel Island,
Santa Barbara Islands (33°46'N, 120°45'W) in
2,072 m.
ACKNOWLEDGMENT
This paper was prepared with the aid of
funds received from the Department of
the Navy, Biology Branch, Office of Naval
Research, ONR grant N00014-91-J-1402,
Biological Studies on Marine Boring and
Fouling Mollusks.
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270 Bulletin Museum of Comparative Zoologt/, Vol. 157, No. 4
2 mm
Plate 1 . Xylophaga concava Knudsen from Pillsbury, station 526.
1 mm
Figure 1 . Dorsal view of apposed valves showing the concave posterior slope and erect mesoplax. Figure 2. Posterior end of
valve and siphons. Figure 3. Enlargement of the siphonal openings to show the six large cirri on the excurrent siphon and the
double row of small cirri on the incurrent siphon.
Xylophagainae • Turner 271
Plate 2. Xylophaga gerda Turner n. sp. from Gerda, station 499.
Figure 1 . Lateral view of holotype showing the attachment of the posterior adductor muscle through the thin valve, the mesoplax
that does not extend above the umbos, and the fecal cylinder in the excurrent canal. Figure 2. Dorsal view of the holotype
showing the mesoplax. Figure 3. Enlargement of the posterior end of the siphons. Figure 4. Diagrammatic cross-section
through the siphons and the fecal cylinder. Figure 5. Three-quarter view of holotype showing the inflated umbos and the simple
curved plates of the mesoplax. Figure 6. A relatively smooth chimney composed largely of periostracum, with a thin coating of
fecal material. Figure 7. A thick chimney, built in sections with "leaves" of periostracum extending at the anterior end of each
section.
272 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
1 mm
Plate 3. Xylophaga gerda Turner n. sp. from Pillsbury, station 944.
0.5 mm
Figure 1 . Outer view of right valve. Figure 2. Inner view of left valve, showing posterior adductor muscle scar and low umbonal-
ventral ridge. Figure 3. Enlargement of the hinge area of left valve to show the chondrophore and anterior adductor muscle
scar. Figure 4. Outer view of right valve of a young specimen. Figure 5. Inner view of left valve of young specimen showing
the prodissoconch, chondrophore, and muscle scars.
Xylophagainae • Turner
273
Plate 4. Xylophaga grevei Knudsen (Figs. 1-6 from Knudsen, 1961: 176-177).
Figures 1-4. Holotype. Figure 1 . Right side of entire specimen showing siphons and mesoplax. Figure 2. Dorsal view. Figure
3. Enlargement of the siphons showing the cirri around the incurrent aperture (35) and the excurrent aperture (6). Figure 4.
Internal view of left valve of the holotype showing the broadly oval posterior adductor muscle scar set high on the posterior
slope. Figures 5, 6. Ventral and dorsal view of the mesoplax. Figure 7. Small specimen carrying two young from Galathea,
station 444.
274 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Plate 5. Xylophaga clenchi Turner and Culliney.
Figures 1-3. Side, dorsal, and anterior views of the holotype, showing the widely space ridges on the anterior slope of specimens'
boring in soft wood and the position of young and the dorsal surface. Figures 4, 5. Inner and outer views of left valve of a
specimen from Atlantis II, station 124, showing the muscle scar and large number of ridges on the anterior slope of a specimen
from hard wood.
Xylophagainae • Turner
275
5 mm
Plate 6. Xylophaga clenchi Turner and Culliney.
Figures 1, 2. Outer and inner views of the right valve of a large specimen with mesoplax in place, dredged by the Pillsbury,
station 394. Figure 3. Lateral view of an entire specimen from the Tongue of the Ocean showing relative size of si-
phons. Figure 4. Enlargement of the siphons, posterodorsal view, to show the incurrent and excurrent apertures and the cirri
around them.
276 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Plate 7. Xylophaga clenchi Turner and Culliney from /ngo/f Expedition, station 67.
Figure 1. Lateral view of right valve with mesoplax in place and young attached to dorsal surface. Figure 2. Dorsal view of:
same specimen showing the prodissoconch. Figure 3. Lateral view of specimen with extended siphon.
Xylophagainae • Turner 277
Plate 8. Xylophaga clenchi Turner and Culliney.
Figures 1 , 2. From Pillsbury, station 238. Figure 1 . Lateral view of right valve. Figure 2. Dorsal view of opposed valves showing
prodissoconch, four young attached posterior to the umbos, and an atypical elongate, longitudinally folded mesoplax. Figures
3, 4. Specimen from Atlantis II, station 124. Figure 3. Anterior view of specimen with mesoplax bent at a right angle. Figure
4. Dorsal view of same specimen showing prodissoconch and mesoplax.
278 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Plate 9. Xylophaga supplicata Taki and Habe from Tosa Bay, Shikoku, Japan.
Figure 1. Anterior view of opposed valves with mesoplax in place. Figure 2. Lateral view of entire specimen from the right with
mesoplax in place. Figure 3. Dorsal view of opposed valves tipped slightly forward to show the minute tubes at the posterior
end of the mesoplax. Figure 4. Enlargement of mesoplax and umbonal area, looking down into the cavity formed by the
incurving of the umbos. Figure 5. Inner view of left valve showing chondrophore with large tooth. Figure 6. Enlargement
showing chondrophore with large tooth.
Xylophagainae • Turner 279
Plate 10. Xylophaga whoi Turner n. sp.
Figures 1-4. Specimen from Pillsbury, station 944. Figures 1, 2. Outer and inner views of the left valve. Figure 3. Frontal
view of mesoplax with smallest tubes seen. Figure 4. Side view of left plate of mesoplax. Figures 5-12. Holotype. Figure
5. Outer view of right valve. Figure 6. Inner view of left valve. Figure 7. Dorsal view of left plate of mesoplax. Figure 8.
Frontal view of mesoplax. Figures 9, 10. Outer surface of left and right plates of mesoplax with average-size tubes. Figures
11, 12. Inner surface of left and right plates of the mesoplax showing the cun/ature at the dorsal margin and the pore to the
inner surface just below it. Figure 13. Enlargement of the hinge area of the left valve to show the chondrophore with a large
tooth on it postehor dorsal margin, the umbonal reflection, and the inner surface of the mesoplax.
280 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
5 mm
/ .1
-'im
Plate 11. Xylophaga who; Turner n. sp. from Pillsbury, station 394.
Figure 1. Anterior view of entire animal showing mesoplax in place, the anterior adductor muscle, the pedal opening of the
mantle, the retracted foot, and young on the dorsal surface of the valves. Figure 2. Enlargement of the umbonal area, mesoplax,
and attached young.
XYLOPHAGAINAE • Turner 281
5 mm
0.1 mm
Plate 12. Xylophaga profunda Turner n. sp. from U.S. Naval Oceanographic Office test site, Tongue of the Ocean,
Andros Island, Bahama Islands.
Fiqures 1-3 Holotype Figure 1 . Dorsal view showing round ridge posterior to the umbonal-ventral sulcus, the mesoplax, and
atfached young Figure 2. Side view of left valve. Figure 3. Enlarged anterior view of dorsal area to show mesoplax. Figures
4, 5. Dorsal view of paratypes to show range of length-width relationships, concavity of postenor slope, as well as arrangement
and number of young. Figure 6. Enlarged young.
282 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
5 mm
Plate 13. Xylophaga profunda Turner n. sp. from U.S. Naval Oceanographic Office test site Tongue of the Ocean,
Andros Island, Bahama Islands.
Figures 1, 2. Inner view of left valve of two specimens to show differences in intensity of muscle scar impressions, the riblike
ridges extending from the umbonal-ventral ridge, the placement of the mesoplax, and the chondrophore. Figure 3. Enlargement
of the hinge area of the left valve to show the chondrophore. Figures 4-6. Anterior, dorsal, and ventral view of mesoplax.
Xylophagainae • Turner
283
Plate 14. Xylophaga abyssorum Dall.
Figures 1-5. Specimens from U.S. Naval Oceanographic Office test site, Tongue of the Ocean, Andros Island, Bahama Is-
lands. Figure 1 . Outer view of right valve of immature specimen with mesoplax in place and showing ridge posterior to umbonal-
ventral sulcus. Figure 2. Inner view of same valve, showing groove posterior to umbonal-ventral ridge, ligament, and charac-
teristic muscle scar of young specimen. Figure 3. Dorsal view of same valve with mesoplax in place. Figures 4, 5. Mesoplax
of young specimen with large ventral portion. Figure 6. Dorsal view of right valve of holotype of Xylophaga abyssorum for
comparison with Figure 3. Figure 7. Dorsal view of young specimen from Gerda, station 266, with mesoplax developing lobes
on dorsal portion.
284 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Plate 15. Xy lophaga abyssorum DaW.
Figures 1-4. Specimens from Pillsbury, station 944. Figure 1. inner view of right valve showing muscle scars, deep groove
posterior to the umbonal-ventral ridge, and the small chondrophore. Figure 2. Enlargement of hinge area of same speci-
men. Figure 3. Hinge area of left valve. Figure 4. Dorsal view of umbonal area with mesoplax in place. This is the most
elaborate mesoplax observed, having two tubes and a third developing. Figure 5. Dorsal view of specimen from Gerda, station
266, with two tubes formed on each plate of the mesoplax and with two young just posterior to the umbos.
Xylophagainae • Turner 285
Plate 16. Xylophaga abyssorum DaW.
Figures 1-5. Specimen from Pillsbury, station 944. Figure 1. Lateral view of right valve. Figure 2. Anterior view of opposed
valves with mesoplax in place. Figure 3. Posterior view of opposed valves with mesoplax in place and showing the inflated
umbos and ridge. Figure 4. Dorsal view of opposed valves. Figure 5. Enlarged view of mesoplax and umbonal area.
286
Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Plate 17. Xylophaga duplicata Knudsen from Galathea, station 745, Gulf of Panama (all from Knudsen, 1961: 175).
Figure 1 . Lateral view of left side of holotype, shiowing tfie mesoplax in place, standing off from tfie surface of thie valves. Figure
2. Inner view of left valve of a paratype shiowing thie small smooth posterior adductor muscle scar. Figure 3. Left side of
extended siphon of a paratype. Figure 4. Enlarged view of posterior end of the siphon. Figure 5. Dorsal view of the meso-
plax. Figure 6. Ventral view of mesoplax.
Xylophagainae • Turner
287
Plate 18. Xylophaga murao/ca/ Turner n. sp. from U.S. Naval Civil Engineering Laboratory Test Site I.
Figure 1. Lateral view of holotype showing siphons. Figure 2. Inner view of left valve showing muscle scar and simple chon-
drophore. Figure 3. Dorsal view of the two plates of the mesoplax of a mature specimen showing large basal portion. Figure
4. Dorsal view of right plate of mesoplax. Figure 5. Ventral view of left plate of mesoplax. Figure 6. Lateral view of left plate
of mesoplax.
288 Bulletin Mtiseum of Comparative Zoology, Vol. 157, No. 4
1 mm
Plate 19. Xylophaga muraokai Turner n. sp. from U.S. Naval Civil Engineering Laboratory Test Site I.
Figure 1 . Anterior view of fiinge area showing chondrophore and internal ligament. Figure 2. Posterior view of mesoplax fitting
between the umbos. Figure 3. Dorsal view of young specimen with partially developed mesoplax. Figure 4. Mesoplax of
young specimen. Figure 5. Enlargement of posterior end of siphons showing cirri around the siphonal apertures.
Xylophagainae • Turner 289
1 mm
Plate 20. Xylophaga atlantica Richards from about 15 miles off Ipswich, Massachusetts, in 73 m.
Figure 1. Entire specimen with siphons extended, showing relative lengths. Figure 2. Enlargement of the posterior end of the
siphons, lateral view. Figure 3. Enlargement of the posterior end of siphons dorsal view showing the numerous small cirri
surrounding the incurrent siphonal aperture and the larger, less numerous cirri of the excurrent siphon.
290 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
1 mm
Plate 21. Xylophaga washingtona Bartsch.
Figures 1-3. Specimen from U.S. Naval Civil Engineering Laboratory Test Site I. Figure 1. Dorsal view of entire specimen
showing the relative length of the siphons. Figure 2. Dorsal view of siphons from just anterior to the truncation of the excurrent
siphon. Figure 3. Dorsal view of siphons in area of truncation to show contracted aperture of the excurrent siphon of a short
tube, and the short lateral lobes extending from the truncation. Figure 4. Specimen from about 40 miles W of Silver Point,
Oregon. Aperture of incurrent siphon showing cirri.
XylOPHAGAINAE • Turner
291
1 mm
Plate 22. Xylophaga rikuzenica Taki and Habe from off Rikuzen, Honshu, Japan.
Paratype, Museum of Comparative Zoology 194821.
Figure 1 . Inner view left valve showing posterior adductor muscle with herringbone markings. Figure 2. Outer view of left valve
showing broad ventral sulcus. Figure 3. Anterior view of opposed valves showing mesoplax in place, the condyles, and chon-
drophore. Figure 4. Dorsal view of opposed valves showing mesoplax in place, umbonal reflection, and broad, deep umbonal-
ventral sulcus. Figures 5-7. Dorsal, lateral, and ventral views of the mesoplax showing the large ventral portion.
292 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
5 mm
5 mm
Plate 23. Xylophaga depalmai Turner n. sp. from U.S. Naval Oceanographic Office test site off Fort Lauderdale, Florida.
Figure 1. Lateral view of holotype shiowing sipfions with lateral pehostracal sheath containing clusters of irregular glasslike
granules. Figure 2. Dorsal view of holotype showing mesoplax that is only slightly coiled posteriorly. Figure 3. Dorsal view of
specimen with slightly more coiled mesoplax. Figure 4. Siphons showing fringed lappets. Figure 5. Enlargement of
fringe. Figure 6. Inner view of left valve showing muscle scars. Figure 7. Inner view of left valve showing prodisso-
conch. Figure 8. Enlargement of hinge area of left valve showing umbonal reflection and chondrophore. Figure 9. Enlargement
of chondrophore.
Xylophagainae • Turner
293
Plate 24. Xylophaga depa/ma/ Turner n. sp. from U.S. Naval Oceanographic Office test site off Fort Lauderdale, Florida.
Figures 1-5. Specimen with the two plates of mesoplax fused. Figure 1. Outer view of right valve. Figure 2. Inner view of
right valve showing lightly impressed muscle scars. Figure 3. Dorsal view of fused plates of mesoplax. Figure 4. Ventral view
of mesoplax. Figure 5. Lateral view. Figures 6-8. Dorsal, ventral, and lateral view of mesoplax with plates partially fused
ventrally. Figures 9, 10. Dorsal and ventral views of a mesoplax, which is broadened anteriorly and has a reduced ventral
portion. Figures 11-13. Dorsal, ventral, and lateral views of an unusually broad mesoplax with unequal plates and reduced
ventral portion. Figures 14,15. Dorsal and ventral views of a broad mesoplax strongly coiled posteriorly, with a widened median
area that separates the coils. Figures 16, 17. Dorsal and ventral views of a typical mesoplax with only the periostracal portion
of the ventral portion fused.
294 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
1 mm
Plate 25. Xylophaga guineensis Knudsen.
Figures 1-3. Specimen from Pillsbury, station 260. Figure 1. Lateral view of entire animal sfiowing foot, siphons, and perios-
tracal sheath with fine whitish granules. Figure 2. Dorsal view showing siphons, prodissoconch, mesoplax, and loose perios-
tracal sheath with granules. Figure 3. Anterior view showing foot, anterior adductor muscle, and mesoplax in place. Figures
4, 5. Specimen from Atlantique Sud. station 33. Figure 4. Inner view of right valve showing specimen with high flaring posterior
slope, prodissoconch, posterior adductor muscle scar, and deep groove bounding the umbonal-ventral ridge. Figure 5. Outer
view of right valve showing wide anterior slope and narrow, bladelike ridge posterior to the shallow umbonal-ventral sul-
cus. Figures 6-9. Specimens from Atlantique Sud, station 146. Figure 6. Inner view of right valve of specimen with low,
rounded posterior, small posterior adductor muscle scar set very high. Figure 7. Outer view of left valve with very closely set
ridges on anterior slope. Figures 8, 9. Dorsal views to show variation in the mesoplax in place.
Xylophagainae • Turner
295
2 3 4
Plate 26. Xylophaga guineensis Knudsen.
Figure 1. Specimen from Atlantique Sud station 146. Umbonal area showing chondrophore, prodissoconch, and umbonal re-
flection. Figures 2-4. Specimens from Atlantique Sud, station 33. Figure 2. Dorsal view of opposed plates of meso-
plax. Figure 3. Ventral view of right plate of the mesoplax to show long cornucopialike shape. Figure 4. Dorsal view of right
plate of mesoplax.
296 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Plate 27. Xylophaga mexicana Dall.
Figures 1-4. Specimens from "replication reef," Santa Monica Bay, California. Figures 1, 2. Outer and inner views of left valve
of specimen close to the holotype. Figure 3. Lateral view of entire specimen showing inflated mesoplax and siphons. Figure
4. Dorsal view of entire specimen showing mesoplax and expanded foot. Figures 5, 6. Outer and inner view of right valve of
the holotype (from Turner, 1955, pi. 90).
Xyloph AGAIN AE • Turner 297
2 X i/^ 3 ^ ^ 4 ^-" 5
Plate 28. Xylophaga mexicana Dall from "replication reef," Santa Monica Bay, California.
Figure 1. Enlargement of the siphons to show the truncation of the excurrent siphon, the finely fringed lappets, granules em-
bedded along the side of the siphon, and the papillose end of the incurrent siphon. Figures 2, 3. Ventral and dorsal view of
mesoplax. Figures 4, 5. Inner and outer lateral views of mesoplax showing flange that fits down between the umbos.
298 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Plate 29. Xylophaga tipperi Turner n. sp. from U.S. Naval Oceanographic Office test site off Fort Lauderdale, Florida.
Figure 1 . Lateral view of holotype showing foot and extended siphons. Figure 2. Inner view of left valve to show smooth posterior
adductor muscle scar. Figure 3. Dorsal view showing mesoplax in place. Figure 4. Enlargement of posterior end of incurrent
siphon to show the fhnged lappets and the single row of glasslike plaques along the side. Figure 5. Lateral and three-quarters
view of glasslike plaques. Figures 6-9. Mesoplax. Figure 6. Dorsal view of right plate showing flat surface and faint sculp-
ture. Figures 7, 8. Ventral view of hght and left plates. Figure 9. Lateral view of right plate to show the compressed main
portion and the posterior ventral flange that extends down between the umbos.
Xylophagainae • Turner 299
1 mm
Plate 30. Xylophaga bayeri Turner n. sp. from U.S. Naval Oceanographic Office test site off Fort Lauderdale, Florida.
Figure 1. Lateral view of fiolotype showing sipfions and mesoplax in place. Figure 2. Dorsal view of holotype, sfiowing the
lateral extension of the mesoplax. Figure 3. Enlargement of the siphons showing the truncated excurrent siphon and the fringed
lappets. Figure 4. Ventral view of the mesoplax. Figure 5. Dorsal view of the mesoplax.
300 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
1 mm
5 mm
Plate 31 . Xylophaga bayeri Turner n. sp.
Figures 1-3. Specimen from Gerda, station 266. Figure 1. Inner view of valves showing smooth muscle scar; narrow, high
umbonal-ventral ridge, and broad, recurved umbonal reflection that adheres to the valve. Figure 2. Dorsal view of specimen
showing broad umbonal reflection and the mesoplax. Figure 3. Enlargement of the mesoplax, dorsal view. Figures 4, 5.
Specimen from U.S. Naval Oceanographic Office test site off Fort Lauderdale, Florida. Figure 4. Lateral view of a young
specimen with partially developed mesoplax. Figure 5. Dorsal view of a young specimen with partially developed mesoplax.
Xyloph AGAIN AE • Turner 301
5 mm
1 mm
Plate 32. Xylophaga japonica Taki and Habe.
Figures 1-5. Paratype from Tosa Bay, Shikoku, Japan. Figure 1. Outer view of left valve showing thin ridge posterior to
umbonal-ventral sulcus and elongate posterior slope. Figure 2. Dorsal view of mesoplax with distinct concentric sculp-
ture. Figure 3. Ventral view of mesoplax showing flange that fits between umbos. Figure 4. Inner lateral view. Figure 5.
Outer lateral view showing flange. Figures 6-11. Specimens from Anton Bruun, station 23. Figure 6. Outer view of left
valve. Figure 7. Inner view of right valve showing chondrophore and smooth posterior adductor muscle scar. Figure 8. Outer
view of mesoplax of a small specimen. Figure 9. Inner view of mesoplax of a small specimen. Figure 10. Outer lateral view
of mesoplax of a small specimen. Figure 1 1 . Inner lateral view of mesoplax of a small specimen.
302 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Plate 33. Xylopholas a/tena/ Turner from Gerda, station 66.
Figures 1-3. Holotype. Figure 1. Lateral view of entire animal with the anterior adductor muscle relaxed so that the mesoplax
is flattened and not visible, showing siphonal plates and young carried on ventral surface. Figure 2. Ventral view showing the
mesoplax held in place by periostracum, the ventrally carried young, and the recurving of the siphonal plates. Figure 3. Dorsal
view showing the mesoplax in place and the umbonal reflection. Figure 4. Enlargement of posterior end with the incurrent
siphon projecting beyond the siphonal plate. Figure 5. Posterior end with left plate removed to show the muscle that extends
into the cavity of the plate to which the siphonal retractor muscles attach. Figure 6. Lateral view of a very small specimen,
contracted antehorly so that the plates of the mesoplax are folded upward. Figure 7. Outer view of siphonal plate. Figure 8.
Inner view of siphonal plate.
Xylophagainae • Turner
303
1 mm
Plate 34. Xylopholas altenai Turner from Gerda, station 66.
Figure 1. Outer view of rigfit valve sfiowing beaked portion of the anterior slope. Figure 2. Inner view of right valve showing
the large posterior adductor muscle scar, pedal retractor scar, prodissoconch, and ligament. Figure 3. Inner view of upper part
of left valve to showing the chondrophore and the mesoplax in its periostracal membrane. Figure 4. Dorsal view of the two
plates of the mesoplax.
304 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
Plate 35. Xyloredo noo/ Turner from U.S. Naval Oceanographic Office test site, Tongue of the Ocean, Bahama Islands.
Figure 1. Inner view of left valve showing divided muscle scar. Figure 2. Outer view of left valve. Figure 3. Outer view of left'
valve of holotype showing the flaring posterior dorsal margin. Figure 4. Inner view of right valve of holotype showing the divided
posterior adductor muscle scar, the deep groove separating the disc from the posterior slope, and the umbonal reflection. Figure
5. Anterior view of opposed valves, showing the chondrophore and internal ligament. Figure 6. Dorsal view showing the thin
mesoplax.
XylOPHAGAINAE • Turner 305
1 mm
Plate 36. Xyloredo ingolfia Turner from /ngo/f Expedition, station 67.
Figure 1 . Lateral view of an entire animal, showing the periostracal sheath, the extended anal and siphonal canals, and the short
siphons. Figure 2. Dorsal view of an entire animal, showing the periostracal sheath, the extended anal and siphonal canals,
and the short siphons. Figure 3. Lateral view of a very young specimen showing the large prodissoconch and the produced
beaked portion of the anterior slope. Figure 4. Lateral view of left valve with periostracal sheath attached. Figure 5. Dorsal
view showing the produced, recurved beaks, the umbonal reflection, and the mesoplax with only the central portion of each plate
calcified. Figure 6. Inner view of right valve with well-marked posterior adductor muscle scar. Figure 7. Inner view of left valve
with lightly marked scar and large prodissoconch.
306 Bulletin Museum of Comparative Zoology, Vol. 157, No. 4
1 mm
Plate 37. Xyloredo ingolfia Turner from /ngo/f Expedition, station 67.
Figure 1 . Inner view of left valve showing large prodissoconch, strong umbonal-ventral ridge, reduced posterior slope, and lightly
impressed posterior adductor muscle scar. Figure 2. Outer view of left valve showing the produced beak and rounded low
posterior slope.
XYLOPHAGAINAE • Turner
307
Plate 38. Xyloredo nace// Turner from U.S. Naval Civil Engineering Laboratory Test Site I (submersible test unit 1-4).
Figure 1 . Lateral view of holotype. Figure 2. Lateral view of specimen partially dissected from the wood shiowing the calcereous
tube with the anterior periostracal margin and the papillose periostracal sheath covering the animal between the tube and the
valves. Figure 3. Anterior-lateral view of entire specimen showing foot and mesoplax. Figure 4. Inner view of left valve showing
muscle scars and chondrophore. Figure 5. Outer view of left valve.
308
Bulletin Mtiseum of Comparative Zoology, Vol. 157, No. 4
INDEX
Taxon names are printed in italics. Page numbers in bold refer to illustrations; those in italics refer tc
species descriptions; t indicates table.
abyssonim, Xylophaga, 4, 12, 22, 2-3-24, 61-63
africana, Xylophaga, 4, 17-18
altenai, Xylopholas, 41—42, 80-81
atlantica, Xylophaga, 4, 27-28, 67
aurita, Xylophaga, 5, 31
hayeri n. sp., Xylophaga, 5, 12, 24, 38, 38-3.9, 77-78
bruuni, Xylophaga, 4, 19, 20, 21
clenchi, Xylophaga, 4, 13, 14, 16, 17-19, 28, 52-55
concava, Xylophaga, 4, 14-15, 15, 16, 26, 48
depalmai n. sp., Xylophaga, 5, 31-34, 35, 38, 40, 70-
71
variation of mesoplax in, 11—12, 32
distribution
of Xylophagainae, 2
of Xylophaga species, 2, 6 (map)
dorsalis, Xylophaga, 5, 12, 38, 39
dorsal plates. See mesoplax
duplicata, Xylophaga, 4, 25, 26, 36, 64
erecta, Xylophaga, 4, 14
foliata, Xylophaga, 4, 19
food chain, 14
galatheae, Xylophaga, 4, 22
gerda n. sp., Xylophaga, 4, 14, 15-16, 49-50
globosa, Xylophaga, 5, 9, 36, 39
grevei, Xtjlophaga, 4, 16—17, 51
growth series, 12
guineensis, Xylophaga, 5, 32, 3-^-35, 40, 72-73
hadalis, Xylophaga, 4
indica, Xylophaga, 5, 40
ingolfia, Xyloredo, 44, 44—45, 84
japonica, Xylophaga, 5, 24, 37, 39^0, 79
Lignopholas, 3
lobata, Xylophaga, 4, 22, 24
Martesia, 3
mesoplax
as character used in grouping, 3-5
variation in, 11-12
mexicana, Xylophaga, 5, 32, 35^37, 38, 40, 74-75
muraokai n. sp., Xylophaga, 4, 12, 25—27, 46, 65—66
murrayi, Xylophaga, 4, 18
naceli, Xyloredo, 44, 45, 45^6, 85
nooi, Xyloredo, 13, 43-44, 46, 82
obtusata, Xylophaga, 4, 20, 21
panamensis, Xylophaga, 4, 18
Pholadidea, 40
pholadids, 3, 41
praestans, Xtjlophaga, 5, 12, 31
profunda, Xylophaga, 13, 14, 24
profunda n. sp., Xylophaga, 13, 14, 21-23, 24, 59-60
reproduction, 3
rikuzenica, Xylophaga, 5, 30—31, 69
siphons, 3—5
supplicata, Xylophaga, 4, 18, 19-20, 20, 21, 22, 56
Teredinidae, 41, 43
teredinids, 2, 3, 43
teremachi, Xylophaga, 5
tipperi n. sp., Xylophaga, 5, 32, 37, 37—38, 76
tonilini, Xylophaga, 5
tubulata, Xylophaga, 4, 20, 21
tumerae, Xylophaga, 5
variation
due to substrate, 6, 7t, 7-10, 10-11
in growth series, 12
in mesoplax, 11-12
washingtona, Xylophaga, 5, 27, 28, 28-30, 31, 68
variation due to substrate in, 6, 7t, 7-10, 10-11
whoi n. sp., Xylophaga, 4, 20-21, 22, 57-58
wolffi, Xtjlophaga, 4, 16, 18
Xylophaga,
distribution of, 2, 6
groups in, 3—5
nomenclature of parts of, 4
Xylopholas, 2, 40-41, 42
Xyloredo Turner, 2, 41, 42-43
(US ISSN 0027-4100)
Sulletln OF THE
Museum of
Comparative
Zoology
THE SOLAS SPIDERS OF
THE GENUS MASTOPHORA
(ARANEAE: ARANEiDAE)
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VOLUME 157, NUMBERS
5 February 2003
(US ISSN 0027-4100)
PUBLICATIONS ISSUED
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MUSEUM OF COMPARATIVE ZOOLOGY
HARVARD UNIVERSITY
Breviora 1952-
bulletin 1863-
Memoirs 1865-1938
JoHNSONiA, Department of Mollusks, 1941-1974
Occasional Papers on Mollusks, 1945-
SPECIAL PUBLICATIONS.
1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and
Evolution of Crustacea. 192 pp.
2. Turner, R. D., 1966. A Suivey and illustrated Catalogue of the Tere-
dinidea (Mollusca: Bivalvia). 265 pp.
3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino-
derms. 284 pp.
4. Eaton, R. J., 1974. A Flora of Concord from Thoreau's Time to the
Present Day. 236 pp.
5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology
and Evolutionary Biology: Essays in Honor of Ernest E. Williams.
725 pp.
6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp.
Other Publications.
Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Culf of Maine.
Reprinted 1964.
Brues, C.T., A. L. Melander, and F M. Carpenter, 1954. Classification of
Insects. {Bulletin of the M. C. Z, Vol. 108.) Reprinted 1971.
Creighton, W. S., 1950. The Ants of North America. Reprinted 1966.
Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In-
ternational Symposium on Natural Mammalian Hibernation. {Bulletin
of the M. C. Z, Vol 124.)
Orinthological Gazetteers of the Neotropics (1975-).
Peters Check-list of Birds of the World, vols. 1-16.
Proceedings of the New England Zoological Club 1899-1947. (Complete
sets only.)
Proceedings of the Boston Society of Natural History.
Price list and catalog of MCZ publications may be obtained from Publica-
tions Office, Museum of Comparative Zoolog)'^, Harvard University, Cambridge,
Massachusetts 02138, U.S.A.
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Plate 1 . Upper left, Mastophora diablo new species (Argentina, photo J. Abalos). Upper right, M. stoweinew species (Kentucky,
photo K. F. Haynes and K. V, Yeargan). Middle left, M. yearganinew species (Kentucky, photo K. F. Haynes and K. V. Yeargan).
Lower left, M. alvareztoroi Ibarra and Jimenez new species (Texas, photo M. Stowe). Lower right, M. fasciata Relmoser (Costa
Rica, photo W. Eberhard).
THE BOLAS SPIDERS OF THE GENUS MASTOPHORA
(ARANEAE: ARANEIDAE)
HERBERT W. LEVP
Abstract. Of 48 species of bolas spiders {Masto-
phora) found in the Americas, 22 are new. Of that
number, nine new species and six previously kno\\ai
species are North American. The North American
Mastophora bisaccata is a group of species of similar
appearance. We can expect additional finds of new
species of these rare and specialized spiders. The spe-
cies range from New Hampshire in the United States
to central Argentina. No species are known from
tropical Amazon or the northwestern states of the
United States. The greatest abundance of species is
in warm temperate areas of southeastern North
America and southern Brazil and northern Argentina.
Agatostichtis is synonymized \\'ith Mastophora. With-
in this paper, Ibarra and Jimenez describe a new spe-
cies from Chiapas and Texas. Evidence from palpal
moi"phology indicates a relationship of Kaira with
Taczanoivskia and Mastophora, suggesting that insect
attractants may have evolved only once.
Epeiroides fasciolata, erroneously placed in Mas-
tophora, is a Kaira, the male of Kaira altiventer The
related Asian genus Euglijptila is synonymized with
Ordgarius.
INTRODUCTION
Female bolas spiders spend die day
resting on leaves and branches, usually
mimicking bird droppings and sometimes
berries, snails, or leaf buds. At night, the
bolas spider feeds on male inoths attracted
by the spider's scent; the scent mimics the
sex attractant of the female moth. This
pheromone was first suggested by Hutch-
inson (1903). Evidence of this pheroixione
was found by Eberhard (1977), and chem-
ical analyses were conducted by Stowe et
al. (1987) and Gemeno et al. (2000). An
approaching inale inoth is caught with a
silken thread bearing a viscid drop, the bo-
' Museum of Comparative Zoology, Harvard Uni-
versity, 26 Oxford Street, Cambridge, Massachusetts
02138-2902.
las, hurled at the moth. Moths stick to the
bolas, whereas detachable wing scales per-
iTiit moths to escape from most orb webs.
The unusual behavior of Mastophora, first
obsei-ved by Hutchinson (1903), has at-
tracted the attention of researchers, in-
cluding taxonomists (Eberhard, 1981;
Stowe, 1986; Stowe et al, 1987; Yeargan,
1988, 1994, 1997). Unlike most American
orb weavers, the genus Mastophora has
been revised in the past. That is, the de-
scribed species were compared and illus-
trated and keys were ixiade. Mello-Leitao
(1931) first reviewed all known species,
and in the same year Canals (1931) revised
all Argentinean species. Both authors re-
lied on differences in the shape of the
horns on the cephalothorax. The presence
of horns is a character of the genus and is
difficult to use for differentiating species.
Genitalia were not illustrated until Gertsch
(1955) revised the North American spe-
cies. Unfortunately, Gertsch's illustrations
were poorly labeled. Those with legends
indicating an internal view of the epigyn-
um actually were the cleared posterior of
the epigynum. Gertsch correctly reported
that the diversity of egg sacs from Florida
suggested overlooked species.
Most of our knowledge of the biology of
these spiders comes froiu a few species,
mostly from North America. All late-instar
and adult female bolas spiders spin a hor-
izontal line composed of multiple threads,
and then attach a bolas to it. The bolas
consists of one, rarely several, balls of
sticky glue drops on a line. A moth attract-
ed by the spider's scent is caught by a
swing of the leg holding the bolas; the ad-
Bull. Mus. Comp. ZooL, 157(5): 309-382, Februaiy, 2003
309
310 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
hesive is strong enough to hold moths. The
moth is wrapped and usually the spider
builds a new bolas and continues hunting
before eating. After approximately 20 min-
utes or more of hunting, when the bolas
has not been used, the spider pulls it back
and ingests the silk and glue and spins a
new bolas.
The bolas of spiders in the genus Mas-
tophora is held with the first leg and
swung at prey. Members of the Australa-
sian genus Ordgarius and the African ge-
nus Cladonielea use the second leg and
whirl the bolas (Stowe, 1986; Yeargan,
1994; Leroy et al, 1998).
Only male moths are attracted. Stowe et
al. (1987) showed that M. cornigera pro-
duces several of the pheromone compo-
nents produced by females of the moth
prey species. Gemeno et al. (2000) showed
that M. phrynosorna produces prey pher-
omone components in proportions that
represent an attractive blend. Attractants
are released only while hunting (Gemeno
et al., 2000). In one study, M. dizzydeani
captured 2.2 moths per night, although a
moth approached the spider as it hunted
about once every 6 minutes (Eberhard,
1981). Different species of moths that
mate at different times and that produce
entirely different pheromones are caught
at different times during the night. Lists
of moth species captured were reported by
Stowe (1988), Yeargan (1994), and Stowe
et al. (1995).
Early instars of Mastophora of both sex-
es and the minute adult males rest on the
edges of leaves and feed mostly on male
nematoceran flies, primarily Psychodidae
(moth midges), which also are attracted by
scent (Yeargan and Quate, 1996, 1997).
The flies are captured with the first two
pairs of legs, without the use of silk. The
legs are armed with rows of strong setae
(Figs. 5, 6). The spiderlings do not feed on
each other (Stowe, 1986; Yeargan, 1994).
In later instars, females lose these bristles
and start to use a bolas.
North American species have only 150—
250 eggs in each of one to five brown egg
sacs (Figs. 445—465). Each egg sac is the
size of the female. Mastophora cornigera
makes more egg sacs. Female Mastophora
in the northeastern states die in autumn.
More eggs have been reported from other
species. Mastophora extraordinaria was
found to produce 530 eggs (Brethes, 1909)
and M. dizzydeani produced 826 eggs
(Eberhard, 1981). Oviposition of M.
hiitchinsoni (a North American species)
takes place in fall and spiderlings emerge
in May (Yeargan, 1988). This differs for M.
cornigera. Clutch sizes for M. hiitchinsoni
ranged froni 178 to 275. The sex ratio ap-
proached one to one (an unusual excep-
tion was observed in one egg case of M.
phrynosorna, see below). Males and fe-
males are similar in size at hatching but
females grow to be much larger than
males. Males mature in June, at about 1.7
mm total length, two months before fe-
males become mature.
The scent may come from the integu-
ment (Lopez, 1998). The horns of the car-
apace of Mastophora contain midgut di-
verticula (Lopez et al., 1985). The silk
glands were described by Lopez andi
Stowe (1985).
The females rest in exposed places dur-
ing the day with legs drawn in, often on a
small pad of silk. All species are ciyptic
and uncommon, and difficult to find when
present. The clustered egg sacs are sus-
pended by strong threads on branches,
and are noticed more often than the spi-
ders, particularly in deciduous forests after
leaves have fallen. When a spider is picked
up, it rolls in the hand rather than holding
on, and when first disturbed may regur-
gitate fluid that has a pungent odor (Eber-
hard, 1981). A summary of research was
reported by Yeargan (1994).
METHODS AND ACKNOWLEDGMENTS
The collections of the following institu-
tions and individuals were used.
AMNH American Museum of Natural
History, New York (N. Platnick,
L. Sorkin)
Mastophora • Levi
311
Natural Histoiy Museum, Lon-
don, United Kingdom (P. Hill-
yard, J. Margerison)
California Academy of Scienc-
es, San Francisco, California
(C. Griswold, D. Ubick)
Canadian National Collections,
Ottawa, Canada (C. Dondale)
Clemson University Arthropod
Collection, Clemson, South
Carolina (J. Moore, D. Carna-
gey)
Cornell University collection,
kept in AMNH (N. Platnick, L.
Sorkin)
Denver Museum of Nature
and Science, Denver, Colorado
(P. Cushing)
D. Ubick, San Francisco, Cali-
fornia
El Colegio de la Frontera Sur,
Tapachula, Chiapas, Mexico
(G. Ibarra)
Facultad de Ciencias, Seccion
Entomologia, Montevideo,
Uruguay (M. Simo)
Fundacion Miguel Lillo, Tu-
cuman, Argentina (S. Z. Turk,
J. A. L. Haedu)
Florida State Collection of Ar-
thropods, Gainesville, Florida
(G. B. Edwards)
Institute Butantan, Sao Paulo,
Brazil (A. Brescovit)
Illinois Natural History Sui'vey,
Urbana, Illinois (C. Favret)
Institut Royal des Sciences Na-
turelles de Belgique, Brussels,
Belgium (L. Baert)
J. Beatty, Carbondale, Illinois
J. Kaspar, Oshkosh, Wisconsin
J. Murphy, London, United
Kingdom
K. V. Yeargan, Lexington, Ken-
tucky
Museo Argentine de Ciencias
Naturales, Buenos Aires, Ar-
gentina (M. E. Galiano, C. L.
Scioscia)
Museu de Ciencias Naturais,
Fundayao Zoobotanica do Rio
Grande do Sul, Porto Alegre,
Rio Grande do Sul, Brazil (E.
H. Buckup, M. A. L. Marques)
MCP Museu de Ciencias, Pontificia
Universidade Catolica do Rio
Grande do Sul, Porto Alegre,
Rio Grande do Sul, Brazil (A.
A. Lise)
MCZ Museum of Comparative Zo-
ology, Cambridge, Massachu-
setts
MKS M. K. Stowe, Gainesville, Flor-
ida
MLJ M. L. Jimenez, La Paz, Mexico
MLP Museo de Universidad Nacion-
al. La Plata, Argentina (C.
Ituarte, L. A. Pereira)
MNHN Museum National d'Histoire
Naturelle, Paris, France (C.
Bollard)
MNRJ Museu Nacional, Rio de Janei-
ro, Brazil (A. B. Kury)
MUSM Museo de Historia Natural,
Universidad Nacional Mayor
de San Marcos, Lima, PeiTi
(Diana Silva D.)
MZAQ Museu, Departamento de
Zoologia da Escola Superior de
Agricultura "Luis de Queiroz,"
Piraciba, Sao Paulo State, Bra-
zil (G. J. de Moraes)
MZSP Museu de Zoologia, Universi-
dade de Sao Paulo, Sao Paulo,
Sao Paulo, Brazil (E. M. Can-
cello, R. Pinto da Rocha)
NHMW Naturhistorisches Museum, Vi-
enna, Austria (J. Gruber)
NMB Naturhistorisches Musuem,
Basel, Switzerland (A. Hanggi)
NMP Natal Museum, Pietermaritz-
burg. South Africa (D. A. Bar-
raclough, C. Conway)
OSU Ohio State University, Marion,
Ohio (R. A. Bradley)
QMB Queensland Museum, Bris-
bane, Queensland, Australia
(R. J. Raven)
TAMU Texas A&M University, College
312 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
1
USNM
ZMUC
Station, Texas (A. Dean, E. Ril-
ey)
National Museum of Natural
History, Smithsonian Institu-
tion, Washington, D.C. (J.
Coddington, L. Lopardo)
Zoological Museum of the Uni-
versity of Copenhagen, Copen-
hagen, Denmark (N. Scharff)
Brazilian colleagues Erica H. Buckup
and Arno Lise encouraged me to revise
Mastophora. The revision was made pos-
sible with the help of numerous South
American colleagues with loan of collec-
tions, including many old specimens, and
advice. Especially helpful were C. Scioscia
and the late M. Galiano of Buenos Aires,
and the Brazilians E. H. Buckup, A. Bres-
covit, R. Pinto da Rocha, G. J. de Moraes,
and A. B. Kuiy. I also thank G. S. Oxford,
K. V. Yeargan, M. K. Stowe, W. Eberhard,
J. Leroy, and A. Leroy for help and advice
and Eric J. Olson for translating difficult
Spanish. L. Leibensperger was helpful in
the laboratoiy. L. R. Levi and the editor,
B. L. Clauson, polished the writing. W.
Eberhard read the introduction and im-
proved the wording. K. Yeargan and M. K.
Stowe read the manuscript and suggested
many improvements and corrections.
Much appreciated are the comments of
two anonymous readers of the manuscript.
Gifts of Mastophora to the MCZ collec-
tions were received from C. Hieber, G.
Ibarra (from Chiapas, Mexico), J. M. Maes
(from Nicaragua), M. K. Stowe, and K. V.
Yeargan. G. Ibarra N. and M. L. Jimenez
sent specimens of a new species with com-
plete descriptions and finished illustra-
tions. Mastophora alvareztoroi is de-
scribed here with the names of Ibarra and
Jimenez as authors.
The project was started with the help of
National Science Fooundation grant BMS
75-05719. The Wetmore-Golles Fund cov-
ered publication costs in part.
For examination of spiders, I followed
the procedures described by Levi (1993a).
For examination and illustration of the
small and difficult genitalia oi Mastophora,
additional methods were employed.
After illustrating ventral and posterior
views of the female, the epigynum was cut
off from the body and the soft tissues were
carefully removed (Fig. 2). The epigynum
was placed on the dry end of a dish with
white paraffin, and extra alcohol was re-
moved with a bit of tissue. The epigynum
was then placed in a conical pit in the par-
affin (Fig. 3) containing two drops of Hoy-
er's medium.- Placement was so that the
anterior end faced the bottom; the flat
posterior was parallel with the surface
(Fig. 3). The viscous nature of Hoyers me-
dium makes it an ideal clearing medium
and keeps the epigynum in position, al-
though presumably other clearing agents,
such as oil of cloves or glycerol, could be
used. The genitalia were carefully exam-
ined and illustrated from at least one spec-
imen of each species; several epigyna were
cleared for abundant and variable species.
Male spiders for which determination
was known were placed in a drop of tinc-
ture of iodine and left overnight before ex-
amining them the next morning in 80%
alcohol. Tincture of iodine gives a good
stain, which may not be permanent, and it
is easy to obtain in pharmacies. Tincture
of iodine is isotonic with alcohol and does
not expand the palpus or wai"p sclerites.
Gertsch (1955) in his revision did not'
intei"pret the two original descriptions by
Banks (1898) of Ordgarius ohesus and O.
coiyulenta. The holotypes were placed in
the California Academy of Science and
were destroyed in the 1906 earthquake.
The specimens, with others, were given to
Banks from the Marx Collections after
George Mai-x died. Banks (1898) pointed
out at the time:
- Hoyers medium is made by dissolving 15 g of
gum arabic (clear flakes) in 25 ml of distilled water
at room temperature. Seventy-five grams of chloral |
hydrate is added, and the mixture is allowed to stand ■
for 1-2 days, until all solids have dissolved. Five mil-
liliters of glycerol is then added and the mixture is
filtered through glass wool and stored in a glass-stop-
pered botde.
Mastophora • Levi 313
Anyone familiar with Dr. Marx's methods of work
will not be surprised to learn that many of the spec-
imens, when sent to me, bore no locality label
whatever. Doubtless he knew where they came
from, but left no clew [sic] that others might use.
Some of the species were numbered, and by ex-
amining several of his series of numbers it was pos-
sible to find localities and his name for the species.
However, Banks published illustrations of
the two species that can be identified if
one disregards the localities stated.
DISCUSSION
Determination of members of the genus
Mastophora is the most difficult of araneid
spiders. By using morphological charac-
ters, 48 presumed species were separated,
of which 22 species are new. Eleven of
these new species are known from a single
female. However, additional specimens
were found of species earlier described
from only a single specimen.
All females are between 7 and 17 mm
and most are approximately 12 mm total
length. Most of the species that lack
humps on the abdomen seem to be found
in North America. A few species of Mas-
tophora are slightly smaller and have dif-
ferent tubercles or horns on the carapace;
these were placed in Agatostichus in the
past, but males and females do not differ
in other characters from Mastophora.
Adult males of all species are approxi-
mately 1.7 mm total length. Males of re-
lated genera in Asia, Australia, and Africa
are the same length. Males of the same
species as females with abdominal huinps
coming from different egg sacs may have
humps or not; females without humps may
have males with or without humps. The
males can be matched to females only by
raising them from an egg sac from a de-
termined female. Some males apparently
emerge from the egg sac as mature adults
(M. cornigera and perhaps M. gasteracan-
thoides), but most take at least two instars
to mature. The males of only a few species
are known, because only W. Eberhard,
Mark K. Stowe, and Ken V. Yeargan have
raised individuals. The males of most
southern South American species remain
unknown. The palpi of different species
are sui"prisingly similar and the determi-
nation of males presents a challenge.
Immatures coming out of the egg sac
have visible median tubercles and horns
on the carapace and are approxinately 1.2—
1.6 mm total length. They may have ab-
dominal humps.
Some egg sacs can be determined.
Those of M. hutchinsoni are unique and
are attached by their base (Figs. 453, 454),
whereas all others are hanging. Some have
a thick stalk (M. bisaccata; Fig. 446), extra
long flaps (M. phrynosoma; Figs. 449,
450), or may lack flaps altogether (M. cor-
nigera; Fig. 455). However, not enough
determined egg sacs were available to
make a key.
The epigyna of females are much re-
duced. Females are separated by their col-
oration and shape of the abdomen, as well
as by their genitalia. However, with so few
specimens available of most species, gen-
eralizations on appearance are a guess.
The most common North American spe-
cies, M. bisaccata, was found to be a group
of species with similar abdominal mark-
ings. Gertsch (1955) missed this because
he relied on coloration and did not care-
fully examine the genitalia of all speci-
mens. Members of the M. bisaccata spe-
cies group also differ slightly in average
size, with M. bisaccata being the largest.
Gertsch (1955) and Yeargan (1994) sus-
pected M. bisaccata to be a group of spe-
cies.
The epigynum lacks a scape and has no
ventral features permitting the palpus to
be held in place (Figs. 12, 19, 26). A pos-
terior edge (Figs. 12, 19, 26) may be pre-
sent. Whatever diagnostic features exist
are on the posterior face, which has two
slits, a plate between, and a plate to the
side (Figs. 13, 20, 27). The plates are usu-
ally weakly sclerotized. The slits lead into
a ventral, or sometimes dorsal, atrium
(Fig. 4), which empties into the seminal
receptacles (Fig. 4). The slits vary in di-
rection in different species (Figs. 13, 27).
The slits may be in depressions or may
314 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
have unique sculpturing along the edges,
or a lip (Fig. 20). I considered these to be
useful characters for determination of spe-
cies.
The palpi of males are lightly sclero-
tized. A median apophysis, a radix, the em-
bolus, and a terminal apophysis holding
the embolus are present (Fig. 7). The con-
ductor, a structure that arises from the te-
gulum facing the embolus, is absent, al-
though it seems to be present in Clado-
nielea species (Fig. 443). The different
species of Ordgarius and Cladornelea show
greater morphological differences from
each other than do those of Mastophora.
Different Ameiican species show some
variation in genitalia and morphology in
different specimens. Perhaps this is asso-
ciated with the rarity of individuals and
wide distribution of species.
All books on venoms list M. gasteracan-
thoides among the venomous American
spiders (Schmidt, 2000). These citations all
come from Escomel (1918), who de-
scribed the venom from specimens found
in grapes in southern Peru. Farmers work-
ing in grapes were bitten on the hands and
legs. He considered the bites to be "cu-
taneo-hemolytique gangrenieux" and gave
detailed description of the signs, which
were necrotic skin lesions at the site of the
bite. B. A. Houssay and J. J. Carbonnel
checked the specimens at the time. In bor-
rowed material, Escomel's specimens were
found in both the Buenos Aires and in the
Paris museums. Examination showed them
to be a new species of Mastophora (M. e.s-
comeli), close to M. gasteracanthoides. No
other evidence of human envenomation
appeared with any labels on specimens of
other species examined. Gertsch (1955)
believed that the responsibility for the
bites should be awarded to some other spi-
der or arthropod. I suspect that M. esco-
meli, unlike others, is more aggressive and
more readily bites human skin than other
species. No recent reports exist of ven-
omous bites from M. escomeli. Mastophora
escomeli may have been abundant in 1917
and then become rare again. Escomel
(1918) also found that extracts from eggs,
injected into guinea pigs, were toxic.
Relationship. Mastophora Holmberg,
1876, shares carapace outgrowths with
Australasian Ordgarius Keyserling, 1886
(Figs. 422-433), and African Cladornelea
Simon, 1895 (Figs. 4M-A4.4:). Both Ord-
garius and Cladornelea species handle the
bolas with the second leg and, unlike Ma.s-
tophora, which swings the bolas in a pen-
dulumlike motion, they whirl the bolas,
Ordgarius when a moth approaches, and
Cladornelea for 15 minutes at a time when
hunting, at approximately 150 rotations
per minute (Leroy et al., 1998).
The carapace outgrowths also are
shared with the African Acantharachne
Tullgren, 1910,^ Madagascan Coelossia Si-
mon, 1895, Madagascan Exechocentrus Si-
mon, 1889, and immatures of Euglijptila
Simon, 1908, from northern Vietnam
(Tonkin). Euglijptila is synonymized below
with Ordgarius. The males of these and
their habits are not known, but females
were illustrated and described by Emerit
(1980, 2000). The genus Agatostichus Si-
mon, 1895, is synonymized below with
Mastophora, and Dicrostichus Simon,
1895, has been synonymized with Ordgar-
ius by Davies (1988). The Mastophora,
Ordgarius, Dicrostichus, Cladornelea
group is absent from European, Mediter-
ranean, and central Asian faunas.
When males are found, African Acan-
tharachne and Coelossia should probably
be synonymized with Cladornelea or Ord-
garius. However, Exechocentrus differs in
having a long eye projection, a long me-
dian tubercle, and a pair of long, posterior
tubercles on the carapace (Emerit, 1978,
1980, 2000).
Eberhard (1981), Stowe (1986), and i
Yeargan (1994) studied the relationships
with other genera. Scharff and Coddington
' Roevver (1942) and Platnick (2001) cited the ge-
nus under the name Acantharanea Strand, 1929. The
name Acantharachne Tullgren, 1910, is not preoc-
cupied, as thought by Strand (Neave, 1939a: 9; Bon-
net, 1955: 124).
Mastophora • Levi 315
constructed a cladogram (1998). The re-
sults of Eberhard (1981), Stowe (1986),
and Yeargan (1994) are summarized in Ta-
ble 1; examination of these results showed
a close relationship of Mastophora to Tac-
zanowskia Keyserling, 1880. Members of
the genus Mastophora seem related to
genera lacking carapace tubercles (Tacza-
nowskia, Celaenia, Kaira, Cijrtaracline,
Poecilopaclujs, and PasUobiis; Table 1).
Robinson and Robinson (1975) first sug-
gested that the web of Pasilohiis was in-
termediate between orb webs and bolas.
My own studies of genitalia of Taczanows-
kia (Levi, 1997) showed that Taczanowskia
is related to Celaenia and Kaira. The distal
pocket of the epigynal scape of Tacza-
nowskia correlates with the large hook on
the median apophysis of the palpus of the
male (Levi, 1997, fig. 19). Unequal claw
lengths and armed femora, which are syn-
apomoiphies, relate Taczanowskia and Ce-
laenia. The denticles next to a tooth on the
side of the median apophysis of Tacza-
nowskia also are found in Kaira species
and Metepeira. Such median apophysis
denticles are unique to several genera and
I consider such a row of denticles as a syii-
apomoi-phy of Kaira, Metepeira, and Tac-
zanowskia. One of the synapomoi-phies of
most genera allied to Araneus is a spine or
tooth on the median apophysis. In con-
trast, males that have a paramedian apoph-
ysis in the palpus rarely have a tooth or
spine on the median apophysis (e.g., Al-
paicla, Eriophora, Ocrepeira, Acacesia,
Cijrtophora, and many others). The shape
of the median apophysis of the palpus of
Pasilobiis also probably is derived from a
median apophysis similar to that of Kaira.
The presence of a spine on the median
apophysis of all these genera with carapace
outgrowths indicates a distant relationship
with Araneus. All these genera have a ter-
minal apophysis (Fig. 7) in the palpus,
which was erroneously labeled as a con-
ductor by some authors. The evidence
from the study of genitalia thus shows that
attraction of insects in Kaira and Masto-
phora most likely evolved only once, not
twice as thought previously (Stowe, 1986;
Table 2). The homology of secreting
glands of the insect attractant in Masto-
phora and Kaira remains uncertain.
TAXONOMIC SECTION
Mastophora Holmberg
Mastophora Holmberg, 1876: 112. Type species M.
extraordinaria Holmberg by monotypy. The gender
of the name is feminine. Neave, 1940: 55. It is not
preoccupied as claimed by Bonnet, 1957: 1995.
Mello-Leitao, 1931: 65. Canals 1931: 17. Roewer,
1942: 900. Gertsch, 1955: 223. Platnick, 2001.
Heterocephala Holmberg, 1876: 143. Type species H.
conifera Holmberg by monotypy. The gender of the
name is feminine. Neave, 1939b: 634. It is not pre-
occupied.
Glyptocranium Simon, 1895: 885. Type species G.
comlgenim Hentz designated by Simon. Neave,
1939b: 484. The gender of the name is neuter.
Bonnet, 1957: 1995. First synonymized by Brethes,
1909.
Agatostichus Simon, 1895: 885. Type species A. leii-
cacantha Simon by original designation and mon-
otypy. Neave, 1939a: 86. Gertsch, 1955: 250. NEW
SYNONYMY.
AgathosticJuis: — Simon, 1895: 473. Roewer, 1942:
900. Bonnet, 1955: 181. Platnick, 1998, 2001.
Note. Simon spelled the generic name
Agatostichus with and without "h," but the
first revisor of the genus, Gertsch (1955),
spelled the genus without "h." Subsequent
users must follow the first revisor (Inter-
national Code of Zoological Nomenclature,
art. 24.2 [International Commission on
Zoological Nomenclature, 1999]). The ge-
nus is synonymized here because the type
species M. leucacantha has long median
tubercles on the carapace (Fig. 316). The
second species described, M. leucahulha
Gertsch, lacks these but has a tubercle be-
tween the posterior median eyes and is
relatively small (Fig. 288). Gertsch thought
small size was diagnostic for the genus, but
this is not the case for the third species,
M. alvareztoroi, and also not for the type
species. Tubercles also are found between
eyes in M. coiyiilenta (Figs. 336, 337).
A revision of all species of the genus was
made by Mello-Leitao (1931) and a revi-
sion of Argentinian species was made by
Canals (1931). Both authors had shared in-
formation. I could not find dates of pub-
316 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
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Mastophora • Levi
317
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318 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
lication of the two 1931 articles, but Mel-
lo-Leitao's must have been pubhshed first,
because Canals cites the page numbers of
Mello-Leitao. Gertsch (1955) revised the
North American species, and first illustrat-
ed the genitalia of the species of his re-
gion.
Diagnosis. Mastophora differs from oth-
er araneid genera by having the carapace
with tubercles and having horns (Fig. 8),
and lacking macrosetae on legs (Figs. 5, 6,
421). Early instars and males have rows of
setae on distal articles of first and second
legs (Figs. 5, 6). Mastophora differs from
Cladoinelea and Ordgaritis, which may
have horns (Fig. 423), by using the first
pair of legs to handle the bolas; Clado-
melea and Ordgarius use the second pairs.
Description. Female. Carapace color
variable, often uniform orange to strongly
marked. Legs usually not banded. Abdo-
men often darker anteriorly, lighter pos-
teriorly, venter often with a median white
square or rectangle (black in M. hutchin-
soni and some M. phrijnosoma). Carapace
about as wide as long. Eyes small, sub-
equal. Anterior median eyes slightly larger,
lateral eyes smallest. Median ocular trap-
ezoid always longer than wide, widest at
posterior median eyes, rarely almost
square. Lateral eyes and medians usually
on a bulge. Clypeus higher than two an-
terior median eye diameters. Carapace
veiy high, with median tubercles, a pair of
horns, usually biforked, and laterally with
medium to less large tubercles (Figs. 8, 9),
sometimes with short white setae. Chelic-
erae with two or three teeth on the ante-
rior margin, one on the posterior margin
{M. hisaccata has two or three teeth; Fig.
1). Legs without macrosetae, early instars
with row of setae on legs as in mature
males (Fig. 5). Length of first patella and
tibia about 1.1 to 1.8 times width of cara-
pace. Second leg longer than first, third
shortest. Abdomen wider than long usually
with a pair of humps, and a pair of scler-
otized discs between humps and anterior
margin (Fig. 10), sometimes with addition-
al tubercles or scattered clumps of setae
or setose. Some species (M. alvareztoroi,
M. felis, and M. haijwardi) have carapace
and abdomen hirsute (Figs. 107—109). The
tubercles on the carapace and on horns are
not perfectly symmetrical and show indi-
vidual differences. The abdominal humps
of some species (M. gasteracanthoides) are
of variable length.
Male. Mature after only two instars or
emerging as adult from egg sac (M. cor-
nigera), all about 1.7 mm total length.
Dirty orange color, sometimes with white
spots on carapace or abdomen. Carapace
as wide as long with eyes on bulges as in
female. Carapace with two median tuber-
cles and minute horns. Legs without ma-
crosetae but with row of soft setae on first
two pairs of legs, slightly shorter than in
females (Figs. 5, 6). Total length of first
patella and tibia about 1.1 times width of
carapace. Abdomen with or without
humps, often in the same species, regard-
less of the presence of humps on the fe-
male abdomen.
Female genitalia. The epigynum has
plates on the ventral side, and differs only
slightly from other species in posterior
view (Figs. 2, 13, 20, 27) but has lost all
copulatory structures. The posterior has
two slits with shadows of atria.
Male genitalia. Palpus of male has a
pointed median apophysis, an embolus
with a simple terminal apophysis (Fig. 7),
but no distinct conductor.
One group of Mastophora is distinct:
the species close to M. gasteracanthoides
{corpulenta, rabida, esconieli, ohtusa, felis,
hohnbergi, reimoseri, satan, and diablo).
Their carapace is high with vertical sides
(Fig. 408); the sides of the thorax have tu-
bercles (Fig. 409), with short, white setae
between; and the abdomen may have high,
tube-shaped horns (Fig. 413). The abdo-
men of all is dark and species can be sep-
arated only by studying the genitalia. They
have been referred to in Spanish as the
cat's head spiders (arana cabeza de gato)
because of their resemblance to the head
of a cat, as can be seen in Figure 421.
Mastophora • Levi
319
F/^B.\ cS'' .o^^
FR. GUIANA
GALAPAGOS
Map 1. Approximate number of species in various American regions.
Relationship. Relationships with other
genera are discussed above.
Natural History. Mastophora all are un-
common and difficult to find. Females rest
on branches or leaves of trees, usually 1.5—
3 m high. M. K. Stowe (personal com-
munication) has seen egg cases 10 m up
and thinks that M. hisaccata seeks out
branches high up in trees. They often are
found on trees and bushes in orchards,
gardens, or along fences. M. K. Stowe
(personal communication) reported that
most species in Florida are found in for-
ests.
Distribution. Mastophora are only
found in America (Map 1).
Misplaced Species. Mastophora fasciol-
ata erroneously placed in Mastophora
(Levi, 1991: 180) is a Kaira. See below.
Key to Female Mastophora
1. Abdomen without humps (Figs. 10, 24,
88, 130); swellings may be visible in
profile (Fig. 11) 2
— Abdomen with distinct humps or tuber-
cles (Figs. 137, 145, 298, 331, 401) ... 15
2(1). North Anierica (Map 2) 3
- South America (Maps 3, 4) 11
3(2). Abdomen subtriangular, with anterior
320 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
lateral swellings (Fig. 88); posterior of
epigynum with lateral and ventral lips
(Fig. 91); eastern United States (Map
2D) phrynosoma
— Abdomen without anterior lateral swell-
ings (Figs. 10, 17, 24); epigynum with-
out posterior lips (except apalachicola)
4
4(3). Posterior of epigynum with slits framed
by lateral lips (Fig. 20); southeastern
United States (Map 2C) apalachicola
Slits without lips (Figs. 13, 27) 5
5(4). Slits short, dieir length distant from ven-
tral margin (Fig. 80); eastern United
States (Map 2A) yeargani
— Slits approaching close to ventral margin
(Figs. 13, 27) 6
6(5). Dorsum of abdomen with bisaccata pat-
tern, anterior orange to gray with
spaces and lines (Figs. 42, 53, 65) 7
— Abdomen marked otherwise (Figs. 10,
24) 9
7(6). Slits approaching each other ventrally,
atria their diameter or less apart (Figs.
56, 57); eastern United States (Map
2B) bisaccata
— Slits parallel or separating ventrally
(Figs. 45, 68) 8
8(7). Shts parallel (Fig. 45), atria visible in
ventral view (Figs. 44); Florida (Map
2G) alachua
— Atria slightly separating ventrally (Figs.
68-70), atria not visible ventrally
(Figs. 67); eastern United States (Map
2C) .stowei
9(6). Atria separating from each other ven-
trally (Figs. 34, 35); legs ringed; Vir-
ginia to Florida (Map 2A) timuqua
— Atria approaching each other (Figs. 13,
27); Florida ." 10
10(9). Abdomen black on anterior and sides
(Figs. 24, 25); atria anterior of seminal
receptacles, seminal receptacles their
diameter distant from ventral border
(Fig. 28); southern Florida (Map 2A)
felcla
— Abdomen with only anterior median
area black (Fig. 10); atria at level with
large seminal receptacles, seminal re-
ceptacles less dian their diameter dis-
tant from ventral border (Fig. 14);
Florida (Map 2C) .satsiima
11(2). Abdomen hirsute (Figs. 109, 110), slits
and atria separating ventrally (Figs.
112, 113); Tucuman, Argentina (Map
3C) liaijwardi
— Abdomen with few setae (Figs. 102,
116); atria otherwise (105, 119) 12
12(11). Abdomen with many black spots and
dark lines (Figs. 130, 131); southern
Brazil (Map 3B) carpogastra
— Abdomen marked otheiAvise (Figs. 102,
116, 123) 13
13(12). Abdomen with light longitudinal lines
(Figs. 102, 103); atria approaching
each other ventrally (Fig. 105); Santa
Catarina, southern Brazil (Map 3C) --
- catarina
— Abdomen marked otherwise (Figs. 116,
123) 14
14(13). Abdomen marked with pair of black
rings, open anteriorly (Fig. 116); Sao
Paulo, Brazil (Map 3C) conimbatai
— Abdomen with two black discs (Fig.
123); Lara, Venezuela (Map 3A) lara
15(1). Abdomen with many dorsal or lateral
humps (Figs. 267, 298, 300, 310, 331,
332) 16
— Abdomen with one pair of humps (Figs.
155, 171), rarely median area of ab-
domen swollen (Fig. 325) 19
16(15). Humps anterior on each side of abdo-
men (Fig. 267); Santiago del Estero,
Argentina (Map 3E) abalosi
— Humps or tubercles dorsally (Figs. 298,
310, 331) 17
17(16). Carapace with short tubercles (Figs. 329,
330); abdomen with numerous small
dorsal tubercles (Figs. 331, 332):
northern Argentina to Buenos Aires
Province (Map 3G) conifera
— Carapace with large tubercles (Figs. 297,
309) 18
18(17). Carapace with median tubercles spine-
shaped (Fig. 309); Panama (Map 2F)
soberiana
— Carapace with median tubercles cone-
shaped (Fig. 297); Texas to Chiapas
(Map 2F) alvareztoroi
19(15). Carapace median tubercles same size or
longer than horns (Figs. 288, 316, 323)
20
— Carapace with median tubercles smaller
than horns (Figs. 136, 381) 22
20(19). Median tubercles spine-shaped, very
long (Fig. 316); Bahia to Rio de Ja-
neiro, Brazil (Map 3G) leiicacantha
— Median tubercles othei"wise (Figs. 288,
323) 21
21(18). Median tubercles cone-shaped (Fig.
323); Rio Grande do Sul, Brazil (Map
3G) hrescoviti
— Carapace tubercles large, rounded on
dark carapace (Fig. 288); Texas, Mex-
ico (Map 2F) leucabulba
22(19). Carapace with sides vertical (Figs. 366,
380), sides widi tubercles (Figs. 367,
381); Mexico to South America (Map
4) 39
— Carapace with sides slanting (Figs. 135,
169); sides with few or no tubercles
(Figs. 136, 170; Maps 2, 3) 23
Mastophora • Levi 321
Map 2. Distribution of Mastophora species of North and Central America.
322 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
— r
^X^
l^
^^'
dizzydeani^, y6ji\-X^<^ ^\
^ lara
lilC-,'7a
cranion ^
longiceps t
pesqueiroU
pickeli ^
piras #
^ ! V/ abalosi ▼
~" comma ▼
melloleitaoi •
yacare A
ypiranga ■
f5^:;f^
extraordinaria
'^T^W^TT^'
brescoviti ▼
conifera •
leucacantha U
Map 3. Distribution of Mastophora species of South America.
Mastophora • Levi
323
North America and Cuba (Map 2) 24
Central and South America (Map 3) 28
Cuba (Map 2G); abdomen with anterior,
lateral swellings (Figs. 145, 146)
vaquera
Continental North America; abdomen
v\ithout lateral swellings (Figs. 137,
155) ': 25
Posterior of epigynum with slits and atria
approaching each other ventrally
(Figs. 159, 160); northeastern United
States (Map 2E) hutchinsoni
Slits and atria otherwise (Figs. 140, 174,
188) .__ 26
Posterior of epigynum with slits framed
by a lip on each side (Fig. 188); south-
eastern United States (Map 2E) archeri
Slits \\dthout lips (Figs. 140, 174) 27
Slits parallel and approach ventral bor-
der (Fig. 174); eastern United States
to California and Honduras (Map 2G)
cornigera
Slits short, their length apart from ven-
tral border (Fig. 140); southern Flori-
da Seminole
Central America and northern South
America to Peru 29
Southern South America, from Pernam-
buco State, Brazil, in the north 30
Humps of abdomen broad swellings
(Fig. 196); slits on posterior of epigyn-
um approaching each other (Fig. 199);
Costa Rica to Venezuela (Map 3A) ---
fasciata
Humps of abdomen narrow (Fig. 207);
slits separating ventrally (Fig. 210);
Colombia to Pena (Map 3A) .. dizzydeani
Humps of abdomen extended to a point
(Figs. 271, 272); Santiago del Estero,
Argentina (Map 3E) comma
Humps distally rounded (Figs. 277, 284)
31
Each side of abdomen with a pair of nar-
row dark-framed longitudinal, white
marks (Figs. 239, 240); humps small
(Fig. 240); Rio Grande do Sul, Brazil
(Map 3D) pesqueiro
Abdomen without such marks 32
Posterior of epigynum with a lip on each
side between sHts and ventral margin
(Fig. 263); Uruguay (Map 3E) ijacare
Posterior of epigynum without such lips
(Figs. 279, 286) 33
Epigynal slits appear forked (Figs. 235,
236); carapace horns unusually thick
and laid back (Figs. 230, 231); Sao
Paulo, Brazil (Map 3D) longiceps
Epigynal slits simple (Figs. 249, 256);
horns small (Figs. 245, 252) 34
Epigynal slits separating ventrally (Figs.
228, 249) 35
- Epigynal slits and atria approaching each
other ventrally (Figs. 256, 286) 36
35(34). Abdomen with a pair of dorsal stippled
dark spots (Fig. 225); Pernambuco
State, Brazil (Map 3D) cranion
- Anterior of abdomen black (Fig. 246);
Minas Gerais, Sao Paulo states, Brazil
(Map 3D) piras
36(34). Anterior of abdomen light, light area
bordered posteriorly by dark (Fig.
283); epigynum as in Fig. 286; south-
ern Brazil, northern Argentina (Map
3F) extraordinaria
- Anterior of abdomen black (Figs. 218,
253, 276) 37
37(36). Atria their diameter or less apart (Figs.
256, 257); Rio de Janeiro to Santa Ca-
tarina, Brazil (Map 3E) ypiranga
- Atria more than their diameter apart
(Figs. 221, 222, 279, 280) ._._ 38
38(37). Atria with median knobs facing each oth-
er (Fig. 280); Parana to Buenos Aires,
Argentina (Map 3E) melloleitaoi
- Atria without kjiobs (Fig. 222); Pernam-
buco, Brazil (Map 3D) pickell
39(22). Central America (Map 4A); posterior of
epigynum with dark patch dorsally in
each of pair of depressions (Fig. 341)
coiytdenta
- Other regions 40
40(39). Galapagos (Map 4A); abdomen wdth only
tiny humps (Fig. 346); epigynum slits
each with a loop ventrally (Figs. 348,
349) rabida
- Other regions; abdomen with larger
humps 41
41(40). Peru (Map 4A); posterior of epigynum,
between slits, with median area having
a bulge on each side (Fig. 355) escoineli
- Other regions 42
42(41). Chile (Map 4B); posterior of epigynum
with a dark spot dorsally in each ad-
jacent depression (Fig. 415) - -
gasteracanthoides
- Brazil to Argentina 43
43(42). Abdomen with humps placed on swollen
area (Figs. 364, 365); Pernambuco
State, Brazil (Map 4C) obtiisa
- Humps not placed on swollen area (Figs.
368, 369) 44
44(43). Dark patch (atria) placed dorsal or mid-
dle of seminal receptacles (Figs. 392,
404) 45
- Atria absent or placed ventrally of sem-
inal receptacles (Figs. 372, 379, 386)
46
45(44).
Posterior of epigynum with dark patch
(atria) dorsal within a depression
(Figs. 392, 394, 395); Pernambuco,
Brazil, to central Argentina (Map 4B)
satan
324 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Map 4. Distribution of Mastophora species of the M. gasteracanthoides group.
- Atria lateral, placed outside of depres- - Slits without such bend (Figs. 371, 378)
sion (Fig. 404); northern, central Ar- 47
gentina (Map 4D) diahlo 47(46). Posterior of epigynum with slits separat-
46(44). Posterior of epigynum with slits with a ing ventrally and with lateral lip (Figs.
ventral bend (Fig. 385); Paraguay 371, 372); Rio de Janeiro, Sao Paulo,
(Map 4C) reimoseri Brazil (Map 4C) felis
Mastophora • Levi
325
Slits parallel (Fig. 378); Paraguay, San-
tiago del Estero (Map 4C) holmbergi
Key to Known North American Male
Mastophora
Space surrounded by median apophysis,
in ectal view of palpus, longer than
wide (Figs. 50, 62, 74, 164, 168) 2
Space surrounded by median apophysis
wider than long (Figs. 39, 85, 96, 99,
179, 182); terminal apophysis usually
shorter than embolus (Figs. 37, 83, 94)
5
Terminal apophysis shorter than embo-
lus (Fig. 48); median apophysis of pal-
pus very short (Figs. 48-50); Florida
(Map 2G) alachua
Terminal apophysis as long or longer
than embolus (Figs. 7, 60) 3
Base of embolus large (Figs. 60, 61);
eastern United States (Map 2B)
bisaccata
Base of embolus small (Figs. 72, 73) 4
Base of median apophysis rounded
(Figs. 162, 166); in ectal view, narrow
part of median apophysis shorter than
base (Figs. 164, 168); northeastern
United States (Map 2E) hutchlnsoni
Base of median apophysis angular (Fig.
72, 73); in ectal view, narrow part of
median apophysis as long or longer
than width of base (Fig. 74); eastern
United States (Map 2C) stowei
Terminal apophysis almost as long as
embolus (Figs. 94, 177, 180) 6
Terminal apophysis about half length or
less of embolus (Figs. 83, 191) 7
Base of median apophysis longer than
wide (Figs. 177, 180); southern United
States to California and Central Amer-
ica (Map 2G) cornigera
Base of median apophysis short (Figs.
94, 98); eastern United States (Map
2D) phrynosoma
Base of median apophysis large, touch-
ing embolus (Figs. 83, 84); eastern
United States (Map 2A) yeargani
Base of median apophysis small (Figs.
37, 191) 8
Length of narrow part of median apoph-
ysis as wide as base in ectal view (Fig.
193); Gulf Coast, Kansas (Map 2E) -
archeri
Length of narrow part of median apoph-
ysis longer than width of base (Fig.
39); Virginia to North Carolina (Map
2A) tiinitcjita
Kaira altiventer{0. P. -Cambridge), new
combination
Epeiroides fasciolata O. P.-Cambridge, 1889: 15, pi.
8, fig. 5, 6. Male from Bugaba, Panama, in
BMNH, examined; now lost. KeyserUng, 1893: 309,
pi. 16, fig. 228, 6. Male from Guatemala.
Kaira altiventer O. P.-Cambridge, 1889: 56, pi. 3, fig.
13, ?. Female from Veragua [Veraguas Prov.], Pan-
ama, in BMNH, examined. Levi, 1993b, 213, figs.
3-22, 9, (?, NEW SYNONYMY.
Aranea fasciolata: — F P.-Cambridge, 1904: 519, pi.
51, fig. 5, S . Claims that Keyserling's specimen is
lost and probably was misidentified according to F.
P-Cambridge, 1904.
Note. Epeiroides fasciolata is a Kaira.
My unpublished illustration of the holo-
type of E. fasciolata shows the distinct
large median apophysis tooth at the base
of the flagella, the characteristic curved,
long, soft conductor, and the drop-shaped,
sclerotized terminal apophysis of Kaira al-
tiventer (Levi, 1993b, figs. 20, 21).
I examined the type in 1967, when vis-
iting the BMNH, and made a drawing of
the palpus of the male, thinking errone-
ously that the species is a Mastophora, But
the palpus is not that of Mastophora. Nei-
ther O. P.-Cambridge, F. R-Cambridge, or
Keyserling showed carapace tubercles. I
overlooked the species when revising
Kaira. Since 1967, the holotype has been
misplaced and cannot be found.
Mastophora satsuma new species
Figures 8-14; Map 2C
Holotype. Female holotype from Riverview, 11 mi.
[17.6 km] SE of Tampa, on Highway 301, Hills-
borough Co., Florida, on satsuma. Citrus nobilis
(tangerine tree), 23 Aug. 1966 (E. R. Simmons), in
FSCA. The specific name is a noun in apposition
after the tree on which the holotype was collected.
Description. Female holotype. Carapace
orange-brown. Chelicerae, labium, endites
light brown. Sternum grayish orange. Cox-
ae and distal leg articles brown. Abdoinen
dorsum whitish with dark gray frame hav-
ing a lobe extending posteriorly to midline
(Fig. 10); venter gray with white square.
Carapace, with few tubercles (Figs. 8, 9)
and short white setae. Abdomen without
humps (Fig. 10). Total length 9.6 mm.
326 Bulletin Museum of Coinparative Zoology, Vol. 157, No. 5
Carapace 4.4 mm long, 4.0 wide in tho-
racic region, 2.4 wide at lateral eyes. First
femur 4.1 mm, patella and tibia 5.2, meta-
tarsus 3.5, tarsus 1.0. Second patella and
tibia 4.0 mm, third 2.3, fourth 3.6. Length
of first patella and tibia 1.1 times width of
carapace.
Males are not known.
Variation. The epigynum is asymmetri-
cal: the left slit is more curved than the
right one and the left seminal receptacles
are larger than the right ones (Fig. 14).
Both seminal receptacles are oval.
Diagnosis. Mastophora satsuma is dis-
tinguished from M. felda (Figs. 22-28) by
being smaller, by differences in dorsal pat-
tern (Fig. 10), by having larger seminal re-
ceptacles (Fig. 14), and also by the larger
depression in the midline of the epigynum
(Fig. 14).
Distribution. Central Florida (Map 2C).
Specimens Examined. No other specimens have
been found.
Mastophora apalachicola new species
Figures 15-21; Map 2C
Holotype. Female holotype from ravine, Bristol, Cal-
houn Co., Florida, 29 Dec. 1939 (A. F Archer), in
AMNH. The specific name is a noun in apposition
after the name of the river at the locality.
Description. Female holotype. Carapace
contrastingly marked, sides dark brown,
dorsum light brown anteriorly, pair of
forks lightest brown (Figs. 15, 16). Chelic-
erae yellow-white with a dark patch on
sides. Labium, endites dark brown. Ster-
num anterior light, posterior dark brown.
Coxae dusky brown, fourth darkest. Distal
leg articles yellow-white, femora and pa-
tellae with brown bands. Abdomen white
(Fig. 17), dorsum with a pair of black
spots, venter with white square containing
three pairs of black dots. Carapace with
few tubercles, with veiy large forked horns
and with short white setae on sides (Figs.
15, 16). Median eyes on bulge, lateral eyes
on bulges. Abdomen without humps and
with large distinct dorsal pair of discs (Fig.
17). Total length 8.8 mm. Carapace 3.5
mm long, 3.4 wide in thoracic region, 2.2
wide at lateral eyes. First femur 3.3 mm, 1
patella and tibia 4.4, metatarsus 3.2, tarsus
1.0. Second patella and tibia 3.3 mm, third
1.8, fourth 3.0. Length of first patella and
tibia 1.3 times width of carapace.
Males are not known.
Variation. Total length of females 8.8 to
9.0 mm. The specimen from Levy County
has the atria larger and more spherical
than those of the holotype (Fig. 21); the
one from Hamilton Co. has the atria larger
and the seminal receptacles much larger.
The illustrations were made from the ho-
lotype.
Diagnosis. Mastophora apalachicola is
distinguished from others by the contrast-
ing carapace coloration (Figs. 15, 16), the
abdomen lacking humps, lacking dorsal
color pattern, and having large dorsal discs
(Fig. 17). The horns (Fig. 15) are larger
than those of M. timuqua (Fig. 29) and M.
satsuma (Fig. 8). The epigynum, unlike
that of similar species, has a lip on each
side; the slits are in a slight depression
(Fig. 20). The epigynal slits and atria are
almost parallel (Figs. 20, 21).
Distribution. South Carolina to north-
ern Florida (Map 2C).
Paratiipes. SOUTH CAROLINA Anderson Co.:
Simpson Agric. Exp. Station, 16 Aug. 1974, 1 imm.
(R. Paigler, CUAC). FLORIDA Hamilton Co.: nr.
White Springs, Big Shoals State Forest, 25 Nov. 1991,
1? (M. K. Stowe 2116, FSCA). Levy Co.: Manatee
Springs State Park, 10 Nov. 1992, 1? (M. K. Stowe
2114,"mCZ).
Mastophora felda new species
Figures 22-28; IVIap 2A
Holotype. Female holotype from near Felda, Hendry
Co., Florida, in orange grove, 8 March 1993 (D.
Smith), in FSCA. The specific name is a noun in
apposition after the type locality.
Description. Female holotype. Carapace ^
dark orange-brown. Chelicerae dusky \
brown. Labium, endites dusky brown.
Sternum brownish orange. Coxae orange-
brown, lighter than sternum and legs. Dis-
tal leg articles dark orange-brown. Abdo-
men anterior, sides, and venter gray (Fig.
24), center and posterior whitish; venter
Mastophora • Levi
327
Figures 1-7. Mastophora. 1-4, female. 1, M. bisaccata, left tip of clielicera and fang from posterior. 2, epigynum, diagram-
matical. 3, dish with paraffin to examine epigyna. 4, M. diablo, epigynum cleared, in posterior view, showing ducts. 5-7, male.
5, M. gasteracanthoides. 6, M. bisaccata. 7, M. gasteracanthoides left palpus without cymbium, mesal view.
Figures 8-14. M. satsuma new species, female. 8, 9, carapace and chelicerae. 8, frontal. 9, lateral. 10, 11, carapace and
abdomen. 10, dorsal. 11, lateral. 12-14, epigynum. 12, ventral. 13, posterior. 14, posterior, cleared.
Figures 15-21. M. apalachicola new species, female. 15, 16, carapace and chelicerae. 15, frontal. 16, lateral. 17, 18, carapace
and abdomen. 17, dorsal. 18, lateral. 19-21, epigynum. 19, ventral. 20, posterior. 21, posterior, cleared.
Figures 22-28. M. felda new species, female. 22, 23, carapace and chelicerae. 22, frontal. 23, lateral. 24, 25, carapace and
abdomen. 24, dorsal. 25, lateral. 26-28, epigynum. 26, ventral. 27, posterior. 28, posterior, cleared.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
328 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
with median white square. Carapace with
few tubercles and with short, white setae
(Figs. 22, 23). Median eyes on bulge, lat-
eral eyes on bulges. Abdomen without
humps (Fig. 24). Total length 12.3 mm.
Carapace 5.3 mm long, 5.4 wide in tho-
racic region, 3.0 wide at lateral eyes. First
femur 5.2 mm, patella and tibia 6.7, meta-
tarsus 4.6, tarsus 1.2. Second patella and
tibia 5.1 mm, third 2.7, fourth 4.8. Length
of first patella and tibia 1.2 times width of
carapace.
Males are not known.
Variation. The holotype has only seven
eyes; it lacks the left posterior median eye.
Diagnosis. Mastophora is distinguished
from M. bisaccata by the different color-
ation of the abdomen (Fig. 24), by the epi-
gynum having only a thin rim in ventral
view (Fig. 26), and by the small dorsal
knobs in the depression on the posterior
of the epigynum (Fig. 27). The atria bend
toward each other (Fig. 28), but are far-
ther apart than those of M. bisaccata (Fig.
57).
Distribution. South-central Florida
(Map 2A).
Specimens Examined. No other specimens have
been found.
Mastophora timuqua new species
Figures 29-39, 445; Map 2A
Holotype. Female holotype from Devil's Millhopper
State Park, Gainesville, Alachua Co., Florida, 19
Nov. 1983 (M. K. Stowe 107A), in MCZ. The spe-
cific name is a noun in apposition after an extinct,
northern Florida Indian tribe.
Description. Female holotype. Carapace
light brown, with sides and eye areas dark-
er brown and white mark in center (Figs.
29, 30). Chelicerae, labium, endites
brown. Sternum light brown. Coxae lighter
than sternum, distal leg articles with dark
brown rings. Abdomen gray with anterior
darker (Fig. 31), venter with indistinct
white square. Thorax with short white se-
tae, a distinct narrow line around margin;
lacking large tubercles on sides of thoracic
region (Figs. 29, 30). Median eyes on a
bulge, lateral eyes on bulges. Abdomen
without humps. Total length 8.5 mm. Car- <
apace 3.5 mm long, 3.4 wide in thoracic
region, 1.8 wide at lateral eyes. First femur
2.9 mm, patella and tibia 4.1, metatarsus
2.7, tarsus 0.8. Second patella and tibia 3.3 i
mm, third 1.8, fourth 2.7. Length of first ^
patella and tibia 1.2 times width of cara-
pace.
Male allotype. Carapace orange with
white median patch. Sternum orange.
Coxae, legs lighter orange. Abdomen
dusky orange. Abdomen with two adjacent
humps. Total length 1.6 mm. Carapace
0.88 mm long, 0.79 wide in thoracic re-
gion, 0.53 wide at lateral eyes. First femur
0.78 mm, patella and tibia 0.78, metatarsus
0.45, tarsus 0.28. Second patella and tibia
0.69 mm, third 0.40, fourth 0.55. Length
of first patella and tibia same as width of
carapace.
Note. Males were raised from egg sac of
M. timuqua.
Variation. The illustrations were made
from the female holotype and male allo-
type.
Diagnosis. Mastophora timuqua is dis-
tinguished from both M. bisaccata and M.
pisgah by lacking dorsal abdominal pattern
(Fig. 31), by having ringed legs, by having
the posterior of the epigynum with atria
separated, and by having a ridge in the
midline (Fig. 34).
The male has a wide space enclosed by
the median apophysis (Fig. 39) and differs
from the male of M. coryiigera by the prox-
imal position of the radix (Fig. 37).
The egg sac is shown in Figure 445.
Distribution. North Carolina to north-
ern Florida (Map 2A).
Paratijpes. NORTH CAROLINA Moore Co.:
reared from egg sac in spring 1941, 3c? (J. Perry, M.
K. Stowe 2113" MCZ). FLORIDA AZrtc/zt/a Co.: Dev-
ils Millhopper State Park, reared spring 1992, S al-
lotype, 11 paratypes (M. K. Stowe 2101, MCZ,
AMNH); spring 1992, raised 3cJ (M. K. Stowe 2106,
FSCA). Levy Co.: Manatee Springs State Park, 5 Nov.
1987, 19 (M. K. Stowe 2111, AMNH). Hillsborough
Co.: Pinecrest Alderman Ford County Park, 9 May
1988, egg sac (M. K. Stowe 21050, FSCA).
Mastophora • Levi
329
Figures 29-39. Mastophora timuqua new species. 29-35, female. 29, 30, carapace and chelicerae. 29, frontal. 30, lateral. 31,
32, carapace and abdomen. 31, dorsal. 32, lateral. 33-35, epigynum. 33, ventral. 34, posterior. 35, posterior, cleared. 36-39,
male left palpus, stained. 36, apical. 37, mesal. 38, ventral. 39, octal.
Figures 40-50. M. alachua new species. 40-46, female. 40, 41, carapace and chelicerae. 40, frontal. 41, lateral. 42, 43,
carapace and abdomen. 42, dorsal. 43, lateral. 44-46, epigynum. 44, ventral. 45, posterior. 46, posterior, cleared. 47-51, male
left palpus, stained. 47, apical. 48, mesal. 49, ventral. 50, ectal.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
330 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Mastophora alachua new species
Figures 40-50; IVIap 2G
Holotype. Female holotype from Devil's Millhopper
State Park, Gainesville, Alachua Co., Florida, 19
Nov. 1983 (Mark K. Stov^/e 107B), in MCZ. The
species is named after the type locality.
Description. Female holotype. Carapace
orange. Chelicerae, labium, endites or-
ange. Sternum orange. Coxae and distal
leg articles orange. Abdomen orange -with
a dorsal, anterior dusky area containing
some bare patches, and a transverse gray
line posteriorly (Fig. 42); venter with white
square. Carapace granular rather than tu-
bercular, sides of carapace with short
white setae; no tubercles on lateral thorac-
ic region (Fig. 41). Abdomen subtriangu-
lar, without humps (Fig. 43). Total length
8.0 mm. Carapace 3.4 mm long, 3.4 wide
in thoracic region, 2.2 wide at lateral eyes.
First femur 3.5 mm, patella and tibia 4.5,
metatarsus 3.2, tarsus 1.0. Second patella
and tibia 3.5 mm, third 1.8, fourth 2.9.
Length of first patella and tibia 1.3 times
width of carapace.
Male allotype. Carapace dusky orange
with median white patch. Sternum, legs,
abdomen dusky orange. Abdomen with
pair of adjacent humps. Total length 1.6
mm. Carapace 0.78 mm long, 0.67 wide in
thoracic region, 0.52 wide at lateral eyes.
First femur 0.67 mm, patella and tibia
0.78, metatarsus 0.42, tarsus 0.11. Second
patella and tibia 0.65 mm, third 0.38,
fourth 0.54. Length of first patella and tib-
ia 1.1 times width of carapace.
Note. The male was raised from an egg
sac.
Variation. Total length of females 8.0-
9.2 mm, males 1.6—1.7. The illustrations
were made from female holotype, correct-
ed with the paratypes. The male illustrated
was the only one available, the allotype,
whose palpus was expanded, and thus the
median apophysis (Figs. 47-50) may not
be at the same angle as in the contracted
palpus.
Diagno.sis. Mastophora alachua is distin-
guished by the abdomen, slightly triangu-
lar in shape and having markings like those
of M. hisaccata (Fig. 42), and by the epi-
gynum, in ventral view showing two dark
areas, the atria, on the posterior margin,
and a narrow double margin (Fig. 45), and
on the posterior, parallel slits, slightly clos-
er ventrally, and between, next to each
other, two shallow U-shaped shadows (Fig.
45).
Distribution. Northern Florida (Map
2G).
Paratypes. FLORIDA Alachua Co.: Devils Mill-
hopper State Park, 21 Nov. 1983, 1$ (M. K. Stowe
106, MCZ); no date, prob. 1984, allotype 6 (M. K.
Stowe 2102, MCZ); Gainesville, 1 Nov.'l990, 1$ (M.
K. Stowe 2115, FSCA).
Mastophora bisaccata (Emerton)
Figures 51-62, 446; Map 2B
Cyi-tarachne bisaccata Emerton, 1884: 325, pi. 34,
fig. 11, ?, pi. 38, fig. 12, egg sac. Female holotypes
from beech tree. New Haven, Connecticut, in
MCZ, examined.
C. multilineata Atkinson, 1888: 546. Two syntypes
presumably from near Chapel Hill, North Carolina,
lost. First synonymized by Banks (1910).
Ordgarius bisaccata: — Keyserling, 1892: 42, pi. 2, fig.
35, ? . McCook, 1894: 198, pi. 12, figs. 2, 3, 9 .
Gh/ptocraniiim bisaccatum: — Bonnet, 1957: 1996.
Ordgarius obesiis Banks, 1898: 250, pi. 15, fig. 9, ? .
Two female syntypes from La Chuparosa [Chupar-
rosa, San Luis Potosi], Mexico, in CAS, destroyed.
NEW SYNONYMY.
Mastophora bisaccata: — Mello-Leitao, 1931: 71.
Roewer, 1942: 900. Kaston, 1948: 232, figs. 737-
740. Gertsch, 1955: 242, pis. 3-5, pi. 6, figs. 1, 4:
text figs. 19-23, .35, 43, 44. $, S. Platnick, 1997:
513. Platnick, 2001.
Note. Atkinson did not tell how Cyrtar-
achne multilineata differs from hisaccata,
although he mentioned the latter species.
The large size, total length 11 and 13 mm,
abdomen 13 and 15 mm wide, suggest he
had M. bisaccata. The name was first syn-
onymized by Banks (1910).
Ordgarius ohesus differs from hisaccata,
according to Banks, by being larger in size
and having the cephalothorax truncate.
The illustration shows the dorsal abdomi-
nal pattern of M. bisaccata. The size is
within the range of M. hisaccata and the
carapace, unlike most Mastophora, is trun-
cate. There is no doubt that this was M.
Mastophora • Levi
331
Figures 51-62. Mastophora bisaccata (Emerton). 51-58, female. 51, 52, carapace and chelicerae. 51, frontal. 52, lateral. 53,
54, carapace and abdomen. 53, dorsal with male. 54, lateral. 55-58, epigynum. 55, ventral. 56, posterior. 57, 58. posterior,
cleared. 57, (Virginia). 58, (Florida). 59-62, male left palpus, stained. 59, apical. 60, mesal. 61, ventral. 62, ectal.
Figures 63-74. M. stowei new species. 63-70, female. 63, 64, carapace and cfielicerae. 63, frontal. 64, lateral. 65, 66, carapace
and abdomen. 65, dorsal, with male. 66, lateral. 67-70, epigynum. 67, ventral. 68, posterior. 69, 70, posterior, cleared. 69,
(Florida). 70, (North Carolina). 71-74, male left palpus, stained. 71, apical. 72, mesal. 73, ventral. 74, ectal.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
332 Bulletin Museum of Comparative Zoologij, Vol. 157, No. 5
bisaccata. The locality of this specimen is
doubtful, because it came from the Marx
collection (see Banks, 1898).
Description. Female from Rhode Is-
land. Carapace orange-brown. Sternum
dark orange-yellow. Legs orange yellow,
first femur darker ventrally. Abdomen
white, anterior dorsally gray with charac-
teristic pattern (Fig. 53), venter with white
square. Carapace with very small tuber-
cles, and small eye projections (Figs. 51—
53). Abdomen without huinps (Fig. 53).
Total length 14.2 mm. Carapace 5.2 mm
long, 5.4 wide in thoracic region, 2.8 wide
at lateral eyes. First femur 4.4 mm, patella
and tibia 6.3, metatarsus 4.3, tarsus 1.2.
Second patella and tibia 4.7 mm, third 2.7,
fourth 4.6. Length of first patella and tibia
1.2 times width of carapace.
Male from Arkansas. Carapace orange-
brown v^th white line in middle, branch-
ing posteriorly into tubercles (Fig. 6). Ster-
num white. Lateral eyes smaller than ine-
dian. Legs colorless yellowish. Abdomen
white. Abdomen with slight, indistinct tu-
bercles (Fig. 6). Total length 1.8 mm. Car-
apace 0.88 mm long, 0.81 wide in thoracic
region, 0.55 wide at lateral eyes. First fe-
mur 0.92 mm, patella and tibia 0.93, meta-
tarsus 0.48, tarsus 0.27. Second patella and
tibia 0.79 mm, third 0.47, fourth 0.59.
Length of first patella and tibia 1.1 times
width of carapace.
Note. Examined males include males
raised from egg sacs by K. Yeargan and
males collected in the same locality as fe-
males.
Variation. The holotype has the epigynal
slits farther apart than in the specimen il-
lustrated. Total length of females 9.0—15.3
mm. Males may have humps on the ab-
domen. The illustrations v^ere made from
the female holotype and males from Ar-
kansas and Ohio. An egg sac collected with
female at Bushnell, Florida, lacked flaps
and was smooth.
Diagnosis. The female of M. bisaccata is
distinguished from the similar M. alachua,
M. stowei, and M. yeargani by being larg-
er, and having the atria approaching each
other in posterior view of the epigynum
(Figs. 56-58).
The male has a narrow space encircled
by the median apophysis in ectal view (Fig.
62), and the median apophysis has a longer
base (Fig. 60) than in M. hutchinsoni.
The egg sac has a heavier stalk than in
other Mastophora species (Fig. 446).
Natural History. Mastophora bisaccata
has been collected from bittersweet in
Connecticut, field of cemeteiy in Illinois,
and on dogwood in Florida. Females rest
under leaves, and sometimes neighboring
leaves are stitched together; the females
may look like leaf galls in Florida (M.
Stowe, personal correspondence); in
North Carolina they may resemble tree
snails (Atkinson, 1888).
Distribution. Eastern United States
(Map 2B).
Specimens Examined. CONNECTICUT Litchfield
Co.: Kent, on bittersweet, Sept. 1937, 19 (AMNH).
NEW YORK Nassau Co.: Long Island: Sea Cliff, IcJ
(MCZ). NEW JERSEY Elizabeth Co.: Rosalie Park,
25 Sep. 1910, 1? (AMNH). Middlesex Co.: New
Brunswick, July 1930, 19 (AMNH). PENNSYLVA-
NIA Westmoreland Co.: 4.8 km S Rector, 13 Sep.
1966, 19 (B. Vogel, DMNS). OHIO Jackson Co.:
Oak Hill, 19 (R. A. Reller, OSU). Logan Co.: Can-
twell Cliffs, 8 Sep. 1935, 19 (OSU). Butler Co.:
Bachelor Woods, Oxford, 11 July 1998, IS (D. M.
Golden, OSU). DISTRICT OF COLUMBIA Wash-
ington, Sept., IS (Fox, cue, AMNH); summer
1935, 19 (H. E. Ewing, USNM). VIRGINIA Falls
Church, 19, 1 (J (MCZ). Powhatan Co.: Powhatan,
Sep. 1984, 19 (A. Moreton, MCZ); 1985, 19 (A.
Moreton, MKS). KENTUCKY /e.s.sa?mne Co.: imm.,
IS raised spring 1995 (K. V. Yeargan, KVY). SOUTH
CAROLINA Oconee Co.: Clemson College, 19
(MCZ). Lexington Co.: Batesburg, 19 (MCZ).
GEORGIA Fulton Co.: Adanta, 2 Aug. 1937, 19 (F.
Figures 75-85. Mastophora yeargani new species. 75-81 , female. 75, 76, carapace and chelicerae. 75, frontal. 76, lateral. 77,
78, carapace and abdomen. 77, dorsal. 78, lateral. 79-81, epigynum. 79, ventral. 80, posterior. 81, posterior, cleared. 82-85,
male left palpus, stained. 82, apical. 83, mesal. 84, ventral. 85, ectal.
Mastophora • Levi 333
Figures 86-99. M. phrynosoma Gertsch. 86-92, female. 86, 87, carapace and chelicerae. 86, frontal. 87, lateral. 88, 89,
carapace and abdomen. 88, dorsal, with male. 89, lateral. 90-92, epigynum. 90, ventral. 91, posterior. 92, posterior, cleared.
93-99, male left palpus, stained. 93-96, (Kentucky). 97-99, (Florida). 93, apical. 94, 97, mesal. 95, 98, ventral. 96, 99, octal.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
334 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
W. Fattig, AMNH). FLORIDA Alachua Co.: Devils
Millhopper State Park, July, Aug. 1978-1981, 59 (M.
K. Stowe, MKS); June, July 1981, 79 (M. K. Stowe,
MKS); 19 Nov. 1983, 1 9 ; Gainesville, 26 July 1980,
penult. (C. Hieber, MCZ); High Springs, Oct. 1935,
19 (H. H. Simpson, USNM). Indian River Co.: Se-
bastian, Apr. 1944, 19 (G. Nelson, MCZ). Lake Co.:
Umatilla, Sep. 1954, 19 (M. W. Tyler, AMNH). Sum-
ter Co.: Bushnell, 24 Oct. 1979, 19 (W. Edv^ards,
FSCA). ILLINOIS Franklin Co.: Zeigler, 10 May
1928, 19 (J. K. Carlovic, MCZ). Jackson Co.: Little
Grand Canyon, S. Murphysboro, 5 Sep. 1971, Id (N.
Magnuson, JAB). Williamson Co.: Canterville, field of
cemeteiy, 14 Oct. 1978, 19 (R. Reith, JAB). MIS-
SOURI St. Louis Co.: 17 Feb. 1940, IS paratypes of
M. archen (W. M. Gordon, AMNH). ARKANSAS
Carroll Co.: Berryville, July 1942, IS (C. Wilton).
MISSISSIPPI Harrison Co.: Gulfport, 19 (AMNH).
Mastophora stowei new species
Plate 1; Figures 63-74, 447; Map 2C
Holotype. Female holotype from American Entomo-
logical Institute, Gainesville, Alachua Co., Florida,
29°36.0'N, 82°22.0'W, 7 Dec. 1987 (M. Stowe
07001), in MCZ. The species is named after tlie
collector, Mark Stowe, who has contributed much
to our knowledge of Mastophora.
Description. Female holotype. Carapace
orange-brown, dusky in eye region (Fig.
65). Chelicerae light orange, labium, en-
dites dusky orange. Sternum orange. Legs
light orange, dusky dorsally. Abdomen dor-
sum whitish with gray pattern anteriorly,
having distinct white spots and streaks
(Fig. 65); venter with white square. Cara-
pace with indistinct tubercles; horns al-
iTiost rectangular in anterior view (Figs. 63,
64). Abdomen without humps and slightly
pointed posteriorly (Fig. 65). Total length
8.0 mm. Carapace 3.5 mm long, 3.4 wide
in thoracic region, 2.1 wide at lateral eyes.
First femur 3.4 mm, patella and tibia 4.5,
metatarsus 2.8, tarsus 1.0. Second patella
and tibia 3.4 mm, third 2.0, fourth 3.2.
Length of first patella and tibia 1.3 times
width of carapace.
Male allotype. Carapace gray-orange
with a white median patch. Sternum or-
ange. Legs orange, dusky dorsally. Abdo-
men dusky orange, with humps. Total
length 1.7 mm. Carapace 0.80 mm long,
0.68 wide in thoracic region, 0.48 wide at
lateral eyes. First femur 0.67 mm, patella
and tibia 0.78, metatarsus 0.41, tarsus 0.29.
Second patella and tibia 0.65 mm, third
0.39, fourth 0.53. Length of first patella
and tibia 1.1 times width of carapace.
Note. Males were raised from the egg
sac of the holotype.
Variation. Total length of females 6.3—
10.5 mm. Males may lack humps. The il-
lustrations w^ere made from the female ho-
lotype, except Figure 70 from a North
Carolina specimen, and the male from the
allotype.
Diagnosis. Mastophora stowei is distin-
guished from M. hisaccata by being small-
er in size and having the atria of the epi-
gynum ventrally departing from each other
(Figs. 68—70). The egg sac is shown in Fig-
ure 447.
Natural History. Females have been
collected from trees along farm fences in
Kentucky, and on a carpet of silk on a ma-
ple leaf in Virginia.
Distribution. Widespread in the eastern
United States (Map 2C).
Paratypes. CONNECTICUT Hartford Co.: Rain-
bow nr. Windsor, 9 Aug. 1939, 19 (A. de Caprio,
USNM). OHIO Logan Co.: Old Mans Cave, 12 Sep.
1924, IS (OSU). VIRGINIA Arlington Co.: Arling-
ton, 7 Sept. 1953, 19 (K. V. Krombein, AMNH). IL-
LINOIS Pope Co.: Dixon Spring State Park, 7 Sept.
1974, 19 (J. A. Beatty, JAB). KENTUCKY Faijette
Co.: Lexington, Oct. 1997, 1 9 , egg sac (K. V. Yeargan,
KVY); 6 Aug. 1998, 14c? (K. Yeargan, KVY); Cold
Stream farm fence, 16 Oct. 1998, 39 (K. Yeargan,
KVY). Jessamine Co.: 3 Aug. 1995, IS (K. Yeargan,
KVY). NORTH CAROLINA Haijwood Co.: Canton,
19 (Holden, MCZ). GEORGIA 19 (MNHN 210).
FLORIDA Alachua Co.: Gainesville, male allotype
and 1 penultimate paratype from egg sac of holotype,
Figures 100-106. Mastophora catarina new species, female. 100, 101, carapace and ctielicerae. 100, frontal. 101, lateral. 102,
103, carapace and abdomen. 102, dorsal. 103, lateral. 104-106, epigynum. 104, ventral. 105, posterior. 106, posterior, cleared.
Figures 1 07-1 1 3. /W. tiaywardi Biraben, female. 1 07, 1 08, carapace and chelicerae. 1 07, frontal. 1 08, lateral. 1 09, 1 1 0, carapace
and abdomen. 109, dorsal. 110, lateral. 111-113, epigynum. Ill, ventral. 112, posterior. 113, posterior, cleared.
Mastophora • Levi
335
Figures 114-120. M. corumbatai new species, female. 114, 115, carapace and chelicerae. 114, frontal. 115, lateral. 116, 117,
carapace and abdomen. 116, dorsal. 117, lateral. 118-120, epigynum. 118, ventral. 119, posterior. 120, posterior, cleared.
Figures 121-127. M. laranew species, female. 121, 122, carapace and chelicerae. 121, frontal. 122, lateral. 123, 124, carapace
and abdomen. 123, dorsal. 124, lateral. 125-127, epigynum. 125, ventral. 126, posterior. 127, posterior, cleared.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
336 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
hatched 27 Apr. to 15 May 1988, preserved 18 July
1988 (M. Stowe 20181, 20183, MCZ, AMNH,
FSCA). TEXAS Harrison Co.: Marshall, 5 July 1991,
19 (S. G. Wellso, JK).
Mastophora yeargani new species
Plate 1; Figures 75-85; 448; Map 2A
Holotype. Female holotype from Coldstream Farm
fence, Lexington, Kentucky, 26 Oct. 1998, male al-
lotype and 11 male and 4 female paratypes
emerged 24 May 1999, preserved in fall (K. V.
Yeargan), in MCZ. The species has been named
after the collector, who has contributed much to
our knowledge of Mastophora ecology.
Description. Female holotype. Carapace
light brown. Chelicerae, labium, endites
yellow. Sternum yellow underlain by white
pigment granules. Coxae and distal leg ar-
ticles yellow. Abdomen anterior of dorsum
gray with w^hite marks anteriorly (Fig. 77),
posterior w^hite, venter whitish with white
square. Carapace shiny. Abdomen with
pair of very slight dorsal swellings. Total
length 10.0 mm. Carapace 4.2 mm long,
4.2 wide in thoracic region, 2.3 wide at lat-
eral eyes. First femur 3.9 mm, patella and
tibia 5.4, metatarsus 3.8, tarsus 1.2. Sec-
ond patella and tibia 4.0 mm, third 2.4,
fourth 3.7. Length of first patella and tibia
1.3 times width of carapace.
Male allotype. Carapace brown with
white triangle in center. Chelicerae, labi-
um, endites orange. Sternuna orange. Cox-
ae and distal leg articles orange. Abdonien
whitish with pair of humps. Total length
1.7 mm. Carapace 0.78 mm long, 0.78
wide in thoracic region, 0.52 wide at lat-
eral eyes. First femur 0.87 mm, patella and
tibia 0.88, metatarsus 0.52, tarsus 0.34.
Second patella and tibia 0.75 mm, third
0.39, fourth 0.60. Length of first patella
and tibia 1.1 times width of carapace.
Note. Males have been raised from egg
sac of female M. ijeargani (by K. Yeargan).
Variation. Total length of females 10.0—
11.5 mm. The illustrations were made
from the holotype and allotype.
Diagnosis. Mastophora yeargani is dis-
tinguished from M. hisaccata and M. stoiv-
ei, which have similar abdominal markings,
by the short slits of the epigynum, which
are almost parallel and their length distant
from the ventral margin (Figs. 80, 81). The
female also lacks duskiness on the dorsum
of the femora and gray pigment on the
clypeus, both of which are present in M.
stowei.
The male differs from M. hisaccata (Fig.
62) by having a longer median apophysis
(Fig. 85) and from M. cornigera (Figs. 177,
179) by having only a short terminal
apophysis in the palpus (Figs. 82, 83), and
by having the median apophysis of a dif-
ferent shape, and the embolus wider (Figs.
83, 84).
Distribution. From New York to Ken-
tucky (Fig. 448).
Paratypes. NEW YORK nr. New York City, on
Amalanchier s^., 19 (AMNH). KENTUCKY Mercer
Co.: Feb. 1995 egg sacs, 29 Sept. 1995, 7 imm., 46
(K. V. Yeargan, KVY). Garrard Co.: Feb. 1995, egg
sacs. Sept. 1995. 6 imm.. 16c? (K. V. Yeargan, KVY).
Mastophora phrynosoma Gertsch
Figures 86-99, 449, 450; Map 2D
Mastophora phrynosoma Gertsch, 1955: 245; pi. 6,
fig. 5, text figs. 24-27, 31, 9 . Female holotype from
Burlington, North Carolina, in AMNH, examined.
Brignoli. 1983: 273. Platnick, 2001.
Description. Female holotype. Carapace
orange-brown. Sternum orange-brown.
Legs orange-brown, indistinctly ringed.
Abdomen anteriorly gray, posteriorly white
(Fig. 88), venter with white square. Cara-
pace with tubercles very small (Figs. 86,
87). Median eyes on bulge, lateral eyes on
bulges. Abdomen subtriangular with a
Figures 128-134 Mastophora carpogastra Mello-Leitao, female. 128, 129, carapace and chelicerae. 128, frontal. 129, lateral.
130, 131, carapace and abdomen. 130, dorsal. 131, lateral. 132-134, epigynum. 132, ventral. 133, posterior. 134, posterior,
cleared.
Figures 135-141. M. seminole new species, female. 135, 136, carapace and chelicerae. 135, frontal. 136, lateral. 137, 138,
carapace and abdomen. 137, dorsal. 138, lateral. 139-141, epigynum. 139, ventral. 140, posterior. 141, posterior, cleared.
Mastophora • Levi 337
135 ^.- ^ ^ " %x_ 136
Figures 142-152. M. vaquera Gertsch, female. 142, 143, carapace and chelicerae. 142, frontal. 143, lateral. 144, left first femur
and patella, mesal. 145, 146, carapace and abdomen. 145, dorsal. 146, lateral. 147-152, epigynum. 147, 150 ventral. 148, 151,
posterior. 149, 152, posterior, cleared. 147-149, (holotype from Matanzas). 150-152, (Santiago).
Scale lines. 1.0 mm; genitalia, 0.1 mm.
338 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
swelling on each anterior lateral side and
indistinct sclerotized discs (Fig. 88). Total
length 12.3 mm. Carapace 4.6 mm long,
4.4 wide in thoracic region, 2.7 wide at lat-
eral eyes. First femur 4.6 mm, patella and
tibia 6.5, metatarsus 4.7, tarsus 1.3. Sec-
ond patella and tibia 4.5 mm, third 2.4,
fourth 4.1. Length of first patella and tibia
1.5 times width of carapace.
Male from Kentucky. Carapace orange-
brown, median triangle enclosing horns
and two tubercles lighter. Sternum orange.
Legs orange-brown. Abdomen both sides
orange-white. Carapace slightly rugose,
with two median tubercles and four pos-
terior horns. Abdomen with small dorsal
humps. Total length 1.8 mm. Carapace
0.81 mm long, 0.80 wide in thoracic re-
gion, 0.55 wide at lateral eyes. First femur
0.91 mm, patella and tibia 0.93, metatarsus
0.46, tarsus 0.31. Second patella and tibia
0.78 mm, third 0.45, fourth 0.60. Length
of first patella and tibia 1.2 times width of
carapace.
Note. Males have been raised by M. K.
Stowe and K. V. Yeargan from egg sacs that
were determined to be from M. phryno-
soma.
Variation. Total length of females 8.3—
12.3 mm, males 1.5—1.7. The lateral swell-
ings of a specimen from Falls Church are
less distinct. The carapace of a female
from Missouri is blackish brown, sternum
and coxae black, legs ringed, and abdom-
inal venter black. The venter of several in-
dividuals is black. The illustrations were
made from the female holotype and from
the reared male.
Diagnosis. Mastophora phrijnosonia is
distinguished by the subtriangular shape of
the abdomen, the small, dorsal sclerotized
discs (Fig. 88), and by the posterior of the
epigynum having a lip surrounding the
slits on three sides (Figs. 90—92).
The palpus of the male has the space
enclosed by the median apophysis \vider
than long and the prong straight (Figs. 96,
99), and has the base of the median
apophysis almost as wide as long (Figs. 94,
98). It differs from that of M. archeri by
having a longer terminal apophysis (Figs.
93, 94), and wider radix (Figs. 95, 98).
The egg sac is distinct, having longer
flaps than in other species (Figs. 449, 450).
Natural History. A female was collected
on elm bush, 1.5 m high, in Texas; in Mis-
souri, a female was collected hanging on a
silk strand on canebrake (Anindinaria gi-
gantea) at night at 0200 h. Males were col-
lected crawling on a table in the labora-
toiy, and in low branches of hawthorn in
Ohio. Immature spiderlings attract psy-
chodid flies {Psychoda phalaenoides)
(Yeargan and Quate, 1996). The female al-
ways rests on the upper leaf surfaces,
which often accumulate a silk pad that
may be visible under the spider. The spi-
ders look like bird droppings in Florida
(M. Stowe, personal communication). The
moths captured in Kentucky and Florida,
obseived by M. K. Stowe and K. V. Year-
gan, were reported by Yeargan (1994).
An egg sac raised by K. V. Yeargan pro-
duced only male spiderlings. A suggestion
has been made that the sex chromosome
system might be different from that of oth-
er spiders (G. Oxford, in letter). Perhaps
this skewed sex ratio was due to the mor-
tality of females.
Distribution. Eastern United States
(Map 2D).
Specimens Examined. CONNECTICUT ISlew Ha-
ven Co.: Mount Carmel, 4 Sept. 1946, 19 (K. M.
Somerman, INHS 4748). Meriden Co.: South Meri-
den, Oct. 1945, 1? (H. L. Johnson, USNM). NEW
YORK New York City, on Prunu.s, 15 (AMNH).
OHIO Cuyahoga Co.: Sagmore Picnic, Sagmore
Hills, Buckeye Trail area, 16 Sept. 1999, 19 (K. Brad-
ley OSU). MARYLAND Anne Arundel Co.: Annap-
olis, 21 Sep. 1941, 19 (M. H. Muma, AMNH). Hotv-
ard Co.: Colombia, Snowden River Parkway, 26 Sep.
1994, 19 (M. Harden, USNM). Montgomenj Co.:
Cabin John, 10 Nov. 1943, egg sacs (I. N. Hoffman,
USNM). VIRGINIA Falls Church, 29 (N. Banks,
MCZ). KENTUCKY Fayette Co.: Lexington, Raven
Run, 1989, 19 (K. V. Yeargan, KVY); 17 May 1999,
egg sacs with only 6 3 (K. V. Yeargan, KVY, MCZ).
GEORGIA Fulton Co.: Atlanta, 21 Aug. 1944, 19 (F.
W. Fattig, AMNH). FhORlD A Alachua Co.: Gaines-
ville, Apr. 1988, 13 (M. K. Stowe, MKS); Devils
Millhopper State Park, 15 July 1980, 19 (M. K.
Stowe, MKS); spring 1992, 23 (M. K. Stowe, MKS);
spring 1994, 8c? (M. K. Stowe, MKS). ALABAMA
Mastophora • Levi
339
Monroe Co.: Randons Creek, 19 Oct. 1941, 1$ (A.
F. Archer, AMNH). INDIANA 19 (A. Petrunkevitch,
NHMW). ILLINOIS Jackson Co.: Carbondale, in
woods at night, 4 Aug. 1967, 19 (J. M. Nelson,
AMNH). MISSOURI Wayne Co.: Markliam Spring,
Mark Twtiin National Forest, 14 Oct. 2000, 19 (E.
L. Quinter, AMNH). St. Louis Co.: 17 July 1940, IS
paratyjje of M. archeri (W. M. Gordon, AMNH).
TEXAS Walker Co.: Huntsville State Park, on elm
bush, 27 Sep. 1987, 19 (W. R. Martin, TAMU).
Mastophora catarina new species
Figures 100-106; Map 3C
Holotijpe. Female from Pinhal, Est. Santa Catarina,
Brazil, Dec. 1948 to Jan. 1949 (A. Mailer), in
AMNH. The specific name is a noun in apposition
after the locality.
Description. Female holotype. Carapace
dark orange-brown (Fig. 102). Sternum
dark orange. Chelicerae dark in front, or-
ange on sides. Endites, labium, sternum,
coxae orange; coxae lighter than sternum.
Legs dark browai. Abdomen gray-brown,
dorsally with three pairs of white longitu-
dinal lines (Fig. 102); venter with a white
square. Eyes distinct. Lateral eyes 0.8 di-
ameter of median eyes. Abdomen without
humps (Fig. 102). Carapace, legs, and ab-
domen with some long white hair. Total
length 7.2 mm. Carapace 3.3 mm long, 3.2
wide in thoracic region, 1.8 wide at lateral
eyes. First femur 3.5 mm, patella and tibia
4.4, metatarsus 3.0, tarsus 1.0. Second pa-
tella and tibia 3.3 mm, third 1.7, fourth
2.7. Length of first patella and tibia 1.4
times width of carapace.
Males are not known.
Diagnosis. Mastophora catarina is sep-
arated from others by the abdomen lack-
ing humps and having six white lines (Fig.
102), by the epigynum having the atria ap-
proaching each other (Figs. 105, 106), and
by the paired notches at the edge of the
epigynum (Fig. 106).
Distribution. Santa Catarina State, Bra-
zil (Map 3C).
Specimens Examined. No other specimens have
been found.
Mastophora haywardi Biraben
Figures 107-113; Map 3C
Mastophora haijwardi Biraben, 1946: 327, figs. 1—3,
9 . Female holotype, from Tucuman, Argentina, in
MLP, examined. Brignofi, 1983: 274. Platnick,
2001.
Description. Female holotype in poor
condition. Carapace brown, with a lighter
square area in thoracic region (Fig. 109),
and with long white setae. Sternum lighter.
Legs brown. Abdomen brown (the holo-
type has an injuiy, a large transverse gash
anteriorly), posterior and sides darker gray
than anterior (Fig. 109), with some long
white setae and indistinct humps visible
only from side; venter light. Total length
11.0 mm. Carapace 5.5 mm long, 4.7 wide
in thoracic region, 3.1 wide in cephalic re-
gion. First femur 4.7 mm, patella and tibia
7.6, metatarsus 5.8, tarsus 1.5. Second pa-
tella and tibia 5.2 mm, third 3.0 (from Bir-
aben, 1946), fourth 4.5. Length of first pa-
tella and tibia 1.3 times width of carapace.
Males are not known.
Diagnosis. This species is separated
from others by being the only setose spe-
cies without distinct humps (Fig. 109), and
by having the slits and the atria in the pos-
terior view of the epigynum ventrally sep-
arated (Figs. 112, 113).
Distribution. Known only from Tucu-
man, Argentina (Map 3C).
Specimens Examined. No other specimens have
been collected.
Mastophora corumbatai new species
Figures 114-120; Map 3C
Holotype. Female holotyj^e from Corumbatai, Est.
Sao Paulo [40 km N Rio Claro], Brazil, 15 July
1935 (Syilvio Bariau), in IBSP no. 1203A. The spe-
cific name is a noun in apposition after tlie locality.
Description. Female holotype. Carapace
red-browii with white rim (Fig. 116). Che-
licerae, labium, endites, sternum, coxae,
proximal ends of femora orange. Distal leg
articles brown. Abdomen light brown, with
colorless butterfly-shaped light area dor-
sally and pair of large dark, open rings
(Fig. 116); venter light brown with white
square. Carapace with various sized, drop-
let-shaped tubercles, sides with long white
setae, some curled (Figs. 114, 115). Ab-
domen without humps, with scattered
340 Bulletin Museum of Comparative Zoologij, Vol. 157, No. 5
long, white setae (Fig. 116). Total length
14.0 mm. Carapace 5.6 mm long, 5.5 wide
in thoracic region, 3.0 wide at lateral eyes.
First femur 5.5 mm, patella and tibia 7.5,
metatarsus 4.9, tarsus 1.6. Second patella
and tibia 5.3 mm, third 3.2, fourth 4.8.
Length of first patella and tibia 1.4 times
width of carapace.
Males are not known.
Diagnosis. Mastophora conimhatai is
distinguished by carapace tubercles that
look like oil droplets (Figs. 114, 115), by
lack of humps and distinctive color pattern
on abdomen (Fig. 116), and by the epi-
gynum with its posterior median plate pro-
jecting, almost scapelike in ventral view
(Figs. 118-120).
Distrihution. Santa Catarina State, Bra-
zil (Map 3C).
Specimens Examined. No other specimens have
been collected.
Mastophora lara new species
Figures 121-127; Map 3A
Holotype. Female holotype from Hato Arriba, 1,400
m, Lara, Venezuela, May 1970 (J. M. Osorio), in
FSCA. The specific name is a noun in apposition
after the locality.
Note. The type locality Hato Arriba is
probably at or near Quebrada Arriba,
1,600 m, 10°14'N, 70°32'W, close to the
border with Falcon, Zulia, and Lara, 52
km W Carora.
Description. Female holotype. Carapace
dark brown, lighter behind, in center and
in area on sides. Chelicerae, labium, en-
dites orange-brown. Sternum orange-
brown. Coxae and distal leg articles or-
ange-brown, distally darker. Abdomen gray
with anterior dorsal pattern of spots and
two large brown discs, each dissected by a
lateral light line (Fig. 123); venter light
brown with white longitudinal rectangle.
Carapace with tubercles and scattered
long white setae (Figs. 121, 122). Legs
with some long white setae. Abdomen
without humps, blunt behind, each side
slightly swollen, long white setae on each
side anteriorly (Fig. 123). Total length 11.5
mm. Carapace 5.0 mm long, 5.0 wide in
thoracic region, 2.7 wide at lateral eyes.
First femur 5.2 mm, patella and tibia 6.7,
metatarsus 4.5, tarsus 1.3. Second patella
and tibia 5.1 mm, third 3.0, fourth 4.5.
Length of first patella and tibia 1.3 times
width of carapace.
Males are not known.
Diagnosis. Mastophora lara differs by
the high almost conical carapace (Figs.
121, 122), by the pattern on the abdomen
(Fig. 123), and by the posterior of the epi-
gynum having a pair of depressions (Figs.
125-127).
Distribution. This species is known only
from the type locality in west-central Ve-
nezuela (Map 3A).
Specimens Examined. No other specimens have
been collected.
Mastophora carpogastra Mello-Leitao
Figures 128-134, 451; IVIap 3B
Mastophora carpogastra Mello-Leitao, 1925: 460.
Two female syntypes from Rio de Janeiro, Brazil,
in MNRJ, 672, examined. Mello-Leitao, 1931: 72,
fig, 3, 15, $.
Gh/ptocraniinn fagoides Vellard, 1926: 327, figs., 9,
egg sac. Female holotype from Butantan, Sao Pau-
lo, Brazil, in IBSF, lost. First synonymized with car-
pogastra by Mello-Leitao (1931).
Mastophora carpogastera: — Roewer, 1942: 900.
Glyptocranium carpogastrum: — Bonnet, 1957: 1996.
Mastophora carpogaster: — Platnick, 1993: 447. Plat-
nick, 2001.
Note. Roewer (1942) changed the name
carpogastera and listed fagoides as a syn-
onym of M. corpulenta Banks. The name
change carpogaster of Platnick (1993,
2001) is not needed because previous re-
visors (Mello-Leitao, 1931; Gertsch, 1955)
kept the original spelling.
Description. Female from Sao Paulo.
Carapace reddish brown. Chelicerae, la-
bium, endites brown. Sternum light
brown. Legs brown. Abdomen white with
symmetrical brownish black patches and
less distinct lines dorsally (Fig. 130); ven-
ter with a median white square containing
eight black spots, sides white with black
spots. Carapace glossy above, sides with
long white setae, some setae on clypeus
(Fig. 129). Legs with long white setae. Ab-
Mastophora • Levi
341
domen oval with scattered setae and barely
visible pair of dorsal tubercles (Fig. 131).
Total length 16.5 mm. Carapace 6.4 mm
long, 6.0 wide in thoracic region, 3.3 wide
behind posterior lateral eyes. First femur
5.3 mm, patella and tibia 7.6, metatarsus
5.7, tarsus 1.6. Second patella and tibia 5.5
mm, third 3.3, fourth 5.2. Length of first
patella and tibia 1.3 times width of cara-
pace.
Males are not known.
Variation. Mello-Leitao (1925) de-
scribed the species as orange to raspberry-
red when alive. Total length of females
11.2-18.0 mm. The female holotype of M.
carpogastra is 20 mm total length. The il-
lustrations were made of specimens from
Sao Paulo.
Diagnosis. The spots and black lines and
lack of humps of the abdomen (Fig. 130)
distinguish the species from all others. In
the cleared epigynum, the atria separate
ventrally (Figs. 133, 134).
The egg sac lacks flaps and has a short
stalk (Fig. 451).
Natural History. The spider mimics a
berry. According to Vellard (1926) the spe-
cies has a preference for orange trees. It
makes three to five egg sacs, each a little
sphere with four white spots below the
midline, 10 mm in diameter, hanging on a
stalk about 3 mm long. The venom is not
active (for mammals?). From the collec-
tions available, M. carpogastra seems lo-
cally more common than other Mastopho-
ra species.
Distrihution. Southeastern Brazil, from
Rio de Janeiro State to Rio Grande do Sul
(Map 3B).
Specimens Examined. BRAZIL Bahra [?Bahia], 1?
(MNHN 18669). Rio de Janeiro: Rio de Janeiro, 2$
(NHMW); Aug. 1937, 1 imm. (Mello-Leitao, MACN
515). Sao Paulo: Agua da Figueira Maracai, 9 Feb.
1967, 15 (G. Brisolla, IBSP 2064); Barueri, Apr.
1963, 19, 5 egg sacs (K. Lenko, MZSP 3069); 16 Mar.
1966, 1 imm. (K. Lenko, MZSP 5265); Diadema,
June 1986, 1$ (R. Snignani, IBSP 4992); Embu,
Sept. 1982, 19, egg sac (A. L. Prestes, IBSP 3501);
Jandira, May 1980, 19 (C. Luiz, IBSP 963); Osasco,
25 Mar. 1974, 19 (F. Ramirez, IBSP 2721); Pacaem-
bu, Nov. 1942, 19 (Braudas, MZSP 357); Perdizes,
26 Aug. 1951, 19 (H. Camargo, MZSP 7496); Ribei-
rao Pires, May 1975, 19 (F. B. Lopes, IBSP 3591);
Rio Claro, July 1941, 19 (R Pereira, MZSP 4552); 4
May 1942, 1 imm. (Clareteano, MZSP 4402); Saco-
ma, 7 Sep. 1943, 1 9 (J. Lima, MZSP 4403); Sao Pau-
lo, July 1921, 19 (MZSP 8068); Feb. 1928, 19 (J.
Lima, MZSP 8070); 31 Jan. 1934, 19 (M. Oliveira,
IBSP 1920); 26 Feb. 1936, 19 (S. Remetente, IBSP
3598); 3 Oct. 1951, 19 (R. Vieira, IBSP 586); 17 Mar.
1955, 19, 2 egg sacs (J. Navas, IBSP 1176); June
1960, 2 9 (J. London, IBSP 1552); June 1960, 19 (L.
Zodiygansky, IBSP 1516); 21 Dec. 1960, 19 (W. An-
drade, IBSP 1618), 10 July 1962, 19 (R. R. Guidug-
lin, IBSP 1775); 21 Dec. 1961, 19 (F. V. Boas, IBSP
1620); 18 July 1962, 19 (E. Botelho, IBSP 1776); 1
Apr. 1963, 19 (Merck Co., IBSP 1834); 2 Aug. 1965,
19 (S. Remetente, IBSP 1949); 16 Dec. 1971, 19 (E
Rafael de Simone, IBSP 309); June 1975, 19 (J. S.
Gomes, IBSP .3594); Feb. 1976, 19 (M. Uchiyama,
IBSP 3597); July 1975, 19 (G. P Treu, IBSP 3595);
Feb. 1976, 19 (E. I. Yamane, IBSP 17789); Feb.
1982, 19, egg sac (D. Zammataro, IBSP 3025); 26
Feb. 1982, 19, egg sac (M. C. Franco, IBSP 14279);
22 Jan. 1986, 19 (E. Steiner, IBSP 8471); 21 Mar.
1986, 19 (D. R. Bizzachi, IBSP 14424); 1 Feb. 1991,
19 (J. Batista, IBSP 14398); Oct. 1992, 19 (C. M.
Nerici, IBSP 5827); 7 Oct. 1996, 19 (A. Fallatti,
IBSP 14192); 11 Aug. 1997, 19 (S. M. Carnelho,
IBSP 14012); 18 Mar. 1998, 19 (A. Pastore, IBSP
16208); Brooklin, Feb. 1962, 19 (L. Travassos, MZSP
4347); Hato Museu Paulista, Feb. 1951, 19 (C. Ra-
bello, MZSP 6609); Ipiranga, Nov. 1906, Oct. 1912,
2 9 (Ihering, L. M. Torre, MZSP 3047); Mar. 1924 (J.
Lima, MZSP 3048); 15 Mar. 1961, 19, 5 egg sacs
(Almeida and Gautero, MZSP 4359); 4 May 1961,
19, 2 egg sacs (N. G. Oliveira, MZSP 4340); 23 Jan.
1984, 1 imm. (G. R. F. Brandao, MZSP 435); Magi
das Gruzes, Ranch das Garmelitas, June 1976, 19 (G.
Torrus, S. Filho, MZSP 11433). Santa Catarina: Ga-
eador, 1982, 19 (D. Lorenzato, IBSP 3539). Rio
Grande do Sul: Porto Alegre, 19,1 egg sac (P. Buck,
MNRJ 1831); 7 July 1986, 19 (S. Oresco, MGN
15236); S. Leopoldo, 24 May 1964, 19 (G. Valle,
MZSP 4233); 14 Oct. 1965, 19 (G. Valle, MZSP
5422).
Mastophora seminole new species
Figures 135-141; IVIap 2G
Holotype. Female holotype from Hollendale, Bro-
ward Co., Florida, 11 June 1987 (W. Birch), in
FSGA. The specific name is a noun in apposition
after the name of the local Indian tribe.
Description. Female holotype. Carapace
olive-brown. Chelicerae, labium, endites
olive-brown. Sternum orange-olive. Coxae
and distal leg articles olive, distally darkest.
Abdomen anteriorly gray, posteriorly light
gray with dark pattern on each side (Fig.
342 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
137); in anterior view, dark area forming a
triangle as wide as humps above, pointed
to carapace below, each side white. Venter
olive gray with white longitudinal rectan-
gle. Carapace glossy and with tubercles
(Figs. 135, 136). Abdomen with a pair of
dorsal humps and triangular (Figs. 137,
138). Total length 11.6 mm. Carapace 4.4
mm long, 4.3 wide in thoracic region, 2.7
wide at posterior lateral eyes. First femur
3.8 mm, patella and tibia 5.2, metatarsus
3.7, tarsus 1.2. Second patella and tibia 3.8
mm, third 2.2, fourth 3.3. Length of first
patella and tibia 1.2 times width of cara-
pace.
Males are not known.
Diagnosis. Mastophora seminole has a
humped, somewhat triangular abdomen
(Figs. 137); the epigynum is as in the
humpless M. ijeargani, with the slits dor-
sally at some distance from the ventral
borders of large shallow depressions (Figs.
140, 141).
Distribution. Southern Florida (Map
2G).
Specimens Examined. No other specimens have
been collected.
Mastophora vaquera Gertsch
Figures 142-152, 452; Map 2G
Mastophora vaquera Gertsch, 1955: 240: figs. 15—18,
9 . Female holotype from Torriente, Matanzas,
Cuba, in AMNH, examined. Brignoli, 1983: 274.
Platnick, 2001.
Description. Female holotype. Carapace
orange-brown, bald dorsally with some
white setae on sides. Sternum, legs or-
ange-brown. Median eyes on bulge, lateral
eyes on bulges. Abdomen light orange-
brown with darker area anteriorly between
swellings (Fig. 145); venter with white
square. Abdomen with humps and an an-
terior, lateral swelling on each side (Figs.
145, 146). First femur with distal, anteri-
orly small tubercles (Fig. 144). Total
length 10.5 mm. Carapace 3.6 mm long,
3.5 wide in thoracic region, 2.3 wide at lat-
eral eyes. First femur 3.2 mm, patella and
tibia 4.7, metatarsus 3.2, tarsus 1.1. Sec-
ond patella and tibia 3.4 mm, third 2.0,
fourth 3.1. Length of first patella and tibia
1.3 times width of carapace.
Males are not known.
Variation. Total length of females 8.5—
10.5 mm. The tubercles on the femur of
the Cuabitas specimens are smaller, less
distinct. Although the shape of the abdo-
men and their coloration are similar, the
epigyna of the two specimens differ (Figs.
147-152). The illustrations (Figs. 142-
149) were made from the female holotype.
Diagnosis. Mastophora vaquera is dis-
tinguished from others by the shape of the
abdomen, humps, and two pairs of swell-
ings, and by the dorsal oval depressions of
the opening slits on the posterior of the
epigynum (Figs. 148, 151).
The egg sac has only minute flaps (Fig.
452).
Distribution. Cuba (Map 2G).
Specimens Examined. CUBA Santiago: Cuabitas,
Oriente [20°04'N, 75°48'W], 20 Aug. 1949, 19
(AMNH). Holquin: Banes, 1-3 Aug. 1955, egg sac (A.
F. Archer, AMNH).
Mastophora hutchinsoni Gertsch
Figures 153-168, 453, 454; IVIap 2E
Cyrtarachne cornigera: — McCook, 1890: 98, 99, fig.
81, egg sacs. Kaston, 1948: 231, figs. 741, 742, 2039
( misidentification ) .
MastopJiora hutchinsoni Certsch, 1955: 2.36, pi. 6, fig.
3, text figs. 10-14, 39, 47, 48, 9, 3. Female holo-
type, from Somers [Westchester Co.], New York
State, in AMNH, not examined. Brignoli, 1983:
273. Yeargan, 1988: 524. Gemeno et al., 2000:
1235. Haynes et al., 1996: 76. Platnick, 2001.
Description. Female from Virginia. Car-
apace brown, sides and posterior much
darker than median and cephalic region,
short setae on sides and no rim. Abdomen
with transverse black band with character-
istic light lines (Fig. 155); venter black in-
cluding behind spinnerets and anterior of
pedicel, with four white spots on each side
between epigynum and spinnerets (Figs.
157). Abdomen with two humps (Figs.
155, 156). Total length 7.3 mm. Carapace
3.3 mm long, 2.7 wide in thoracic region,
2.3 wide at lateral eyes. First femur 2.5
mm, patella and tibia 3.6, metatarsus 2.8,
tarsus 0.8. Second patella and tibia 2.7
Mastophora • Levi
343
Figures 153-168. Mastophora hutchinsoni GerXsch. 153-160, female. 153, 154, carapace and chelicerae. 153, frontal. 154,
lateral. 155, 156, carapace and abdomen. 155, dorsal, with male. 156, lateral. 157, abdomen, ventral. 158-160, epigynum. 158^
ventral. 159, posterior. 160, posterior, cleared. 161-168, male left palpus, stained. 161-164, (New Hampshire). 165-168, (Ken-
tucky). 161, 165, apical. 162, 166, mesal. 163, 167, ventral. 164, 168, ectal.
Figures 169-182. M. cornigera (Hentz). 169-175, female. 169, 170, carapace and chelicerae. 169, frontal. 170, lateral. 171,
1 72, carapace and abdomen. 171, dorsal, with male. 1 72, lateral. 1 73-1 75, epigynum. 1 73, ventral. 1 74, posterior. 1 75, posterior,
cleared. 176-182, male left palpus, stained. 176-179, (California). 180-182, (Texas). 176, apical. 177, 180 mesal 178 181
ventral. 179, 182, ectal.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
344 Bulletin Museutn of Comparative Zoology, Vol. 157, No. 5
mm, third 1.6, fourth 2.4. Length of first
patella and tibia 1.3 times width of cara-
pace.
Male from New Hampshire. Prosoma
beige with two median tubercles and forks.
Abdomen whitish with paired humps. To-
tal length 1.7 mm. Carapace 0.86 mm
long, 0.71 wide in thoracic region, 0.53
wide at lateral eyes. First femur 0.80 mm,
patella and tibia 0.91, metatarsus 0.41, tar-
sus 0.27. Second patella and tibia 0.80
mm, third 0.42, fourth 0.63. Length of first
patella and tibia 1.3 times the width of car-
apace.
Note. The male was matched with fe-
males because of its northerly collecting
site. They have also been collected at sites
for females in Kentucky and South Caro-
lina.
Variation. Total length of females 6.2—
10.4 mm. The males may lack humps. The
illustrations were made from a female
from Rhode Island, the inale from south-
ern New Hampshire, and a male from
Kentucky.
Diagnosis. The humps of the abdomen
(Fig. 155) and the black median coloration
of the venter, sometimes containing two
longitudinal white patches (Fig. 157), sep-
arate the species from M. bisaccata. Un-
like M. cornigera, which has humps, the
slits on the posterior of the epigynum ap-
proach each other ventrally (Figs. 158,
160), as they do in M. bisaccata.
The palpus (Figs. 161—168) is similar to
that of M. bisaccata (Figs. 59-62), with a
narrow space surrounded by the median
apophysis in ectal view (Figs. 164, 168) but
differs in the shape of the round base of
the median apophysis (Figs. 162, 166).
The egg sacs are unique compared with
those of other species. They are attached
by the broad base with the funnel facing
away from the attachment (Figs. 153, 154).
Natural History. In Ohio, females were
found in a house yard on a variety of low
branches of a variety of trees, including
crab apple, redbud, hawthorn, burr oak
(R. Bradley, in letter). Spiders were col-
lected primarily from hackberry (Celtis oc-
cidentalis L.) and wild cherry (Pninus ser-
otina Ehrgard) in Kentucky (Yeargan,
1988), on an abandoned apple tree in Vir-
ginia, in a peach orchard in South Caroli-
na, and in an apple orchard in Illinois.
Prey caught in Kentucky were reported by
Yeargan and Quate (1996).
Although the egg sacs look like hazel-
nuts or a small broken branch, the species
may be found by searching for the egg sacs
in autumn and winter after the leaves have
fallen.
Distribution. Northeastern United
States from New Hampshire and South
Carolina to Minnesota (Map 2E).
Specimens Examined. NEW HAMPSHIRE Hills-
borough Co.: Ponemah [in Amherst], Aug. 1912, 13
(E. B. Bryant, MCZ). MASSACHUSETTS 19 (San-
born, MCZ). Middlesex Co.: Pepperell, 15 July 1978,
1 imm. 6, 1.5 mm (F. J. Murphy, JM). RHODE IS-
LAND Providence, 19 (N. Banks, MCZ). CON-
NECTICUT Hartford Co.: Rocky Hill, 23 Oct. 1940,
19 (A. Morgan, USNM). NEW JERSEY Bur?mgton
Co.: Riverton, Sept. 1926, 2 imm., 13 (R. J. Sim,
OSU). Hunterdon Co.: White House Station, Sept.
1917, 19 (J. J. Brochon, USNM). Morris Co.: Dover,
8 Dec. 1950, egg sacs (USNM). OHIO Delaware Co.:
5 km W of Delaware Dam, 40.37°N, 83.10°W, Oct.
2001, 29, egg sacs (R. Bradley, OSU). DISTRICT
OF COLOMBIA Washington, yard, 18 Oct. 1920, 1 9
(USNM). VIRCINIA Falls Church, 59, 2 imm. (N.
Banks, MCZ). Augusta Co.: Augusta, 14 June 1976,
13 ,6 Oct. 1976, 1 9 (J. R McCaffrey, CNC). SOUTH
CAROLINA Anderson Co.: Simpson Agric. Exp. Sta-
tion, 10 July 1978, Ic?; 13 Aug. 1979, 19; 19 Sep.
1978, 19 (G. Lee, CUAC). KENTUCKY Fayette Co.:
Lexington, 7 Sep. 1990, 19 (K. V. Yeargan, MCZ); 20
Sep. 1990, 19 (K. V. Yeargan, KVY); 2 Oct. 1999, 19
(K. V. Yeargan, MCZ). Clark Co.: Feb. 1996, egg sac,
43 (K. V. Yeargan, KVY). TENNESSEE Ashburn, 30
mi. N. Nashville, 17 July 1933, 13 (W Ivie, AMNH).
MICHIGAN Livingston Co.: E. S. George Reserve,
22 July 1951, 1 imm. (H. K. Wallace, FSCA). IN-
DIANA Putnam Co.: Greencastle, 19 (N. Banks,
MCZ). ILLINOIS Champaign Co.: Univ. Illinois ap-
ple orchard, Sept. 1993, 19 (S. D. Gaimari, INHS).
Jackson Co.: 8 km S Carbondale on avocado plant,
Oct. 1976, 19 (JAB). MINNESOTA Hennepin Co.:
Minneapohs, 1 Nov. 1931, 19 (W J. Gertsch,
AMNH).
Mastophora cornigera (Hentz)
Figures 169-182, 455; Map 2G
Epeira cornigera Hentz, 1850: 20, pi. 3, fig. 8, 9 .
Immature female holotype from Alabama, de-
stroyed. Hentz, 1875: 123, pi. 14, fig. 8, 9 .
Mastophora • Levi
345
Cijrtarachne bicurvata Becker, 1879: 77, pi. 2, figs.
16-19, 9 . Female holotype from peach tree, Don-
aldsonville, Louisiana, in IRSNB, examined. First
synonymized by Marx (1890).
Ciji'tarachne comigera: — Keyserling, 1880: 300, pi. 4,
fig. 4, 9. McCook, 1890: 98, fig. 80 (in part).
Ordgarius comigerus: — Marx, 1890: 541. McCook,
1894: 197, pi. 12, fig. 1, 9.
Glijptocraniuin cornigenim: — Simon, 1895: 882, 885.
Bonnet, 1957: 1996.
Mastophora comigera: — Mello-Leitao', 1931: 70, figs.
9, 20. Gertsch, 1955: 233, pi. 6, fig. 2, text figs. 1-5,
37, 41, 42, 9, 3. Platnick, 1997: 513. Platnick,
2001.
Description. Female from Alabama.
Carapace evenly colored orange. Chelic-
erae, endites, labium orange. Sternum or-
ange. Legs orange, darker above. Abdo-
men with black caps on humps, anterior
gray with light lines, posterior light (Fig.
171); venter with a white square. Carapace
granular with many small tubercles and
dark spots on sides (Figs. 169, 170). Ab-
domen with pair of humps (Fig. 171). To-
tal length 12.0 mm. Carapace 5.6 mm
long, 4.6 wide in thoracic region, 3.0 wide
at lateral eyes. First femur 4.2 mm, patella
and tibia 6.0, metatarsus 4.0, tarsus 1.2.
Second patella and tibia 4.7 mm, third 2.6,
fourth 4.3. Length of first patella and tibia
1.3 times width of carapace.
Male from California. Carapace beige
with median dorsal white band including
median horns, no tubercles. Sternum, legs
golden yellow. Abdomen white, anteriorly
dusky; venter dark yellow. Carapace gran-
ulate. Total length 1.7 mm. Carapace 0.88
mm long, 0.79 wide in thoracic region,
0.52 wide behind posterior lateral eyes.
First femur 0.66 mm, patella and tibia
0.79, metatarsus 0.40, tarsus 0.28. Second
patella and tibia 0.68 mm, third 0.41,
fourth 0.54. Length of first patella and tib-
ia 1.1 times width of carapace.
Note. Males came from California, an
area from which only one species of Mas-
tophora, M. cornigera, is known.
Variation. Total length of females 8.8—
14.0 mm, males 1.6—1.7. The illustrations
were made from the female from Alabama
(Figs. 169—175) and males from California
(Figs. 176-179) and Texas (Figs. 180-182).
Adult males may lack humps.
Diagnosis. Unlike most other North
American species, M. cornigera has dis-
tinct humps on the abdomen, often with a
black cap or slightly sclerotized (Fig. 173),
and the epigynum differs from that of M.
hutchinsoni (Figs. 159, 160) by having al-
most parallel slits, only slightly converging
ventrally, on the posterior face of the epi-
gynum (Figs. 174, 175), and by lacking the
lip surrounding the slits as in M. archeri
(Fig. 188).
The male differs from other North
American species by having the space
within the curl of the median apophysis in
ectal view wider than long (Figs. 179, 182),
and from M. ijeargani by the base of the
embolus, which, in mesal view, is longer
than wide (Figs. 177, 180).
The egg sac has small flaps or none and
a relatively wide stalk (Fig. 455).
Natural History. Unlike other North
American species, M. cornigera is active in
California all year. Also, unlike other
North American species, the males
emerge from the egg sac as mature indi-
viduals. This species was found on cycad
leaf in full sun in San Diego, and on ]at-
ropha curcas euphorbia in Nicaragua.
Distribution. From Kentucky and Ten-
nesse west to California and south to Cen-
tral America (Map 2G).
Specimens Examined. KENTUCKY Fayette Co.:
Univ. Kentucky Maine Chance farm, Sept. 1996, 1 9
(K. V. Yeargan, KVY). TENNESSEE Ashburn, 30 mi.
N Nashville, 17 July 1933, 19 (W. Ivie, AMNH). AL-
ABAMA Mobile Co.: Mobile, 19 (N. Banks, MCZ);
1932, 19 (H. P. Loding, MCZ). LOUISIANA East
Baton Rouge Par: Baton Rouge, Apr 1916, 19 (New-
ell, MCZ). Orleans Par: New Orleans, 1918, 19 (H.
E. Hubert, USNM); 1 Oct. 1935, 19 (T. E. Snyder,
MCZ); 26 Sep. 1936, 19 (J. N. Cowanloch, USNM).
TEXAS Travis Co.: Shellberg Tract, 30°25'N,
97°52'W, 18-19 Apr. 1994, 3S (Dunlap et al,
TAMU). Galveston Co.: Texas City, 1921?, 29 (S. W.
Bilsing, MCZ). San Patricio Co.: Welder Wildlife
Refuge, 11.8 km NE Sinton, 17 Oct. 1967, 16 (C.
Parrish, CAS). Hidalgo Co.: Edinburg, Oct. 1934, 39,
13; 1935, 19 (S. Mulaik, AMNH); 7 Dec. 1935, 16
(M. Welch, AMNH), 2 imm., 16 (S. Mulaik,
AMNH); 18 km SE Pharr, Santa Ana Wildlife Ref-
uge, 1 Oct. 1977, 19 (O. Alirenholtz, AMNH); 20
346 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Dec. 1983, 1? (M. K. Stowe, MKS); 27 June 1984,
19 (M. K. Stowe, MKS); Mercedes, 10 Apr. 1986,
egg sac, 70 imm., 593 (D. A. Dean, TAMU). Cain-
eron Co.: Brownsville, 9 Apr. 1986, hatched 2 June
1986, 1 egg sac, 64 imm., 62c? (D. A. Dean, TAMU);
Harlingen, Feb. 1980, 1?, 64 imm., 63(? (C. W. Ag-
new et al., TAMU); E of Harlingen, 3 Jan. 1936, 1 9
(L. I. Davis, M. Stegmeier, AMNH). ARIZONA Mar-
icopa Co.: Phoenix, Apr. 1941, 1? (AMNH). CALI-
FORNIA Contra Costa Co.: Walnut Creek, Sydney
Drive, July 1992, 19, 13 (T. Trosin, J. Fraser, CAS).
Santa Clara Co.: Palo Alto, 1914, 19 (H. Heath,
MCZ). Santa Barbara Co.: Santa Barbara, 5 Oct.
1948, 19 (H. Shantz, AMNH). Lo.s Angele.s Co.:
Claremont, 19 (N. Banks, MCZ); Glendale, 19,
many 3 (C. E. Hutchinson, AMNH, MCZ); Los An-
geles, 19 (MNHN, 3059); 10 Oct. 1942, 53 (J. H.
Branch, AMNH); East Los Angeles, 1943, 19 (C.
Cowles, AMNH); Malibu, Nov. 1968, 19,3 egg sacs
(USNM); Westwood Village, Aug.-Oct. 1942, 19 (P
Verrity et al., FSCA). Orange Co.: Santa Ana, 43 (R.
K. Bishop, USNM); San Juan Capistrano, 25 Sept.
1952, 19, 7 egg sacs (R. E. Ryckman, AMNH). San
Bernardino Co.: San Bernardino, 1880, 19 (J. B. Par-
ish, MCZ). San Diego Co.: San Diego, 29 (USNM),
4 Oct. 1974, 19 (D. Bishop, USNM); San Diego,
Vista, 8 June 1989, 1 imm. (J. W. Schott, MCZ); Chu-
la Vista, 5 Dec. 1981, 19 (H. V. Weems, FSDA); 10
mi. NE Ramona, 22 July 1982, 19 (J. Halstead, DU);
Lakeside, 19 (C. Kingeiy, USNM); Feb. May 1968,
egg sacs, 3 3 (C. Kingeiy, USNM). MEXICO Baja
California Sun 44 km W La Paz, 0.2 km S km 44,
on Highway 1, 31 Dec. 1978, 19 (D. Weissman, R.
Love et al, CAS). NICARAGUA Managua: Mateare,
12 Sep. 1995, 19 (C. Grimm, M. Maes, MCZ).
Mastophora archeri Gertsch
Figures 183-193, 456; Map 2E
Mastophora archeri Gertsch, 1955: 239: figs. 6—9, 36,
45, 46, 9, 3. Female holotype from Fruitland
Park, Florida, in AMNH, examined. Brignoli, 1983:
273. Platnick, 2001.
Description. Female holotype. Carapace
orange-brown. Sternum dark orange. Legs
orange-brown, indistinctly ringed. Abdo-
men anteriorly gray, posteriorly white; ven-
ter with white square. Carapace with short
tubercles (Figs. 183, 184). Median eyes on
bulge, lateral eyes on bulges. Abdoinen
with small humps (Fig. 185). Total length
11.5 mm. Carapace 4.2 mm long, 4.3 wide
in thoracic region, 2.7 wide at lateral eyes.
First femur 3.8 mm, patella and tibia 5.6,
[metatarsus 3.5, tarsus 1.5, after Gertsch,
1955]. Second patella and tibia 4.1 mm.
third 2.3, fourth 3.7. Length of first patella
and tibia 1.3 tiiues width of carapace.
Male allotype. Carapace orange, darkest
on sides, a median white line and median
of forked tubercles white. Sternum or-
ange. Legs orange. Abdomen orange-
white without marks. Carapace with two
small asymmetrical tubercles in addition to
forked tubercles. Abdomen with indistinct
humps. Total length 1.7 mm. Carapace
0.86 mm long, 0.78 wide in thoracic re-
gion, 0.53 wide at lateral eyes. First femur
0.79 mm, patella and tibia 0.87, metatarsus
0.44, tarsus 0.33. Second patella and tibia
0.74 mm, third 0.54, fourth 0.71. Length
of first patella and tibia 1 . 1 times width of
carapace.
Note. A male from Gainesville, Florida,
was raised from the egg sac. The match of
the male allotype is uncertain.
Variation. Total length of females 9.4—
14.8 mm. The illustrations were made
from the female holotype (Figs. 183—189)
and the raised male from Gainesville (Figs.
190-193).
Diagnosis. Mastophora archeri is distin-
guished from M. cornigera by the smaller
tubercles on the carapace (Figs. 183, 184)
and left and right lip on the posterior of
the epigynum (Figs. 188, 189), and from
M. hutchinsoni by having a white square
on the venter and by the sculpturing of the
epigynum.
The palpus of the male has the space
surrounded by the median apophysis wid-
er than long (Fig. 193) and differs fro in
M. phrynosoma and M. hutchinsoni by
having only a minute terminal apophysis
(Figs. 190, 191). It differs from M. phrij-
nosonia by the narrow radix (Fig. 191).
The egg sac has small flaps and a rela-
tively long stalk of median thickness (Fig.
456).
Natural History. Collected from Myrica
in hammock woods in Alabama.
Distribution. Southern United States
from South Carolina, Florida, and Ala-
bama to Kansas (Map 2E).
Paratypes. ALABAMA Baldivin Co.: Lagoon, 29
Mastophora • Levi 347
Figures 183-193. Mastophora archeri Gertsch. 183-189, female. 183, 184, carapace and chelicerae. 183, frontal. 184, lateral.
185, 186, carapace and abdomen. 185, dorsal, with male. 186, lateral. 187-189, epigynum. 187, ventral. 188, posterior. 189,
posterior, cleared. 190-193, male left palpus, stained. 190, apical. 191, mesal. 192, ventral. 193, ectal.
Figures 194-204. M. fasciata Reimoser. 194-200, female. 194, 195, carapace and chelicerae. 194, frontal. 195, lateral. 196,
1 97, carapace and abdomen. 1 96, dorsal, with male. 1 97, lateral. 1 98-200, epigynum. 1 98, ventral. 1 99, posterior. 200, posterior,
cleared. 201-204, male left palpus, stained. 201, apical. 202, mesal. 203, ventral. 204, ectal.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
348 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Sep. 1949, IcJ allotype (A. F. Archer). KANSAS
Douglas Co.: 22 Sept. 1948, 1? (R. H. Beaner,
AMNH).
Specimens Examined. SOUTH CAROLINA
Charleston, Oct. 1941, 19 (E. B. Chamberlain,
USNM). FLORIDA Alachua Co.: Devils Millhopper
State Park, reared spring 1992, 43 (M. K. Stowe,
MKS). Hillsborough Co.: Thonotosassa, 11 Dec.
1969, 1$ (D. A. Vaughn, FSCA). ALABAMA Mobile
Co.: Mobile, 12 Nov. 1939, 15 (A. C. Cole, AMNH);
Mt. Vernon, Oct. 1941, 19 (H. R Loding, AMNH).
Mastophora fasciata Reimoser
Plate 1; Figures 194-204, 457; Maps
2F, 3A
Mastophora fasciata Reimoser, 1940: 356, fig. 8, 9 .
Female holotype, from Orosi, Prov. Cartago, cen-
tral plain [12 km SE Cartago], Costa Rica, in
NMW, examined. Roewer, 1942: 900. Platnick,
2001.
Mastophora pickeli occidentalis Schenkel, 1953: 29.
Female holotype from Pozon [Falcon], Venezuela,
in NMB, examined. Brignoli, 1983: 274. Platnick,
2001.
Note. Mastophora p. occidentalis has the
same small lobe on the posterior margin
of the epigynum, and the same shadows
on the posterior face as in M. fasciata.
They also share the very broad humps of
the abdoiTien and the posterior light band.
Description. Feinale from Costa Rica.
Carapace light orange-brown. Sternum
brown. Legs lighter brown. Abdomen
white with black and gray marks (Fig.
196); venter light brown, center barely
lighter than sides. (Reimoser described a
posterior, transverse, yellow-red band,
which has disappeared from the holotype,
but is light in other specimens.) Humps
broad (Fig. 196). Holotype total length
11.5 mm. Carapace 4.7 mm long, 5.0 wide
in thoracic region, 3.0 wide at lateral eyes.
First femur 4.3 mm, patella and tibia 6.3,
metatarsus 4.4, tarsus 1.2. Second patella
and tibia 4.6 mm, third 2.7, fourth 4.0.
Length of first patella and tibia 1.3 times
width of carapace.
IVIale from Costa Rica. Carapace yellow-
brown with white, central mark covering
median tubercles. Legs light yellow-
brown. Abdomen dorsally white, ventrally
yellow-brown. Eyes without pigment. Car-
apace rugose with posterior median forked
tubercles. Abdomen subtriangular without
humps. Palpal patella with one iTiacroseta.
Total length 1.6 mm. Carapace 0.74 mm
long, 0.65 wide in thoracic region, 0.46
wide behind posterior lateral eyes. First
feinur 0.55 mm, patella and tibia 0.65,
metatarsus 0.35, tarsus 0.26. Second pa-
tella and tibia 0.59 mm, third 0.28, fourth
0.48. First patella and tibia as long as
width of carapace.
Note. Males were collected with females
at San Antonio de Escazu.
Variation. Total length of females 11.5—
14.5 mm. Both males and shriveled fe-
males have a triangular abdomen, whereas
that of a well-fed female is more rounded.
The illustrations were made from the fe-
male holotype (Figs. 194—200) with a spec-
imen from Puntarenas Province, Costa
Rica.
Diagnosis. The female is distinguished
from other species by the many small car-
apace tubercles (Figs. 194, 195), by the
broad humps of the abdomen (Fig. 196),
and by the ventrally converging slits of the
epigynum (Figs. 199, 200).
The male has a median apophysis that
is longer than that of other species (Figs.
202, 204) and more rounded than that of
M. leucabulba (Figs. 293, 295) and M. al-
vareztoroi (Fig. 306, 307). The median
apophysis is almost as long as the diameter
of the bulb (Fig. 204).
The egg sac lacks flaps (Fig. 457).
Natural History. JVIales are mature
when they leave the egg sac.
Distribution. The species is known from
Costa Rica and Venezuela (IVIaps 2F, 3A).
Figures 205-215. Mastophora dizzydeani Eberhard. 205-211, female. 205, 206, carapace and chelicerae. 205, frontal. 206,
lateral. 207, 208, carapace and abdomen. 207, dorsal, with male. 208, lateral. 209-211, epigynum, 209, ventral. 210, posterior.
211, posterior, cleared. 212-215, male left palpus, stained. 212, apical. 213, mesa!. 214, ventral. 215, ectal.
Mastophora • Levi
349
209
222 - —
A,
"^XM/y
223
Figures 216-222. M. pickeli Mello-Leitao, female. 216, 217, carapace and chelicerae. 216, frontal, 217, lateral. 218, 219,
carapace and abdomen. 218, dorsal. 219, lateral. 220-222, epigynum. 220, ventral. 221, posterior. 222, posterior, cleared.
Figures 223-229. M. cranion Mello-Leitao, female. 223, 224, carapace and chelicerae. 223, frontal. 224, lateral. 225, 226,
carapace and abdomen. 225, dorsal. 226, lateral. 227-229, epigynum. 227, ventral. 228, posterior. 229, posterior, cleared.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
350 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Specimens Examined. COSTA RICA San Jose: San
Jose, 1? (Tristan, MCZ); San Antonio de Escazu,
1,350 m, May 1980, 19 (W. Eberhard S31, MCZ); 7
Sept. 1980, imm., 5, many S (W. Eberhard SJl-26,
MCZ); 11 Oct. 1980, 1$ (W. Eberhard S31, MCZ).
Puntarenas: Parrita, 30 m, 12 Jan. 1987, 39, 2(? (W.
Eberhard FN9-40ff, MCZ).
Mastophora dizzydeani Eberhard
Figures 205-215; Map 3A
Mastophora dizzijdeani Eberhard, 1981: 144, figs.
1-9, 9 , S . Female holotype from field of Melendez
campus of the Universidad del Valle on the south-
ern edge of Cali, Colombia, in MCZ, examined.
Platnick, 1989: 340. Platnick, 2001.
Description. Female holotype. Carapace
dark brown. Sternum orange-brown. Legs
brown. Abdomen white anteriorly, dorsally
■with transverse gray band, posteriorly
dusky white (Figs. 207, 208); venter with
a median white square. Carapace heavily
sclerotized, grooves on sides, many tuber-
cles flat and wide (Figs. 205, 206). Abdo-
men subtriangular with pair of dorsal
humps (Fig. 207). Total length 13.3 mm.
Carapace 5.7 mm long, 5.3 wide in tho-
racic region, 3.3 wide at lateral eyes. First
femur 4.8 mm, patella and tibia 6.7, meta-
tarsus 4.7, tarsus 1.5. Second patella and
tibia 5.0 mm, third 2.8, fourth 4.4. Length
of first patella and tibia 1.2 times width of
carapace.
Male allotype from eastern edge of Lago
Calima. Yellowish white with longitudinal
white thoracic mark, and dorsum of ab-
domen white with a couple of indistinct
gray patches anteriorly. Height of clypeus
about 1.3 diameters of anterior median
eye. Abdomen with a pair of humps. Pal-
pal patella with one weak macroseta. Total
length 1.6 mm. Carapace 0.75 mm long,
0.74 wide in thoracic region, 0.55 wide at
lateral eyes. First femur 0.71 mm, patella
and tibia 0.72, metatarsus 0.41, tarsus 0.26.
Second patella and tibia 0.65 mm, third
0.36, fourth 0.51. Length of first patella
and tibia about same length as width of
carapace.
Note. Males and females have been col-
lected together at Cali, Colombia.
Variation. Total length of females 10.8-
13.3 mm. The illustrations were made
from specimens from Cali.
Diagnosis. Females are distinguished by
the wide heart-shaped abdomen with two
humps (Fig. 207) and the distinctly shaped
dark marks around the slits on the poste-
rior of the epigynum (Fig. 210). The atria
separate from each other ventrally (Fig.
211).
Males have a smaller median apophysis
(Fig. 213) than that oi M. fasciata, in ectal
view surrounding a rounded space (Fig.
215).
Natural History. The spiders rest on ex-
posed sites: on the barb of a barbed wire,
fence posts, and upper surface of leaves.
Specimens also were collected on a guay-
oba tree in a yard and in sugar cane areas.
Moths caught include Spodoptera fmgi-
perda (a sugar cane pest) and Leucania sp.
(Eberhard, 1977, 1981).
Distribution. Colombia to northern
Peru (Map 3A).
Specimens Examined. COLOMBIA Valle: S of
Cali, on plants, 6 June 1948, 19 (E. M. Poulsen,
ZMUC); nr. Cali, Jan. 1977, imm. 9 (W. Eberhard,
MCZ); Aug. 1977, 19, AS (W. Eberhard, MCZ);
Lago Calima, 1,400 m, 19 Nov. 1977, 19 (W. Eber-
hard EG 3-20, MCZ); Rio Tulua, 1,100 m, Aug.
1977, 2S (W. Eberhard, MCZ). PERU Piura: Mal-
lagra, Rio Cliira, 8 June 1941, 29, imm. (D. L. Friz-
zell and H. E. Frizzell. AMNH. CAS).
Mastophora pickeli Mello-Leitao
Figures 216-222; Map 3D
Mastophora pickeli Mello-Leitao, 1931: 73, figs. 6, 18,
24, 25, 9 . Female holotype from Tapera, Pemam-
buco, Brazil, in MNRJ, 395, examined. The specific
name is a noun in apposition after the locality.
Roewer, 1942: 901. Platnick, 2001.
Glijptocranium pickeli: — Bonnet, 1957: 1998.
Note. Vanzolini and Papavero (1968)
listed three localities with the name Tap-
ero in Pernambuco. I assume this locality
is the only one also listed in the Index to
Map of Hispanic America (American Geo-
graphical Society of New York, 1944), a
railroad station, west of Recife.
Description. Female holotype. Cara-
pace, chelicerae, labium, endites, brown.
Sternum patchy orange-brown. Coxae and
Mastophora • Levi
351
distal leg articles brown. Abdomen black
anteriorly enclosing white streaks, white
posteriorly (Fig. 218); ventrally with indis-
tinct white square on gray. Carapace with
shallow tubercles, glossy, with short setae
on sides (Figs. 216, 217). Abdomen with-
out setae, with a pair of wide humps bear-
ing distinct smaller humps dorsally (Figs.
218, 219) and slight swellings on side (Fig.
219). Total length 9.5 mm. Carapace 4.4
mm long, 3.7 wide in thoracic region, 2.4
wide at lateral eyes. First femur 3.4 mm,
patella and tibia 4.8, metatarsus 3.3, tarsus
0.9. Second patella and tibia 3.4 mm, third
2.0, fourth 3.2. Length of first patella and
tibia 1.1 times width of carapace.
Males are not known.
Diagnosis. This species is distinguished
by distinct small humps on a larger swell-
ing of the abdomen (Figs. 218, 219) and
the epigynum with atria approaching each
other (Figs. 221, 222). It differs from M.
ypiranga, which has a siinilar epigynum,
by having the carapace, viewed from an-
terior, wider and more swollen, the ante-
rior median eyes facing slightly laterally
and ventrally, and the forked tubercles laid
back, with their tips facing posteriorly
(Fig. 217).
Distribution. Known only from the type
locality (Map 3D).
Specimens Examined. No other specimens have
been found.
Mastophora cranion Mello-Leitao
Figures 223-229; Map 3D
Mastophora cranion Mello-Leitao, 1928: 49, pi. 1, 9 .
Female holotype from Tapera, Est. Pemambuco,
Brazil, in MNRJ no. 00394, examined. Mello-Lei-
tao, 1931: 72, figs. 2, 14, 9. Roewer, 1942: 955.
Platnick, 2001.
GJyprocraniiini cranion: — Bonnet, 1957: 1997.
Note. For locality information, see note
under M. pickeli.
Description. Female holotype. Carapace
orange-brown, black pigment between an-
terior median eyes. Chelicerae, labium,
endites brown. Sternum uneven white and
brown. Coxae and distal leg articles light
brown. Abdomen white with a pair of dor-
sal dark spots, each consisting of stippled
black dots (Fig. 218); venter whitish with
white square, spinnerets brown. Carapace
with long white setae on sides and on clyp-
eus and many small tubercles (Figs. 223,
224). Abdomen with wide humps, anterior
edge with long setae (Figs. 225, 226). Total
length 10.8 mm. Carapace 4.7 mm long,
4.3 wide in thoracic region, 2.8 wide at lat-
eral eyes. First femur 4.0 mm, patella and
tibia 6.3, metatarsus 4.7, tarsus 1.1. Sec-
ond patella and tibia 4.3 mm, third 2.4,
fourth 3.7. Length of first patella and tibia
1.5 times width of carapace.
Males are not known.
Diagnosis. Mastophora cranion is distin-
guished from others by black spots on the
wide humps of the abdomen (Figs. 225,
226), its long white setae, and its epigyn-
um with the atria in line with the outer
margin of the seminal receptacles (Fig.
229).
Distribution. Known only from the type
locality (Map 3D).
Specimens Examined. No other specimens have
been found.
Mastophora longiceps Mello-Leitao
Figures 230-236; Map 3D
Mastophora longiceps Mello-Leitao, 1940: 57, fig. 5,
? . Female holotype from Ilha Sao Sebastiao, Est.
Sao Paulo, Brazil, in MNRJ, examined. Brignoli,
1983: 273. Platnick, 2001.
Glyptocranium longiceps: — Bonnet, 1957: 1998.
Description. Female holotype. Carapace
beige in center, dark brown on sides. Che-
licerae, labium, endites, sternum orange-
brown. Coxae and distal leg articles or-
ange-brown. Abdomen contrastingly
marked black and white (Figs. 232, 233);
venter gray with a pair of light patches.
Carapace shiny with short setae on sides,
both black and white; forked horns thick,
fingerfike (Figs. 230, 231). Legs with short
black and white setae. Eyes indistinct
without dark pigment. Abdomen bald with
wide humps (Figs. 232, 233). Total length
13.0 mm. Carapace 7.5 mm long, 6.0 wide
in thoracic region, 3.8 wide at lateral eyes.
First femur 6.3 mm, patella and tibia 9.2,
352 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
metatarsus 6.4, tarsus 1.6. Second patella
and tibia 6.5 mm, third 3.5, fourth 5.6.
Length of first patella and tibia 1.5 times
width of carapace.
Diagnosis. The thick horns (Figs. 230,
231) and wide median plate on the pos-
terior of the epigynum (Fig. 235) separate
this species from M. melloleitaoi (Figs.
274-280).
Distribution. Known only from the type
locality (Map 3D).
Specimens Examined. No other specimens have
been found.
Mastophora pesqueiro new species
Figures 237-243; Map 3D
Holotype. Female holotype from Pesqueiro, Monte-
negro, Rio Grande de Sul, Brazil, 14 June 1977 (M.
F. Beurmann), in MCN no. 5730. The specific
name is a noun in apposition after the locality in
Montenegro.
Description. Female holotype. Carapace
light brown to dark red-brown, glossy, with
two lines of white setae posteriorly on each
side and a white edge. Chelicerae, labium,
endites, sternum, coxae light brown. Distal
leg articles light brown, darker ventrally,
with white setae. Abdomen marked with
two darker framed white patches on each
side and a transverse dusky line anteriorly
in a median white area (Figs. 239, 240);
venter light brown with white square. Car-
apace glossy with posterior horns pointed
and two small median tubercles (Figs. 237,
238). Abdomen with small, pointed humps
(Figs. 239, 240). Total length 10.5 mm.
Carapace 4.7 mm long, 4.6 wide in tho-
racic region, 2.3 wide at lateral eyes. First
femur 4.3 mm, patella and tibia 6.9, meta-
tarsus 4.6, tarsus 1.4. Second patella and
tibia 5.0 mm, third 2.7, fourth 4.7. Length
of first patella and tibia 1.5 times carapace
width.
Males are not known.
Diagnosis. Mastophora pesqueiro is dis-
tinguished by the conical horns on the
smooth carapace (Figs. 237, 238), the four
white marks on the abdomen, and the dis-
tinctly shaped humps (Figs. 239, 240). The
epigynum has a median bulge on its pos-
terior face (Figs. 241-243).
Distribution. Known only from the type
locality (Map 3D).
Specimens Examined. No other specimens have
been found.
Mastophora piras new species
Figures 244-250; IVlap 3D
Holotype. Female holotype from Emas, Pirassunga,
Est. Sao Paulo, Brazil, 2 Nov, 1952 (Pietracatelli,
Werner, and Dionisio), in MZSP, 4339. The specific
name is an arbitraiy combination of letters.
Description. Female holotype. Carapace
dark brown. Chelicerae light brown. La-
bium, endites, sternum light brown. Coxae
and distal leg articles brown with indistinct
darker blotches. Abdomen anterior black
enclosing some white anterior loops, pos-
terior white (Figs. 246, 247); venter with
indistinct white square on gray-brown.
Carapace heavily sclerotized, shiny, with
sides thinner with a punctuate pattern of
tiny tubercles. Abdomen triangular heart-
shaped with lateral bulges and distinct
small humps (Figs. 246, 247). Total length
13.8 mm. Carapace 6.5 mm long, 5.9 wide
in thoracic region, 3.3 wide at lateral eyes.
First femur 5.3 mm, patella and tibia 7.7,
metatarsus 4.8, tarsus 1.6. Second patella
and tibia 5.5 mm, third 3.3, fourth 4.8.
Length of first patella and tibia 1.2 times
width of carapace.
Males are not known.
Variation. Total length of females 13.0-
13.8 mm. The paratype has some long,
white setae on sides of carapace, clypeus.
Figures 230-236. iVIastophora longiceps Mello-Leitao, female. 230, 231, carapace and chelicerae. 230, frontal. 231, lateral.
232, 233, carapace and abdomen. 232, dorsal. 233, lateral. 234-236, epigynum. 234, ventral. 235, posterior. 236, posterior,
cleared.
Figures 237-243. M. pesqueiro new species, female. 237, 238, carapace and chelicerae. 237, frontal. 238, lateral. 239, 240,
carapace and abdomen. 239, dorsal. 240, lateral. 241-243, epigynum. 241, ventral. 242, posterior. 243, posterior, cleared.
Mastophora • Levi 353
Figures 244-250. M. piras new species, female. 244, 245, carapace and chelicerae. 244, frontal. 245, lateral. 246, 247, car-
apace and abdomen. 246, dorsal. 247, lateral. 248-250, epigynum. 248, ventral. 249, posterior, 250, posterior, cleared.
Figures 251-257. M. yplranga new species, female. 251, 252, carapace and chelicerae. 251, frontal. 252, lateral. 253, 254,
carapace and abdomen. 253, dorsal. 254, lateral. 255-257, epigynum. 255, ventral. 256, posterior. 257, posterior, cleared.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
354 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
legs, and anterior of abdomen; the abdo-
men is all dark, perhaps because of poor
preservation. The illustrations were from
the holotype.
Diagnosis. Mastophora pirns is distin-
guished from M. raelloleitaoi by the heart-
shaped abdomen with small humps (Figs.
246, 247) and by the wider median area of
the epigynum in posterior view (Fig. 249).
Distribution. Minas Gerais and Sao Pau-
lo states of southern Brazil (Map 3D).
Paratypes. BRAZIL Minas Gerais: Gavemador Va-
ladares, 14/15 Oct. 1981, 1 imm. (N. Sorkin, T. Spitz-
man, AMNH). Sao Paulo: Sao Bernardo, Nov. 1926,
19 (H. Bakkenist, MZSP 8069).
Mastophora ypiranga new species
Figures 251-257; IVIap 3E
Holotype. Female holotype from Ypiranga, Cap. [city
of Sao Paulo], Est. Sao Paulo, Brazil, 1898, in
MZSP, 5791. The specific name is a noun in ap-
position after the locality.
Description. Female holotype. Carapace
orange-brown. Chelicerae yellow-brown.
Labium, endites, sternum light brovwi.
Coxae and distal leg articles light brown.
Abdoinen anterior black, posterior white
(Figs. 253, 254); venter grayish white with
white square. Carapace with low tubercles,
with some short white setae on sides (Figs.
251, 252). First femur with S-shaped cur-
vature. Abdomen with two humps (Figs.
253, 254). Total length 12.7 mm. Carapace
4.7 mm long, 4.2 wide in thoracic region,
2.7 wide at lateral eyes. First femur 3.7
mm, patella and tibia 5.4, metatarsus 3.3,
tarsus 1.1. Second patella and tibia 3.8
mm, third 2.2, fourth 3.6. Length of first
patella and tibia 1.2 times width of cara-
pace.
Males are not known
Variation. Total length of females 9.7-
12.7 mm. The illustrations were made
fro in the holotype.
Diagnosis. Mastophora ypiranga differs
from M. pickeli and M. melloleitaoi by the
shape of the abdomen (Figs. 253, 254) and
by the narrower, almost triangular median
plate of the epigynum and the atria ap-
proaching each other (Figs. 256, 257).
Other differences are that the carapace ap-
pears less swollen when viewed from an-
terior, the anterior median eyes are di-
rected forward, and the horns are erect
with the tips pointing dorsally (Fig. 252).
The anterior of the abdomen, which over-
hangs the carapace, is white.
Distribution. Minas Gerais to Santa Ca-
tarina states of southern Brazil (Map 3E).
Paratypes. BRAZIL Minas Gerais: Vicosa, 1931,
1? (E. J. Hambleton, AMNH). Sao Paulo: Sao Paulo,
July 1928, 19 (R. Cassalo, IBSP 4530). Santa Catar-
ina: June 1919, 19 (Luederwaldt, MZSP 8067).
Mastophora yacare new species
Figures 258-264; IVlap 3E
Holotype. Female holotype from Rincori del Yacare,
Artigas, Uruguay, 20 Jan. 1957, in FCMU. The spe-
cific name is a noun in apposition after the locality.
Description. Female holotype. Carapace
light brown. Chelicerae, labium, endites
brown. Sternum orange, underlain by
white pigment. Legs brown, femora light-
est. Abdomen black over humps, whitish
posteriorly (Figs. 260, 261); venter light
brown with white square containing two
rows of three black spots; spinnerets dark
brown. Abdoinen with a pair of wide, dor-
sal humps (Figs. 260, 261). Total length
11.5 mm. Carapace 4.1 mm long, 4.0 wide
in thoracic region, 2.6 wide behind pos-
terior lateral eyes. First femur 3.5 mm, pa-
tella and tibia 5.1, metatarsus 3.5, tarsus
1.1. Second patella and tibia 3.8 mm, third
Figures 258-264. Mastophora yacare new species, female. 258, 259, carapace and chelicerae. 258, frontal. 259, lateral. 260,
261, carapace and abdomen. 260, dorsal. 261, lateral. 262-264, epigynum. 262, ventral. 263, posterior. 264, posterior, cleared.
Figures 265-268. M. abalosi Urtubey and Baez, female, after authors. 265, horns of carapace, frontal. 266, carapace and
chelicerae, lateral. 267, carapace and abdomen, dorsal. 268, epigynum, postehor.
Figures 269-273. M. comma Baez and Urtubey, female, after authors. 269, horns of carapace. 270, carapace and chelicerae,
lateral. 271, 272, carapace and abdomen. 271, dorsal. 272, lateral. 273, epigynum, posterior.
Mastophora • Levi 355
Figures 274-280. M. melloleitaoi Canals, female. 274, 275, carapace and chelicerae. 274, frontal. 275, lateral. 276, 277,
carapace and abdomen. 276, dorsal. 277, lateral, 278-280, epigynum. 278, ventral. 279, posterior. 280, posterior, cleared.
Figures 281-287. M. extraordinaria Holmberg, female. 281, 282, carapace and chelicerae. 281, frontal. 282, lateral. 283, 284,
carapace and abdomen. 283, dorsal. 284, lateral. 285-287, epigynum. 285, ventral. 256, posterior. 287, posterior, cleared.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
356 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
2.0, fourth 3.3. Length of first patella and
tibia 1.3 tiiTies width of carapace.
Males are not known.
Diagnosis. Mastophora yacare is distin-
guished from M. melloleitaoi by its epigyn-
uni and the atria, which separate and face
sideways (Fig. 264).
Distribution. Known only from the type
locality (Map 3E).
Specimens Examined. No other specimens have
been found.
Mastophora abalosi Urtubey and Baez
Figures 265-268, 458; Map 3E
Mastophora abalosi Urtubey and Baez, 1983: 3, figs.
1-11, 9. Female holotype and seven paratypes
from the city of Santiago del Estero, Argentina, in
the Inst. Animales Venenosos, Santiago del Estero,
unavailable. Platnick, 1989: 340. Platnick, 2001.
Description. Female (after Urtubey and
Baez, 1983). Carapace brown. Chelicerae
yellowish. Labium, endites yellow-white.
Sternum spotted orange-yellow. Sternum
dark orange. Legs yellowish. Abdomen an-
terior clear yellow followed by dark band
that covers humps (Fig. 267), posteriorly
pale yellow; venter with light yellow. Car-
apace with numerous tubercles, setae only
on sides; tubercles with light tips. Abdo-
men with humps (Fig. 267). Total length
12.5 mm. Carapace 5.6 mm long, 5.4 wide
in thoracic region. First femur 4.8 mm, pa-
tella and tibia 6.8, metatarsus 5.1, tarsus
1.3. Second patella and tibia 5.2 mm, third
2.6, fourth 4.8. Length of first patella and
tibia 1.3 times width of carapace.
Males are not known.
Diagnosis. Mastophora abalosi is distin-
guished from M. extraordinaria by some
morphological characters (Urtubey and
Baez, 1983). The posterior of the epigyn-
um was illustrated (Fig. 268) and shows
the atria slightly to the lateral of the slits.
The egg sac has flaps (Fig. 458).
Distribution. Santiago del Estero in
northern Argentina (Map 3E).
Specimens Examined. No specimens were avail-
able.
Mastophora comma Baez and Urtubey
Figures 269-273, 459; Map 3E
Ma.stophora comma Baez and Urtubey, 1985: 3, figs.
1—9, 9 . Female holotype from the city of Santiago
del Estero, Argentina, and seven paratypes in the
Instituto de Animales Venenosos, Santiago del Es-
tero, unavailable. Platnick, 1989: 340. Platnick,
2001.
Description. Female (after Baez and Ur-
tubey, 1985). Carapace brown. Chelicerae
yellow. Sternum orange. Legs orange-yel-
low. Abdomen dusky yellow covered with
many tiny maroon spots with white setae,
spotted with brown spots and a few darker
maroon spots, and posteriorly with a few
transverse bands (Fig. 271); venter yellow,
spinnerets maroon. Intense yellow area
between epigastric groove and spinnerets,
with a few small maroon spots. Carapace
with tubercles (Figs. 269, 270). Abdomen
with a pair of dorsal humps tipped by nip-
ples (Figs. 271, 272). Total length 12.0
mm. Carapace 3.7 mm long, 4.8 wide in
thoracic region. First femur 3.9 mm, pa-
tella and tibia 6.4, metatarsus 5.0, tarsus
1.2. Second patella and tibia 4.6 mm, third
2.4, fourth 3.9. Length of first patella and
tibia 1.3 times width of carapace.
Males are not known.
Diagnosis. This species differs from oth-
ers by the unusual stnacture of the abdo-
menal humps (Fig. 272) and by the two
parallel slits on the posterior face of the
epigynum (Fig. 273).
The egg sac lacks flaps and has a short
stalk (Fig. 459).
Distribution. Santiago del Estero in
northern Argentina (Map 3E).
Specimens Examined. No specimens of this species
were available.
Mastophora melloleitaoi Canals
Figures 274-280; Map 3E
Mastophora Mello-Leitaoi Canals, 1931: 20, figs. 1-4,
pi. 2, fig. 4, 9 . Female holotype, from Rosas, Prov.
Buenos Aires, Argentina, in MACN, examined.
Mastophora mello-leitdo: — Mello-Leitao, 1931, 73:
figs, 5, 17, 9 .
Mastophora mello-leitaoiae: — Roewer, 1942: 900.
Glyptocranium melloleitaoi: — Bonnet, 1957: 1998.
Mastophora melloleitaoi: — Platnick, 2001.
Mastophora • Levi
357
Note. Roewer's (1942) spelling appears
to be an error.
Description. Female from Balcarce.
Carapace light brown, not shiny. Chelic-
erae, endites, labium light brown. Sternum
light brown. Legs brown. Abdomen light
brown, anterior gray. Near midline ante-
riorly a light V-shaped mark, followed by
a light upside-down V. Carapace bald, ex-
cept for some setae posteriorly. Abdomen
humps wide (Figs. 276, 277). Total length
13.0 mm. Carapace 5.5 mm long, 4.8 wide
in thoracic region, 3.2 wide at lateral eyes.
First femur 4.2 mm, patella and tibia 6.3,
metatarsus 4.2, tarsus 1.3. Second patella
and tibia 4.5 mm, third 2.7, fourth 4.2.
Length of first patella and tibia 1.3 times
width of carapace.
Males are not known.
Variation. Total length of females 9.0—
12.8 mm. The total length of the holotype
is 12.8 mm, carapace 4.6 mm wide, patella
and tibia 5.8. Humps may have black caps.
The illustrations (Figs. 274—278) were
made from a female from Balcarce; Fig-
ures 279 and 280 are from the holotype.
Diagnosis. The female differs from M.
extraordinaria in that the carapace is not
shiny (Fig. 274), in having the anterior of
the abdomen dark (Figs. 276, 277), and in
having the slits on the posterior of the epi-
gynum almost parallel (Fig. 279). Masto-
phora melloleitaoi also lacks the dark me-
dian dorsal area on the posterior of the
epigynum present in M. extraordinaria
(Fig. 286). If cleared, each atrium shows a
small median lobe (Fig. 280) not present
in other species.
Distribution. From Parana State, south-
ern Brazil, to Buenos Aires Province, Ar-
gentina (Map 3E).
Specimens Examined. BRAZIL Rio de Janeiro: Vas-
souras, Fazenda de Sao Sebastiao, March 1871, 1?
(B. P. Mann, MCZ). Parana: Curitiba, 21 Jan. 1965,
1 imm. (C. Valle, MZSP 4326). Rio Grande do Sul:
Rodeio Bonito, Bage, 10 Feb. 1967, 1 9 (C. de Oliv-
eira, MCN 473); Santa Maria, 9 May 1973, 19 (D.
Link, MCN 01659). ARGENTINA Cordoba: Cruz
Alta, 15 Aug. 1948, 19 (J. R Duret, MACN). Bueno.s
Aires: Ascension, 19 (B. Gerschman, MACN); Bal-
carce, 16 Feb. 1950, 19 (Cuccioli, MACN); Burzaco,
19 (F C. S., MACN); San Isidro, 19 Apr. 1949, 19
(N. Konmilev, MACN); Zelaya, 2 imm. (J. Pereyra,
MACN 523).
Mastophora extraordinaria Holmberg
Figures 281-287, 460; Map 3F
Mastophora extraordinaria Hohnberg, 1876: 113. Fe-
male from Buenos Aires, Argentina, lost. Brethres,
1909: 163, figs. 1, 2, 9, egg sacs. Canals, 1931: 17,
figs. 1-5, pi. 1, fig. 1. Mello-Leitao, 1931: 70, figs.
7, 19. Roewer, 1942: 900. Platnick, 2001.
Glyptocranium extraordinaria: — Bonnet, 1957: 1997.
?MaMophora cinerea Mello-Leitao, 1943: 105, fig. 4,
imm. Iiumature holotype from Cordoba, Argenti-
na, in MLP examined. DOUBTFUL NEW SYN-
ONYMY.
Mastophora intermedia Mello-Leitao, 1945: 240, figs.
14-17, 9 . One female holotype from Pindapoy, Mi-
siones Prov., Argentina, in MLP, examined. Brig-
noh, 1983: 274. NEW SYNONYMY.
Note. Holmberg described the size of
the prominences of the abdomen and the
two brown spots.
Mastophora cinerea is a light-colored
immature with slightly elongate humps. Its
placement is doubtful. (Do immature M.
extraordinaria have longer humps?)
Mastophora intermedia has humps
rounded, as wide as long, and has the same
internal genitalia as does M. extraordinar-
ia. The differences are a slight median
notch of the posterior rim of the epigynum
in ventral view, and absence of the median
dark area posterior in the epigynum (Fig.
286).
Description. Feinale from Chascomus.
Carapace dark brown with narrow white
rim. Legs brown, slightly ringed. Abdomen
white with a pair of dorsal black patches
(Fig. 283). Carapace shiny on tips with low
tubercles and three grooves on each side.
Abdomen with few setae on sides with one
pair of humps; venter with white square.
Total length 12 mm. Carapace 4.8 mm
long, 4.6 wide in thoracic region, 2.8 wide
at posterior lateral eyes. First femur 3.9
mtn, patella and tibia 5.5, metatarsus 3.5,
tarsus 1.2. Second patella and tibia 4.5
mm, third 2.7, fourth 4.1. Length of first
patella and tibia 1.1 times \\ddth of cara-
pace.
Males are not known.
358 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Variation. The epigyiium is variable in
width and die abdominal humps may be
wider or higher. Total length of females
9.5—14 mm. The illustrations were made
from the female from Chascomus.
Diagnosis. Compared with M. mellolei-
taoi, the carapace has low, flat tubercles;
the horns of the carapace are relatively
smooth and shiny (Figs. 281, 282); and the
bulge bearing the median eyes is indis-
tinct. The anterior of the abdomen is usu-
ally white (Fig. 284). The median plate of
the epigynuin is flat dorsally, sclerotized,
and darker than ventrally (above on Fig.
286); it is raised in the middle in M. nieJ-
loleitaoi. The atria approach each other
(Fig. 287), unlike those of M. nielloleitaoi
(Fig. 280).
Natural History. The egg sac is drop-
shaped, 11 mm wide, almost the size of
the female abdomen (Fig. 460). A female
from Gonzales Catan was found on a citrus
tree. An immature in Uruguay was col-
lected at night "from regular web."
Distribution. From Rio Grande do Sul
State, southern Brazil, to Buenos Aires
Pro\4nce, west to Chaco and Cordoba
provinces, Argentina (Map 3F).
Specinwns Examined. BRAZIL Rio Grande de Sul:
Canela, 12 Feb. 1966, 1? (A. Lise, MCN 0119); Gar-
ibaldi, 30 Oct. 1974, 1$ (O. Simonaggio, MCN
2381). URUGUAY Montevideo, 1$ (J. Canosa,
MACN 4182); 1? (E. Cordero, MNRJ 14011); Pun-
tas Arroyo Laureles, Tacuarembo, 1 imm. (FCMU
293); Treinta y Tres, 20 Aug. 1971, 1? (FCMU 295).
ARGENTINA Chaco: Saenz Pena, Sept. 1933, 1?
(B. Ohneiser, MACN 31331). Cordoba: Cordoba, 29
(M. J. Viana. AMNH, MACN 1106); Calamuchita.
Dec. 1940, 4? (J. M. Viana, MACN 1005); Dec.
1941, 29 (J. M. Viana, MACN 1106); Agus do Oro,
Mar. 1940, 1 imm. (J. A. De Carlo). Buenos Aire.s:
Buenos Aires, 19 (Scly, MNHN 23388); Chascomus,
Oct. 1934, 19 (I. Dor, MACN 35983); Cap. Federal,
1 imm. (E. Pizarro, MACN 12782); Florencio Varela,
1 May 1949, 19 (O. de Ferrarini, MLP 13549); Gon-
zales Catan, 6 June 1949, 1 9 (Touson); Hurlingham,
Jan. 1954, 19 (Giai, MACN 4359); La Plata, 19 (M.
Biraben, MLP 16178); 19 (M. Biraben, BMNH);
Moreno, Feb. 1939, 19 (Schiapelli, Gerschman,
MACN); 14 Nov. 1943, 19 (S. M. Doello Jurado,
MACN 1361); Rosas, 1930, 19 (J. B. Daguerre,
MACN 4185).
Mastophora leucabulba (Gertsch),
new combination
Figures 288-295; IVIap 2F
Agatostichus leucabulba Gertsch, 1955: 250: figs. 34,
38, 40, 9 . Female holotype from Harlingen, Texas,
in AMNH, destroyed.
Agathostichus leucabulba: — Brignoli, 198.3: 255.
Agathostichus leucabulbus: — Platnick, 2001.
Note. Platnick's change of spelling is not
required.
Description. Female (after Gertsch,
1955). Carapace reddish brown except for
yellow rim and orange median patch en-
closing tubercles; tubercles tipped white.
Chelicerae brownish at base. Labium, en-
dites yellowish. Sternum yellowish. Coxae
yellowish and distal leg articles yellowish
to light brown with faint brown markings
on first leg. Abdomen yellowish to whitish
above with dusky flecks, black around the
base (Fig. 289). Carapace with woolly se-
tae, one large bulb behind median eyes, a
large median bulb flanked by a pair of
small bulbs on the side, behind bifid
horns, sides with dark warts and cephalic
area coarsely roughened, hidden by mat of
wooly hairs. Lateral eyes on connate tu-
bercles, median eyes on large elevated tu-
bercle at the posterior edge of which is a
small tubercle. Abdomen with long
humps. Total length 6.7 mm. Carapace 2.8
mm long, 2.9 wide in thoracic region. First
femur 3.6 mm, patella and tibia 4.9, meta-
tarsus 3.6, tarsus 1.0. Second patella and
tibia 3.4 mm, third 1.9, fourth 2.6. Length
of first patella and tibia 1.7 tinies width of
carapace.
Diagnosis. Mastophora leucabulba is
distinguished by the enlarged tubercle be-
tween the median eyes (Figs. 288, 290)
and the large blunt tubercles and dark col-
oration of the carapace (Figs. 289, 290).
Doubtful males, considered and labeled
M. cornigera by W. Ivie, and listed under
that species by Gertsch, may be this spe-
cies (Figs. 292-295).
Distribution. Southern Texas to Hon-
duras (Map 2F).
Specimens Examined. TEXAS Duval Co.: San Di-
Mastophora • Levi
359
Figures 288-295. Mastophora leucabulba Gertsch. 288-291 , female, after Gertschi. 288, carapace and ctielicerae, lateral. 289,
290, carapace and abdomen. 289, dorsal. 290, lateral. 291, epigynum, posterior. 292-295, left palpus of presumed male. 292,
apical. 293, mesal. 294, ventral. 295, ectal.
Figures 296-307. M. alvareztoroi Ibarra and Jimenez new species. 296-303, female. 296, 297, carapace and chelicerae. 296,
frontal. 297, lateral. 298-300, carapace and abdomen. 298, dorsal. 299, ventral. 300, lateral. 301-303, epigynum. 301, ventral.
302, posterior. 303, posterior, cleared. 304-307, male. 304, carapace, chelicera and righit palpus. 305, dorsal. 306-307, left
palpus. 306, mesal. 307, ventral.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
360
Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
ego, 29 Apr. 1895, 2 c?, imm. doubtful determination
(USNM). Wilson Co.: Floresville, 29 Apr. 1895, 1
imm., 46, doubtful determination (AMNH). MEXI-
CO Tamaulipas: 64 km S Linares, imm., 1.9 mm long
(Gertsch, 1955, AMNH, destroyed, not examined).
HONDURAS E of Tela Beach, 6 July 1929, imm. (A.
M. Chickering, MCZ).
Mastophora alvareztoroi
Ibarra and Jimenez new species
Plate 1; Figures 296-307; Map 2F
Holotijpe. Female holotype from Rancho Alejandria,
Municipio Estacion Juarez, Chiapas, Mexico, 25
Sept. 1975 (M. Alvarez del Toro), in MCZ. The
species has been named after the collector, the late
Miguel Alvarez del Toro, who dedicated his life to
the study and protection of the Chiapas fauna and
is the author of a book on Chiapas spiders (Alvarez
del Toro, 1992).
Agathostichus sp. Alvarez del Toro, 1992: 17.3, fig.
121, 9, photo.
Description. Female from Rosaria Iza-
pa. Carapace horns white, sides dark
brown with eyes, clypeus, and rim yellow-
ish (Figs. 296-298). Chelicerae yellowish
with a pair of dark patches. Labium, en-
dites dark brown, distally light. Sternum
dark brown. Coxae and distal leg articles
yellowish; first femur \\4th fine black ring
and distally with ventral dark patch. Ab-
domen yellow, dark brown on anterior bor-
der, and brown on some hump tips (Fig.
298); venter light brown with white
square; a dark brown patch on epigynal
area, and no pigment in spinneret area.
Carapace hirsute with cone-shaped white
tubercles (Figs. 296, 297). Posterior me-
dian eyes 0.8 diameter of anterior medi-
ans, laterals 0.7. Anterior median eyes 1.7
diameters apart. Ocular trapezoid rectan-
gular, slightly wider than long. Chelicei"ae
with one anterior tooth and three posterior
teeth. Abdomen heart-shaped, three pairs
of dark spots dorsally and sparse black se-
tae, hirsute with tufts on white setae on
dorsum. A pair of dorsal humps, followed
by a series of four median, conical humps
and several smaller lateral humps on each
side. Setae, also tufts of white setae on
small circular and convex white areas, scat-
tered on dorsum of abdomen, with scat-
tered dark brown setae, especially on an-
terior half, also with a few feather-shaped
symmetrically arranged dark brown setae
(Fig. 298). Total length 8.9 mm. Carapace
3.4 mm long, 3.3 wide in thoracic region,
1.7 wide at lateral eyes. First femur 4.3
mm, patella and tibia 5.5, metatarsus 4.2,
tarsus 1.1. Second patella and tibia 3.8
mm, third 2.3, fourth 3.5. Length of first
patella and tibia 1.6 times width of cara-
pace.
Male weakly sclerotized, perhaps just
molted. Carapace reddish brown except
for white median light patch enclosing
white horns. Chelicerae dark yellow. Ster-
num dark brown. Coxae, distal leg articles
yellowish. Abdomen whitish. Carapace el-
evated behind, second midline tubercle
slightly longer than first, lateral horns
about half length of median horns, sepa-
ration between median and lateral horns
greater than width of laterals at base; each
transverse horn with translucent seta on
tip; carapace with woolly white setae. Che-
licerae with three posterior median teeth
and one posterior tooth. Abdomen with
two dorsal paired humps; four anterior,
four median, and two posterior feather-
shaped setae (Fig. 305). Total length 1.7
mm. Carapace 0.8 mm long, 0.8 wide in
thoracic region. First femur 1.1 mm, pa-
tella and tibia 1.1, metatarsus 1.1, tarsus
1.03. Second patella and tibia 0.8 mm,
third 0.4, fourth 0.6. Length of first patella
and tibia 1.4 times width of carapace.
Note. Males and females were matched
Figures 308-314. Mastophora soberiana new species, female. 308, 309, carapace and chelicerae. 308, frontal. 309, lateral.
310, 311, carapace and abdomen. 310, dorsal. 311, lateral. 312-314, epigynum. 312, ventral. 313, posterior. 314, posterior,
cleared.
Figures 315-321. Mastophora leucacantha (Simon), female. 315, 316, carapace and chelicerae. 315, frontal. 316, lateral. 317,
318, carapace and abdomen. 317, dorsal. 318, lateral. 319-321, epigynum. 319, ventral. 320, posterior. 321, posterior, cleared.
Mastophora • Levi 361
Figures 322-328. M. brescoviti new species, female. 322, 323, carapace and chelicerae. 322, frontal. 323, lateral. 324, 325,
carapace and abdomen. 324, dorsal. 325, lateral. 326-328, epigynum. 326, ventral. 327, posterior. 328, posterior, cleared.
Figures 329-335. M. conifera (Holmberg), female. 329, 330, carapace and chelicerae. 329, frontal. 330, lateral. 331, 332,
carapace and abdomen. 331, dorsal. 332 lateral. 333-335, epigynum. 333, ventral. 334, posterior. 335, posterior, cleared.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
362 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
because they were collected in the same
area in Chiapas, have similar cone-shaped
carapace tubercles and horns, and both
sexes have feather-shaped setae.
Variation. Living females look all woolly.
Total length of females 8.6—8.9 mm, cara-
pace 3.3-4.0 mm wide. An immature, 4.3
mm total length, is similar to the adult fe-
male in relative proportions, horns, and
pointed abdomen, but has fewer abdomi-
nal humps (only the pair and the four in
midline). As in males of the genus, a row
of long setae is present on the anterior of
the first and second metatarsi and tarsi.
The illustrations (Figs. 298-300) were
made from the paratype collected with the
holotype.
Diagnosis. Mastophora alvareztoroi is
distinguished from M. leucahulha by the
shape and humps of abdomen (Figs. 298—
300), by the cone-shaped tubercles on the
carapace (Figs. 296, 297), by having feath-
er-shaped setae (Figs. 298, 305), and by
the epigynum having atria bent toward
each other (Figs. 302, 303).
Natural History. Some specimens came
from coffee groves in Finca Irlanda and
Rosario Izapa. The specimens were be-
tween 1 and 2 m above the ground. Mark
Stowe (in correspondence) wrote that this
fuzzy spider bears a strong resemblance to
congregations of fuzzy scale insects in the
same habitat in Texas.
Distribution. Southern Texas and Chia-
pas, Mexico (Map 2F).
Paratypes. MEXICO Chiapas: Rancho Alejandria,
Municipio Estacion Juarez, 25 Sept. 1975, 1$ (M.
Alvarez del Toro, MLJ); Finca Irlanda, 870 m,
15°10'N, 92°21'W, 65 km NNW of Tapachula, 12
Aug. 1987, Id (G. Ibarra, A. Garcia, M. Moreno,
ECOTAR); Rosario Izapa, Municipio Tuxtla Chico,
430 m, 14°59'N, 92°09'W, 18 km ENE of Tapachula,
25 Aug. 1995, 1? (A. Ventura, ECOTAR).
Specimens Examined. TEXAS Hidalgo Co.: Santa
Ana Natl. Wildlife Refuge, 18 Dec. 1983, 19 (M. K.
Stowe 112, MKS). MEXICO Chiapas: Finca Ham-
burgo, ca. 15°10'N, 92°19'W, 950 m, Municipio de
Tapachula, 16 Nov. 1994, 1 imm. (G. Ibarra, ECO-
TAR).
Mastophora soberiana new species
Figures 308-314; Map 2F
Holotype. Female holotype from Pipeline Road, Ca-
nal Zone [in Soberiana National Park, Panama], 26
July 1976 (Y. Lubin, G. G. Montgomery), in MCZ.
The specific name is a noun in apposition after die
locality.
Description. Female holotype (in poor
physical condition). Carapace yellowish,
sides of thorax dark brown, a wide, light
rim and tubercles with white pigment
(Figs. 308, 309). Chelicerae, labium, en-
dites yellowish. Sternum brown. Coxae,
distal leg articles yellowish. Abdomen
white dorsally (Fig. 310), white ventrally,
except for yellowish genital area and spin-
nerets. Carapace with spine-shaped tuber-
cles and white setae (Figs. 308, 309). Ab-
domen with three median humps and nu-
merous small lateral humps, with tufts of
white setae (Figs. 310, 311). Total length
8.3 mm. Carapace 3.7 mm long, 3.4 wide
in thoracic region, 1.8 wide behind pos-
terior lateral eyes. First femur 4.7 mm, pa-
tella and tibia 5.8, tarsi lost. Second patella
and tibia 4.2 mm, third 2.3, fourth 3.5.
Length of first patella and tibia 1.7 times
width of carapace.
Males are not knowii.
Diagnosis. This species is distinguished
from M. alvareztoroi by the longer cara-
pace tubercles and by having the atria of
the epigynum located dorsally (Fig. 314).
Distribution. Panama (Map 2F).
Specimens Examined. No other specimens have
been found.
Mastophora leucacantha (Simon),
new combination
Figures 315-321; Map 3G
Agatostichus leucacantha Simon, 1895: 885, fig. 947,
carapace. Immature holotype from Rio Salobro,
Baliia, Brazil, in MNHN, 8486, examined. Mello-
Leitao, 1931: 67. Gertsch, 1955: 250.
Agatho St i chits leucacantha: — Simon, 1897: 473.
Roewer, 1942: 900.
Agathostichus leiicacanthus: — Bonnet, 1955: 182.
Platnick, 2001,
Note. In his publications, Simon (1895,
1897) did not include the female symbol
Mastophora • Levi
363
as he did for other species, indicating that
the specimen was immature. Roewer
(1942) and Gertsch (1955) cited it as fe-
male.
Platnick (2001) considered the name of
Simon (1895) to be a nomen nudum be-
cause the description was shared by the
genus and species. But this is vaHd for
19th century descriptions (International
Code of Zoological Nomenclature, art.
12.2.6 [International Commission on Zoo-
logical Nomenclature, 1999]).
Description. Female from the Organ
Mountains. Carapace yellowish white with
white marks and median white tubercles,
brown triangle on posterior slope; anterior
point of triangle dark between forked tu-
bercles (Fig. 315). Chelicerae yellowish.
Labium dark brown, endites yellowish.
Sternum dark brown. Coxae and distal leg
articles yellowish white. Abdomen yellow-
ish white with a small black mark on each
side (Fig. 317); venter yellowish white with
white square surrounded by a gray line;
dorsum and sides with indistinct gray
marks. Carapace with woolly setae and
long tubercles (Figs. 315-317). Median
eyes on a swelling, each lateral pair on a
swelling. Median ocular trapezoid almost
square. Chelicerae with three anterior
teeth, one posterior tooth. Legs with white
setae. Abdomen with a pair of dorsal
humps and tufts of white setae (Figs. 317,
318). Total length 8.3 mm. Carapace 3.7
mm long, 3.6 wide in thoracic region, 2.2
wide at lateral eyes. First femur 3.6 mm,
patella and tibia 4.6, metatarsus 3.3, tarsus
1.0. Second patella and tibia 3.6 mm, third
2.1, fourth 3.2. Length of first patella and
tibia 1.3 times width of carapace.
Males are not known.
Variation. The immature holotype, total
length 4.0 m, has carapace tubercles short-
er and lacks some lateral ones; venter of
abdomen with black square. The illustra-
tions were made from the adult female
from Organ Mountains. The tufts of setae
on the abdomen were prominent on the
female from Organ Mountains when first
examined (by the author in 1969), but
have mostly been lost as result of handling.
Diagnosis. Mastophora leucacantha is
distinguished by the long median horn that
is almost as long as the carapace, and dif-
fers from M. alvareztoroi by having the ab-
domen rounded behind, whereas M. al-
vareztoroi has the abdomen lobed behind,
and differs from M. leucabulba by having
all tubercles behind the eyes and from M.
soberiana by the shape of the epigynum
(Fig. 320).
Distribution. Bahia to Rio de Janeiro
states, Brazil (Map 3G).
Specimens Examined. Rio de Janeiro: Cachoeirin-
ha, Montaigne Orgues [Serra Orgaos, Organ Moun-
tains], 1902, 1? (E. R. Wagner, MNHN 26035).
Mastophora brescoviti new species
Figures 322-328; Map 3G
Holotype. Female holotype from Jardim Botanico,
Porto Alegre, Rio Grande do Sul, Brazil (A. D.
Brescovit), in MCN no. 26135. The species has
been named after the collector and arachnologist
A. D. Brescovit.
Description. Female holotype. Carapace
with symmetrical white lines on head re-
gion, sides of cephalic area dark brown,
sides of thorax light browai with dark
streaks and speckles, and many downy
white setae (Figs. 322, 323). Chelicerae
patchy brown. Labium, endites, sternum
dark brown. Coxae light brown and distal
leg articles yellowish with narrow brown
rings. Abdomen light browii, darker ante-
riorly between humps and pedicel, darker
patches on each side, humps darkest, with
bunches and individual white setae (Fig.
324); venter light brown. Lateral eyes on
bulges. Carapace with median tubercle
longest. Abdomen dorsally with a pair of
humps and a median swelling bearing
white setae (Figs. 324, 325). Total length
9.2 mm. Carapace 3.7 mm long, 3.4 wide
in thoracic region, 1.8 wide behind pos-
terior lateral eyes. First femur 4.0 mm, pa-
tella and tibia 5.1, metatarsus 3.7, tarsus
1.2. Second patella and tibia 3.8 mm, third
2.3, fourth 3.3.
Males are not known.
364 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Diagnosis. Mastophora hrescoviti is dis-
tinguished by the long posterior, median
tubercle, and unusual shape of the abdo-
men with a median swelling behind the
humps (Figs. 324, 325). The posterior of
the epigynum has a pair of diagonal swell-
ings (Fig. 327) not present in other spe-
cies.
Distribution. Only known from Porto
Alegre, Brazil (Map 3G).
Specimens Examined. No other specimens have
been found.
Mastophora conifera (Holmberg)
Figures 329-335; Map 3G
Heterocephala conifera Hohnberg, 1876: 143. Female
from Boradero [Prov. Buenos Aires], Argentina,
lost.
Mastophora conifera: — Canals, 1931: 18, figs. 1-5, pi.
1, fig. 2. Mello-Leitao, 1931: 71, figs. 4, 16. Roewer,
1942: 900. Platnick, 2001.
Glyptocranium coniferum: — Bonnet, 1957: 1996.
Description. Female from Tigre [in poor
condition]. Carapace, sternum, legs or-
ange-brown. Abdomen anteriorly black,
posteriorly lighter gray, with some black
streaks (Figs. 331, 332); venter black with
a pair of white spots. Carapace with tips
of horns thin, setae on sides of thoracic
area (Figs. 329, 330). Anterior median
eyes largest, laterals smallest. Abdomen
with numerous dorsal tubercles (Figs. 331,
332). Total length 12.0 mm. Carapace 5.3
mm long, 4.6 wide in thoracic region, 2.8
wide at lateral eyes. First femur 4.3 mm,
patella and tibia 5.3, metatarsus 3.8, tarsus
broken. Second patella and tibia 4.3 mm,
third 2.4, fourth 3.7. Length of first patella
and tibia equals 1.2 times width of cara-
pace.
Males are not known.
Diagnosis. Mastophora conifera is dis-
tinguished by the tubercular abdomen
(Figs. 331, 332) and by the epigynum,
which in posterior view has a pair of de-
pressions containing short, ventrally con-
verging slits (Fig. 334).
Distribution. Santa Fe and Buenos Ai-
res provinces, Argentina (Map 3G).
Speciinens Examined. ARGENTINA Santa Fe: Co-
lonia Macias, Nov. 1942, imm. shriveled (J. M. Viana,
MACN). Buenos Aires: Tigre, 1902, 1$, once dried
up (J. Brethes, MACN 5896).
Mastophora corpulenta (Banks)
Figures 336-342, 461 ; IVIaps 2F, 4A
Ordgarius coijiulentus Banks, 1898: 251, pi. 15, fig.
8. Female holotype from San Jose del Cabom Baja
California, Mexico, in CAS, destroyed. Neotype
here designated the holotype of M. lenca.
Mastophora corpulenta: — Roewer, 1942: 900. Plat-
nick, 2001.
Mastophora lenca Gertsch, 1955: 247, figs. 28-30, 32,
33, 9 . Female holotype from Zamorano [PZambra-
no], Honduras, in AMNH, examined. Brignoli,
1983: 274. Platnick, 2001. NEW SYNONYMY.
Glyprocraniurn corpulentum: — Bonnet, 1957: 1997.
Note. Banks (1898) described and pic-
tured (fig. 8) elongated tubercles on the
abdomen and lateral tubercles on the side
of the carapace. The only North American
or Central American species known to
have both these characters is M. lenca. The
type locality of Ordgarius corpulentus is
uncertain, because, as Banks himself
pointed out, the collection was handled by
C. Marx before being turned over to
Banks after Marxs death, and Mai"x locality
labels are confused.
Description. Female holotype. Carapace
dark brown, with short white setae not
covering tubercles; tubercles with light
tips (Figs. 336, 337). Sternum dark orange.
Legs dark brown. Median eyes on bulge,
lateral eyes on bulges. Abdomen gray, with
long humps (Fig. 339); venter with white
square. First tarsus with S-shaped curva-
Figures 336-342. Mastophora corpulenta (Banks), female. 336, 337, carapace and chelicerae. 336, frontal. 337, lateral. 338,
339, carapace and abdomen. 338, dorsal. 339, lateral. 340-342, epigynum. 340, ventral. 341, posterior. 342, posterior, cleared.
Figures 343-350. M. rabida new species, female. 343, 344, carapace and chelicerae. 343, frontal. 344, lateral. 345, 346,
carapace and abdomen. 345, dorsal. 346, lateral. 347-350, epigynum. 347, ventral. 348, posterior. 349, posterior, cleared. 350,
seminal receptacle, median.
Mastophora • Levi
365
352
Figures 351-361, M. escomeli new species. 351-357, female. 351, 352, carapace and chelicerae. 351, frontal. 352, lateral.
353, 354, carapace and abdomen. 353, dorsal. 354, lateral. 355-357, epigynum. 355, ventral. 356, posterior. 357, posterior,
cleared. 358-360, male left palpus, stained. 358, apical. 359, mesal. 360, ventral. 361, ectal.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
366 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
ture. Total length 11.0 mm. Carapace 5.4
mm long, 5.2 wide in thoracic region, 2.8
wide behind posterior lateral eyes. First
femur 4.7 mm, patella and tibia 7.6, meta-
tarsus 5.4, tarsus 1.4. Second patella and
tibia 4.8 mm, third 2.4, fourth 4.1.
Males are not known.
Diagyiosis. Mastophora corpulenta is
distinguished from M. diablo by the small
dorsal knobs in the depression of the pos-
terior of the epigynum (Fig. 341).
The egg sac is fig-shaped, lacks lateral
flaps, and has a thick stalk (Fig. 461).
Distribution. Central America (Map
4A).
Specimens Examined. NICARAGUA Leon, Aban-
gasca, 13 Dec. 1994, 1 subadult 9 (J. M. Maes,
MCZ).
Mastophora rabida new species
Figures 343-350; Map 4A
Holotype. Female holotype and immature female
paratype from Rabida Island, Galapagos Islands,
Ecuador, 12 May 1981 (Y. D. Lubin, 319), in MCZ.
The specific name is a noun in apposition after the
locality
Note. The female holotype is a penulti-
mate instar, ready to molt. The exuvium is
loose above the epigynum. The epigynum
is mature but not sclerotized.
Description. Female holotype. Carapace
dark orange. Chelicerae, labium, endites
orange. Sternum orange with white pig-
ment. Coxae and distal leg articles dark
dusky orange. Abdomen white with some
faint gray marks (Fig. 345); venter with
white square. Carapace appearing downy,
covered with tubercles; tubercles with
hght tips (Figs. 343, 344). Median eyes on
bulge, lateral eyes on bulges. Abdomen
slightly wider than long with small humps
(Figs. 345, 346). Total length 7.7 mm. Car-
apace 3.1 mm long, 2.9 wide in thoracic
region, 1.8 wide at lateral eyes. First femur
2.8 mm, patella and tibia 4.2, metatarsus
3.2, tarsus 1.0. Second patella and tibia 3.0
mm, third 1.7, fourth 2.8. Length of first
patella and tibia 1.4 times width of cara-
pace.
Males are not known.
Diagnosis. Mastophora rabida is distin-
guished by the ventral loops of the slits on
the posterior of the epigynum (Figs. 348,
349).
Natural Historij. From Y. Lubin (per-
sonal correspondence): "#319. nocturnal
araneid on orb web. During day sits on
twigs. 1 female, 1 juvenile. [NB: niaybe it
was the juvenile on an orb web? I didn't
specify in the notes. YL]. #510. Tagus
Cove, Isabella [This is a mangrove area]."
Notes from field book 19: 30: "on Croton
bush, hanging from thread with legs 3,4.
Legs 1,2 held outwards, flexed. No bolas.
Spider stretches legs forward when I hum,
then moves to edge of leaf and adopts
same posture."
Distribution. Galapagos Islands (Map
4A).
Paratypes. Galapagos Islands: Isabella Island, Ta-
gus Cove, on Croton scoideri at night, 13 May 1983,
1 imm. (Y. Lubin 510, MCZ).
Mastophora escomeli new species
Figures 351-361; Map 4A
Glyptocraniuni gasteracanthoides: — Escomel, 1918,
136 (misidentification).
Holotype. Female holotype from Valle de Majes, nr.
Arequipa, Depto. Arequipa, Peru, 1920 (E. Esco-
mel), in MNHN. The species has been named after
the collector and author of a paper on tlie venoms
of Mastophora.
Note. A similar specimen examined by
Figures 362-365. Mastophora obtusa Mello-Leitao, immature female. 362, 363, carapace and chelicerae. 362, frontal. 363,
lateral, 364, 365, carapace and abdomen. 364, dorsal. 365, lateral.
Figures 366-372. M. felis Piza, female. 366, 367, carapace and chelicerae. 366, frontal. 367, lateral. 368, 369, carapace and
abdomen. 368, dorsal. 369, lateral. 370-372, epigynum. 370, ventral. 371, posterior. 372, posterior, cleared.
Figures 373-379. M. holmbergi Canals, female. 373, 374, carapace and chelicerae. 373, frontal. 374, lateral. 375, 376, cara-
pace and abdomen. 375, dorsal. 376, lateral. 377-379, epigynum. 377, ventral. 378, posterior. 379, posterior, cleared.
Mastophora • Levi 367
^^\K^'-^^'^'',-~r^v 366 ^^"^
Figures 380-386. M. reimoseri new species, female. 380, 381 , carapace and chelicerae 380, frontal. 381 , lateral. 382, 383,
carapace and abdomen. 382, dorsal. 383, lateral. 384-386, epigynum 384, ventral 385, posterior. 386, posterior, cleared.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
368 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Canals (MACN) was labeled M. satan,
presumably because of its long first legs.
Description. Female holotype. Carapace
orange-brown. Chelicerae brown. Labium,
endites, sternum orange. Coxae orange,
distal leg articles brown. Abdomen brown-
ish, underlain by some white patches
(Figs. 353, 354); venter with white square.
Carapace with many large tubercles, tu-
bercles on sides; tubercles with light tips,
with short white setae between, but not
covering tubercles, and longer white setae
on sides (Figs. 351, 352). Median eyes on
a bulge; lateral eyes on bulges. First tarsus
with S-shaped cuivature. Abdomen with
long humps (Figs. 353, 354). Total length
13.5 mm. Carapace 5.0 mm long, 5.0 wide
in thoracic region, 3.0 wide at lateral eyes.
First femur 4.6 mm, patella and tibia 7.7,
metatarsus 6.3, tarsus 1.6. Second patella
and tibia 4.8 mm, third 2.7, fourth 4.3.
Length of first patella and tibia 1.5 times
width of carapace.
Male of uncertain affiliation. Color and
shape as in other males. Total length 1.7
mm. Carapace 0.83 mm long, 0.81 wide in
thoracic region, 0.54 wide at lateral eyes.
First femur 0.72 mm, patella and tibia
0.81, metatarsus 0.49, tarsus 0.13. Second
patella and tibia 0.71 mm, third 0.39,
fourth 0.54. Length of first patella and tib-
ia 1.0 times width of carapace.
Note. Affiliation of males with females is
uncertain.
Variation. Total length of females 12.0—
13.5 mm. Length of first patella and tibia
1.5—1.7 times width of carapace. The fe-
male from lea is more setose, with long
setae on the legs and many shorter setae
on the abdomen. The depressions of the
female epigynum are larger and the bulge
between the slits is less distinct. The illus-
trations were made from the female ho-
lotype.
Diagnosis. Mastophora esconieli is dis-
tinguished from M. gasteracanthoides by
having a swelling between the slits on the
posterior of the epigynum (Fig. 356) and
having the atrium ventral to the seminal
receptacles (Fig. 357). In M. gasteracan-
thoides, the slits are in a shared depression
and the atria are dorsal to the seminal re-
ceptacles (Figs. 415, 416).
Natural Historij. Specimens were found
on grapevines near Arequipa, and were
known to readily bite grape workers as
they pruned the plants, causing necrotic
lesions.
Distribution. Dry coastal region of Peru
(Map 4A).
Paratypes. PERU Lima: Lomas de Lachay, 26 May
1996, 19 (N. Llerana Martinez, MUSM). Arequipa:
Arequipa, 1912, 29 (E. Escobal, MNHN); 19 (E.
Escobal, MACN 4198).
Specimens Examined. PERU lea: lea, 1992, 1 9
(Cascavilca-Rubio, MACN). La Libertad: Cerro
Campana, N Trujillo, 23 May 1989, many c?, imm.
(A. Salas, MUSM).
Mastophora obtusa Mello-Leitao
Figures 362-365; Map 4C
Mastophora obtusa Mello-Leitao, 1936; 134, fig. 2,
imm. Immature holotype from Pesqueira, Pernam-
buco, Brazil, in MNRJ, 41845, examined. Roewer,
1942: 955. Platnick, 2001.
Glyptocranium obtusuin: — Bonnet, 1957: 1998.
Note. Pesqueira is located in the note
with the description of M. pickeli.
Description. Female holotype. Carapace
reddish brown with white rim. Chelicerae,
labium, endites brown. Sternum light
brown. Coxae and distal leg articles brown.
Abdomen very light brown; venter with
white square. Carapace with short white
setae on sides (Figs. 363, 364). Abdomen
high with a pair of humps. Total length 4.8
mm. Carapace 1.8 mm long, 1.7 wide in
thoracic region, 1.2 wide at lateral eyes.
First femur 1.6 mm, patella and tibia 2.3,
metatarsus 1.3, tarsus 0.5. Second patella
and tibia 1.7 mm, third 0.8, fourth 1.8.
Length of first patella and tibia 1.4 times
width of carapace.
Diagnosis. Although the type is imma-
ture, M. obtusa is distinguished from many
other species by the high abdomen and
the humps on a joined swelling (Fig. 364).
The high sides of the carapace, the shape
of the tubercles, and the lack of pigment
pattern on the abdomen suggest that the
species belongs to the M. gasteracanthoi-
Mastophora • Levi
369
des group of species. Perhaps this is an im-
mature of M. satan.
Specimens Examined. No other specimens have
been found.
Mastophora felis Piza
Figures 366-372; Map 4C
Mastophora felis Piza, 1976: 83, fig. 1. Female holo-
type from Piracicaba, Sao Paulo, Brazil, in MZAQ
no. A0105, examined. Brignoli, 1983: 273. Platnick,
2001.
Note. The holotype was embedded in
difficult-to-remove fungal mycehum.
Description. Female holotype. Carapace
dark brown with tips of tubercles light and
a thin white rim, each posterior median
eye in a light patch. Chelicerae brown. La-
bium, endites brown. Sternum browii.
Coxae and distal leg articles orange-brown.
Abdomen brownish gray (Figs. 368, 369);
venter with a median white square. Cara-
pace with long tubercles, the median of
the horn's base with multiple tubercles
(Figs. 366, 367), covered with short white
setae between tubercles. Median and lat-
eral eyes on bulges. Legs with white setae.
Abdomen with a pair of long dorsal humps
(Fig. 369). Total length 13.0 mm. Cara-
pace 6.3 mm long, 6.4 wide in thoracic re-
gion, 3.8 wide at lateral eyes. First femur
5.8 mm, patella and tibia 10.6, metatarsus
8.8, tarsus 2.3. Second patella and tibia 6.7
mm, third 3.6, fourth 5.5. Length of first
patella and tibia 1.6 times width of cara-
pace.
Males are not known.
Variation. Total length of females 11.3-
13.0 mm. Length of first patella and tibia
1.4—1.6 times width of carapace. The illus-
trations were made from the female ho-
lotype.
Diagnosis. Mastophora felis is distin-
guished from all others ha\ang long wide
humps and carapace with tubercles on
sides and by having the atria of the epi-
gynum ventral to the seminal receptacles
(at 11 and 1 h in Fig. 372) and the slits
with a lateral lip (Fig. 371). The carapace
tubercles are longer than those of M. satan
and the posterior eyes are on light patches.
Distribution. Rio de Janeiro and Sao
Paulo states, Brazil (Map 4C).
Paratijpes. BRAZIL Rio de Janeiro: Santo Antonio,
Rio Bonito [22°42'S, 42°37'W], 1933, 1? (S. Reme-
tente, IBSP 418). Sao Paulo: PCampinas, July 1982,
1? (C. Froelich, IBSP 4968).
Mastophora holmbergi Canals
Figures 373-379; IVIap 4C
Mastophora Holmbergi Canals, 1931: 22, figs. 1-5, pi.
3, fig. 5; pi. 4, figs. 7, 8, ? . Female from km 701,
Santiago del Estero, in MACN, 24133 [7140], ex-
ainined.
Mastophora holmbergi: — Mello-Leitao, 1931, 7.3, figs.
10, 21, ?. Roewer, 1942, 900. Platnick, 2001.
Glijptocranium holmbergi: — Bonnet, 1957, 1997.
Description. Female holotype. Carapace
dark red-brown to black, light transverse
band in front of posterior median eyes.
Sternum orange. Legs orange-brown. Ab-
domen with faint pattern (Figs. 375); ven-
ter with white square. Carapace with many
tubercles with light tips especially on sides,
and a few downy setae; lateral eyes on
bulges (Figs. 373, 374). First tarsus slightly
S-shaped. Abdomen with narrow humps.
Total length 11.0 mm. Carapace 5.2 mm
long, 5.3 wide in thoracic region, 3.0 wide
at lateral eyes. First femur 5.7 mm, patella
and tibia 10.0, metatarsus 9.2, tarsus 2.0.
Second patella and tibia 6.2 mm, third 3.2,
fourth 5.0. Length of first patella and tibia
1.9 times width of carapace.
Males are not known.
Variation. Total length of females 11.0—
15.3 mm. The illustrations were made
from the female holotype.
Diagnosis. Unlike Mastophora reimo-
seri, M. hohnbergi has long first legs. In
the epigynuin the atria are visible in ven-
tral view (Fig. 377) and the slits are par-
allel, but at their ventral ends the slits
bend toward each other (Fig. 378).
Distribution. Paraguay, to Santiago del
Estero, Argentina (Map 4C).
Specimens Examined. PARAGUAY Rca. del . . . [il-
legible], Nov. 1940, 1$ (Cranwell-Giai, MACN
1630).
370 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Mastophora reimoseri new species
Figures 380-386; Map 4C
Holotijpe. Female holotype from Asuncion, Paraguay,
1908 (E. Reimoser), in NHMW. The species has
been named after the collector and Austrian arach-
nologist, E. Reimoser.
Description. Female holotype. Speci-
men faded. Carapace yellow-brown. Che-
licerae, labium, endites yellow-brown.
Sternum, legs golden brown. Carapace
with many tubercles and short setae, clyp-
eus with longer setae (Figs. 380, 381). Me-
dian eyes and lateral eyes on bulges. First
femora bent at their distal ends with long
setae at point of greatest curvature. Ab-
domen with distinct humps and short se-
tae, base of huinps with longer setae (Figs.
382, 383). Total length 8.5 mm. Carapace
4.0 mm long, 4.0 wide in thoracic region,
2.3 wide at lateral eyes. First femur 3.7
mm, patella and tibia 6.2, metatarsus 4.5,
tarsus 1.2. Second patella and tibia 4.0
iTim, third 2.1, fourth 3.5. First patella and
tibia 1.6 times width of carapace.
Males are not known.
Diagnosis. The species is distinguished
by the musical note— shaped marks of the
slits in the epigynum (Fig. 385), and by
having shorter legs than M. hohnbergi.
Distribution. Known only froin Asun-
cion, Paraguay (Map 4C).
Specimens Examined. No other specimens have
been found.
Mastophora satan Canals
Figures 387-398, 462, 463; Map 4B
Mastophora satan Canals, 1931: 25, figs. 1-5, pi. 3,
fig. 6, 9 . Female holotype from La Rioja, Argen-
tina, in MACN, 5260, examined. Mello-Leitao,
1931: 73, figs. 11, 22. 9. Roewer, 1942: 901. Plat-
nick, 2001.
Glijptocranium satan: — Bonnet, 1957: 1998.
Description. Female from Cordoba.
Carapace dark brown with many short
light setae and narrow white riin. Sternum
red-brown. Legs red-brown with long
white setae. Abdomen whitish with dusk-
iness on humps and sides (Fig. 389); ven-
ter with white square. Carapace with many
tubercles, lateral eyes on bulges, median
eyes on bulge (Figs. 387, 388). Abdomen
humps veiy long (Figs. 389, 390). Total
length 14.0 mm. Carapace 6.4 mm long,
6.6 wide in thoracic region, 3.5 wide at lat-
eral eyes. First femur 7.0 mm, patella and
tibia 11.5, metatarsus 10.0, tarsus 2.2. Sec-
ond patella and tibia 7.0 mm, third 3.3,
fourth 5.7. Length of first patella and tibia
1.7 times width of carapace.
Male from Rio Grande do Sul, Brazil.
Carapace beige with a triangular white
patch. Sternum, legs beige. Abdomen
whitish. Carapace with four tubercles, ab-
domen with two humps. Total length 1.7
mm. Carapace 0.92 min long, 0.81 wdde in
thoracic region, 0.58 wide at lateral eyes.
First femur 0.87 mm, patella and tibia
0.92, metatarsus 0.49, tarsus 0.31. Second
patella and tibia 0.74 mm, third 0.44,
fourth 0.55.
Note. The association of male and fe-
male is uncertain. The male is placed with
the most common species in the area; also,
a female of the species has been collected
at the collecting site.
Variation. Total length of females 9.7-
17.5 mm. The holotype is 11.2 mm total
length, carapace 5.7 mm wide and long,
the first patella and tibia 9.7. Length of
first patella and tibia 1.4—1.6 tiines cara-
pace width in feinales from Brazil, 1.7 in
specimens from Uruguay, 1.5 from Men-
doza, 1.6 from La Pampa, 1.5 from Entre
Rios. The illustrations were made of a fe-
male from Cordoba Province, Argentina, a
female from La Rioja Province (Fig. 394),
a female from Tucuinan Province (Fig.
395), and of the male from Rio Grande do
Sul, Brazil.
Diagnosis. Mastophora satan is separat-
ed from M. gasteracanthoides and M. dia-
blo by the long first tibia and metatarsus,
each 9.7 inm or longer. The epigynum dif-
fers from that of M. diablo by showing the
atria as a dark patch in the dorsal slope of
a depression (Figs. 392, 394, 395), whereas
in M. satan, atria are outside and lateral to
the depression (Figs. 404, 406). The epi-
gynum is similar to that of M. gasteracan-
thoides but differs in ventral view, where
Mastophora * Levi
371
Figures 387-398. Mastophora satan Canals, 387-395, female. 387, 388, carapace and chelicerae. 387, frontal. 388, lateral.
389, 390, carapace and abdomen. 389, dorsal, with male. 390, lateral. 391-395, epigynum. 391, ventral. 392, 394, 395, posterior.
393, posterior, cleared. 392, 393, (Cordoba). 394, (holotype from La Rioja). 395, (Tucuman). 396-398, male left palpus. 396,
mesal. 397, ventral. 398, ectal.
Figures 399-407. M. diablo new species, female. 399, 400, carapace and chelicerae. 399, frontal. 400, lateral. 401, 402,
carapace and abdomen. 401, dorsal. 402, lateral. 403-407, epigynum. 403, ventral. 404, 406, posterior. 405, 407, posterior,
cleared. 404, (Chaco). 406, 407, (Santiago del Estero).
Scale lines. 1.0 mm; genitalia, 0.1 mm.
372 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
M. gasteracanthoides has a distinct sclerite
a short distance anterior to the posterior
edge (Fig. 414), whereas M. satan has the
whole median area shghtly sclerotized
(Fig. 391).
The median apophysis of the palpus of
the male (Fig. 398) is more cui-ved than
that of M. gasteracathoides (Fig. 420).
The egg sac lacks flaps and has only a
short stalk (Figs. 462, 463).
Distribution. From Pernambuco State,
eastern Brazil, to La Pampa Province, Ar-
gentina (Map 4B).
Specimens Examined. BRAZIL Pernambuco: Tap-
era, 1? (B. Pickel, MNRJ 391). Bahia: Feiora de San-
tana, July 1994, 1? (S. D. Cunha, IBSP 16246). Sao
Paulo: Santo Andre, 16 June 1965, 1? (L. Daga,
IBSP 1931); Seminario Santa Terezinha, Tiete, 5 May
1953, 1$ (IBSP 887); Sao Jose do Rio Preto, 9 Feb.
1964, 1? (Vizotto, MZSP 3471); 1 June 1964, 1 egg
sac, 19 (Vizotto, MZSP 3470). Santa CataHna: Blu-
menau, 19 (NHMW). Rio Grande do Sid: 1 9 (P
Buck, MNRJ 41644); Rodeio Bonito, Boge, 5 June
1980, 19 (E. W. Aguiar, MCN 9103); Canela, 10 Feb.
1966, 29 (A. A. Lise, MCN 0752); Sao Leopoldo, 5
Mar. 1965, 19 (C. Valle, MZSP 4797); Porto Alegre,
12 Apr. 1926, 19 (R. Gliesch, MNRJ .392); Parque
Zoologico, Sapucaia do Sul, 9 Dec. 1985, 29 (A. E.
Tovares, MZSP 14079); Porto Alegre, 17 Mar. 1955,
1 imm. (T. de Lema, MCN 01628)^ 16 June 1963, 1 9
(A. Lise, MCN 01820); 6 Oct. 1988, 1 9 (R. Villanova,
MCP 105); Belem Velho, Porto Alegre, 17 July 1979,
1 9 (V. Mott, MCN 2608); Morro Santana, Porto Ale-
gre, 5 May 1984, 13 (S. M. Silva, MCN 12204); 13
Sept. 1984, 19 (A. A. Lise, MCN 29426); Santa Ma-
ria, Aug. 1986, 19 (MCP 10340); Santo Antonio da
Patulha, 30 Oct. 1980, 19 (T. K. Moreira, MCN
9456). URUGUAY Villasboas, 1953, 19 (L. Lecour,
FCMU); Paso del Cerro (Artigas), May 1956, 19 (C.
Fuques, FCMU); Artigas, Sept. 1959, 19 (C. Fuques,
FCMU); Mar. 1965, 19 (C. Fuques, FCMU 1965);
Ruta 3, Salto, 3 Aug. 2001, 1 9 (V. Vazquez, Williams,
FCMU 562); Cuareim, Espinillares, Artigas, 12 Mar.
1956, 19 (C. Fuques, FCMU 296). ARGENTINA
Misiones: 1940, 19 (Exp. Zotla. Armanini, MACN
2050). Catamarca: Sijan, Nov. 1964, 19 (Aliumada,
MACN). La Rioja: 2 9 (Sr. Giacomelli, MACN 4186).
Tucumdn: Tucuman, 12 Dec. 1984, 19 (FMLT2159).
Santiago del Estero: Santiago del Estero, 20 Apr.
1958, 1 9 (J. W. Abalos, MACN); La Banda, 1958, 1 9
(J. Abalos, MACN); July 1958, 19 (D. Bravo,
MACN); Tabla Redonda, Depto. Banda, 23 Dec.
1959, 19, egg sacs, imm. (J. W. Abalos, MACN).
Mendoza: Mendoza, 1907, 19 (E. Reimoser,
NHMW); San Rafael, Feb. 1940, 19 (D. Pereyra,
MACN 1799). Cordoba: Mina Clavero, April 1973,
19 (Stirbel, MACN). Entre Rios: Concordia, 1931,
1 9 (MNRJ 57953). Santa Fe: Santa Fe, 1931, 1 9 (M.
Biraben, MNRJ 522). Buenos Aires: Ireneo Portela,
30 April 1922, 19 (Sclieimer, MACN). La Pampa:
General Pico, Feb. 1952, 19 (Williamson, MACN);
30 Mar. 1958, 29, doubtful determination (William-
son, MACN).
Mastophora diablo new species
Plate 1 ; Figures 399-407, 464; IVIap 4D
Holotype. Female holotype from Colonia Benitez,
Chaco, Argentina, Sept. 1959 (Bachmann) in
MACN, no. 54.32.
Mastophora gasteracanthoides: — Mello-Leitao, 1931:
69, in part. Canals, 1931: 19 (misidentified gaster-
acanthoides Nicolet).
Note. Most specimens of this species
had been misidentified as M. gasteracan-
thoides in collections.
Description. Female holotype. Carapace
brown, with many downy short setae and
narrow white rim. Sternum brown. Legs
brown with long white setae. Abdomen
brownish white; venter with white square.
Abdomen humps long (Figs. 401, 402).
Total length 13.0 mm. Carapace 5.3 mm
long, 5.8 wide in thoracic region, 3.2 wide
at lateral eyes. Abdomen 15 mm high.
First femur 5.5 mm, patella and tibia 8.6,
metatarsus 6.3, tarsus 1.3. Second patella
and tibia 5.7 mm, third 3.0, fourth 4.8.
Length of first patella and tibia 1.5 times
width of carapace.
Males are not known.
Variation. Total length of females 10.3—
16.7 mm. Length of first patella and tibia
1.3—1.5 times width of carapace. Figures
399—405 were made from the female ho-
lotype; Figures 406 and 407 were from
specimens from Santiago del Estero.
Diagnosis. Differs from M. satan in hav-
ing first tibia less than 9 mm total length
and from both M. satan and M. gastera-
canthoides by having the atria outside and
lateral to the depressions in posterior view
of the epigynum (Figs. 404-^07).
The egg sac lacks flaps and has a heavier
stalk (Fig. 464) than that of M. satan.
Natural History. A large syiphid fly (14
mm total length) was collected in Moreno,
Buenos Aires, with one adult.
Mastophora • Levi 373
-^'\f^
Figures 408-421. Mastophora gasteracanthoides (Nicolet). 408-416, 421, female. 408, 409, carapace and chelicerae. 408,
frontal. 409, lateral. 410-413, carapace and abdomen. 410, 412, dorsal, witfi male. 411, 413, lateral. 410, 411, (Santiago). 412,
413, (Chilian). 414-416, epigynum. 414, ventral. 415, posterior. 416, posterior, cleared. 417-420, male left palpus. 417, apical.
418, mesal. 419, ventral. 420, ectal. 421, female, frontal view with right legs.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
374 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Distribution. Northern and central Ar-
gentina to La Pampa Province (Map 4D).
Paratypes. ARGENTINA Formosa: Las Lomitas,
Oct. 1966, 1? (A. Vogt, MACN). La Rioja: 2 9 (Prof.
Gomez, MACN 4187). Tuciimdn: El Timbo, 27 May
1952, 29 (J. Campos, FMLT 994); Los Bosques, 19
(BMNH); Dept. Burruyaco, El Haranjo, 13 Jmie
1964, 19 (M. Ines Cortez, FML 1693). Santiago del
Estero: El Zanjon, 5 June 1960, 1 9 (J. W. Abalos,
MACN); Fernandez, 4 Apr. 1960, 1 9 (J. W. Abalos,
MACN); Frias, 2 Oct. 1970, 19 (J. W. Abalos,
MACN); La Banda, 24 May 1958, 19 (J. Areas,
MACN); Los Juries, March 1959, 19 (L. Remedi,
MACN); Santiago del Estero, 18 Sept. 1958, 19 (D.
Luna, MACN); 1959, 1 9 (J. W. Abalos, MACN). Cor-
rientes: Bella Vista, Nov. 1944, 19 (Silberman,
MACN). Entre Ri'o.s: Basavilbaso, 19 (U. Podesta,
MACN 4189). Santa Fe: Roldan, June 1943, 19 (Es-
cuek 230, MACN). Cordoba: Argliello, Feb. 1946,
1 9 (J. A. De Carlo, MACN 1657); Bajo Grande, 1 9
(MLP 15563). Buenos Aires: Castela, 27 July 1958,
19 (Ing. Favret); Moreno, Jan. 1946, 19 (R. D.
Schiapelli, MACN 1658). La Fampa: Mira Pampa,
April 1949, 19 (C. Vigliorcho, MLP 16638); General
Pico, March 1951, 19 (C. Ballani, MLP 1.3642).
Mastophora gasteracanthoides (Nicolet)
Figures 7, 408-421, 464; Map 4B
Epeira gasteracanthoides Nicolet, 1849: 485, pi. 5,
fig. 7a, b, 9 . Specimens from gardens and fields of
central provinces, Santiago, Chile, in MNHN, lost.
Glyptocranium gasteracanthoides: — Simon, 1895:
882, fig. 946, 9 . Bonnet, 1957: 1997.
Mastophora gasteracanthoides: — Porter, 1918: 139.
Roewer, 1942: 901. Archer, 1963: 19. Platnick,
2001.
Mastophora gasteracanthoides oxalidis Archer, 1963:
16. Female holotype, males, and imm. from Loma
de Peiluelas, 6 km al sur de La Serena, Coquimbo,
in AMNH, examined. Platnick, 2001. NEW SYN-
ONYMY.
Note. The short humps of recently col-
lected specimens from Santiago (Figs. 410,
411) differ from those of the specimens
illustrated by Nicolet (1849) and Simon
(1895), which have higher humps. Older
specimens kept in MNHN from Chilian
(Figs. 412, 413) had longer humps but
genitalia and carapace similar to those of
recently collected specimens.
The subspecies named by Archer (1963)
had no diagnosis.
Description. Female from Santiago.
Carapace brown, tubercles lighter. Chelic-
erae lighter. Sternum brown. Legs lighter
brown. Abdomen brownish white with
some asymmetrical darker patches and
humps (Figs. 410, 411); venter with white
square. Total length 12 mm. Carapace 4.8
mm long, 4.8 wide in thoracic region, 2.5
wide at lateral eyes. First femur 4.0 mm,
patella and tibia 6.3, metatarsus 4.8, tarsus
1.3. Second patella and tibia 4.3 mm, third
2.6, fourth 4.0. Length of first patella and
tibia 1.3 times width of carapace.
Male. Carapace brown, with white me-
dian band covering the two median tuber-
cles and the pair of horns. Sternum, legs
brown. Abdomen dusky white anterior,
venter with indistinct white spots. Abdo-
men without humps (Fig. 7). The palpus
has one weak patellar seta. Total length 2.3
mm. Carapace 0.9 mm long, 0.9 wide in
thoracic region, 0.7 wide at lateral eyes.
First femur 0.9 mm, patella and tibia 1.0,
metatarsus 0.6, tarsus 0.3. Second patella
and tibia 0.9 mm, third 0.5, fourth 0.7.
Length of first patella and tibia 1.1 times
width of carapace.
Note. Males came out of an egg sac col-
lected with females.
Variation. Total length of females 9.6—
13.5 mm. The illustrations of the female
are made from a female from Santiago, ex-
cept Figures 412 and 413, which are of a
female from Chilian.
Diagnosis. Mastophora gasteracanthoi-
des differs from M. cliahlo by having the
atria show as dark spots in the posterior
slope of the depression in posterior view
of the epigynum (Fig. 415), and from M.
Figures 422-433. Ordgarius magnificus (Rainbow). 422-428, female. 422, 423, carapace and chelicerae. 422, frontal. 423,
lateral. 424, 425, carapace and abdomen. 424, dorsal, with male. 425, immature, lateral. 426-428, epigynum. 426, ventral. 427,
posterior. 428, posterior, cleared. 429-433, male. 429, carapace and chelicerae, lateral. 430, dorsal. 431-433, left palpus. 431,
mesal. 432, ventral. 433, ectal.
Mastophora • Levi 375
Figures 434-444. Cladomelea akermani Hewitt. 434-440, female. 434, 435, carapace and chelicerae. 434, frontal. 435, lateral.
436, 437, carapace and abdomen. 436, dorsal, with male. 437, lateral. 438-440, epigynum. 438, ventral. 439, posterior. 440,
posterior, cleared. 441-444, male. 441, dorsal. 442-444, left palpus. 442, mesal. 443, ventral. 444, ectal.
Scale lines. 1.0 mm; genitalia, 0.1 mm.
376 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
satan by having shorter first legs and a dis-
tinct sclerotized plate on the epigynum,
anterior of the posterior margin, in ventral
view (Fig. 414).
The palp of the male has a median
apophysis straighter (Fig. 420) than that of
M. satan (Fig. 398).
The egg sac (Fig. 465) lacks flaps and
has a thin stalk.
Distribution. Central Chile (Map 4B).
Specimens Examined. CHILE 19 (MNHN, 114).
Coquimho: 6 km S of La Serena, 23—30 Nov. 1961,
19, egg sac with 3 and inim. ? (A. F. Archer,
AMNH). Valparaiso: Quilput [Quilpue], 1904, 19 (C.
Porter, MNHN 23457). Metropolitana: Santiago, 4
June 1947, 1 imm. (R. Donoso. AMNH); Feb. 1955,
19 (I. Pedag., AMNH); Quilicura, Oct. 1979, 19 (L.
Peiia, AMNH), Aug.-Sept., 2 imm. (L. E. Peiia,
AMNH); Renca, W of Santiago, 2 Oct. 1984, 19 (L.
E. Peiia, AMNH); Los Espejo, 6 Nov. 1973, 19 (C.
L. Cartagena, MCZ). Nuhle: Chilian, 19 (Delfin,
MNHN, 23520).
Ordgarius Keyserling
Onlgariiis Keyserling, 1886: 114. Tsqae species by
monotypy, M. monstrosus Keyserling, 1886, from
Queensland, Australia. Neave, 1940: 453. Roewer,
1942: 902. Bonnet, 1958: 3200. Davies, 1988: 318,
figs. 36, 37, 9, S.
Eiigli/ptila Simon, 1908: 151. Type species designated
by Bonnet, 1956, E. acanthonotata from Tonkin
[northern Vietnam]. Neave, 1939b: 325. Roewer,
1942: 903. Bonnet, 1956: 1810. NEW SYNONY-
MY.
Note. Two species of Euglijptila were
described by Simon, both from immatures.
Euglyptila acanthonotata is in MNHN,
lost; the other, E. nimthorax, 2.5 mm total
length was found, described, and illustrat-
ed by Emerit (1980). It is an immature of
Ordgarius sexspinosus Thorell, 1894, total
length 14 mm, and is illustrated by Yin,
1997: 384. NEW SYNONYMY.
Ordgarius magnificus (Rainbow)
Figures 422-433, 466
Dicrostichus magnificus Rainbow, 1897: 523, pi. 17,
fig. 8, 9 . Holotype from Mount Kembla, New
South Wales, Australia, not examined. Roewer,
1942: 900.
Ordgarius magnificus: — Davies, 1988: 316.
Description. Female. Carapace yellow-
ish, dark browii in groove between ce-
phalic and thoracic areas, a dark band
above eyes, tubercles white. Sternum light
orange. Coxae and distal leg articles light
orange. Abdomen yellowish white with
ventral white square. Carapace with me-
dian eyes on stalk, with horns quite small
(Figs. 422, 423). Eyes subequal, median
eye area almost square. Chelicerae fang
groove with three anterior teeth, one small
posterior tooth. Abdomen narrowing to
posterior end, with tubercles (Figs. 424,
425). Total length 13.5 mm. Carapace 6.3
mm long, 6.7 wide in thoracic region, 3.3
wide at lateral eyes. First femur 6.0 mm,
patella and tibia 7.3, metatarsus 4.7, tarsus
1.4. Second patella and tibia 7.0 mm, third
3.9, fourth 5.8. Length of first patella and
tibia 1.1 times width of carapace.
Male. Carapace beige, white pigment
spots at base of spines. Sternum dark
brown. Legs light beige. Dorsum of ab-
domen white, venter black. Carapace (Fig.
429), abdomen as in Figure 430. Row of
setae on tarsi as in male Mastophora. Pal-
pal patella with no setae. No endite tooth,
no coxal hook. Total length 1.7 mm. Car-
apace 0.88 mm long, 0.81 wide in thoracic
region, 0.52 wide at lateral eyes. First fe-
mur 0.78 mm, patella and tibia 0.89, meta-
tarsus 0.57, tarsus 0.30. Second patella and
tibia 0.78 mm, third 0.39, fourth 0.59.
Length of first patella and tibia 1.1 times
width of carapace.
Figures 445-468. Egg sacs of Mastophora species, including species of Ordgarius and Cladomelea. 445, M. timuqua. 446, M.
bisaccata. 447, M. stowei. 448, M. yeargani. 449, 450, M. phrynosoma. 449, (Kentucl<y). 450, (Florida). 451, M. carpogastra.
452, M. vaquera. 453, 454, M. hutchinsoni. 453, (after Kaston, 1981). 454, (New Jersey). 455, M. cornigera. 456, M. archeri.
457, M. fasciata. 458, M. abalosi (after Urtubey and Baez, 1983). 459, M. comma (after Baez and Urtubey, 1985). 460, M.
extraordinaria. 461, M. corpulenta. 462, 463, M. satan. 462, (Santiago del Estero). 463, (Rio Grande do Sul). 464, M. diablo.
Mastophora • Levi 377
468 -
Cladomelea akermani
465, M. gasteracanthoides. 466, Ordgarius magnificus (after Davies, 1988). 467, O. monstrosus (after Davies, 1988). 468,
Cladomelea akermani.
Scale lines. 1.0 mm; all except 466 and 467 are approximately the same magnification.
378 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
Genitalia. The epigynum like Mastopho-
ra in having only a posterior lobe ventrally
(Fig. 426) and having two slits and diag-
nostic sculpturing on the posterior face
(Fig. 427) and having indistinct atria (Fig.
428). The palpus of the male has no con-
ductor but has the tip of the tegulum
sclerotized (at 11 h in Fig. 433) and the
median apophysis more sclerotized than
that oiMastophora (Figs. 431-433).
Variation. The female examined came
from Olderley, Brisbane, Queensland, the
male from Mulgowie, SE Queensland
(QMB).
Natural History. Unlike most Masto-
phora, this species ties leaves together and
may have a diurnal retreat. The female
uses the second leg to swing a bolas. The
egg sac of O. niagnificus is spindle-shaped
(Fig. 466), that of O. nionstrosus resem-
bles those of Mastophora and has minute
flaps (Fig. 467).
Distribution. Australia.
Cladomelea Simon
Cladomelea Simon, 1895: 886, figs. 949, 950. Type
species by original designation Cijrtarachne longi-
pes O. P.-Cambridge, 1877: 559, from West Africa.
Neave, 1939a: 750. Roewer, 1942: 900. Bonnet,
1956: 1097.
Note. Cladomelea longipes is veiy simi-
lar to C. akermani.
Cladomelea akermani Hewitt
Figures 434-444, 468
cladomelea akermani Hewitt, 1923: 63, figs. 4, 5, 9 .
Female holotype from Pietermaritzburg, Natal, not
e.xamined. Roewer, 1942: 500. Leroy, Jocque, and
Leroy, 1998: 1, 5, S.
Description. Female. Carapace light or-
ange-brown, distal ends of projections
black. Chelicerae, labium, endites orange.
Sternum light orange-brown. Coxae or-
ange-brown; legs brown. Dorsum of ab-
domen whitish with a pair of brown tu-
bercles (Fig. 436); venter whitish with a
median, transverse, white rectangle. Car-
apace with median eyes on a bulge, three
projections and long white setae, no horns
(Figs. 434, 435). Height of clypeus equals
about five diameters of anterior median
eye. Abdomen widest in middle, dorsum
with numerous rounded tubercles, not
completely symmetrical (Figs. 436, 437).
Total length 15.5 mm. Carapace 5.4 mm
long, 5.2 wide, 2.3 wide at lateral eyes.
First femur 6.7 mm, patella and tibia 10.8,
metatarsus 8.4, tarsus 1.4. Second patella
and tibia 7.3 mm, third 3.3, fourth 4.7.
Length of first patella and tibia 1.9 times
width of carapace.
Male. Carapace, labium, endites, ster-
num dark brown. Coxae, distal leg articles
light. Dorsum of abdomen maculated
black, gray, and white (Fig. 441); venter
dark brown. Carapace rugose, without tu-
bercles, posterior area swollen. Height of
clypeus equals 1.8 diameters of anterior
median eye. Endite without tooth. Palpal
patella without macroseta. First coxa with-
out hook. Row of setae on tarsi, as in male
Mastophora. Abdomen widest in middle,
dorsum sclerotized with three humps and
two pairs of sclerotized discs (Fig. 441).
Total length 1.6 mm. Carapace 0.94 mm
long, 0.72 wide, 0.51 wide at lateral eyes.
First femur 0.80 mm, patella and tibia
0.91, metatarsus 0.48, tarsus 0.34. Second
patella and tibia 0.70 mm, third 0.41,
fourth 0.57. Length of first patella and tib-
ia 1.3 times width of carapace.
Genitalia. The epigynum is as in Mas-
tophora, having only a posterior lip ven-
trally (Fig. 438) and having two slits and
diagnostic sculpturing on the posterior
face (Fig. 439) and tiny atria (Fig. 440).
The palpus of the male has a distinct con-
ductor supporting the embolus (at 1 h in
Fig. 443) and the median apophysis more
sclerotized than that of Mastophora (Figs.
442-444).
Variation. A second female from Pieter-
maritzburg, Natal (AMNH), examined had
the height of the clypeus only four diam-
eters of the anterior median eye. Female
examined and illustrated from Umgeni
Valley Reservation, Kwa— Zulu— Natal,
South Africa, the male came from Umgeni
Valley project near Howick, South Africa
(NMP).
Mastophora • Levi
379
Natural History. The spider is found in
grasslands on grass of the Kwa—Zuki— Natal
area around Pietermaritzburg, South Af-
rica (Leroy et al., 1998). The egg sac is
drop-shaped and attached to a grass blade
(Fig. 468). The female handles the bolas
with a second leg and swings it in a hori-
zontal plane.
Distribution. South Africa.
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382 Bulletin Museum of Comparative Zoology, Vol. 157, No. 5
INDEX
Valid names are printed in italics. Page numbers refer to main references, starred page numbers to illus-
trations.
abalosi, 355*, 356, 377*
Acacesia, 315
Acantharachne, 314
Acantharanea, 314
Agathostichus, 315
Agatostichus, 315
akermani, 375*, 377*, 378
alachua, 329*, 330
Alpaida, 315
altiventer, 325
alvareztoroi, 359*, 360
apalachicola, 326, 327*
Araneus, 315
archeri, 346, 347*, 377*
bicurvata, 345
bisaccata, 327*, 330, 331*, 377=1
bisaccatum, 330
brescoviti, 361*, 363
cai-pogaster, 340
carpogastera, 340
carpogastra, 327*, 340, 377*
caipogastrum, 340
catarina, 335*, 339
Celaenia, 315-317
Cladomelea, 314, 316-18, 378
Coelossia, 314
comma, 355*, 356, 377*
conifera, 361*, 364
coniferus, 364
comigera, 343*, 344, 377*
cornigera, 342
cornigerum, 345
cornigerus, 345
corpulenta, 364, 365*, 377*
coi^pulentum, 364
coi-pulentus, 364
corumbatai, 335*, 339
cranion, 349*, 351
Cyrtarachne, 315-317
Cyrtophora, 315
diablo, 327*, 371*, 372, 377*
Dicrostichus, 314
dizzy deani, 349*, 350
Eriophora, 315
escomeli, 365*, 366
Euglyptila, 376
Exechocentrus, 314
extraordinaria, 355*, 357, 377*
fagoides, 340
fasciata, 347*, 348, 377*
fasciolata, 318, 325
felda, 326, 327*
felis, 367*, 369
gasteracanthoides, 327*, 373*, 374, 377*
gasteracanthoides, 366, 372
gasteracanthoides oxalidis, 374
Glyptocranium, 315
hayivardi, 335*, 339
Heterocephala, 315
holmbergi, 367*, 369
hutchinsoni, 342, 343*, 377*
intermedia, 72
Kaira, 315-317, 325
lara, 335*, 340
lenca, 364
leucabulba, 358, 359*
leucabulbus, 358
leucacantha, 361*, 362
leucacanthus, 363
longicep.s, 351, 353*
magnificus, 375*, 376, 377*
Mastophora, 315-317
melloleitaoi, 355*, 356
Metepeira, 315
monstro.su.s, 377*
multilineata, 330
obesus, 330
obtusa, 367*, 368
obtusum, 368
occidentalis, 348
Ocrepeira, 315
OrdgaHus, 314, 316, 317, 376
Pasilobus, 315-318
pesqueiro, 352, 353*
phrynosoma, 333*, 336, 377*
pickeli, 349*, 350
pickeli occidentalis, 348
piras, 352, 353*
Poecilopachys, 315-317
rabida, 365*, 366
reimoseri, 367*, 370
satan, 370, 371*, 377*
satsuma, 325, 327*
Seminole, 337*, 341
soberiana, 361*, 362
stowei, 331*, 334, 377*
Taczanowskia, 315—317
timuqua, 328, 329*, 377*
vaquera, 337*, 342, 377*
yacare, 354, 355*
yeargani, 333*, 336, 377*
ypiranga, 353*, 354
i
I
Mb
SuLUttn
seum
Comparative
Zoology
A MONOGRAPHIC REVISION OF THE
ANT GENUS PRISTOMYRMEX
(HYMENOPTERA: FORMICIDAE)
MINSHENG WANG
iBRAR
OCT 2 7 2003
HARVARD
•JNIVERSIT^
HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSEI IS, U.S.A.
VOLUME 157, NUMBER 6
10 April 2003
(US ISSN 0027-4100)
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1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and
Evolution of Crustacea. 192 pp.
2. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere-
dinidea (Mollusca: Bivalvia). 265 pp.
3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino-
derms. 284 pp.
4. Eaton, R. J., 1974. A Flora of Concord from Thoreau's Time to the
Present Day. 236 pp.
5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology
and Evolutionary Biology: Essays in Honor of Ernest E. Williams.
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6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp.
Other Publications.
Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine.
Reprinted 1964.
Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of
Insects. {Bulletin of the M. C. Z, Vol. 108.) Reprinted 1971.
Creighton, W. S., 1950. The Ants of North America. Reprinted 1966.
Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In-
ternational Symposium on Natural Mammalian Hibernation. {Bulletin
of the M. C. Z, Vol. 124.)
Orinthological Gazetteers of the Neotropics (1975-).
Peter's Check-list of Birds of the World, vols. 1-16.
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© The President and Fellows of Harvard College 2003.
A MONOGRAPHIC REVISION OF THE ANT GENUS PRISTOMYRMEX
(HYMENOPTERA: FORMICIDAE)
MINSHENG WANG^
CONTENTS
Abstract 383
Introduction - 383
Collections 385
Measurements and Indices 385
A Brief Histoiy of the Genus Pristomynnex ... 386
Genus Pristomynnex Mayr 387
List of Pristomynnex Names wdth Synonymies 392
Key to the World Species of Pristomynnex
(Workers) 393
The punctatus Group - 404
Pristomynnex divisus sp. n. 404
Pristomynnex fossulatus (Forel) 406
Pristomynnex piilcher sp. n. 408
Pristomynnex punctatus (F. Smith) 410
Pristomynnex rigidus sp. n. 415
The cribrarius Group 417
Pristomynnex cribrarius Arnold ._._ 418
The qimdridens Group 420
Pristomyrmex africanus Karavaiev 423
Pristomyitnex bicolor Emeiy stat. n. 425
Pristomyrmex brevispinosus Emery 428
Pristomynnex collinus sp. n. 432
Pristomymiex costatus sp. n. 434
Pristomynnex curvidus sp. n. 437
Pristomyrmex eduardi Forel 440
Pristomynnex erytliropygiis Taylor 441
Pristomy nnex flatus sp. n. 443
Pristomy rmex foveolatus Taylor 446
Pristomynnex hirsutus sp. n. 449
Pristomynnex longispinus sp. n. 450
Pristomynnex modestus sp. n. 452
Pristomynnex nitidissimus Donisthorpe 453
Pristonu/nnex occultus sp. n. _ 455
Pristomyrmex orbiceps (Santschi) 456
Pristonujrmex quadridens Emery 459
Pristomynnex quadridentatus (Andre) 463
Pristomyrmex quindentatus sp. n. 467
Pristomynnex sulcatus Emery stat. n 469
Pristomynnex thoracicus Taylor 473
Pristomynnex trachylissus (F. Smith) 474
Pristonujrmex trogor Bolton 476
' Museum of Comparative Zoology, Harvard Uni-
versity, Cambridge, Massachusetts 02138.
Pristomynnex wheeleri Taylor 478
Pristomyrmex wilsoni Taylor — 481
The trispinosus Group 483
Pristomyrmex bispinosus (Donisthorpe) 484
Pristomynnex browni sp. n. 485
Pristomyrmex trispinosus (Donisthoipe) 488
The levigatus Group 489
Pristomynnex acerosus sp. n. 491
Pristomynnex boltont sp. n. __ 492
Pristomyrmex coggii Emery 493
Pristomynnex inermis sp. n. 496
Pristomynnex largus sp. n. 497
Pristomynnex levigatus Emery 499
Pristomyrmex longus sp. n. 502
Pristomymiex lucidus Emery — 503
Pristomynnex mandibularis Mann 505
Pristomymiex minusculus sp. n. 507
Pristomymiex obesus Mann 509
Pristomyrmex simplex sp. n. 512
The profundus Group 514
Pristomynnex profundus sp. n. 515
The umbripennis Group 516
Pristomymiex fuscipennis (F Smith) 517
Pristomynnex picteti Emeiy 518
Pristomynnex pollux Donisthoi-pe 521
Pristomyrmex reticulatus Donisthorpe 524
Pristomymiex umbripennis (F Smith) 525
Nomen Nudum 528
Pristomyrmex parvispina Emery 528
Acknowledgments 539
References 539
Abstract. The ant genus Pristomynnex is revised
as a whole for the first time. The genus is redefined,
and seven species groups are erected and discussed.
Illustrations are present for all 52 species. A key to
the worker caste is provided. Twenty-one new species
are described: 20 from the Oriental region and one
from Mauritius. Thirteen names are newly synony-
mized, and two former infraspecific taxa are elevated
to species rank.
INTRODUCTION
Pristornijrrnex, an ant genus of moderate
size, contains 52 living species, but fossils
Bull. Mus. Comp. Zool., 157(6): 383-542, April, 2003 383
384 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
have not been discovered. Pristomijmiex
occurs primarily in the Oriental region,
but six endemic species are present in the
eastern rainforest of Australia and five en-
demic species in Africa. In addition, in
Mauritius there are three native species,
one of which also occurs on Reunion Is-
land. Lastly, one species, Pristomijmiex
punctatus, has invaded temperate China,
Korea, and Japan. This species has also
been detected at two entry ports in the
United States and thus shows potential for
spread via human commercial actions.
Pristomijmiex belongs to the subfamily
Myrmicinae. It possesses a raised trans-
verse ridge or a few toothlike prominences
on the dorsal labrum in all female castes,
including workers, ergatoid queens, and
queens. This character is also shared by
the myrmicine genera Acanthomyrmex,
Myrmecina, and Perissomymiex. As a re-
sult, these four living genera are grouped
together in the tribe Myrmecinini (Bolton,
1994, 1995, personal communication;
Brown, 1971). Pristomyrmex is unique in
the tribe because it is the only genus pos-
sessing 11 antennal segments in all three
female castes and 12 segments in the male.
Most Pristomyrmex species dwell in the
rainforest, foraging as predators or scav-
engers. An Asian species, P. punctatus,
however, occurs in open and disturbed
habitats (e.g., bare hills, agricultural areas,
and beaches). These ants prefer to nest in
soil, litter, or rotten wood; in rotten parts
of living trees; in dead standing trees; or
around plant roots.
Pristomyrmex is of great interest be-
cause it exliibits several unusual biological
and evolutionary phenomena. The absence
of moi"phologically normal queens and re-
production primarily by unmated workers
in P. punctatus {=P. pungens) is a highly
unusual life history in the Formicidae. It
has attracted much attention from those
who hope to obtain insight into the nature
of reproductive conflict within colonies
since, in this species, reproductive division
of labor occurs among moi"phologically
identical workers (Itow et al., 1984; Mu-
zutani, 1980, 1982; Peeters, 1993; Tsuji,
1988a,b,c, 1990a,b, 1994, 1995; Tsuji and
Ito, 1986). Ergatoid queens, a special
wingless female caste morphologically in-
termediate between the queen and the
worker, are present in at least four species:
P. punctatus, P. africanus, P. wheeleri, and
P. mandibularis; two of them (P. africanus
and P. wheeleri) possess both queen and
ergatoid queen castes. Character displace-
ment, showing that two species possess a
greater difference in sympatric than allo-
patric populations, has also been reported
in this genus by Taylor (1965). In addition,
simulating death, slowness of movement,
and nocturnal foragers are also recorded
in Pristomyrmex (Donisthorpe, 1946; Tay-
lor, 1965; Weber, 1941). Colony size varies
greatly among species, ranging from about
a dozen to several thousand workers (Don-
isthorpe, 1946; Itow et al, 1984; Mann,
1919; Taylor, 1965, 1968).
Although Pristomyrmex is biologically
promising, the taxonomic foundation of
the genus is poor. Much of the literature
on Pristomyrmex is more than 50 years old
and consists of isolated descriptions of spe-
cies or infraspecific forms. Only a handful
of papers present more comprehensive
studies of the Australian and African su-
bfaunas, respectively (Bolton, 1981; Taylor,
1965, 1968). The tropical Asian region,
however, containing the bulk of the de-
scribed taxa, has been in taxonomic chaos,
for many years obscuring a better under-
standing of the evolution and radiation of
this interesting group.
This survey takes the whole Pristomyr-
mex into consideration. I believe that only
after that the entire genus covering all zoo-
geographical regions is comprehensively
investigated can a full set of characters to
define the genus be summarized, the spe-
cies groups correctly erected, and the re-
lationships between species properly ana-
lyzed and then the possible origin and the
evolution of the genus hypothesized.
I present a detailed description of the
taxonomic characters for the worker caste
of each species. These characters not only
Revision of the Ant Genus Pristomyrmex • Wang 385
are useful for the species identity but also
provide important information for a fur-
ther study on the phylogeny within the ge-
nus. I also include illustrations and de-
scriptions of males for many species as
possible. This was done for three reasons.
First, two species (P. pollux and P. reticu-
latus) were described, each from a single
male speciiTien, many years ago. Without
examining other available males, I would
not be able to assign these two species to
their appropriate species group, and the
discovery of other new species would then
be impeded. Second, the males of inost
ant genera are very poorly characterized
and thus cannot be curated properly in
museum collections. Finally, I feel that
these inales contain some clues for the
study of the phylogenetic relationships of
the genus.
COLLECTIONS
AMNH American Museum of Natural
Histoiy, New York, N.Y., U.S.A.
ANIC Austrahan National Insect Col-
lection, Canberra City, Australia
BMHH Bishop Museum, The State Mu-
seum of Natural and Cultural
History, Honolulu, Hawaii,
U.S.A.
BMNH Natural Histoiy Museum, Lon-
don, U.K.
CASC California Academy of Sciences,
San Francisco, California, U.S.A.
IZAS Institute of Zoology, Academy of
Sinica, Beijing, China
IZUA Institute of Zoology, Ukrainian
National Academy of Sciences,
Kiev, Ukraine
LACM Natural History Museum of Los
Angeles County, Los Angeles,
California, U.S.A.
MCSN Museo Civico di Storia Naturale
"Giacomo Doria", Genoa, Italy
MCZC Museum of Comparative Zool-
ogy, Harvard University, Cam-
bridge, Mass., U.S.A.
MHNG Museum d'Histoire Naturelle,
Geneva, Switzerland
MNHA
MNHN
MNHU
NACA
NHMB
NHMV
NHPS
OXUM
SAMC
USNM
Museum of Nature and Human
Activities, Sanda, Hyogo, Japan
Museum National d'Histoire
Naturelle, Paris, France
Museum fiir Naturkunde der
Humboldt-Universitat zu Berlin,
Berlin, Gennany
National Arthropod Collection,
Mount Albert Research Center,
Auckland, New Zealand
Naturhistorisches Museum, Ba-
sel, Switzerland
Naturhistorisches Museum, Vi-
enna, Austria
Naturhistoriska Piksmuseet,
Stockliolm, Sweden
Oxford University Museum, Ox-
ford, U.K.
South African Museum, Cape
Town, South Africa
National Museum of Natural
History,
U.S.A.
Washington, D.C.
MEASUREMENTS AND INDICES
Head Width (HW). Maximum width of
head, in full-face view, excluding the eyes
(Fig. 1).
Head Width Inchiding the Eyes (HWE).
Maximum width of head across the eyes,
in full-face view. This measurement is
used only in the male.
Head Length (HL). Length of the head
in full-face view, excluding the inandibles
(Fig. 1), measured from the midpoint of a
straight line across the occipital margin to
either the apex of the median tooth (if it
is present) of the anterior clypeal niargin
or the midpoint of a line connecting the
apexes of the two lateral teeth (if the me-
dian tooth is absent) of the anterior clypeal
margin or the midpoint of the anterior
clypeal ixiargin (if the anterior margin lacks
any teeth).
Cephalic Index (CI). HW/HL X 100.
Scape Length (SL). Length of the an-
tenna! scape, including the lamella encir-
cling the base of the scape but excluding
the basal condyle (Fig. 1).
Scape Index (SI). SL/HW X 100.
386 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 1-3. Measurements illustrated for this study (worker of Pristomyrmex longispinus sp. n.
Entire body, lateral view; 3: Dorsal view of alitrunk, petiole, and postpetiole.
1: Head, full-face view; 2:
Pronotal Width (PW). Maximum width
of the pronotum in dorsal view (Fig. 3).
Alitnink Length (AL). Diagonal length
of the alitrunk in lateral view, from the an-
teriormost point of the pronotum to the
apex of the metapleural lobe (Fig. 2).
Eye Length (EL). Maximum length of
the eye.
Total Length (TL). TLl + TL2 + TL3
(see Fig. 2). (Note: The measurements of
TL do not deal with those individuals
whose gasters are abnormally contracted
or prolonged or whose petioles are raised
too high or too low.) TLl: A line mea-
sured from the apex of the closed man-
dibles to the midpoint of a straight line
across the occipital margin, in full-face
view. TL2: A straight line from the anter-
iormost point of the pronotum to the
point at which the posterior margin of
postpetiole meets the uppermost point of
an articulation. TL3: A line from the an-
terior-uppermost point of the articulation
to the apex of gaster.
Pronotal Spine Length (PSLl). A
straight distance from the base to the apex
of pronotal spine (see Fig. 2).
Propodeal Spine Length (PSL2). A
straight distance from the base to the apex
of propodeal spine (see Fig. 2).
Po.stpetiole Index (PPI). PFW/PPL X
100 (PPW: maximum width of the post-
petiole in dorsal vies; PPL: length of the
postpetiole in dorsal view).
All measurements are taken in milli-
meters.
Note: For the Australian species, my
measurements differ slightly from Taylors
(1965, 1968). For the maximum measur-
able width of head, I have excluded the
eyes, which were included by Taylor.
A BRIEF HISTORY OF THE GENUS
PRISTOMYRMEX
The genus Pristomijnnex, when erected
by Mayr (1866), contained one species, P.
pungens Mayr ( = a junior synonym of P.
Revision of the Ant Genus Pristomyrmex • Wang 387
ptinctatus (F. Smith)), a member of the
present punctatus group. At that time,
Mayr did not reahze that five species de-
scribed by F. Smith (1858, 1860, 1861,
1863, 1865), that is, Mijrmica fuscipennis,
Myrinica punctata, Mijnnica trachylissa,
Mijrniica unibripennis, and Solenopsis lae-
vis, also belonged in the new genus. Mayr
(1866) provided a description of the genus,
which actually was siiTiply derived from
some characters of the species P. puncta-
tus (F. Smith). A second member of the
punctatus group was introduced by Sants-
chi (1916) when he transferred Tetramo-
rium (Xiphomyrmex) fossulatum Forel to
Pristomyrmex.
The content of the genus Pristonujrmex
expanded for the first tiiTie when Mayr
(1886) transferred Myrmica trachylissa F.
Smith to Pristonujrniex. Pristonujnnex tra-
chylissus is now a ineinber of the quaclri-
dens group. After that, many species of the
quadridens group were discovered (Bol-
ton, 1981; Donisthoipe, 1949c; Emeiy,
1887, 1895, 1897, 1900; Forel, 1914; Ka-
ravaiev, 1931, 1933; Taylor, 1965, 1968),
and several more naines were added to the
group when Odontomynnex Andre and
Hijlidris Weber were designated by Forel
(1915) and Brown (1953) as a subgenus
and a synonym of Pristomyrmex, respec-
tively. But, Mann (1919) found no evi-
dence supporting the subgenus Odonto-
mi/nnex.
The iTiembers of the third species group
(i.e., umbripennis group) o£ Pristonujrniex
were recognized first by Emeiy He de-
scribed a new species {PHstonujnnex pic-
teti) in 1893 and transferred a species
{Myrmica fuscipennis F Smith) to Pristo-
myrmex in 1901. Donisthoi"pe expanded
the umbripennis group: He transferred
Myrmica umbripennis F. Smith and Sole-
nopsis laevis F. Smith to Pristomyrmex and
described Pristoinynnex pollux and Pris-
tomyrmex reticulatus (1932, 1946, 1949a).
When Emery (1897) described Pristo-
myrmex coggii, Pristonujnnex levigatus,
and Pristomyrmex lucidus, representatives
of the present levigatus group w^ere added
to the genus for the first time. Pristomyr-
mex cribrarius, the sole member of the
cribrarius group, was described by Arnold
(1926). Lastly, Brown (1971) synonymized
the genus Dodous Donisthorpe, adding
species belonging to the present trispino-
sus group to the genus. Thus, the genus
Pristomyrmex became clearly delimited
and assumed its modern form.
GENUS PRISTOMYRMEX MAYR
Pristomyrmex Mayr, 1866: 903. Tyjae species: Pristo-
myrmex pungens Mayr, op. cit.: 904 [=Mynnica
'punctata F. Smith, I860: 108; =Pristomymiex
punctatus (F. Sinith)]; by monotypy.
Odontomynnex Andre, 1905: 207. Type species:
Odontomyrmex quadridentatus Andre, op. cit.:
208; by monotypy. [As a subgenus, thus synonym,
o( Pristomyrmex by Forel, 1915: 53.]
Hylidris Weber, 1941: 190. Type species: Hylidris
myersi Weber, loc. cit. {=Pristomynnex africaniis
Karavaiev); by original designation. [Synonymy by
Brown, 1953: 9.]
Dodous Donisthorpe, 1946: 145. Type species: Do-
dous trispinostis Donisthorpe, loc. cit.; by original
designation. [Synonymy by Brown, 1971: 3.]
Diagnosis of worker, queen, and erga-
toid queen. Combination of the following
asterisked four characters (i.e., characters
2, 7, 11, and 29 in the worker caste) sep-
arating Pristomyrmex froin other myrmi-
cine genera.
Definition: Worker. Possessing the fol-
lowing combination of characters:
1. Small (TL 1.74, HL 0.46, HW 0.46)
to large-sized (TL 7.06, HL 1.68, HW
1.74) iTionomorphic myrmicine ants.
*2. Mandible soineM^hat subtriangular;
masticatory margin of mandible with three
to five teeth, which have one or the other
of the following six basic arrangements:
(1) the strongest apical + the second
strongest preapical + the smallest
third + the acute basal tooth, dia-
stema lacking, as in levigatus group
and in profundus group, or
(2) the strongest apical + the second
strongest preapical + two smaller
teeth of similar size, diastema indis-
tinct or lacking, as in umbripennis
group, or
(3) the strongest apical + the second
388 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
strongest preapical + a shorter
(first) diastema (sometimes the first
diastema is not distinct) + a small
denticle + a longer (second) diaste-
ma + a small basal denticle, as in
both P. bispmcsus and P. trispino-
SU.S, or
(4) the apical + the preapical + a lon-
ger diastema + a small denticle + a
shorter diastema (sometiines the
second diastema is indistinct) + a
sinall basal denticle, as in P. browni,
or
(5) the strongest apical + the second
strongest preapical + a distinct di-
astema + a basal tooth (which is
sometimes formed by the fusion of
the two small teeth) or two (or
three) small teeth of similar size, as
in punctatus group, cribrarius
group, and most members of the
qiiardridens group, or
(6) the strongest apical + the second
strongest preapical + an intercalary
tooth + a very short diastema (or
this diastema indistinct) + two small
teeth of similar size, as shown in P.
trachijlissu.s.
3. Basal margin of mandible with a
broad-based triangular or an acute and
prominent tooth, or only curved, not form-
ing tooth, or almost straight.
4. Median part of clypeus shieldlike,
projecting posteriorly between the bases of
the antennae; lateral parts of clypeus in
front of antennal insertions usually re-
duced to ridges but rarely (in the two Ori-
ental species P. clivisus and P. pulcher) de-
veloped so that the antennal fossae do not
reach the lateral anterior margins of clyp-
eus.
5. Anterior clypeal margin usually with
a median tooth and one to tliree pairs of
lateral denticles (or crenulate shapes) but
sometimes the median tooth rudimentaiy
(as in some species of the levigatiis group)
and sometimes anterior clypeal margin
lacking any distinct denticles (as in P. pro-
fundus, P. divisus, and P. pidcher).
6. Ventral surface of clypeus with a me-
dian tooth or two lateral teeth, or with a
transverse ridge, or without any ridge or
tooth.
*7. Dorsal lab rum with a raised trans-
verse ridge or a few toothlike proininenc-
es, present on the anterior portion of la-
brum in most species.
8. Palp formula 1,2, 1,3, 2,2, 2,3, 4,3, or
5,3.
9. Frontal lobes absent in punctatus
and trispinosus groups or weak, as in lev-
igatiis, profundus, and qiiadridens groups,
or somewhat expanded, as in unibripennis
group; as a result, the articulations of the
antennae are mostly or entirely exposed in
full-face view.
10. Frontal carinae usually developed,
extending to the level of the posterior mar-
gins of eyes, but sometimes frontal carinae
absent or very short, as in the trispinosus
group, in P. trogor, and in P. longispinus.
*11. Antennae with 11 segments; apical
three segments forming a distinct club.
12. Base of each antennal scape encir-
cled by a narrow lamella, except in P. pro-
fundus; this lamella usually with a broad
and deep notch on the center of dorsal
surface in the unibripennis group but en-
tire in the other species groups.
13. Antennal scrobes usually absent or
weakly developed, but in P. profundus, the
scrobes are deep and well developed.
14. Eyes present in all known species,
situated approxiinately at the midlength of
the sides of the head; usually moderate-
sized, but small in the several species (P.
boltoni, P. coggii, P. longus, P. eduardi, P.
picteti, and P. pollux).
15. Ahtrunk usually lacking dorsal su-
tures, but in the three species of the trispi-
nosus group, a promesonotal suture or im-
pression present.
16. Pronotum unarmed, or armed with
a pair of tubercles, teeth, or spines of vary-
ing sizes.
17. Mesonotum usually unarmed, but
with a pair of thick, blunt, and digitlike
short prominences in P. trispinosus, and
sometimes weakly tuberculated in P. bispi-
no.sus and P. browni.
Revision of the Ant Genus Pristomyrmex • Wang 389
18. Propodeum armed with a pair of
teeth or spines, except in P. inennis.
19. Metapleural lobes usually subtrian-
gular, or each with a blunt-rounded to
semicircular apex, but indistinct in P. pro-
fundus.
20. Fore tibial spurs pectinate. Middle
and hind tibiae sometimes without any
spur, sometimes with either simple or hair-
like spurs.
21. Propodeal spiracles circular and
high-positioned on the lateral surfaces of
the propodeum.
22. Metapleural gland bullae large, sep-
arated from the propodeal spiracles, and
positioned above the posterior lower cor-
ners of propodeum.
23. Petiole in profile nodiform or
wedge-shaped, pedunculate, usually with a
long anterior peduncle.
24. Subpetiole sometiixies without a
ventral process, sometiiTies bearing a nar-
row semitranslucent lamella. In P. acero-
sus, a pinlike process is present.
25. Postpetiole in profile nodiform,
usually rounded dorsally.
26. Petiole spiracle, postpetiole spira-
cle, and first gastral spiracle visible.
27. Dorsal surfaces of head and alitrunk
smooth, or possessing either scattered fo-
veolate punctures, or foveolate-reticulate
sculpture, or developed rugo reticulum, or
regular striate sculpture. Gaster unsculp-
tured.
28. Dorsal surfaces of head and alitrunk
usually with nuinerous hairs, but only a
few hairs present on the dorsal alitrunk in
P. fossulatus, P. orbiceps, and P. trogor.
Petiole and postpetiole each usually with
one to three pairs of hairs, but sometimes
more pairs of hairs present; sometiixies
petiole and postpetiole lacking hairs. First
gastral tergite usually without hairs or vdth
a few sparse hairs, but sometimes first gas-
tral tergite covered with numerous, evenly
distributed, erect or suberect hairs.
*29. Anterior clypeal margin lacking a
median seta at the midpoint of the margin,
instead usually having two to three pairs of
long, forward-projecting hairs flanking the
midpoint of margin.
30. Sting slender and long.
Female. Usually alate, but in some spe-
cies (P. punctatus, P. niandibularis) , only
ergatoid queens have been found. In some
species (P. wheeleri, P. africanus), both
alate and ergatoid queens exist.
Alate Queen. Characters similar to those
of worker in the structure and shape of
mandible, palp formula, clypeus, frontal
lobes, frontal carinae, antennae, meta-
pleural lobes, tibial spurs, petiole node,
postpetiole, and sting as well as in the
sculpture of body. But larger, with slightly
or much larger eyes, than in the conspe-
cific worker; three ocelli present. The ali-
trunk with wings and flight sclerites; well-
marked dorsal sutures present. Pronotal
spines usually absent, but in some species,
the pronotuin is arined with a pair of teeth
that are much shorter than in conspecific
worker; propodeal teeth or spines usually
shorter than those of conspecific worker.
Wing venation as shown in Figures 4—5.
On the forewings, the marginal cell (see
Holldobler and Wilson, 1990: 9) is always
open; R -\- Sc thick (for the explanation of
symbols used, see Brown and Nutting,
1950); A short, far from the anal angle; A,
Cu-A, M/2+3 usually reduced to vestigial
lines distally; cross-vein m-cii and r-m ab-
sent; cross-vein cu-a usually present but
sometimes broken in larger species (such
as P. picteti, P. umbripennis) and some-
times rudimentary or very weak in some
samples of a few smaller species (e.g., P.
orbiceps, P. lucidus); Ir absent; anal lobe
usually indistinct in smaller species but
present in larger species. Hind wings with-
out anal lobe. (Note: The venation of the
both fore and hind wings of alates, in Pris-
tomijnnex, is rather stable, with only slight
variations within the different species. For
example, on the forewings, A^2+3, some-
times becomes an almost entirely vestigial
line, but sometimes it is distinct and rather
long; jR.s/4 + Rs/5 is rather thick and long
in some larger species but thin and short
in some sinaller species).
390 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 4-7. General forewing and hindwing venation of alate queens and males of Pristomyrmex. 4; Forewing of alate queens;
5: Hindwing of alate queens; 6: Forewing of males; 7: Hindwing of males.
[The forewing and hindwing venation of the alate queens of the following 13 Pristomyrmex species was examined: P. brevispi-
nosus, P. collinus, P. orbiceps, P. quadridens, P. quindentatus, P. sulcatus, P. levigatus, P. lucidus, P. obesus, P. fuscipennis,
P. picteti, P. pollux, and P. umbripennis. The males of 16 Pristomyrmex species were examined; P. brevispinosus, P. flatus, P.
trogor, P. longispinus, P. orbiceps, P. quadridens, P. quadhdentatus, P. sulcatus, P. browni, P. trispinosus, P. obesus, P.
levigatus, P. picteti, P. pollux, P. umbripennis, and P. punctatus.]
ErgatoicI Queen. General characters, in-
cluding the pronotal proininences and size
of body, similar to those of the conspecific
worker. Ocellus present (one ocellus in P.
mandibularis but three ocelli in P. punc-
tatus, P. wheeleri, and P. africanu.'i); apter-
ous, but mesonotum more convex than in
conspecific worker; pro-mesonotal suture
present in P. mandibularis but represented
by an impression in P. punctatus, P. whee-
leri, and P. africanus.
Male. Possessing the following combi-
nation of characters (summarized accord-
ing to 54 specimens falling into at least 17
species):
1. Small to moderate size (TL 2.40—
6.04, HL 0.48-0.94, HW 0.51-0.98, HWE
0.62—1.10), usually sinaller than the con-
specific queen.
2. Head, in full-face view, across and in-
cluding the eyes, usually broader than long
(Figs. 261-269).
3. Mandibles vestigial, very small,
rounded or toothlike, far from meeting, as
indicated by an arrow in Figure 262.
4. Anterior ixiargin of labrum broadly
concave at center; dorsum of labrum with-
out any transverse ridge or toothlike prom-
inences (see Figs. 262, 264, 269).
5. Eyes very large, well developed, and
convex, situated at the sides of head.
6. Antennae filiform, 12 segments, lack-
ing a lamella encircling the base. Scapes
short, usually distinct shorter than the
maxiinum length of eye; of the other 11
funicular seginents, the first segment
shortest, the apical segment longest, the
remaining nine seginents much longer
than their broad.
7. Three ocelli conspicuous and well
developed, situated on the vertex of the
head.
8. Antennal sockets set back from the
posterior margin of the clypeus.
9. Antennal scrobes absent.
10. Frontal carinae absent or very short
and weak.
11. Frontal lobes absent so that the ar-
ticulations of the antennae are coinpletely
exposed in full-face view.
12. Palp formula as in the conspecific
worker in seven species exaiTiined (i.e., P.
punctatus, P. quadridens, P. curvulus, P.
brevispinosus, P. stdcatus, P. picteti, and P.
pollux).
13. Clypeus convex in the middle, not
projecting posteriorly upward between the
bases of antennae; its shape transverse, or
Revision of the Ant Genus Pristomyrmex • Wang 391
somewhat semicircular; its anterior margin
entire, without any denticles, usually rath-
er straight but sometimes arched.
14. Cheeks very short.
15. Alitrunk robust, with wings, well-
developed flight sclerites, and well-marked
sutures.
16. Pronotum narrow in middle, over-
hung by mesoscutum in lateral view, lack-
ing any annaments.
17. Mesonotum well developed, con-
sisting of a large mesoscutum, a rather
large mesoscutellum, and two small axillae.
Notauli usually distinct, fonning a Y shape,
but sometiines they show a V shape, and
sometimes they are absent or very weak.
Parapsidal furrows usually absent, but
sometimes they are superficially im-
pressed.
18. Metanotum transverse, narrow,
overhung by mesoscutellum.
19. Propodeum showing a sloping dor-
sal surface; propodeal anxiaixients absent
or present; if present, they are usually
shorter than in the conspecific worker.
20. Metapleural lobes present, sub-
triangular, or toothlike, or blunt-rounded
to semicircular.
21. Venation (Figs. 6—7) as in alate
queen.
22. Legs slender; fore tibial spurs pec-
tinate; middle and hind tibiae usually lack-
ing any spurs but sometimes simple spurs
are present.
23. Petiole with a long or a rather long
anterior peduncle. In dorsal view, sides of
petiole subparallel. Petiole node low, lower
than in the conspecific worker and queen;
subpetiole lacking any lamella or toothlike
projection.
24. Postpetiole node rather low, lower
than in the conspecific worker and queen.
In profile, subpostpetiole usually lacking
any projections, but sometiines bearing a
small tooth.
25. Positions of spiracles on propo-
deum, petiole, postpetiole, and first gastral
seginent similar to those in the conspecific
worker and queen.
26. Usually much less sculptured than
conspecific worker and queen.
27. Numerous hairs present on the en-
tire dorsal surfaces of body.
(Note: The genitalia of males is not dis-
sected.)
The male of Pristomyrmex can be dis-
tinguished within the tribe Myrmecinini
by the following characters:
Pristomyrmex
Antennae: 12 segments
Mandibles: Very sinall, toothlike, not
meeting
Petiole: With a long anterior peduncle
Forewing: Without m-cu cross-vein;
inarginal cell open
Acanthomyrmex
Antennae: 13 seginents
Mandibles: Subtriangular, with six to
eight teeth, ineeting when they are
closed
Petiole: Similar to that of Pristomyrmex
Forewing: ?With m-cu cross-vein; mar-
ginal cell closed
Myrmecina
Antennae: 13 segments
Mandibles: Similar to those of Pristo-
myrmex
Petiole: Without an anterior peduncle
Forewing: Without m-cu cross-vein;
marginal cell closed
Perissomyrmex
Antennae: ?10 segments
Mandibles: Unknown
Petiole: Unknown
Forewing: Unknown
Larva. According to Wheeler and
Wheelers (1954, 1960, 1973, 1976) stud-
ies, the larva of Pristomyrmex has the fol-
lowing combination of characters:
1. Stout and rather short.
2. Head extremely long and narrow.
3. Thorax more slender than abdomen
and forming a neck, which is cuived ven-
trally. Diameter greatest near middle of
abdomen, decreasing gradually toward
head; posterior end rounded.
4. Body without tubercles.
392 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
5. Mandibles subtriangular, without
medial blade; apical tooth curved medially
and usually acute; subapical medial tooth
small.
6. Body hairs numerous, with five or six
types, including anchor-tipped hairs. Head
hair few, short to moderately long.
7. Gula spinulose.
8. Anterior surface of labium densely
spinulose.
9. Palps lateral.
Pupa. Not enclosed in cocoons (Wheel-
er and Wheeler, 1976).
LIST OF PRISTROMYRMEX HAMES
WITH SYNONYMIES
(Currently valid names are in boldface)
acerosus: sp. n.
africanus: Pristomyrtnex africanus
Karavaiev
=beni
=mhomu
=myersi
=primus
aniensis: Pristomyrmex quadridens van
aruensis Karavaiev
= quadridens
beni: Hijlidris myersi subsp. beni Weber
= africanus
bicolor: stat. n.: Pri.stomyrmex trachy-
lissa var. bicolor Emery
=taunis syn. n.
bispinosus: Dodous bispinosus Donis-
thoi"pe
boltoni: sp. n.
brevispinosus: Pristomyrmex brevi-
spinosus Emery
=yaeijamensis syn. n.
brotvni: sp. n.
castaneicolor: Pristomyrmex castaneico-
lor Donisthorpe
=umbripennis
castor: Pristomyrmex castor Donis-
thorpe
= umbripennis
coggii: Pristomyrmex coggii Emery
collinus: sp. n.
costatus: sp. n.
cribrarius: Pristomyrmex cribrarius
Arnold
curvulus: sp. n.
divisus: sp. n.
eduardi: Pristomyrmex eduardi Forel
erythropygus: Pristomyrmex ery-
thropygus Taylor
flatus: sp. n.
formosae: Pristomyrmex brevispinosus r.
sulcatus var. formosae Forel,
1912: 54.
unavailable name
fossulatus: Tetramorium (Xiphomyr-
mex) fossulatum Forel
foveolatus: Pristomyrmex foveolatus
Taylor
fuscipennis: Myrmica fuscipennis F.
Smith
hirsutus: sp. n.
inermis: sp. n.
japonicus: Pristomyrmex japonicus Forel
=punctatus
laevigatus: Hylidris laevigatus Weber
=orbiceps
laevis: Solenopsis laevis F. Smith
= umbripennis
largus: sp. n.
levigatus: Pristomyrmex levigatus
Emery
= mendanai syn. n.
longispinus: sp. n.
longus: sp. n.
lucidus: Pristomyrmex lucidus Emery
mandibularis: Pristomyrmex mandi-
bularis Mann
mbomu: Hylidris myersi subsp. mbomu
Weber
=^ africanus
melanoticus: Pristomyrmex obesus
subsp. melanoticus Mann
=obesus
mendanai: Pristomyrmex mendanai
Mann
=levigatus
minusculus: sp. n.
modestus: sp. n.
myersi: Hylidris myersi Weber
= africanus
nitidissimus: Pristomyrmex nitidissi-
mus Donisthorpe
obesus: Pristomyrmex obesus Mann
^^melanoticus syn. n.
Revision of the Ant Genus Pristomyrmex * Wang 393
=pegasiis syn. n.
occultus: sp. n.
orbiceps: Xiphomyrmex orbiceps Sants-
chi
=laevigatus
orbiculatus: Pristorntjnnex orhiculatiis
Donisthorpe
=quadridens
pan imp unci atus: Pristomyrmex panim-
pnnctatus Emery
=umhripennis
parvispina: Pristorntjnnex parvispina
Emery, 1900: 678. Nomen nu-
dum
pegasns: Pristomyrmex pegasns Mann
—ohesus
picteti: Pristomyrmex picteti Emeiy
=tingiana syn. n.
pollux: Pristomyrmex pollux Donis-
thoi-pe
primus: Hijlidris niyersi subsp. primus
Weber
=africanus
profundus, sp. n.
pulcher: sp. n.
punctatus: Mynnica punctata F. Smith
=japonicus
=pungens syn. n.
pungens: Pristomyrmex pungens Mayr
=punctatus
quadridens: Pristomyrmex quadri-
dens Emery
=aruensis syn. n.
=orbiculatus syn. n.
quadridentatus: Odontomyrmex quad-
ridentatus Andre
=queenslandensis
queenslandensis: Pristomyrmex {Odon-
tomyrmex) quadridentatus var.
queenslandensis Forel
= quadridentatus
quindentatus. sp. n.
reticulatus: Pristomyrmex reticula-
tus Donisthorpe
rigidus: sp. n.
simplex: sp. n.
sulcatus: Stat, n.: Pristomyrmex brevi-
spinosus subsp. sulcatus Emery
taurus: Pristomyrmex taurus Stitz
=bicolor
thoracicus: Pristomyrmex thoracicus
Taylor
tingiana: Pristomijrmex picteti var. tin-
giana Stitz
=picteti
trachylissus: Mynnica trachylissa F.
Smith
trispinosus: Dodous trispinosus Donis-
thoipe
trogor: Pristomyrmex trogor Bolton
umbripennis: Mynnica umbripennis F.
Smith
=castaneicolor syn. n.
=castor syn. n.
=laevis syn. n.
=panimpunctatus syn. n.
wheeleri: Pristomyrmex wheeleri
Taylor
wilsoni: Pristomyrmex wilsoni Taylor
yaeyamensis: Pristomyrmex yaeyamen-
sis Yamane and Terayama
= brevispinosus
Key to the World Species of
Pristomyrmex (Workers)
Note: P. fuscipennis and P. reticulatus, whose
worker castes are presently unknown, are not includ-
ed in the key.
1. Dorsum of alitrunk in profile not arched,
with mesonotum much higher than
propodeal dorsum, that is, a vertical
cliff present between mesonotum
and propodeal dorsuin (Fig. 8). An-
tennal scrobes well developed and
deep. Basal margin of mandible
with a strong tooth adjacent to the
basal tooth of masticatory margin so
that five teeth are set close together
(Fig. 27). Base of antennal scape
without a circling lainella (Fig. 80)
(profundus group; Asia: Sabali)
profundus (p. 515)
Dorsum of alitrunk in profile, excluding
armainents, more or less arched-
shaped, never showing a vertical
cliff between mesonotum and pro-
podeal dorsum (Figs. 9—25). Anten-
nal scrobes absent or shallow. Tooth
on basal margin of mandible either
absent or present; if present, it is on
about midway, not adjacent to the
basal tooth of masticatory margin
(Figs. 28, 29, 35). Base of antennal
scape with a circling lamella (Figs.
81, 82) 2
394 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Table 1. A list of Pristomyrmex species and their biogeographic distribution.
Species Group
Species Name
ptinctatus group
cribrarius group
quadridens group
trispinosus group
levigatus group
profundus group
umhripennis group
Total number of species
Total number of the
endemic species
P. divisus sp. n.
+
P. fossidatus (Forel)
P. pulcher sp. n.
+
*P. punctatus (F. Smith)
+
P. rigidus sp. n.
+
P. cribrarius Arnold
P. africanus Karavaiev
P. bicolor Emery
+
P. brevispinosus Emery
+
P. collinus sp. n.
+
P. costatus sp. n.
+
P. curvuhis sp. n.
+
P. eduardi Forel
+
P. erythropygus Taylor
P. flatus sp. n.
+
P. foveolatus Taylor
P. hirsutus sp. n.
+
P. longispinus sp. n.
+
P. modestus sp. n.
+
P. nitidissimus Donisthorpe
+
P. occultus sp. n.
+
P. orbiceps (Santschi)
P. quadridens Emery
+
P. qiiadridentatus (Andre)
P. qiiindentatus sp. n.
+
P. sulcatus Emery
+
P. thoracicus Taylor
P. trachijlissus (F. Smith)
+
P. trogor Bolton
P. wheeleri Taylor
P. wilsoni Taylor
P. bispinosus (Donisthorpe)
P. browni sp. n.
P. trispinosus (Donisthoi-pe)
P. acerosus sp. n.
+
P. boltoni sp. n.
+
P coggii Emery
+
P. inennis sp. n.
+
P. largus sp. n.
+
P. levigatus Emery
+
P. longus sp. n.
+
P. lucidus Emery
+
P. mandibularis Mann
+
*P minusculus sp. n.
+
P. obesus Mann
+
P. simplex sp. n.
+
P. profundus sp. n.
+
P. fuscipennis (F. Smith)
+
P. picteti Emery
+
P pollux Donisthorpe
+
P reticulatus Donisthorpe
+
P. umbripennis (F. Smith)
+
38
36
Notes: ORI, PAL, AUS, AFR, and lOI are abbreviated, respectively, from the Oriental region, the Palaearctic
region, Australia, Africa, and Indian Ocean Islands. AFR refers to the African continent only. Pristomyrmex
has not been recorded from Madagascar. "*" symbol indicates tliat P. punctatus and P minusculus occur in
the two regions, respectively.
Revision of the Ant Genus Pristomyrmex • Wang 395
Figures 8-25. Alitrunks of Pristomyrmex workers, lateral view. 8: P. profundus sp. n.; 9; P. levigatus Emery; 10: P. inermis sp.
n.; 11: P. minusculus sp. n.; 12: P. picteti Emery; 13: P. po//ux Donisthorpe; 14: P. brevispinosus Emery (non-type); 15: P.
brevispinosus Emery (syntype); 16: P. foveolatus Taylor; 17: P. sulcatus Emery (syntype); 18: P. sulcatus Emery (non-type); 19:
P. quadridentatus (Andre); 20: P. whee/eri Taylor; 21: P. longispinus sp. n.; 22: P. trispinosus (Donisthorpe); 23: P. drown/ sp.
n.; 24: P. fa/co/or Emery; 25: P. wilsoni Taylor. Figure 26. Propodeal spines of the worker of Pristomyrmex browni sp. n., dorsal
396 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 27-39. Mandibles of Pristomyrmex workers. 27: P. profundus sp. n.; 28: P. levigatus Emery; 29: P. mandibularisMann;
30: P. p/cteft' Emery; 31: P. quadridens Emery; 32: P. quindentatus sp. n.; 33: P. quadridentatus {Ar\dre); 34: P. trachylissus {F .
Smith); 35: P. rigidus sp. n.; 36: P. punctatus (F. Smith); 37: P. droM/n/ sp. n.; 38: P. trlspinosus (Donisthorpe); 39: P. bispinosus
(Donisthorpe).
Masticatory margin of mandible with
four teeth; the third tooth, counting
from the apex, smallest, distinctly
smaller than the basal one; diastema
absent between the preapical and
the third tooth (Figs. 28-29) {levi-
gattts group; Asia, Australia) 3
Masticatory margin of mandible with
three to five teeth; if four teeth pre-
sent, then the third tooth, counting
from the apex, similar in size to tlie
basal one; diastema either present
or indistinct between the preapical
and the third tooth (Figs. 30-39) ... 14
Propodeum unarmed (Fig. 10). Petiole
node in profile wedge-shaped (Fig.
45) (Asia: Papua New Guinea)
inermis (p. 496)
Propodeum armed with a pair of teeth
or spines (Figs. 9, 11). Petiole node
in profile nodiform, not wedge-
shaped (Figs. 40-44) 4
Pronotum armed with a pair of teeth
(Fig. 11) (Asia and Pacific Is.: Papua
New Guinea; Indonesia; Pohnpei
Is.; Palau Is.; Yap I.; Tonga Is.; Wal-
lis Is.; found rarely in N. Queens-
land, Australia) .... tninusculus (p. 507)
Pronotum unarmed (Fig. 9) 5
Postpetiole in profile with an arched an-
terior face and a steeply sloping pos-
terior face and the apex of postpe-
tiole pointing posterior-upwardly
(Fig. 43); in dorsal view, postpetiole
usually longer than broad, very rare-
ly about as long as broad. Petiole
node with a single evenly blunt-
rounded apex (Fig. 43). Head
broader; HW mostly >1.00 (Asia:
Papua New Guinea) lucidus (p. 503)
Postpetiole in profile with a somewhat
evenly convex dorsum, lacking an
abruptly steep posterior face (Figs.
40—42, 44) and in dorsal view broad-
er than long. Petiole node in profile
with a distinct anterodorsal angle
(Figs. 40^2, 44). Head narrower;
HW <1.00 6
Revision of the Ant Genus Pristomyrmex • Wang 397
Figures 40-58. Petiole nodes and postpetioles of Pristomyrmex worl<ers. 40A: Dorsal surface of the petiole node of P. obesus
Mann, dorsal view; 41 A: Dorsal surface of the petiole node of P. longus sp. n., dorsal view; 40B, 41 B, 42-58: Petiole nodes
and postpetioles, lateral view: 40B: P. obesus Mann; 41 B: P. longus sp. n.; 42: P. acerosus sp. n.; 43: P. lucidus Emery; 44:
P. mandibularis Mann; 45: P. inermis sp. n.; 46: P. punctatus (F. Smith); 47: P. rigidus sp. n.; 48: P. cribrarius Arnold; 49: P.
quadridens Emery; 50: P. africanus Karavaiev; 51: P. nitidissimus Donisthorpe; 52: P. collinus sp. n.; 53: P. ffafus sp. n.; 54:
P. curvulus sp. n.; 55: P. longispinus sp. n.; 56: P. hirsutus sp. n.; 57: P. sulcatus Emery; 58: P. modestus sp. n.
398 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 59-60. Gasters of Pristomyrmex workers, lateral
view. 59: Gaster of P. simplex sp. n.; 60: Gaster of P. obesus
Mann.
Dorsum of petiole node laterally com-
pressed and in dorsal view distinctly
longer than broad (Fig. 41 A) (Asia:
New Guinea) longus (p. 502)
Dorsum of petiole node not laterally
compressed and in dorsal view
about as broad as or broader than
long, not longer than broad (Fig.
40A) 7
Dorsum of alitrunk unsculptured,
smooth and shining. Dorsum of
head between frontal carinae usu-
ally smooth and shining (Fig. 61),
except for a few punctures border-
ing frontal carinae 8
Dorsum of alitrunk with some scattered
foveolate punctures. Dorsum of
head between frontal carinae with
scattered foveolate punctures or fo-
veolate-reticulate sculpture (Figs.
62-64) 12
10.
11.
12(7).
Eyes smaller, with two to three omma-
tidia in the longest row (Asia: New
Guinea) boltoni (p. 492)
Eyes larger, usually with five to seven
ommatidia in the longest row 9
Larger species, with HW 0.90-0.96 and
HL 0.90-0.90 (Pacific Is.: Pohnpei
I.) largus (p. 497)
Smaller species; HW < 0.80, HL < 0.80
10 I
Ventral surface of petiole with a long I
pinlike process (Fig. 42) (Pacific Is.:
New Hebrides) acerosus (p. 491)
Ventral surface of petiole without a long
pinlike process (Figs. 40, 41, 44) ... 11
Each side of petiole with a longitudinal
carina that separates the tergite
from the sternite (Fig. 40). Basal
margin of mandible with a short,
broad toodi (Fig. 28) (Asia and Pa-
cific Is.: Papua New Guinea, Solo-
mon Is., Nama Is., New Britain Is.)
levigatus (p. 499)
Sides of petiole unsculptured and
smooth, lacking a longitudinal carina
(Fig. 44). Basal margin of mandible
with a prominent tooth (Fig. 29)
(Pacific Is.: Fiji)
mandibularis (p. 505)
Entire first gastral tergite evenly clothed
with numerous erect or suberect
hairs (Fig. 60) (Pacific Is.: Solomon
Is.) obesus (p. 509)
Only a few hairs present usually near the
65
66
67
Figures 61-67. Characters on the sculpture of the dorsal heads of Pristomyrmex workers, full-face view, excluding a portion
of the mandibles. 61 : Smooth head; 62-63: Scattered foveolate punctures; 64: Foveolate-reticulate sculpture; 65: Rugoreticulum;
66: Regular striations around the antenna! fossae and on the genae; 67: Regular striations on the entire dorsum of the head.
Revision of the Ant Genus Pristomyrmex * Wang 399
base of the first gastral tergite (Fig.
59) 13
13. Dorsum of head, except for the scrobes,
with foveolate-reticulate sculpture;
punctures often aUgned so that it
seems that several longitudinal ru-
gae appear between frontal carinae.
Eyes smaller, EL = 0.06-0.08, with
three to four ommatidia in the lon-
gest row (Asia: Papua New Guinea)
coggii (p. 493)
Dorsum of head between frontal carinae
with some scattered foveolate punc-
tures; space between foveolae
smooth. Eyes larger, EL = 0.09—
0.12, usually containing five (rarely
four) ommatidia in the longest row
(Asia: Papua New Guinea)
simplex (p. 512)
14(2). Masticatory margin of mandible with
four teeth, lacking a distinct diaste-
ma (Fig. 30). Lamella, circling the
base of antennal scape, with a broad
and deep notch on the center of the
dorsal surface (Fig. 82). Petiole
node in profile longer than high
(umbripennis group; Asia) 15
Masticatory margin of mandible with
three to five teeth; usually with a
distinct diastema (Figs. 31-33, 35—
39); if (very rarely) diastema indis-
tinct, masticatoiy margin with five
teeth (Fig. 34). Lamella, circling the
base of antennal scape, entire, with-
out a notch on the center of the dor-
sal surface (Fig. 81). Petiole node in
profile usually higher than long 17
15. Eyes larger, usually consisting of 20 or
more ommatidia, containing six to
seven ommatidia in the longest row.
Propodeum with a pair of toothlike
armaments that are shorter than the
distance between their bases. About
one-third of antennal scape usually
laterally compressed near the base
(Asia: Papua New Guinea, Indone-
sia) umbripennis (p. 525)
Eyes smallei", generally consisting of less
than 10 ommatidia, with three to
four ommatidia in the longest row.
Propodeum with a pair of spines
that are longer than the distance be-
tween their bases. Antennal scape
not laterally compressed near the
base 16
16. Propodeal spines longer and strongly up-
curved at their apices (Fig. 13).
Larger species with HL 1.42-1.54,
HW 1.42-1.58 (Asia: W. Malaysia,
N. Borneo) pollux (p. 521)
Propodeal spines shorter and not strong-
ly upcurved at their apices (Fig. 12).
Smaller species with HL 1.04—1.36,
HW 1.02-1.40 (Asia: Papua New
Guinea, Indonesia, Singapore, Ma-
laya, Sabah, Brunei, Philippines) -
picteti (p. 518)
17(14). Pronotum unarmed. Eyes larger, usually
containing seven or more (very rare-
ly six) ommatidia in the longest row.
Dorsal surfaces of head and alitrunk
with well developed rugoreticulum
or many foveolate punctures {punc-
tatus group; Asia, Africa) 18
Pronotum usually with a pair of teeth or
spines; if (very rarely) pronotal teeth
or spines absent, then either eyes
smaller, with two to five ommatidia
in the longest row, or dorsal surfaces
of alitrunk and head between frontal
carinae unsculptured and smooth ... 22
18. Dorsum of alitrunk with well developed
coarse reticulum. Propodeum
armed with a pair of long spines.
Antennal scapes longer, usually
>0.78; one-sixth to one-fifth of the
length of the scapes projecting be-
yond the occipital margin. Palp for-
mula 5,3 19
Dorsum of alitrunk with scattered fove-
olate punctures. Propodeum armed
with a pair of short spines. Antennal
scapes shorter, with the length 0.54-
0.60, only close to the occipital mar-
gin. Palp formula 4,3 (South Africa)
fossulatus (p. 406)
19. Lateral portions of clyj^eus, in front of
antennal fossae, developed and not
reduced to narrow margins (Figs.
73, 74). Anterior clypeal margin
lacking distinct denticles (Figs. 65,
68). Median portion of clypeus not
flat 20
Lateral portions of clypeus reduced to a
narrow margin in front of the anten-
nal fossae (Fig. 75). Anterior clypeal
margin with five to seven denticles.
Median portion of clypeus more or
less flat 21
20. Dorsum of head with scattered foveolate
punctures; spaces between foveolae
smooth (Fig. 63). Frontal carinae
short, not extending to the level of
the posterior margins of eyes in full-
face view. Alitrunk in dorsal view
with a deep longitudinal furrow at
middle (Asia: Philippines) -— -
divisus (p. 404)
Dorsum of head entirely sculptured with
coarse reticulum (Fig. 65). Frontal
carinae long, extending to the level
of the posterior margins of eyes in
400 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 68-72. Characters on the anterior clypeal margins of Pristomyrmex workers. 68: Anterior clypeal margin entire; 69:
Anterior clypeal margin with two teeth; 70: Anterior clypeal margin with three teeth; 71 : Anterior clypeal margin with seven teeth;
72: Anterior clypeal margin with three prominences.
full-face view. Alitrunk in dorsal
view lacking a deep longitudinal fur-
row at the midline (Asia: Malaya) -
pulcher (p. 408)
21(19). Two or more pairs of erect hairs present
on the dorsum of petiole node (Fig.
46). Basal margin of mandible al-
most straight, without a distinct
tooth (Fig. 31). Ventral surface of
clypeus lacking toothlike promi-
nences. Dorsal alitrrmk more or less
depressed, with marginate sides.
Sculpture of the sides of pedicel
segments lighter and finer (wide-
spread in the east and south of Asia;
occasionally intercepted at entry
ports in North America)
- - punctatus (p. 410)
A pair of hairs present on the dorsum of
petiole node (Fig. 47). Basal margin
of mandible with an acute or broad-
based triangular tooth (Fig. 35).
Ventral surface of clypeus usually
Figures 73-75. Lateral portions of the clypei of Pristomyrmex
workers in front of antennal fossae. 73: P. divisus sp. n.; 74:
P. pulcher sp. n.; 75: P. punctatus {F. Smith).
witli two minute tootlilike promi-
nences. Dorsal alitrunk convex, not
depressed. Sculpture of the sides of
pedicel segments more coarse (Asia:
Thailand, Malaya, Sarawak, Sabah,
Brunei, Sumatra) rigidus (p. 415)
22(17). Promesonotal suture present. Propodeal
spines developed and long, in dorsal
view joining together at base and
forming a "fork" (Fig. 26). Alitrunk
in profile with a convex promeson-
otum and a deeply concave propo-
deal dorsum (Figs. 22, 23). Dorsum
of head, at least on the genae and
around the antennal sockets, with
regular striate sculpture, lacking fo-
veolate punctures or rugoreticulum
(Figs. 66, 67) {trispinosus group; In-
dian Ocean Islands) 23
Promesonotal suture absent. Propodeal
armaments in dorsal view usually
well separated at the base and not
resembling a fork. In rare case,
where the propodeal spines are set
close together at tlie base, the dor-
sum of alitrunk, in profile, lacks a
deeply concave propodeum (Fig.
25). Dorsum of head smooth or
sculptured with foveolate punctures
or -with lugo reticulum, but never
showing regular striate sculpture _ __
23. Propodeal spines in dorsal view diver-
gent, in profile almost straight.
Larger species with HW > 1.00, HL
> 1.10, SL > 1.30
Propodeal spines in dorsal view subpar-
25
24
Revision of the Ant Genus Pristomyrmex* Wanp 401
allel (Fig. 26), in profile view bent
at about a right angle near the base
(Fig. 23). Smaller species with HW
0.82-0.90, HL 0.88-1.01, SL 0.80-
0.97 (Indian Ocean Is.: Mauritius,
Reunion I.) brotvni (p. 485)
24. Dorsum of head and alitinmk entirely
covered with regular long coarse
striations (Fig. 67). Mesonotum with
a pair of strong, blunt digitlike
prominences (Fig. 22) (Indian
Ocean Is.: Mauritius)
trispinosus (p. 488)
Dorsum of head smooth and shining,
with rugae only present around the
antennal fossae, on the genae and
sometimes around the centrical disc
of dorsal head (Fig. 66). Dorsum of
alitrunk smooth and shining. Me-
sonotum lacking well-developed
digitlike prominences (Indian
Ocean Is.: Mauritius)
bispinosus (p. 484)
25(22). Sides of postpetiole with several coarse
longitudinal rugae (Fig. 48). In pro-
file view, the posterodorsal and pos-
teroventral comers of the petiole
node right-angled. Palp formula 4,3
{cribrarius group; Africa: Mozam-
bique, South Africa)
cribrarius (p. 418)
Sides of postpetiole unsculptured or at
most with a single longitudinal ruga
(Figs. 49-58). In profile view, the
posterodorsal and posteroventral
corners of the petiole node not
right-angled. Palp formula 1,3 or 2,2
or 2,3 {quadridens group; Asia, Aus-
traha, Africa) 26
26. Mandibular dentition arranged as an api-
cal tooth -I- a preapical -I- a diastema
+ three small denticles of similar
size (Fig. 32). Sometimes, the three
small denticles are fused together so
that they are not clearly visible, but
the length of the masticatoiy margin
covered by the three small denticles
is slightly longer than that of diaste-
ma. Pronotuiu either unarmed or
armed with a pair of short triangular
spines that are shorter than the pro-
podeal spines 27
Mandibular dentition arranged as an api-
cal -t- a preapical -I- a diastema -I-
one or two denticles, and the length
of the masticatory margin covered
by the one or two denticles is dis-
tinctly shorter than that of diastema
(Figs. 31, 33, 35). If (very rarely)
mandibular dentition not as de-
scribed previously but arranged as
an apical -I- a preapical -I- a small
denticle + a very short diastema (or
diastema indistinct) + two small
denticles (Fig. 34), then the prono-
tum is armed with a pair of long ro-
bust spines that are much longer
than propodeal spines (Fig. 24) 29
27. Pronotum unarmed. Eyes smaller, with
three ommatidia in the longest row
(Asia: Sumatra) eduardi (p. 440)
Pronotum armed with a pair of short tri-
angular spines. Eyes larger, usually
with five to SLx (rarely with four) om-
matidia in the longest row 28
28. Dorsal surfaces of head and alitnmk only
with scattered shallow foveolate
punctures; dorsum of aUtrunk with
a smooth and unsculptured median
longitudinal strip (Asia: Indonesia)
quindentatus (p. 467)
Dorsal surfaces of head, except for scro-
bal areas, and alitrunk entirely cov-
ered with well developed coarse ru-
goreticum (Asia: Sarawak, Sabah) —
occultus (p. 455)
29(26). Dorsal surfaces of both alitrunk and
head between frontal carinae either
smooth or with some scattered fo-
veolate punctures but lacking fove-
olate-reticulate sculpture or rugore-
ticulum 30
Dorsal surfaces of both alitrunk and
head with foveolate-reticulate sculp-
ture or rugoreticulum 42
30. Area of dorsal head between frontal ca-
rinae unsculptured and smooth.
Dorsum of alitrunk without foveo-
late punctures 31
Dorsal surfaces of both alitrunk and
head between frontal carinae with
some scattered foveolate punctures
40
31. Alitrunk, in dorsal view, with a transverse
ridge at the appro.ximate position of
metanotal groove (Fig. 19). Anterior
clypeal margin with three strong
teeth (Fig. 70) 32
Alitrunk, in dorsal view, unsculptured,
lacking a transverse ridge at the ap-
proximate position of metanotal
groove. Anterior clypeal margin usu-
ally with five to seven small denti-
cles (Fig. 71) 34
32. First gastral tergite with numerous,
evenly distributed, suberect hairs.
Alitrunk in dorsal view with several
short rugae present at the juncture
between the pronotum and the me-
sonotum (Australia: New South
Wales) erythropygus (p. 441)
First gastral tergite lacking any suberect
402 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 76-79. Characters on the ventral surfaces of the clypei of Phstomyrmex workers. 76: A tooth; 77: Two toothlike
prominences; 78: A short ruga; 79: A long transverse ridge.
33.
34(31)
35.
hairs. Alitnink in dorsal view without
rugae at the juncture between tlie
pronotum and the mesonotum 33
Propodeal spines longer, subequal to or
longer than pronotal spines (Fig.
20). Ventral center of clypeus with a
weak and short ruga (Fig. 78). Head
broader, with HW 0.97-1.34 and CI
103-116 (Austraha: New South
Wales, SE Queensland)
wheeleri (p. 478)
Propodeum with a pair of teeth or short
spines, much shorter than pronotal
spines (Fig. 19). Ventral surface of
clypeus with a long, well-developed
transverse ridge (Fig. 79). Head nar-
rower, with HW 0.80-1.08 and CI
93-101 (Australia: New South
Wales, Queensland)
quadridentatus (p. 463)
Pronotum tuberculate, lacking teeth or
spines (Africa: Ivoiy Coast, Ghana,
Nigeria, Cameroon, Gabon, Congo
and Angola) orbiceps (p. 456)
Pronotum armed v\dth a pair of teeth or
spines 35
Petiole and postpetiole without erect
hairs (Fig. 50). Frontal carinae ab-
sent. Pronotum with a pair of tri-
angular short spines. Ventral surface
of clypeus with two toothlike prom-
inences (Fig. 77) (Africa: Zaire)
trogor (p. 476)
Petiole and postpetiole with at least one
to two pairs of hairs (Figs. 49, 52).
Frontal carinae present and usually
extending to the level of the poste-
rior margins of eyes. In rare cases
Figures 80-82. Lamella, circling the base of the antennal
scape of the Phstomyrmex worker, absent (80), entire (81), or
with a broad and deep notch (82).
where the frontal carinae are very
short or absent, the pronotum is
armed with a pair of well-devel-
oped, long spines that are longer
than the distance between their ba-
ses (Fig. 21). Ventral surface of clyp-
eus either with a transverse ruga, or
with a tooth at center, or without
any ruga or tooth, but never show-
ing two toothlike prominences 36
36. Pronotum with a pair of triangular short
spines, much shorter than the dis-
tance between their bases 37
Pronotum with a pair of long spines, lon-
ger than distance between their ba-
ses (Figs. 21, 25) 38
37. Petiole node in profile lacking distinct
anterior face distinguishable from
the upper surface of peduncle (Fig.
53). Larger species, with HW 0.98-
1.04, HL 0.94-1.02, EL 0.22-0.24
(Asia: Philippines) flatus (p. 443)
In profile, anterior face of petiole node
distinct from the dorsal surface of
peduncle (Fig. 52). Smaller species,
with HW 0.77-0.94, HL 0.82-0.94,
EL 0.14-0.18 (Asia: Philippines) --
collinus (p. 432)
38(36). Propodeal spines short or moderately
long, much shorter and slender tlian
pronotal spines (Fig. 21) 39
Propodeal spines exceptionally long,
subequal in length to or slightly lon-
ger than pronotal spines (Fig. 25)
(Australia: Queensland)
wilsoni (p. 481)
39. Petiole node in profile lacking distinct
anterior face distinguishable from
the upper surface of peduncle (Fig.
55). Clypeus unsculptured, lacking a
median longitudinal carina. Frontal
carinae short, not extending to the
level of the posterior margins of
eyes (Asia: Philippines)
longispinus (p. 450)
In profile, anterior face of petiole node
distinct from the dorsal surface of
peduncle (Fig. 54). Clypeus with a
median longitudinal carina. Frontal
carinae long, extending to the level
of the posterior margins of eyes
(Asia: Philippines) .. curvulus (p. 437)
Revision of the Ant Genus Pristomyrmex • Wanp
403
40(30). Petiole and postpetiole lacking erect
hairs (Fig. 50). Ventral surface of
clypeus with two toothlike promi-
nences (Figs. 77) (Africa: Ghana,
Cameroon, Gabon, Angola, Kenya,
Zaire, Sudan) africanus (p. 423)
Petiole and postpetiole, respectively,
with one to five pairs of erect hairs.
Ventral surface of clypeus either
with a transverse ruga (Figs. 78, 79)
or with a toothlike prominence (Fig.
76) 41
41. Smaller species (HW 0.82-1.02, HL
0.82-1.02, EL 0.14-0.20). Ventral
center of clypeus with a toothlike
prominence. Usually one to two
pairs of hairs present, respectively,
on the dorsal surfaces of petiole
node and postpetiole (Asia and Pa-
cific Is.: Papua New Guinea; Indo-
nesia; Pohnpei Is.)
quadridens (p. 459)
Larger species (HW 1.22-1.24, HL
1.10-1.16, EL 0.24-0.25). Ventral
surface of clypeus with a transverse
ruga, lacking a toothlike prominence
at center. Four to five pairs of short
hairs present, respectively, on the
dorsal surfaces of petiole node and
postpetiole (Fig. 51) (Asia: New
Guinea) nitidissitnus (p. 453)
42(29). First gastral tergite with numerous,
evenly distributed, erect or suberect
hairs. Petiole node with a single
evenly blunt-rounded apex (Fig. 56)
(Asia: Philippines) ... hirsutus (p. 449)
First gastral tergite lacking erect hairs.
Petiole node not showing a single
evenly blunt-rounded apex but with
a higher anterodorsal angle than the
posterodorsal (Fig. 57) 43
43. Masticatory margin of mandibles with
five teeth; diastema very short or in-
distinct between the preapical and
the third tooth (Fig. 34). Basal mar-
gin of mandible with a central,
broadly curved lobe. Anterior clyp-
eal margin with three prominences,
that is, a median tooth and a broad,
low convex lobe on each side (Fig.
72) (Asia: Sarawak, Sabah, Borneo)
trachylissus (p. 474)
Masticatory margin of mandibles with
three to four teeth; diastema distinct
and long, present between the
preapical and the third tooth (Figs.
31, 33). Basal margin of mandible
almost straight, without a distinctly
curved lobe. Anterior clypeal mar-
gin usually with five to seven denti-
cles (Fig. 71) 44
44. Pronotal spines exceptionally long, usu-
ally exceeding 0.40 (very rarely
0.37), usually longer than the dis-
tance between the bases of two pro-
notal spines (Fig. 24) (Asia: Suma-
tra, Java, Malaya, Sarawak, Sabah,
Borneo, Philippines) bicolor (p. 425)
Pronotum armed with a pair of teeth or
spines (<0.32; Figs. 14, 15, 17, 18)
that are always shorter than the dis-
tances that separate their bases 45
45. Propodeal spines long (0.19-0.30), much
longer than the pronotal armaments
(Fig. 16), palp formula 2,3 46
Propodeal spines short (0.04-0.13), usu-
ally shorter than but sometimes
slightly longer than the pronotal ar-
maments (Figs. 14, 15), palp for-
mula 1,3 __._ 47
46. Postpetiole in dorsal view much broader
than long, with PPI 133-150. An-
tennal scapes shorter (SL 0.70-0.82,
SI 81-93) (Australia: Queensland)
foveolatus (p. 446)
Postpetiole in dorsal view slightly broad-
er than long, with PPI 109-121. An-
tennal scapes longer (SL 0.86-0.98,
SI 97-103) (Australia: Queensland)
thoracicus (p. 473)
47(45). Petiole node in profile slightly longer
than high, with the anterodorsal an-
gle on approximately the same level
as or weakly higher than the poster-
odorsal; dorsum and sides of petiole
node with seven to eight foveolate
punctures (Fig. 58) (Asia: Sarawak)
modestus (p. 452)
Petiole node in profile higher than long,
with the anterodorsal angle distinct-
ly elevated above the posterodorsal;
dorsum and sides of petiole node
without foveolate punctures (Fig.
57) 48
48. Ventral surface of clypeus lacking a cen-
tral prominent tooth but usually
bearing a transverse ruga (Asia: Ma-
laya, Singapore, Sarawak, Borneo,
Sabali, Philippines) ... costatus (p. 434)
Ventral surface of clypeus with a central
prominent tooth (Fig. 76) 49
49. Pronotum with a pair of moderately long
spines (0.14—0.20), usually much
longer (sometimes slightly longer)
than the propodeal armaments
(0.07-0.13) (Figs. 18) (Asia: Malaya,
Thailand, Nepal, Burma, China) —
sulcatus (p. 469)
Pronotum with a pair of teeth or short
spines (0.06-0.10), usually similar in
length to or slightly shorter than the
propodeal armaments (0.04-0.12)
1
404 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
(Figs. 14, 15) (Asia: Sumatra, Sula-
wesi, Malaya, Sarawak, Sabah, Thai-
land, Philippines, Taiwan, Japan) -
brevispinosus (p. 428)
THE PUNCTATUS GROUP
Worker. Small to medium sized (HL
0.62-0.98, HW 0.64-0.94, TL 2.62-3.44),
with the following combination of charac-
ters:
(1) Masticatory margin of mandibles
with three to four teeth arranged as the
strongest apical + the second strongest
preapical + a long diastema + two small
basal denticles of similar size (or a broad
basal tooth).
(2) Palp formula 5,3 in four Oriental
species and 4,3 in the single South African
species.
(3) Frontal carinae present.
(4) Lateral portions of clypeus reduced
to a narrow margin in front of the antennal
fossae in three species (P. punctatus, P. rig-
idus, and P. fossulatus) but developed in
the other two species {P. divisus and P.
pulcher).
(5) Frontal lobes indistinct or absent.
(6) Lamella that encircles the base of
antennal scape, entire.
(7) Alitrunk in profile showing a contin-
uous convex dorsum and in dorsal view
lacking sutures.
(8) Pronotum unarmed.
(9) Propodeum with a pair of spines.
(10) Petiole node in profile more or less
wedge-shaped, lacking distinct anterior
face distinguishable from the upper sur-
face of peduncle.
This group has five species. Four occur
in the Oriental region, one of which {P.
punctatus) has spread to warm-temperate
areas in the eastern Palaearctic. The re-
maining species (F. fossidatus) is confined
to South Africa. Pristomyrrnex punctatus is
a unique species within the genus. It is the
only Pristomyrmex extending its range to
the temperate zone, and it is further char-
acterized by the possession of a unique life
history that may preadapt it for dispersal
by natural and human transport.
The punctatus group is closely related
to the cribrarius and quadridens groups
because all female castes of these species
groups possess a distinct diastema after the
preapical tooth on the masticatory margin
of mandibles (except in P. trachylissus,
which has five teeth on the masticatoiy
margin). Though sharing the previously
described mandibular character, the trispi-
nosus group is relatively distant from the
punctatus group because it possesses so
many autapomorphic characters, for ex-
ample, frontal carinae absent, dorsal ali-
trunk with a promesonotal suture or iin-
pression, propodeal spines in dorsal view
showing a "fork", some regular striation
present on the dorsal surfaces of head and
alitrunk.
The punctatus group differs froin the
cribrarius group by lacking a pair of pro-
notal spines in the workers and by showing
the anterior face of the petiole node not
distinct from the upper surface of pedun-
cle in the female castes. The punctatus
group differs from the quadridens group
by possessing palp formulae of 5,3 and 4,3
(the quadridens group possesses palp for-
mulae of 2,2, 1,3 and 2,3).
Pristomyrmex divisus sp. n.
Figures 83-84
Diagnosis (Worker). Lateral portions of
clypeus in front of antennal fossae, devel-
oped, not reduced to margins, so that the
antennal fossae are placed well behind the
anterior clypeal margin; dorsal head only
with scattered foveolate punctures.
Holotype Worker (MCZC). TL 3.06, HL
0.76, HW 0.80, CI 105, SL 0.82, SI 103,
EL 0.22, PW 0.56, AL 0.74. Paratypes,
35 workers (MCZC, BMNH, ANIC,
MHNG).
Worker TL 3.06-3.40, HL 0.72-0.82,
HW 0.74-0.86, CI 98-111, SL 0.78-0.90,
SI 98-110, EL 0.21-0.24, PW 0.53-0.64,
AL 0.72-0.80, PPW 0.26-0.30, PPL 0.18-
0.22, PPI 123-156 (n = 20).
Mandibles with a few longitudinal rugae
but smooth near the masticatory margin.
Dentition of the masticatory margin of
Revision of the Ant Genus Pristomyrmex • Wang 405
Figures 83-84. Pristomyrmex divisus sp. n. 83: Worker head, full-face view; 84: Worker, lateral view.
mandible: the strongest apical tooth + the
second strongest preapical + a long dia-
stema + a broad, truncated basal tooth (or
two minute denticles). A weak minute
prominence present about midway on the
basal margin of mandible. Clypeus with a
strong median longitudinal carina extend-
ing through the frontal area; on each side
of the median clypeal carina, a few addi-
tional rugae are usually present. Anterior
clypeal margin lacking denticles. Median
portion of clypeus higher than frontal area;
lateral portions of clypeus developed, not
reduced to iTiargins. Ventral surface of
clypeus lacking any toothlike prominences
but usually with a few rugae. Palp formula
5,3. Frontal carinae short, not extending to
the level of the posterior margins of eyes.
Antennal scrobes absent. Frontal lobes ab-
sent; thus, the antennal articulations are
completely exposed. Antennal scapes,
when lying on the dorsal head, suipassing
the occipital margin of head by one-sixth
to one-fifth of their length. Eyes large and
prominent, containing 8 to 10 ommatidia
in the longest row. Dorsum of alitrunk in
dorsal view marginated, more or less de-
pressed, and usually with a deep longitu-
dinal furrow at middle. Pronotuixi un-
armed. Propodeal spines well developed,
acute and long, much longer than the dis-
tance between their bases. Metapleural
lobes small, dentiform, and acute. Petiole
in profile with a long peduncle; dorsum of
peduncle, together with the anterior face
of petiole node, forming a long declivity
that reaches the top of petiole node. Ven-
tral surface of petiole lacking any process.
Postpetiole in profile with a convex dor-
sum, in dorsal view somewhat transverse-
rectangular and much broader than long.
Dorsum of head v^th numerous large and
shallow foveolate punctures; space be-
tween foveolae smooth; ventral head with
denser foveolate punctures. Dorsal surface
of alitrunk with reticulate rugae. Petiole al-
ways, and postpetiole usually, with a coarse
longitudinal ridge on each side. In dorsal
view, petiole node and postpetiole each
usually bounded by a rim; dorsums of both
petiole and postpetiole, except for rims,
very smooth and polished. Gaster unsculp-
tured. Dorsal surfaces of head and alitrunk
with numerous erect to suberect short
hairs. A pair of hairs present, respectively,
near the top of both petiole node and post-
petiole. First gastral tergite without hairs.
Two or three pairs of long, forward-pro-
jecting hairs present near the anterior
clypeal margin. Scapes and tibiae with
some erect to suberect short hairs. Color
uniform reddish-brown; appendages
sometimes slightly lighter.
Queen and Male. Unknown.
Comments. This species is so far known
only from the Philippines, and its closest
relative without doubt is P. pulcher, from
Malaysia. The workers of two species share
the following diree characters that are not
seen in the other three members of the
406 Bulletin Museum of Comparative Zoology, Vol. 157, No. 6
Figures 85-86. Pristomyrmex fossulatus (Forel). 85A: Worker head, full-face view; 85B: Showing a short transverse ruga on
the ventral clypeus; 86: Worker, lateral view, hairs omitted from the petiole node and postpetiole.
punctatus group (P. punctatus, P. rigidus,
and P. fossulatus): (1) lateral portions of
clypeus, in front of the antennal fossae, de-
veloped, making the antennal fossae well
behind the anterior clypear margins; (2)
anterior clypeal inargin lacking distinct
denticles; and (3) the median portion of
clypeus not flat but somewhat concave. In
the workers of P. punctatus, P. rigidus, and
P. fossulatus, the anterior clypeal margin is
equipped with five to seven denticles, and
the lateral portions of clypeus in front of
the antennal fossae are reduced to margins
(in other words, the antennal fossae reach
the lateral anterior margins of clypeus),
and the median portion of clypeus is more
or less flat.
The workers of P. divisus are easily sep-
arated from those of P. pulchen The ce-
phalic dorsum shows rugoreticulum in P.
pulcher but scattered foveolate punctures
in P. divisus ; the frontal carinae do extend
to the level of the posterior margins of
eyes in P. pulcher but not so in P. divisus;
a pronounced median longitudinal furrow
is present on the dorsal surface of alitrunk
in P. divisus but absent in P. pulcher
Holotype Worker Philippines: Duma-
guete, 1949, J. W. Chapman.
Paratypes. 18 workers with same data as
holotype; 14 workers, Philippines: Duma-
guete (J. W. Chapman); three workers,
Philippines: Dumaguete, Silliman Univer-
sity, 9. V. 1949 (Domingo Empeso).
Ecological Information. Unknown.
Pristomyrmex fossulatus (Forel)
Figures 85-86
Tetramoriuiti {Xiphomynnex) fossulatum Forel, 1910:
428. Syntype workers. South Africa: Natal, Will
Broak (Wroughton) (MHNG) [examined].
Pristomyrmex fossulatus (Forel) Santschi, 1916: 51.
Diagnosis (Worker). Masticatory margin
of mandible with a long diastema after the
preapical tooth; palp formula 4,3; eyes
wdth 8 to 10 ommatidia in the longest row;
pronotum lacking teeth or spines; dorsal
surfaces of head and alitrunk with scat-
tered foveolate punctures.
Worker TL 2.63-2.92, HL 0.62-0.71,
HW 0.64-0.75, CI 98-106, SL 0.56-0.61,
SI 81-88, EL 0.17-0.18, PW 0.44-0.50,
AL 0.64-0.74, PPW 0.26-0.28, PPL 0.16-
0.19, PPI 147-163 (n = 5).
Mandibles smooth and shining. Denti-
tion of the masticatory margin of iTiandi-
ble: an apical tooth + a preapical tooth +
a long diasteina + a truncated basal tooth.
Basal margin of mandible lacking a tooth-
like prominence or curved lobe. Clypeus
with a strong median longitudinal carina.
Anterior clypeal margin with a median
denticle and two to three others on each
side, but sometimes two lateral denticles
Revision of the Ant Genus Pristomyrmex • Wans.
407
are fused together. Lateral portions of
clypeus reduced to margins, and antennal
fossae reaching the lateral anterior iTiar-
gins of clypeus. Ventral center of clypeus
with a short transverse ruga. Palp formula
4,3. Frontal carinae extending to the level
of the posterior margins of eyes. Antennal
scrobe short, shallow, but distinct, mar-
gined by the frontal carina and a longitu-
dinal ruga. Frontal lobes absent; thus, the
antennal articulations are completely ex-
posed. Antennal scapes, when lying on the
dorsal head, close to the occipital margin
of head. Eyes large, containing 8 to 10 om-
matidia in the longest row. Pronotum with
a pair of blunt tubercles, lacking teeth or
spines. Propodeuin armed with a pair of
spines, about as long as the distance be-
tween their bases. Metapleural lobes sub-
triangular. Petiole node in profile wedge-
shaped, with a triangular apex. Subpetiole
with a narrow flange. Postpetiole in profile
higher than long, with a rounded dorsum,
in dorsal view transverse-rectangular and
much broader than long. Dorsum of head,
except for the scrobal areas, with numer-
ous scattered foveolate punctures. Similar
but sparser punctures present on the dor-
sal surface of alitrunk. Petiole and post-
petiole each usually with a longitudinal
ruga on each side. Gaster unsculptured.
Several pairs of hairs present on the dor-
sum of head beyond the level of the an-
tennal insertions. A row of forward-pro-
jecting hairs present on the anterior clyp-
eal margin. Hairs on the rest of the body
as follows — iTiesonotum (one pair), petiole
(zero to one pair), and postpetiole (one to
two pairs dorsally) — frequently lost by
abrasion (Bolton, 1981: p. 286). First gas-
tral tergite lacking erect or suberect hairs.
Scapes and tibiae with short hairs. Color
reddish-brown; appendages yellow-brown.
Queen. I have not seen the queen of this
species, but Menozzi (1942: 172) gave a
description of this caste
